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Self-organized spatial patterns are increasingly recognized for their contribution to ecosystem functioning. They can improve the ecosystem's ability to respond to perturbation and thus increase its resilience to environmental stress. Plastic pollution has now emerged as major threat to aquatic and terrestrial biota. Under laboratory conditions, we tested whether plastic leachates from pellets collected in the intertidal can impair small-scale, spatial self-organization and byssal threads production of intertidal mussels and whether the effect varied depending on where the pellets come from. Specifically, leachates originating from plastic pellets collected from relatively pristine and polluted areas respectively impaired and inhibited the ability of mussels to self-organize at small-scale and to produce byssal threads compared to control conditions (i.e., seawater without leaching solution). Limitations to natural self-organizing processes and threads formation may translate to a declined capacity of natural ecosystems to avoid tipping points and to a reduced restoration success of disturbed ecosystems., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier B.V. All rights reserved.)

Interspecific and intraspecific diversity are essential components of biodiversity with far-reaching implications for ecosystem function and service provision. Importantly, genotypic and phenotypic variation within a species can affect responses to anthropogenic pressures more than interspecific diversity. We investigated the effects of interspecific and intraspecific diversity on microplastic ingestion by two coexisting mussel species in South Africa, Mytilus galloprovincialis and Perna perna, the latter occurring as two genetic lineages. We found significantly higher microplastic abundance in M. galloprovincialis (0.54 ± 0.56 MP items g -1 WW) than P. perna (0.16 ± 0.21 MP items g -1 WW), but no difference between P. perna lineages. Microbeads were the predominant microplastic (76 % in P. perna, 99 % in M. galloprovincialis) and polyethylene the prevalent polymer. Interspecific differences in microplastic abundance varied across locations, suggesting diverse sources of contamination. We suggest that microplastic ingestion can be species-specific even in organisms that coexist and play similar functional roles within ecosystems., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Environmental filtering (EF), the abiotic exclusion of species, can have first order, direct effects with cascading consequences for population dynamics, especially at range edges where abiotic conditions are suboptimal. Abiotic stress gradients associated with EF may also drive indirect second order effects, including exacerbating the effects of competitors, disease, and parasites on marginal populations because of suboptimal physiological performance. We predicted a cascade of first and second order EF-associated effects on marginal populations of the invasive mussel Mytilus galloprovincialis, plus a third order effect of EF of increased epibiont load due to second order shell degradation by endoliths. Mussel populations on rocky shores were surveyed across 850 km of the south-southeast coast of South Africa, from the species' warm-edge range limit to sites in the centre of their distribution, to quantify second order (endolithic shell degradation) and third order (number of barnacle epibionts) EF-associated effects as a function of along-shore distance from the range edge. Inshore temperature data were interpolated from the literature. Using in situ temperature logger data, we calculated the effective shore level for several sites by determining the duration of immersion and emersion. Summer and winter inshore water temperatures were linked to distance from the mussel's warm range edge (our proxy for an EF-associated stress gradient), suggesting that seasonality in temperature contributes to first order effects. The gradient in thermal stress clearly affected densities, but its influence on mussel size, shell degradation, and epibiosis was weaker. Relationships among mussel size, shell degradation, and epibiosis were more robust. Larger, older mussels had more degraded shells and more epibionts, with endolithic damage facilitating epibiosis. EF associated with a gradient in thermal stress directly limits the distribution, abundance, and size structure of mussel populations, with important indirect second and third order effects of parasitic disease and epibiont load, respectively., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 The Author(s). Published by Elsevier B.V. All rights reserved.)

Dietary tracers, such as bulk stable carbon (δ 13 C) and nitrogen (δ 15 N) isotopes, can be used to investigate the trophic interactions of marine predators, which is useful to assess their ecological roles within communities. These tracers have also been used to elucidate population structure and substructure, which is critical for the better identification of management units for these species affected by a range of threats, particularly bycatch in fishing gears. Off eastern South Africa, large populations of Indo-Pacific bottlenose (Tursiops aduncus) and common dolphins (Delphinus delphis) co-occur and are thought to follow the pulses of shoaling sardines (Sardinops sagax) heading north-east in the austral winter. Here we used δ 13 C and δ 15 N to investigate the trophic interactions and define ecological units of these two species along a ≈800 km stretch of the east coast of South Africa, from Algoa Bay to the coast of KwaZulu-Natal. Common and bottlenose dolphin dietary niche overlapped by 39.7% overall in our study area, with the highest overlap occurring off the Wild Coast (40.7% at Hluleka). Both stable isotopes were significantly enriched in bottlenose dolphins sampled in the western part of our study area (i.e., Algoa Bay and Amathole) compared to eastern animals (i.e., from Hluleka, Pondoland, and KZN). In areas where genetic information is not available or is insufficient, food web tracers (such as stable isotopes) can be used to group individuals based on trophic ecology, which can provide ecological units for management of populations. The distinct isotope signatures found here for bottlenose dolphins can, therefore, be used as management units for conservation efforts in the future., Competing Interests: Declaration of competing interest The authors declare the following financial interests/personal relationships which may be considered as potential competing interests: Stephanie Plon reports equipment, drugs, or supplies was provided by Dr. Leszek Karczmarski. Stephanie Plon reports financial support and equipment, drugs, or supplies were provided by Eastern Cape Parks and Tourism Authority. Stephanie Plon reports financial support was provided by Petroleum Oil and Gas Corporation of South Africa Ltd., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

