Population dynamics of Perna perna in low shore mussel beds were investigated over a 15 month period at six sites along the south coast of South Africa, with particular reference to the effects of wave exposure. The degree of exposure was first quantitatively ascertained using the dissolution of cement blocks to measure average wave force and dynamometers to measure maximum wave force. The mean mass loss of the cement blocks was higher at Diaz Cross and Kwaai Hoek than at Mgwalana, Rufanes and Riet River. No data were available for Fish River. The mean maximum wave force encountered at Diaz Cross, Kwaai Hoek and Fish River was significantly higher (p<O.05) than that at Mgwalana, Rufanes and Riet River. These results allowed the former sites to be classified as exposed and the latter as sheltered. Early recruitment (1-5 mm) occurred throughout the year, but peaked significantly (p<0.05) from January 1995 to May 1996 on both shore types. Although mean recruit density (1-15 mm) was Significantly higher at the exposed (5 896.m⁻²) than the sheltered shores (2 986.m⁻²), some sites did not show this trend. Further investigation revealed that the densities of recruits (1-5 & 6-10 mm) were significantly higher on algae than on mussels (p<0.05). Adult densities (>15 mm) were positively correlated with recruit densities (1-5 mm) for both shore types (p<O.05). In tum, adult density (>15 mm) was significantly lower on exposed (mean of 3 348.m⁻²) than on sheltered shores (mean of 4 796.m⁻²) (p<O.05). Adult mussels on exposed shores had significantly higher mean and maximum lengths than those on sheltered shores (p<O.05). Biomass, which is a product of density and length, showed no significant difference between the two shore types (p>O.05). The effect of exposure on growth rate formed the focal point of this study and was determined using three different approaches. The first technique, mark-recapture, involved filing notches on the growing edges of mussels in the field. After 111 days, mussels were removed and the growth measured. The second approach used internal growth bands to measure growth rates, once the periodicity with which these bands were laid down was established. Thirdly, using Shepherd's length composition analysis (SLCA), growth rates were determined from length frequency distributions in 11 samples taken over 15 months. The general conclusion from all three approaches was that growth rate was twice as fast on the exposed shores as on the sheltered shores (p<O.05). A mean length (averaged from all three methods) of 47.06 mm was attained within the first year of growth at the exposed shores and 22.07 mm at the sheltered shores. There were however considerable differences among these approaches. The mark-recapture method predicted the lowest growth rates, followed by growth band method and lastly SLCA. The mean mortality index (Z.year⁻¹) for mussels was significantly (p<0.05) higher at the exposed shores (Z=1.81) than at the sheltered shores (Z=0.73). Consequently, the percentage survival rates per annum of mussels of all ages (total), 18 days to 6 months (juveniles) and 12 months to mortality (adults) was lower on the exposed than the sheltered shores. The survival rate of juveniles was as low as 0.71% per annum on the exposed shores and 9.29% per annum on the sheltered shores. The adult survival rate of exposed shore mussels was 11.78% per annum, considerably lower than that of sheltered shore mussels, 48.05%. The turnover rate on exposed shores was faster than on sheltered shores as the mean longevities were 2.6 and 6.7 years respectively. In conclusion, these findings showed that the effects of exposure on recruitment, growth and mortality are important in low shore mussel beds.

The effects of small scale (cm) water movement on the settlement and growth rates of the brown mussel Perna perna were investigated on the south-east coast of South Africa (33°28′S, 27°10′E). L-shaped metal baffles attached to the substratum decreased the erosion rates of cement balls and it was concluded that the baffles decreased the water flow rate around cement balls. These L-shaped baffles were then used to decrease water flow rates around mussel patches and pot-scouring pads used as artificial substrata for the settlement of P.perna larvae. Anova indicated that settlement rate varied by date and site while decreased water flow rate significantly increased larval settlement (p<0.05), only on the site and day that had the overall highest number of settlers. Mussels in the low zone had significantly higher growth rates than those in the high zone. Decreased water flow rate significantly increased mussel growth rate in the lower zone (Anova, p<0.05), while it did not have a significant effect on the mussel in the high zone. Thus water flow manipulation increased growth rates in the zone, which already had high growth rate. It was concluded that small-scale (cm) water flow patterns have an effect on both Perna perna settlement and growth rates, but only under specific conditions. Larval settlement rate was significantly increased by water flow manipulation on the site and day that had the highest number of settlers. Growth rates were significantly increased by decreased water flow rate only in the low zone, where growth rates are the highest. Although water flow was manipulated in both zones its effect in the high zone was insignificant (Anova) compared to other factors affecting growth rates at this tidal level.

