Supply-side ecology of the brown mussel, Perna perna: an investigation of spatial and temporal variation in, and coupling between, gamete release and larval supply

Sampling of recruitment-associated variables of Perna perna was done approximately monthly for 14 months at intertidal locations 500 m apart, nested within sites 25 km apart. Paired with intertidal locations were nearshore locations, 600 m to sea. Sampling assessed spawning, densities of larvae in the water column and densities of late plantigrades and juveniles on the shore. Major events in each variable were synchronous over larger scales (10s of kilometres) while subsidiary events were synchronised at smaller scales, varying within sites (100s of metres) or even within locations (metres). This suggests that the processes driving major events operated over large scales while processes operating at much more local scales drove less intense, more localised events. A major spawning event occurred at all locations in May–June 1998. Weaker spawning events occurred at different times in different locations. Larvae were found on 80% of sampling occasions, densities peaking in January–March 1998 and 1999 at all locations. Plantigrades and juveniles showed less clear patterns, with considerable residual variation. There was no sign of strong coupling among variables with few significant direct or cross correlations. The major sources of variability shifted from time to space as one progressed from spawning, to plantigrade density to juvenile density. For spawning, time was the most important source (58%) of heterogeneity and space accounted for little (8%) of the total variance. For larvae and late plantigrades, time was still the most important source of variability (41% and 33%, respectively), but space was a much more substantial component. For juveniles, small-scale (residual) spatial variability dominated total variability (75%). This strongly suggests the importance of hydrography and its effects on variation in delivery of larvae to the intertidal from offshore. These findings also indicate greater spatial heterogeneity as recruits age, reflecting small-scale variations in larval delivery and the increasing importance of post-settlement mortality

Scales of mussel bed complexity: structure, associated biota and recruitment

Hierarchically scaled surveys were carried out on beds of the brown mussel Perna perna (Linnaeus) on the South coast of South Africa. The object was to assess spatial and temporal variations in the complexity of mussel beds and to investigate relationships between mussel bed complexity and mussel recruitment. Complexity was divided into three components: physical complexity; demographic complexity; associated biota. A series of variables within each component were recorded at two different scales (10 and 50 cm) within nested quadrats on three separate occasions. The nested ANOVA design explicitly incorporated spatial scale as levels of the ANOVA. These scales were: shores (areas 1 km in length separated by 25 km); transects (areas 20 m in length separated by 100s of meters); 50×50-cm quadrats separated by meters and 10×10-cm quadrats separated by cm) This approach was intended to generate hypotheses concerning direct associations between recruitment and complexity versus co-variation due external processes. Three main questions were addressed: (1) At what scale does each variable of complexity exhibit greatest significant variation? (2) At these scales is there similar ranking of variables of complexity and recruitment? (3) Within this/these scales, is there any significant relationship between the variables measured and mussel recruitment? On two occasions (Nov. 97 and Mar. 98) the majority of variables showed greatest significant variation at the transect-scale. On a third occasion (Oct. 97) most variables showed greatest significant variation at the quadrat-scale and the site-scale. On all occasions a markedly high percentage of the variation encountered also occurred at the smallest scale of the study, i.e., the residual scale of the ANOVA analyses. Some similarity in the ranking of variables occurred at the transect scale. Within the transect-scale, there was little indication of any relationship between variables of complexity and recruitment. Relationships were inconsistent either among transects or among sampling occasions. Overall, the results suggest that a high degree of variation in mussel bed complexity consistently occurs at very small scales. High components of variance generally also occur at one or more larger scales; however, these scales vary with season. Mussel recruitment does not appear to be directly affected by complexity of mussel beds. Instead it appears external factors may influence both complexity and recruitment independently. In addition recruitment may influence complexity rather than vice versa

The Southern Ocean Group at Rhodes University: seventeen years of biological oceanography in the Southern Ocean reviewed

