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Abstract: An eastward shift in the center of abundance of the South African rock lobster Jasus lalandii occurred during the early 1990s into an area known as East of Cape Hangklip (EOCH). Given (1) the predatory capabilities of J. lalandii, (2) an intricate relationship between the urchin Parechinus angulosus and juveniles of the abalone Haliotis midae, and (3) existing over-exploitation of elements of the ecosystem, the ‘lobster invasion’ has major implications for the benthic ecosystem and associated fisheries. We surveyed the abundance of J. lalandii and the benthic community composition EOCH at six sites (three ‘invaded’ and three ‘non-invaded’ by J. lalandii), in three different depth zones (<5m, 6–12m and 13–20m). At all depths, J. lalandii was significantly more abundant in invaded areas than in non-invaded areas, and benthic communities were significantly different. The high densities of rock lobsters at invaded sites led to cascading effects including elimination of urchins and depletion of grazers, consequent enhancement of macroalgae, and diminishment of encrusting corallines. Non-invaded sites had few lobsters, abundant herbivores, less macroalgae and more encrusting corallines. Elimination of urchins in the invaded area has important implications, as juveniles of the commercially harvested abalone H. midae depend on shelter beneath urchins in this region. Floral species diversity was greater at invaded sites and increased with depth, whereas faunal species diversity was greater at non-invaded sites but also increased with depth. The depths at which strongest effects of J. lalandii were felt coincided with the depth of maximum abundance of the urchin P. angulosus, the abalone H. midae, the kelp Ecklonia maxima and encrusting corallines, with serious consequences for associated fisheries and the benthic community. We argue that the differences between invaded and non-invaded areas are sufficient to recognize that they are alternate stable-states and constitute a regime shift. [Copyright &y& Elsevier]

Abstract: As the alien species that most dominates space along the South African coast, the Mediterranean mussel Mytilus galloprovincialis has radically altered community composition on invaded shores. We experimentally assessed interspecific interactions between this invasive species and dominant indigenous species in conjunction with considering how wave action moderates such interactions. The density of both M. galloprovincialis and the limpet Scutellastra granularis increased with wave action. Conversely, the tube-building polycheate Gunnarea capensis was negatively affected by wave exposure, being most abundant on sheltered shores. The influence of wave action on the indigenous mussel Aulacomya ater, however, remains unclear. M. galloprovincialis outcompeted both G. capensis and A. ater at moderate to high exposure levels, whereas it had both positive and negative effects on S. granularis. It outcompeted adult limpets on primary rock space on semi-exposed and exposed shores, reducing densities of this portion of the population. However, recruitment of S. granularis was facilitated by M. galloprovincialis, as greater numbers recruited to the secondary substratum offered by mussel shells. Again this interaction intensified with wave action. Due to the extremely high density of recruits on secondary space, the net effect of M. galloprovincialis on S. granularis was positive. Thus, wave action not only influences the abundance of individual species, but also mediates both positive and negative interspecific interactions in rocky shore communities, including the impact of alien species such as M. galloprovincialis. [Copyright &y& Elsevier]

Abstract: Grazing is a key factor structuring aquatic communities, and can have both positive and negative effects. Here, we report on the effects of an intertidal estuarine gastropod, Assiminea globulus, on sediment bacteria, microphytobenthos and meiofauna, based on field observations and manipulative experiments. Field observations showed inconsistent patterns of chlorophyll-a concentrations in zones occupied by A. globulus relative to zones lacking this species, and meiofaunal density increased downshore. Bacterial density, however, was always consistently and significantly elevated in zones of A. globulus across all sites sampled, suggesting promotion of bacterial density by this snail. Field experiments were more conclusive, showing a significant negative effect of A. globulus on sediment chlorophyll-a levels at both the surface and at a sediment depth of 2cm, but positive effects on bacterial density at both depths. A. globulus density did not affect overall meiofaunal density, but did affect meiofaunal community structure, generally by depressing the density of harpacticoid copepods but promoting platyhelminthes, and enhancing taxonomic diversity at high density. The complex effects of A. globulus probably reflect the interactive effects of bioturbation, fertilisation, grazing and pelletisation, all of which need to be considered when understanding grazer effects on soft-sediment communities. [Copyright &y& Elsevier]