Certain aspects of the population dynamics of the brown mussel, Perna perna, were examined at 18 sites along the south coast of South African. Specifically the effects of wave exposure and tidal height were examined in relation to mussel size, biomass and density. A single set of samples was removed from each of the 18 sites, over three spring tide cycles. Sites were classified as exposed or sheltered prior to sampling. Principal component analysis (PCA) (based on mussel length data) and length frequency histograms revealed that there was a general decrease in the modal size of the adult mussel cohort with an increase in tidal height. The effects of exposure on mussel size decreased higher on the shore. On the exposed low shore the maximum size of mussels had a mean length of 102.3mm and was significantly larger (ANOVA, p<0.0001) than that for mussels on sheltered shores (86.7mm). The difference between mean maximum lengths of mussels on the mid shore was not so great, exposed sites had a average mean maximum length of 79.9, while on the sheltered shores it was 68.4mm. On the high shore the difference between the average mean maximum lengths at exposed and sheltered sites was only 3.9mm. The fact that the effects of exposure were greatest on the low shore was also borne out in the PCA. In this analysis low shore exposed and sheltered zones separated into two groups with little overlap, mid shore exposed and sheltered zones were positioned next to each other, and exposed and sheltered high shore zones were clumped together. Densities of adult mussels (>l5mm) were calculated as real densities from randomly placed quads i.e. not from areas of 100% cover. Density decreased up the shore; low, mid and high shore zones were significantly different from each other (ANOVA , p<0.0001; followed by multiple range tests). There was no significant difference between the densities of mussels at exposed and sheltered sites within each zone (ANOVA, p=0.7155). Recruit (<l5mm) densities increased with an increase in adult mussel densities, and this relationship was significant at all zones and for both degrees of exposure (regression analysis, p<0.05 in all cases). The regression between recruits and adults was strongest on the mid and high shore exposed sites. There was a general trend towards stronger regressions and greater predictability with an increase in shore height. The presence of free space within the mussel beds and significant regressions between recruit and adult densities indicates that mussel populations are recruit limited. Mean biomass decreased with an increase in shore height and was probably related to the decrease in size and density of mussels at higher shore levels. Exposure did not affect the average biomass within each zone. A fine scale study of the effects of wave exposure, tidal height and substratum type on recruit densities was undertaken at two sites, viz. Diaz Cross and High Rocks. Two shores, one exposed and one sheltered were identified at each of the sites. All shores were classified prior to sampling. Sampling was completed over a 30 day period during peak recruitment, and samples were removed on as many days as sea and tide conditions permitted i.e. daily when possible. The total density of early plantigrades was greater at Diaz Cross than it was at the High Rocks, and this may be related to the local hydrodynamic patterns adjacent to the two sites. Exposure affected the densities of early and late plantigrades on algae on the low shore sites, where greater numbers of recruits were recorded on exposed low shore zones. Densities of plantigrades on the mussel bed and on algae on the mid and high shore were not affected by exposure. Low and mid shore zones usually had greater densities (at 100% cover of substratum) of plantigrades than the high shore zones, this was probably related to lower settlement rates on the high shore as a result of reduced submergence time. Generally greater plantigrade densities were recorded on algal substrata than on the mussel bed. In only one of the 20 comparisons completed was the density of plantigrades greater on mussels than it was on algae. However when the area of the substratum within a zone was taken into account the number of plantigrades in the mussel bed at a zone was often greater than the number on algae within the same zone. In close to half of these comparisons the total numbers of plantigrades were greater on the mussels than on the algae. This was due to the greater area of mussel bed available to recruits. There was no evidence supporting the suggestion of secondary settlement of plantigrades from algae to the mussel bed. The results of this study demonstrate the importance of wave exposure, tidal height and substratum on certain aspects of the ecology of Perna perna. The importance of these factors is demonstrated at both the adult and early recruit stages of this mussel.