This paper reviews the main findings of the Southern Ocean Group at Rhodes University over the last 17 years. A primary contribution has been the development of conceptual models of the physical-biological driving mechanisms that support enormous seasonal populations of land-based top predators at the Prince Edward Islands. Collectively, these models are referred to as the life-support system of the islands. Near-shore subcomponents of the ecosystem, including inshore feeding predators, are largely supported by autochthonous primary production of kelps and localized diatom blooms. These energy sources feed indirectly into top predator populations via the benthic communities. A crucial link is formed by the bottom-dwelling shrimp, Nauticaris marionis, which feeds largely on benthic species and detritus and is eaten by a number of diving seabirds. The frontal systems that lie north and south of the islands are important feeding grounds for offshore feeding birds. A decadal-scale southward shift in the position of the Sub-antarctic Front towards the islands is reflected in increases in populations of these species.Rhodes Centenary issu

Mechanisms of habitat segregation between an invasive (Mytilus galloprovincialis) and an indigenous (Perna perna) mussel: adult growth and mortality

The invasive mussel Mytilus galloprovincialis and the indigenous mussel Perna perna coexist intertidally on the south coast of South Africa through partial vertical habitat segregation: M. galloprovincialis dominates the upper shore and P. perna the lower shore. Recruitment patterns can explain the zonation of P. perna, but not the invasive species. We examined the role of post-recruitment interactions by measuring spatial and temporal differences in adult growth and mortality rates of the two species. Specifically, we tested the hypothesis that interspecific differences in growth and mortality reflect adult distribution patterns. The two study locations, Plettenberg Bay and Tsitsikamma, are 70 km apart with two sites (separated by 300–400 m) per location, each divided into three vertical zones. Growth was measured seasonally using different marking methods in 2001 and 2003. Cumulative adult mortality was measured through summer in 2003/2004. Both species generally grew more slowly upshore, but they showed different effects of season. For P. perna, growth was significantly reduced in winter in the low zone, but unaffected by season in the high zone. For M. galloprovincialis, growth was either unaffected by season or increased in winter, even in the high zone. Thus, growth of P. perna and M. galloprovincialis was reduced under cool winter and warm summer temperatures, respectively; and while growth was more similar between species in summer, M. galloprovincialis grew much faster than P. perna in winter. Mortality of P. perna increased upshore. For M. galloprovincialis, mortality was not zone-dependent and was significantly greater than for P. perna on the low-shore and (generally) across the shore in Tsitsikamma. Both species had higher growth and mortality rates in Plettenberg Bay than in Tsitsikamma. Thus, P. perna seems able to maintain spatial dominance on the low-shore and at certain sites because of higher mortality of M. galloprovincialis. We conclude that seasonality in growth of the two species reflects their biogeographic affinities and that coexistence is possible through pre-recruitment effects that limit the vertical distribution of P. perna and post-recruitment effects that limit M. galloprovincialis

Will the invasive mussel Mytilus galloprovincialis Lamarck replace the indigenous Perna perna L. on the south coast of South Africa?

The mussel Mytilus galloprovincialis is invasive worldwide, has displaced indigenous species on the west coast of South Africa and now threatens Perna perna on the south coast. We tested the hypothesis that Mytilus will replace Perna by examining changes in their distribution on shores where they co-exist. Total cover, adult density, recruit density, recruit/adult correlations and mean maximum lengths of both species were measured in 2001 at two contrasting sites (Plettenberg Bay and Tsitsikamma) 70 km apart, each including two locations 100 m apart. Cover and density were measured again in 2004. Total mussel abundance was significantly lower in Tsitsikamma, and recruit density was only 17% that of Plettenberg Bay. Abundance and cover increased upshore for Mytilus, but decreased for Perna, giving Mytilus higher adult and recruit density and total cover than Perna in the upper zones. Low shore densities of recruits and adults were similar between species but cover was lower for Mytilus, reflecting its smaller size, and presumably slower growth or higher mortality there. Thus, mechanisms excluding species differed among zones. Recruitment limitation delays invasion at Tsitsikamma and excludes Perna from the high shore, while Mytilus is excluded from the low shore by post-recruitment effects. Recruitment limitation also shapes population structure. Recruit/adult correlations were significant only where adult densities were low, and this effect was species-specific. Thus, at low densities, larvae settle or survive better near adult conspecifics. After 3 years, these patterns remained strongly evident, suggesting Mytilus will not eliminate Perna and that co-existence is possible through partial habitat segregation driven by recruitment limitation of Perna on the high shore and post-settlement effects on Mytilus on the low shore