Abstract: Besides the direct impacts of exploitation on target species, indirect effects on non-target species are unavoidable and find expression in changes in community structure. We quantified the effects of experimental harvesting of Mytilus galloprovincialis on intertidal communities on the South African west coast. In the mid- and low-shore, four months of harvesting at intensities greater than F =0.3 and F =0.6 respectively, resulted in significant changes in community composition. These changes were driven by progressively greater spatial dominance by the macroalgae Cladophora flagelliformis, Porphyra capensis and Ulva species as harvesting intensity increased. Four months after cessation of harvesting, community structure had not recovered and even areas subjected to as little as F =0.3 supported significantly altered communities in both zones. The fact that substantial community changes were induced by even low-intensity exploitation is indicative of low resilience to harvesting. The densities and cover of the dominant taxa returned to a pre-harvest state 16 months after the cessation of harvesting, but multivariate analyses indicated that the overall community composition required 32 months for Cover Letter full recovery. Although these communities displayed elasticity within three years, it is recommended that should a M. galloprovincialis fishery be established in the region, harvesting be implemented at a maximum intensity of F =0.3. This approach would maximise yields and protect associated intertidal communities. [Copyright &y& Elsevier]

Abstract: Successful fisheries management is underpinned by an understanding of the processes that underlie the population dynamics of exploited stocks. This study considered the effects of experimental harvesting on recruitment of Mytilus galloprovincialis along the west coast of South Africa, where harvesting of this alien species is being contemplated. In particular, the role of settlement habitat availability in the form of adult mussels was analysed. To track the effects of a spectrum of harvesting intensities, five treatments were implemented: F =0 (i.e. a control), F =0.3, F =0.6, F =0.9 and F =1. At these harvesting intensities 0%, 30%, 60%, 90% or 100% of mussel biomass was removed respectively at the outset of the experiment. A significant negative exponential relationship (p <0.01) was found between M. galloprovincialis recruit density and harvesting intensity, with intensities greater than F =0.3 dramatically reducing recruitment. This pattern was recorded throughout the intertidal zone and remained temporally constant over 2 years. Significant positive linear relationships (p <0.01) between recruit density and adult mussel biomass or density indicate a strong correlation between availability of settlement habitat and recruitment. It is likely that the high recruit density recorded at low harvesting intensities (2000–20000 per 0.01 m2) exceeds the level required for population maintenance. However, if settlement habitat is eliminated or significantly reduced, as was achieved by F =0.3 or above, recruitment may become limiting. Thus, to protect stock replenishment, harvesting of M. galloprovincialis in this region should take place at intensities less than F =0.3. [Copyright &y& Elsevier]

Abstract: Field observations and experimental evidence have shown that bioturbation by the southern African sandprawn Callianassa kraussi may significantly influence the abundance and distribution of the filter-feeding bivalve Eumarcia paupercula and the grazing gastropod Nassarius kraussianus. It was hypothesized that (1) sediment reworking by C. kraussi negatively affects microalgal growth on the sediment surface, leading to a reduction in food intake by N. kraussianus, (2) sediment deposited by C. kraussi will also diminish the food uptake of E. paupercula by interfering with its filtration mechanism. To test these hypotheses, manipulative field and laboratory experiments were undertaken in which N. kraussianus and E. paupercula were added to treatments with and without C. kraussi, and their survival and gut chlorophyll-a content measured. The effects of C. kraussi on sediment erodability and on condition of E. paupercula (tissue mass/shell length) were determined in a second experiment. In the presence of C. kraussi, (1) microalgal consumption by both N. kraussianus and E. paupercula was halved; (2) condition and survival of E. paupercula were significantly reduced but survival of N. kraussianus was unaffected; and (3) sediment erodability was increased. A significant negative relationship was established between sediment erodability and condition of E. paupercula. Evidently C. kraussi exerts a strongly negative influence on the feeding of E. paupercula and N. kraussianus, and this may explain the scarcity of these organisms in areas containing high densities of C. kraussi. [Copyright &y& Elsevier]