Population dynamics of many intertidal organisms are highly influenced by the abundance and distribution of planktonic larvae in the water column and their arrival on the shore. The brown mussel, Perna perna was used to investigate two of the primary processes that affect population size and dynamics, larval availability and settlement, on the south coast of South Africa. Perna perna is a dominant species on rocky shores of the southern and eastern coasts of South Africa. It creates three-dimensional beds that provide habitats for many other species and hence promotes biodiversity. Larval availability and settlement were examined at different spatial and temporal scales using a nested experimental design. To detect possible relationships between larval availability and settlement, the studies were simultaneous. Two sites, 4km apart, were chosen to investigate mussel settlement patterns. Within each site, three locations (300m from each other) were selected. At each location, five artificial settler collectors were placed at approximately 20cm intervals. Collectors were replaced at a range of time intervals, from daily to seasonal, for 16 months. Each intertidal location was paired with an offshore station, 500m from the shore, where larval availability was measured. At each offshore station, three vertical hauls were collected twice a month using a plankton net. Plankton sampling lasted for 14 months and was designed to examine variability on three temporal scales: seasonal, lunar and daily. The results showed no correlation between the distribution of larvae in the water and settlers on the shore. While larvae were abundant in the water at the start of sampling, they became very rare throughout the rest of the study at both sites and all locations. In contrast, distinct peaks of settler abundance were observed during the seasonal settlement study. In addition to the expected, strong temporal variation that emerged from both studies at all time scales, spatial patterns of variability were also observed. While no spatial effect was detected for the larvae in the water column, there was distinct spatial variation in settlement at the location level: some locations always showed higher settlement than others. These results suggest that, on scales of hundreds of meters to kilometers, larval availability and settlement are very unpredictable in time and that differential delivery of larvae occurs from nearshore waters to the shore. Although the effect of the state of the moon (new or full) was not significant in either study, more settlers seemed to arrive on the shore during new moon. Wind direction did not correlate significantly with settlement. However, the dropping of offshore winds and the prevalence of onshore winds, which are characteristic of summer, may be linked to the start of settlement. Nevertheless, further investigations on tidal or lunar cycles and on the influence of wind on surface currents are required to clarify the effects of moon and wind on settlement.

The general biology and seasonality of breeding of intertidal populations of the brown mussel Perna perna in South Africa are reasonably well known, but we have little information on variability either within or among populations. Synchronous spawning offers adaptive advantages to externally breeding animals. Firstly, it enhances fertilization rates and therefore the species' reproductive fitness. Secondly, spawning can also be timed to coincide with environmental conditions conducive to larval settlement and development. In addition, synchronisation of spawning will influence the synchrony of settlement. Synchronisation of larval settlement, in turn, has implications for popUlation biology, as highly pulsed settlement is likely to lead to density-dependant mortality of recruits and uncoupling of adult/recruit densities, while poorly synchronised settlement will not. Generally, sea temperature and food availability are considered the key factors underlying the initiation and the duration of the breeding cycle of mussels. However, there are proximate local cues that trigger the proliferation, maturation and release of gametes. In this study, the hypothesis tested is that factors that control food availability affect gonad development and so influence synchrony among populations. Much of the published work on spawning is based on observations of the presence of larvae in the plankton, or on settlement. A more reliable method correlates the sequence of gonad development throughout the year with changes in length-weight relationships, using histology. This study is also designed to investigate temporal differences in the timing of the breeding cycle between sheltered and exposed sites along the south coast of South Africa by histological analysis of the reproductive tissue (the gonad) and by dry weight/shell length regreSSIOns. To do this, these two techniques were applied to six mussel populations at three III ocalities that were separated on scales of about 10-20km. Within each locality, two study sites were .dentified. One was exposed to strong wave action and one was sheltered. A few hundred meters ;eparated these sites. The first technique used length-weight regressions as an indication of mussel ~ondition. Abrupt decreases in the dry body weight of a hypothetical standard animal were taken to indicate periods of spawning. Regressions were assessed for samples of 40 mussels taken from each site at intervals of 4 weeks over 13 months. The results were analysed using a 3-way ANCOV A, with dry weight as the dependent variable, shell length as a covariate, and site, exposure and month, as independent variables. The second approach used the more reliable and detailed method of assessing the annual reproductive cycle using histological sections of the gonad. Histological sections of gonads from thirty female mussels, sampled monthly from each site, were examined in the laboratory. Each gonad was categorized into one of six arbitrary developmental stages based on ovary morphology. Synchrony in spawning was examined by comparison of gonad developmental stages of individuals within and among populations. The data were analysed by 3-way nested ANOV A with mean gonad index for each population as the dependent variable, month as an independent variable and exposure nested in site. The results obtained from both techniques showed strong synchronisation among different popUlations, regardless of the scales at which they were separated. The data also indicated good synchrony within populations and, again among populations, regardless of the degree of exposure. The results also indicated that the gonad condition varied significantly at each site, exposure level and month. However, there were significant interactions among these three factors. This means that on a broader seasonal scale the six mussel populations were reasonably synchronised, but on finer monthly scales, there were temporal differences in duration of gametogenic events. The implication is that ovary development is cued by environmental factor(s) that operate on scales of at least 7-20 km rather than more localised events that may affect food availablity either through aggregation of food (local hydrography at different localities) or food delivery to the shore (degree of wave action at different sites.) / KMBT_363 / Adobe Acrobat 9.54 Paper Capture Plug-in