Wave exposure effects on population structure and recruitment in the mussel Perna perna suggest regulation primarily through availability of recruits and food, not space

Recruitment and population structure of Perna perna in low shore mussel beds were investigated over 15 months at six sites along the south coast of South Africa. Initial, subjective classification of sites as wave exposed or wave sheltered (three of each) was confirmed using the dissolution of cement blocks to measure average water flux and dynamometers for maximum wave force. Recruitment occurred throughout the year, but recruit (1–5 mm) densities were significantly higher from January to April 1996 on both shore types. Recruit densities were positively correlated with adult (>15 mm) densities for both shore types (P 0.05). Thus wave exposure dramatically affects density, recruitment and mussel size, but not recruitment timing or biomass where there is 100% cover, and mediates a three-way interaction among food supply, larval supply and intraspecific competition for space. In contrast to shores with saturation recruitment, mussel biomass here appears to be limited by recruit supply and constraints of food, especially on sheltered shores, while density is regulated through intraspecific competition for space primarily on exposed shores and at small spatial scales

Contrasting spatial heterogeneity of sessile organisms within mussel (Perna perna L.) beds in relation to topographic variability

We examined the spatial heterogeneity in three sessile rocky shore organisms, the mussel Perna perna, the barnacle Octomeris angulosa (Sowerby) and the red alga Gelidium pristoides (Turn.) at a range of continuous local scales along horizontal transects within mid- and upper mussel beds of South African shores. We also examined the relationships between variability of organisms and topographic features (rock depressions, slope, aspect), and between mussel, barnacle and algal variability over the same scales. To estimate spatial heterogeneity, we analyzed scaling properties of semivariograms using a fractal approach. Relationships between different variables at the different scales were examined by cross-semivariograms. Spatial dependence of P. perna variability increased with spatial dependence of topographic variability, so that scaling regions of mussel and topographic distributions corresponded well. This relationship often improved with larger local scales (mussel cover increased with depressions, steeper slope and aspect towards waves), while at smaller spatial scales, variability in mussel cover was less well explained by variability in topography. The variability of the barnacle O. angulosa exhibited spatial dependence, even on topographically unstructured shores. In contrast, the distribution of the alga G. pristoides revealed high fractal dimensions, showing spatial independence on topographically unstructured shores. Algae also showed a very strong negative relationship with mussels at most local scales, and a negative relationship with barnacles in upper zones, especially at larger local scales. Barnacles may show clear spatial dependence because of hydrodynamics (at larger local scales) and the need to find a future mate in close proximity (at smaller local scales), while algae may show a strong negative relationship with mussels because of competition for space

Temporal scales of variation in settlement and recruitment of the mussel Perna perna (Linnaeus, 1758)

Population dynamics of many intertidal organisms are strongly affected by the abundance and distribution of larvae arriving on the shore. In particular, not only absolute numbers of settlers but also the degree of synchronisation of settlement can have a strong influence on whether density-dependent or density-independent processes shape adult shape populations. Temporal variation in rates of settlement and recruitment of the mussel Perna perna on the south coast of South Africa was investigated using a nested spatial design at different temporal scales. Variability in settlement at spring tides was examined at two temporal scales: lunar (to investigate the effect of state of the moon on settlement) and tidal (to investigate the influence of state of the tide on mussel settlement). Recruitment over neap tides was examined at one temporal scale, fortnight (to investigate the effect of date on mussel recruitment). Strong temporal variation was evident for both settlement and recruitment, but not at all time scales. Distinct peaks of settler/recruit abundance were observed during the lunar and neap tide studies. Recruitment intensity differed over the course of the year, and pulsing of recruitment was generally synchronised among locations. However, the strength of pulsing differed dramatically among locations, giving a significant interaction between fortnight and location. The finest temporal scale, investigated in the tidal study, did not reveal a significant effect of the state of the tide on settlement. The state of the moon (new or full) was not significant as a main factor (p = 0.052), although generally more settlers arrived on the shore during new moon. Phase of the moon appeared to have an effect on settler abundances, but only when and where densities were high