Abstract: The faunal composition of high-shore beds of eelgrass Zostera capensis versus unvegetated sandflats dominated by the sandprawn Callianassa kraussi was investigated by experimentally transplanting plots of eelgrass into sandflats where the sandprawn was either left unmanipulated or eliminated by prior faunal suffocation. After 12 months, multidimensional scaling defined five faunal clusters that were dictated primarily by the presence or absence of eelgrass rather than sandprawns. By 18 months, eelgrass and sandprawns played equal roles in defining faunal clusters. After 30 months, sandprawns had displaced the eelgrass transplants and faunal composition converged on that of sandflat control plots. Most species were either ‘eelgrass-associated’ or ‘sandflat-associated’, but three had unique responses to the treatments: two high-shore species (Assiminea globulus and Hydrobia sp.) that were abundant in eelgrass beds never appeared in the eelgrass transplants, whereas another eelgrass-occupant, Siphonaria compressa, became superabundant on the transplants for 18 months. Unexpectedly, the eelgrass fauna was never more rich or diverse than that of unvegetated sandflats, although abundance was greater in the eelgrass. As hypothesised, the sandflats were disproportionally populated by burrowers and eelgrass by non-burrowers, but a second hypothesis that these habitats should support mainly hard-bodied and soft-bodied animals respectively was rejected. The distinctively different faunas of eelgrass and unvegetated sandflats are seemingly respectively maintained by sediment stabilisation by eelgrass versus bioturbation by sandprawns. The transplants of eelgrass into sandflats developed an eelgrass-associated fauna provided sandprawns were excluded, but ultimately reverted to a sandflat fauna once sandprawns reinvaded. [Copyright &y& Elsevier]

Abstract: Increasing recognition is being given to the concept of ‘ecosystem engineers’, which alter the physical nature of the environment and thereby influence other species by means that are neither competitive nor trophic. This paper examines a case study of such effects, in which negative correlations between the abundances of the sandprawn Callianassa kraussi and the eelgrass Zostera capensis imply mutual exclusion because Z. capensis stabilises sediment and inhibits burrowing by C. kraussi, whereas sandprawn bioturbation smothers eelgrass. To test this, we undertook manipulative experiments in which Z. capensis was transplanted into sandflats where C. kraussi was either left undisturbed or eliminated by defaunation. This demonstrated that (1) Z. capensis thrived and expanded in sandflats in the absence of bioturbation but deteriorated and disappeared if C. kraussi was present. (2) In the short term, introduction of Z. capensis reduced densities of C. kraussi, and in established beds of eelgrass, C. kraussi was rare. (3) The mudprawn Upogebia africana was largely restricted to treatments that contained Z. capensis, and its densities were greater in the absence of C. kraussi than in its presence. The presence of eelgrass and the exclusion of C. kraussi also diminished sediment penetrability, suspension of particles and surface burial rates compared to the situation in undisturbed Callianassa-dominated sandflats. These results confirm a mutually negative interaction between C. kraussi and Z. capensis and strengthen the belief that the underlying mechanisms are the antagonistic effects of bioturbation by C. kraussi versus sediment stabilisation by Z. capensis. U. africana seems to benefit directly from Z. capensis but possibly also benefits indirectly from the exclusion of C. kraussi by Z. capensis, since its filter-feeding mode of life requires sufficient sediment stability to maintain semi-permanent U-tubes. [Copyright &y& Elsevier]

Abstract: The influence of an alien mussel Mytilus galloprovincialis (Lamarck) on an indigenous limpet Scutellastra argenvillei (Krauss) was investigated on the west coast of South Africa by comparing four situations in which limpets occupied rock patches surrounded by mussels: (1) ‘established patches’ of high-density adult limpets; (2) ‘experimentally cleared patches’ from which mussels were manually removed; (3) ‘naturally disturbed patches’ where mussels had been eliminated by wave action; and (4) ‘half-density’ patches formed by thinning established patches. Limpets in established patches were most effective in retarding settlement and lateral invasion by mussels, but all patches shrank due to mussel encroachment. As patches shrank, limpet density fell in established and naturally disturbed patches due to emigration. Limpets living on mussel beds were small, sparse and never achieved sexual maturity; 60% shifted to rock, whereas only 7% moved from rock to mussels. The limpets showed highest fidelity to established patches (79% after 12 months) and lowest fidelity in naturally disturbed patches (20%). Thinning of established patches reduced fidelity to 26%. Mussel beds did not provide a suitable alternative substratum for S. argenvillei after they displaced the limpets from rocks. Moreover, elimination of mussels by waves failed to allow S. argenvillei to re-establish dense, adult populations resembling those prior to arrival of the mussel. Adults of S. argenvillei feed collectively by trapping kelp blades. On the topographically complex mussel beds limpets cannot feed this way. In sum, on exposed shores where M. galloprovincialis achieves maximal recruitment and growth, S. argenvillei is incapable of preventing mussel encroachment and is likely to become completely displaced. Semi-exposed shores do, however, offer a refuge preventing global extinction of the limpet. [Copyright &y& Elsevier]