Predation is a critical ecological process that directly and indirectly mediates population stabilities, as well as ecosystem structure and function. The strength of interactions between predators and prey may be mediated by multiple density dependences concerning numbers of predators and prey. In temporary wetland ecosystems in particular, fluctuating water volumes may alter predation rates through differing search space and prey encounter rates. Using a functional response approach, we examined the influence of predator and prey densities on interaction strengths of the temporary pond specialist copepod Lovenula raynerae preying on cladoceran prey, Daphnia pulex , under contrasting water volumes. Further, using a population dynamic modeling approach, we quantified multiple predator effects across differences in prey density and water volume. Predators exhibited type II functional responses under both water volumes, with significant antagonistic multiple predator effects (i.e., antagonisms) exhibited overall. The strengths of antagonistic interactions were, however, enhanced under reduced water volumes and at intermediate prey densities. These findings indicate important biotic and abiotic contexts that mediate predator-prey dynamics, whereby multiple predator effects are contingent on both prey density and search area characteristics. In particular, reduced search areas (i.e., water volumes) under intermediate prey densities could enhance antagonisms by heightening predator-predator interference effects., Competing Interests: There are no conflicts of interest relating to this study., (© 2021 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Rivers are impacted by pollutants from anthropogenic activities such as urbanisation and agricultural practices. Whilst point source pollution has been widely studied and in some cases remediated, non-point pollutant sources remain pervasive, particularly in developing countries that lack economic and human specialist capacity. Monitoring of pollution levels in many regions is additionally challenged by a lack of robust indicators for nitrogen inputs, however, diatom community indices and analysis of variation in microphytobenthos (MBP) stable isotope analysis variations have potential. The present study investigates variations and utilities in benthic diatom indices and MPB δ 15 N along different river sections (n = 31) of an austral river between two seasons (wet and dry), testing for relationships with key environmental variables (physical, water and sediment), in the context of N monitoring. One hundred and eighteen diatom taxa belonging to 36 genera were identified, with physical (water flow), water (nitrate, P and total dissolved solids) and sediment (B, Ca, Cr, Na, N, P, SOM, Pb and Zn) variables correlating to one or more of the 12 diatom indices presented. In particular, Biological Diatom Index, Biological Index of Water Quality, Central Economic Community, Index of Artois-Picardie Diatom (IDAP) and Sládeček's Index were strongly explained by sediment variables, whilst Descy's Pollution Index and Schiefele and Schreiner's Index were explained by water and physical variables. Whilst MPB δ 15 N were within the "no impact" level in the wet and dry seasons at reference (i.e. unpolluted) sites, all sites located in agricultural or urban areas, and downstream of sewage discharges, had a wider range that encompassed increasing organic impacts ("inorganic impacts" to high "organic impacts"). Temperature and turbidity (negative), as well as dissolved oxygen, waterway width and depth (positive), significantly affected MPB δ 15 N, whilst effects of chemistry variables were less apparent. Overall, we found that MPB δ 15 N signatures were significantly correlated with Trophic Diatom Index, the Specific Pollution Sensitivity Index and the Artois-Picardie Diatom Index, suggesting the utility of diatoms and MPB δ 15 N in assessments of aquatic pollution. In turn, MPB δ 15 N values are strong indicators of N pollution across spatial and seasonal gradients. Thus, the results showed the effects of sediment variables on diatoms to be strong, indicating that sediment rather than water characteristics more strongly structure diatom communities. Thus, sediment variables should be sampled when conducting bioassessment studies., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2020 Elsevier B.V. All rights reserved.)