Environmental domains and range-limiting mechanisms: testing the Abundant Centre Hypothesis using southern African sandhoppers

Predicting shifts of species geographical ranges is a fundamental challenge for conservation ecologists given the great complexity of factors involved in setting range limits. Distributional patterns are frequently modelled to “simplify” species responses to the environment, yet the central mechanisms that drive a particular pattern are rarely understood. We evaluated the distributions of two sandhopper species (Crustacea, Amphipoda, Talitridae), Talorchestia capensis and Africorchestia quadrispinosa along the Namibian and South African coasts, encompassing three biogeographic regions influenced by two different oceanographic systems, the Benguela and Agulhas currents. We aimed to test whether the Abundant Centre Hypothesis (ACH) can explain the distributions of these species’ abundances, sizes and sex ratios and examined which environmental parameters influence/drive these distributions. Animals were collected during a once-off survey at 29 sites over c.3500 km of coastline. The ACH was tested using a non-parametric constraint space analysis of the goodness of fit of five hypothetical models. Distance Based Linear Modelling (DistLM) was performed to evaluate which environmental traits influenced the distribution data. Abundance, size and sex ratio showed different patterns of distribution. A ramped model fitted the abundance (Ramped North) and size (Ramped South) distribution for A. quadrispinosa. The Inverse Quadratic model fitted the size distribution of T. capensis. Beach slope, salinity, sand temperature and percentage of detritus found on the shore at the time of collection played important roles in driving the abundance of A. quadrispinosa. T. capensis was mainly affected by salinity and the morphodynamic state of the beach. Our results provided only some support for the ACH predictions. The DistLM confirmed that the physical state of the beach is an important factor for sandy beach organisms. The effect of salinity and temperature suggest metabolic responses to local conditions and a role in small to mesoscale shifts in the range of these populations

Biogeographical patterns of endolithic infestation in an invasive and an indigenous intertidal marine ecosystem engineer

By altering the phenotypic properties of their hosts, endolithic parasites can modulate the engineering processes of marine ecosystem engineers. Here, we assessed the biogeographical patterns of species assemblages, prevalence and impact of endolithic parasitism in two mussel species that act as important ecosystem engineers in the southern African intertidal habitat, Perna perna and Mytilus galloprovincialis. We conducted large-scale surveys across three biogeographic regions along the South African coast: the subtropical east coast, dominated by the indigenous mussel, P. perna, the warm temperate south coast, where this species coexists with the invasive Mediterranean mussel, M. galloprovincialis, and the cool temperate west coast dominated by M. galloprovincialis. Infestation increased with mussel size, and in the case of M. galloprovincialis we found a significantly higher infestation in the cool temperate bioregion than the warm temperate region. For P. perna, the prevalence of infestation was higher on the warm temperate than the subtropical region, though the difference was marginally non-significant. On the south coast, there was no significant difference in infestation prevalence between species. Endolith-induced mortality rates through shell collapse mirrored the patterns for prevalence. For P. perna, endolith species assemblages revealed clear grouping by bioregions. Our findings indicate that biogeography affects cyanobacteria species composition, but differences between biogeographic regions in their effects are driven by environmental conditions.Agência financiadora Número do subsídio Fundacao para a Ciencia e Tecnologia (FCT-MEC, Portugal) UID/Multi/04326/2019 IF/01413/2014/CP1217/CT0004 South African Research Chairs Initiative (SARChI) of the Department of Science and Technology National Research Foundationinfo:eu-repo/semantics/publishedVersio