Mytilus galloprovincialis is invasive on rocky shores on the west coast of South Africa where it has become the dominant intertidal mussel. The success of this species on the west coast and its superior competitive abilities, have led to concern that it may become invasive on the south coast at the expense of the indigenous mussel Perna perna. On shores where these species co-occur, there appears to be habitat segregation among zones occupied by mussels. M.galloprovincialis dominates the high-shore and P.perna the low-shore, with a mixed zone at mid-shore level. This study examined the factors responsible for these differences in distribution and abundance. The study was conducted in Plettenberg Bay and Tsitsikamma (70km apart) on the south coast of South Africa. Each site included two randomly selected locations (300-400m apart). A third mussel species, Choromytilus meridionalis, is found in large numbers at the sand/rock interface at one location in Plettenberg Bay. Aspects of settlement, recruitment, growth and mortality of juvenile and adult mussels were examined at different tidal heights at each site. Quantitative analysis of mussel population structure at these sites supported the initial observation of vertical habitat segregation. Post-larvae were identified to species and this was confirmed using hinge morphology and mitochondrial DNA analysis. Size at settlement was determined for each species to differentiate between primary and secondary settlement. Adult distribution of C.meridionalis was primarily determined by settlement, which was highly selective in this species. Settlement, recruitment and growth of P.perna decreased with increasing tidal height, while post-settlement mortality and adult mortality increased higher upshore. Thus all aspects of P.perna’s life history contribute to the adult distribution of this species. Presumably, the abundance of P.perna on the high-shore is initially limited by recruitment while those that survive remain prone to elimination throughout adulthood. M.galloprovincialis displayed the same patterns of settlement and recruitment as P.perna. However, post-settlement mortality in this species was consistently low in the low and high zones. Juvenile growth also decreased upshore, suggesting that M.galloprovincialis may be able to maintain high densities on the high-shore through the persistence of successive settlements of slow-growing individuals. The low cover of M.galloprovincialis on the lowshore appeared to be determined by adult interactions. M.galloprovincialis experienced significantly higher adult mortality rates than P.perna in this zone. There were seasonal variations in the competitive advantages enjoyed by each species through growth, recruitment or mortality on the low-shore. In summer, P.perna had higher recruitment rates, faster growth and lower mortality rates, while M.galloprovincialis had slightly higher recruitment rates and faster growth rates in winter. P.perna is a warm water species while M.galloprovincialis thrives on the cold-temperate west coast of South Africa. Therefore both species appear to be at the edge of their optimal temperature regimes on the south coast, which may explain the seasonal advantages of each. Nevertheless, P.perna has maintained spatial dominance on the low-shore suggesting that it may ultimately be the winner in competition between these species. M.galloprovincialis appears to have a refuge from competition with P.perna on the high-shore due to its greater tolerance of desiccation stress, while being competitively excluded from the low-shore. Warm water temperatures coupled with poor recruitment rates at most sites may limit the success of M.galloprovincialis on this coast.

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.

Organisms inhabit environments that have many dimensions, each of which can vary temporally and spatially. The spatial-temporal variations of environmental stressors and disturbances may have major but different effects on indigenous and invasive species, favouring either of them at different times and places. The invasive mussel Mytilus galloprovincialis invaded the South African coast 30 years ago and, on the south coast of South Africa, it now competes and co-exists with the indigenous Perna perna in the lower eulittoral zone (referred to here as the mussel zone) The invasive and indigenous species dominate the upper and the lower mussel zones respectively, while the two co-exist in the mid-zone. My results show that intertidal mussels experience, and respond to, spatial and temporal fluctuations of several biotic and abiotic stressors. The invasive and the indigenous species adopt different strategies when reacting to environmental factors and their physiological and behavioural responses vary in time and in different habitats as different pressures become of overriding importance. Attachment strength of both species decreased in summer and increased in winter, and was higher on the open coast than in bays for both species, showing a strong positive correlation with wave force in time and space. P. perna had significantly higher attachment strength than M. galloprovincialis but, contrary to previous studies, the difference in gonad index between the two species varied according to the habitat. In bay habitats, M. galloprovincialis had a higher maximum reproductive effort than P. perna, however, on the open coast, there was no significant difference between the two species, suggesting that for the invasive species wave action is a limiting factor not only in terms of the attachment strength but also of energy availability for reproductive tissue development. Major spawning events occurred during periods of low wave action while minor spawning coincided with periods of intense hydrodynamic stress. On the open coast, gonad index was negatively correlated with attachment strength for both species while, in bays, there was no correlation between these two factors for either. The two species also showed different behaviour. In the field, M. galloprovincialis moved significantly more than P. perna over a period of six months. The higher mobility of the invasive species was also confirmed in the laboratory where, in general, M. galloprovincialis formed clumps more readily than P. perna. Taken collectively, these results suggest that channelling more energy into attachment strength limits reproductive tissue development and that, while the indigenous species invests more in byssal production, the invasive species adopts a more dynamic strategy looking for aggregation or a safer arrangement. Higher endolithic infestation and a greater expression of heat shock proteins (Hsps) in mussel populations on the open coast than in bays indicate that this habitat is a more stressful environment not only in terms of wave action. Endolith damaged mussels had significantly lower attachment strengths and condition indices than clean mussels, probably due to the need to channel energy into shell repair. The constant shell repair and expression of Hsps typical of open coast populations are energetically demanding processes. These observations suggest that on the open coast, mussels are subjected to more severe energetic constraints than in bay habitats. Wave and sand stress fluctuated seasonally with the former having a greater effect on mussel mortality on the open coast and the latter a higher impact on bay populations. Overall, mussel mortality rates were higher on the open coast than in bays. My results show that populations on the open coast had fewer private haplotypes and less genetic endemism than those inside bays. Gene flow analysis showed the relatively stable bay habitats act as source populations with greater genetic migration rates out of bays than into them. These differences in genetic structure on scales of las of kilometers show that coastal configuration strongly affects selection, larval dispersal and haplotype diversity. Environmental gradients that are key factors in species distribution over large geographical scales can also be responsible for micro-scale distributions. My results show that M. galloprovincialis colonizes the upper mussel zone where temperature is high, but is less tolerant to this stressor and has to maintain a high expression of Hsps. This suggests that temperature is probably a limiting factor in its invasion towards the sub-tropical east coast. There are inter- and intra-specific differences in responses to the environment which highlight the efforts of M. galloprovincialis and P. perna to optimize resource utilization for survival and reproduction. Determining these differences is crucial to understanding patterns of co-existence between competing indigenous and invasive species.