The Mediterranean mussel Mytilus galloprovincialis is the most successful marine invasive species in South Africa. Its presence has had significant ecological consequences on the intertidal communities of the west coast. On the south coast, M galloprovincialis co-exists and competes with the indigenous intertidal mussel Perna perna in the lower balanoid zone, where they show partial habitat segregation. The upper and the lower mussel zones are dominated by M. galloprovincialis and P. perna respectively while they co-occur in the mid zone. In this thesis M. galloprovincialis and P. perna are compared in terms of their population genetics and their ecophysiology. The success of an invader depends on its ability to react to new environmental factors, especially when compared to indigenous species. The distribution and diversity of intertidal species throughout the world are strongly influenced by periodic sand inundation and hydrodynamic stress. Occupying the lower intertidal zone, P. perna is more strongly influenced by sand (burial and sand in suspension) than M. galioprovincialis. Despite this, P. perna is more vulnerable to the effects of sand, showing higher mortality rates under experimental conditions in both the laboratory and the field. M. galioprovincialis has longer labial palps than P. perna, indicating a better ability to sort particles. This, and a higher tolerance to anoxia, explains its lower mortality rates when exposed to burial or suspended sand. Habitat segregation is often explained by physiological tolerances, but in this case, such explanations fail. The ability of a mussel to withstand wave-generated hydrodynamic stress depends mainly on its byssal attachment strength. The higher attachment strength of P. perna compared to M. galioprovincialis and of solitary mussels compared to mussels living within a bed (bed mussels) can be explained by more and thicker byssal threads. M galloprovincialis also has a wider shell, is subjected to higher hydrodynamic loads than P. perna and shows a higher theoretical probability of dislodgement, this is borne out under field conditions. The attachment strength of both species increased from higher to lower shore, in parallel to a gradient of a stronger wave action. Monthly measurements showed that P. perna is always more strongly attached than M. galloprovincialis and revealed seasonal fluctuations of attachment strength for both species in response to wave height. The gonad index of both species was negatively cross-correlated with attachment strength. The results are discussed in the context of the evolutionary strategy of the alien mussel, which directs most of its energy to fast growth and high reproductive output, apparently at the cost of reduced attachment strength. This raises the prediction that its invasive impact will be more pronounced at sites subjected to low or moderate wave action at heavily exposed sites. The potential of a species for invasion is also determined by the ability of the invader to disperse. Population genetics provide indirect information about dispersal through a direct measurement of gene flow. The low genetic divergence (measured as mtDNA) of M. galloprovincialis confirms its recent arrival in South Africa. In contrast, the population genetics structure of P. perna revealed strong divergence on the south-east coast, resulting in a western lineage (straddling the distributional gap of the Benguela System), and an eastern lineage, with an overlap region of the two on the south coast between Kenton-on-Sea and Haga Haga. This genetic disjunction may be caused by Agulhas Current acting as an oceanographic barrier to larval dispersal, or by different environmental selective forces acting on regional populations. Over the last ten years, M. galloprovincialis has shown a decrease or cessation of its spread to the east in exactly the region of the genetic disjunction in P. perna, again suggesting either an oceanographic barrier to larval dispersal, or increasing selection driven by sharp gradients in environmental conditions.

In South Africa, the indigenous mussel Perna perna is threatened by both an invasive species and excessive human exploitation. The Mediterranean mussel Mytilus galloprovincialis is an invasive species that has been introduced to many parts of the world. In South Africa, this species arrived in the 1970s and spread rapidly along the west coast where today it is the dominant mussel species. Along the west coast, M. galloprovincialis is competitively superior in all aspects to the indigenous mussel species, and, as a result, has displaced some of them. On the south coast, M. galloprovincialis found more oligotrophic waters, higher species richness, and a stronger competitor in the indigenous mussel P. perna. The rate of spread of M. galloprovincialis along the south coast has decreased over the last 10 years and the present eastern limit of its distribution in South African is East London. On the south coast, M. galloprovincialis has not yet completely replaced P. perna; instead, the two exhibit spatial segregation, with P. perna dominating the low shore, M. galloprovincialis the high shore and an overlap zone between the two. An experiment on competition was carried out at one site on the south coast. The results showed that, on the low shore, P. perna is a more dominant competitor for space than M. galloprovincialis. Also byssus attachment of the two species differs, P. perna being much stronger than M. galloprovincialis, which suffers high mortality due to wave action on the low shore, especially in monospecific beds. As a result, mortality of M. galloprovincialis through wave action is reduced by the presence of P. perna, which seems to confer protection against dislodgement. However, in the absence of strong wave action, P. perna competitively excludes M. galloprovincialis. Human exploitation along 160 km of coast was examined by sampling mussel populations and using aerial surveys to determine where harvesters were distributed. Collectors did not seem to discriminate between species. The study has shown that higher abundances of mussels were found in protected or inaccessible sites, while in unprotected sites mussels were scarce. Coastal nature reserves are being proven to be effective in protecting mussel populations.