The Mediterranean mussel Mytilus galloprovincialis invaded the shores of South Africa in about the mid-1970s. It was first detected at a harbour in Saldanha on the west coast and apparently arrived accidentally. From there, it spread at a rate of about 115 km·year−1 and now occupies the whole of the west coast of South Africa and at least the southern half of Namibia. It was deliberately introduced from the west coast to the south coast for mariculture. In this case study, we record its effects on intertidal rocky shores, cast in terms of predictions based on (a) the history of its invasions elsewhere, (b) its mode of dispersal, (c) its physiological performance relative to indigenous mussels, (d) the role of wave action as a moderator of competition, (e) the influence of relative body sizes, (f) the projected effects of the mussel on infauna, (g) consumption by higher trophic levels, and (h) rates of parasitism.Several properties of M. galloprovincialis itself, and of the recipient community, conspired to favour the spread and establishment of this alien mussel, including high productivity on the west coast of South Africa, prevalently strong wave action, a sparsity of predators, an absence of parasites, the mussel''s fast growth and high reproductive output, and its possession of a planktotrophic larva. It competitively displaces several species because of its physiological performance. Some of the species gain a substitute substratum on the mussels themselves, but only if they are small enough to live and reproduce on the mussels. M. galloprovincialis has had little effect on infauna, but has provided an additional source of food for higher predators, including the rare and endangered African Black Oystercatcher (Haematopus moquini).Nearly all these effects and conditions were forecasted (or, at the least, explainable with hindsight), but despite these successes in predicting the impacts of M. galloprovincialis, its spread was not only unavoidable but was encouraged by its transfer to the south coast for mariculture. Moreover, there was one completely unpredictable effect of M. galloprovincialis—which led to mass mortalities of a swimming crab. Given the failure of even quite detailed and accurate predictions to allow control of M. galloprovincialis once it arrived, prevention rather than cure must be the prime means of avoiding future unwanted introductions of invasive species. [Copyright &y& Elsevier]

Almost all juveniles of the abalone Haliotis midae shelter beneath the urchin Parechinus angulosus. Field observations showed that all juvenile abalone remained concealed under urchins by day. Even by night, only 10–25% left the protection of urchins to forage. Offered a choice of rocks or urchins in tide pools, the juveniles strongly preferred urchins rather than rocks. In aquaria, both urchins and abalone were active only at night. The frequency with which juvenile abalone left shelter was greatest if rocks constituted the only form of concealment, intermediate when they sheltered under urchins alone and least when the urchins had access to drift kelp. The treatment that provided abalone with both urchins and drift kelp had the lowest rate of urchin movement and the highest microalgal benthic standing stocks, the gut contents of the abalone comprised mainly kelp and their gut fullness indices were higher than in the other treatments. Concealment of juvenile abalone beneath urchins evidently provides them with several benefits. First, when urchins trap drift kelp, the abalone are capable of feeding on it and enhance their uptake of food. Second, this behaviour reduces the frequency with which juvenile abalone leave the shelter of urchins. Third, urchins that trap drift kelp reduce their rates of movement and their consumption of benthic microalgae, and we speculate that this may improve feeding conditions for the early postsettlement “recruit” stages of H. midae, and reduce the risk of urchins “bulldozing” them. Thus, for a complex of reasons, the dependency of P. angulosus on drift kelp provides benefits to abalone that would not exist if this urchin acted as a conventional grazer. [Copyright &y& Elsevier]