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.

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.

Recruitment of planktonic larvae into sedentary benthic populations regulates the population dynamics of marine invertebrates. The processes controlling recruitment, however, are poorly understood, and recruitment remains largely unpredictable, which complicates management of exploited shellfish resources. The mussels Perna perna, Choromytilus meridionalis and Mytilus galloprovincialis, found on the south coast of southern Africa, have planktonic larvae and sedentary adult stages. This thesis examines dispersal, settlement and early post-settlement growth and mortality, and their effect on recruitment and demography of intertidal mussel populations in the region of Algoa Bay on the south coast of southern Africa. Temporal and spatial variation in the body mass, density and size structure of mussels, the distribution of bivalve larvae on plankton grids in the nearshore zone and the distribution of a recently introduced invasive mussel, Mytilus galloprovincialis, were examined between 1989 and 1992. Furthermore, data on hourly or daily changes in wind strength and direction, air and sea surface temperatures and low and high tide levels in the study region, were obtained. There were 3-4 peaks in spawning (characterised by an abrupt decline in weight) and settlement activity annually. These peaks varied in exact timing, intensity and duration between sites and over time. However, at a site, spawning was followed by settlement 4-8 weeks later, and there was a significant (P < 0.05) direct correlation between spawning intensity prior to the appearance of a new cohort and the cohort density (settlement intensity). The stochastic spatial and temporal variation in breeding activity was superimposed on a more general pattern of a higher intensity of spawning and settlement in Algoa Bay than on the open coast, and a higher settlement intensity on coastal sandstone shores than on dune rock shores. Spawning was more frequent in winter and spring, and the probability of spawning and settlement peaked around the spring and autumn equinox, if temperature and wind conditions were suitable. Larval behaviour had little effect on their dispersal in the well-mixed nearshore region. Larvae were passively dispersed by currents, and their dispersal range and direction depended on prevailing winds and local topography. The sharp decline in density of recruit and adult M. galloprovincialis with increasing distance from the point of introduction, showed that some larvae were carried by wind generated currents over moderately long distances (-100 km). However, since most (76 %) M. galloprovincialis recruited within 4 km of the parent population, it is possible that larvae become trapped in small gullies and crevices around rocky shores, and have a limited dispersal range. This could explain the link between local patterns of spawning and settlement. The distribution and abundance of settlers on the shore was influenced by larval behaviour and the availability of settlement, substrata. Larvae preferred to settle primarily on foliose coralline algae and migrate to the adult mussel bed when they were larger (0.60-7 mm), but larvae also settled directly on adult mussels, possibly because the amount of coralline algae was limited. Both direct and secondary settlement were considered to be important in maintaining mussel populations since the rate of settlement was low(generally < 60 000.m-2). Cohort analyses showed that prior to maturity post-settlement growth (- 30 mm in 10 months) and mortality rates (60-100%) were high, but varied. When settlement intensity was low this variability uncoupled the relationship between spawning and recruitment intensity. Multiple regression analysis showed that together reproductive effort (gamete output), settlement intensity, growth and mortality prior to maturity, accounted for 76 % of the variance in recruitment into mature adult populations. The low settlement rate coupled with the short life span of mussels « 3 years), meant that populations underwent marked spatial and temporal variations in structure and abundance as settlement intensity varied, but there were consistent general differences between mussel populations on dune rock and sandstone shores in Algoa Bay and on the open coast. It was concluded from these results that, spawning intensity and post-settlement growth and mortality, rather than dispersal, regulated recruitment and the structure and abundance of intertidal P. perna and C. meridionalis populations along the south coast of southern Africa. On the basis of these results it is recommended that species with limited dispersal, variable recruitment and high natural mortality, such as P. perna, should be conserved by protecting a small part of the population in reserves, and controlling utilisation outside reserves to minimize disturbance to local brood stocks. Furthermore, since the potential for reseeding adjacent exploited areas is limited, several small reserves placed at regular intervals along the coast would be more effective than a single large reserve.