Predation is a critical factor that mediates population stability, community structure, and ecosystem function. Predatory natural enemies can contribute to the regulation of disease vector groups such as mosquitoes, particularly where they naturally co-occur across landscapes. However, we must understand inter-population variation in predatory efficiency if we are to enhance vector control. The present study thus employs a functional response (FR; resource use under different densities) approach to quantify and compare predatory interaction strengths among six populations of a predatory temporary pond specialist copepod, Lovenula raynerae, from the Eastern Cape of South Africa preying on second instar Culex pipiens complex mosquito larvae. All individuals from the sampled populations were predatory and drove significant mortality through per capita predation rates of 0.75-1.10 mosquitoes/h at maximum densities over a 5-h feeding time. Individuals from all copepod populations exhibited Type II FRs with no significant differences in attack rates. On the other hand, there were significant differences in handling times, and therefore also maximum feeding rates (maximum experimental prey density: 32), suggesting possible genetic differences among populations that influenced predation. Owing to a widespread distribution in arid landscapes, we propose that predatory calanoid copepods such as L. raynerae play a key regulatory role at the landscape scale in the control of disease vector mosquito populations. We propose that these ecosystems and their specialist biota should thus be conserved and enhanced (e.g., via selective breeding) owing to the ecosystem services they provide in the context of public health., (© The Author(s) 2020. Published by Oxford University Press on behalf of Entomological Society of America. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Increased sedimentation and siltation associated with anthropogenic environmental change may alter microbial biofilms and the carbohydrates they produce, with potential bottom-up effects in these ecosystems. The present study aimed to examine to what extent carbohydrate (associated with biofilm exopolymer) concentration and benthic algal biomass vary among different sediment types (size-structure categories) using a microcosm experiment conducted over a period of 28 days. Substrate treatment and time had a significant effect on the total chlorophyll-a concentrations, whilst a significant interaction was present in the case of total sediment carbohydrates. Total sediment carbohydrates did not relate significantly to chlorophyll-a concentrations overall, nor for any substrate treatments owing to a non-significant 'chlorophyll-a × substrate' interaction term. The diatom community characteristics across sediment sizes were unique for each treatment in our study, with unique dominant diatom taxa compositions within each sediment size class. The finest sediment particle-size (<63 μm) may be the least stable, most likely due to lower binding. We anticipate that the current study findings will lead to a better understanding of how different sediment types due to sedimentation and siltation will impact on primary productivity and the composition of diatom communities in aquatic systems., Competing Interests: Declaration of competing interest All authors declare that no conflict of interest exists., (Copyright © 2019 Elsevier B.V. All rights reserved.)

Protecting the structural and functional integrity of lotic ecosystems is becoming increasingly important as many ecological systems face escalating pressures from human population growth and environmental impacts. Knowledge on the functional composition of macroinvertebrates in austral temperate streams is generally lacking hindering the design and implementation of water management and restoration goals. Therefore, this study examined the effects of urban land-use activities on the benthic macroinvertebrate functional feeding guild structure among different stream orders in an austral river system (Bloukrans River) in the Eastern Cape Province of South Africa. Water quality and macroinvertebrate community data were collected across two seasons from 18 sites in two different stream order categories (i.e. 1, 2 + 3), following standard methods. We separated macroinvertebrates into functional feeding groups (FFGs), which we then used to assess the effects of riparian condition on FFG organization. Linear mixed effects model (LMM) results demonstrated that total dissolved solids (TDS), canopy cover, phosphate and channel width were the key variables that described the major sources of variation in macroinvertebrate FFGs. Based on FFG proportions, collector-gatherers were the most abundant in the Bloukrans River and represented 71.3% of the macroinvertebrate assemblages. The FFG ratios indicated that all the eighteen sites were strongly heterotrophic (i.e. streams received additional sources of energy from leaf litter and other organic matter), showed below expected linkage with riparian input and stable substrates were limited. The FFG ratios offered some insights into the overall functioning of Bloukrans River system. Our results highlight the importance of including macroinvertebrate functional diversity as a complementary approach to assess the ecological integrity in management and restoration plans of river systems., (Copyright © 2019 Elsevier B.V. All rights reserved.)

The consideration of nutrients in pollution dynamics is important for environmental management and conservation. Developing countries are yet to appreciate the aquatic ecosystem pollution impacts on their economies and as such, information on water pollution dynamics is limited. This study assessed the spatio-temporal dynamics of nutrient loading and retention in stream water and sediments in the Bloukrans River system, Eastern Cape province, South Africa over the course of the wet and dry season. Sediment and water samples were analysed for total phosphorus (TP) and nitrogen (TN) concentrations, and were used in combination with river flow discharge, to determine nutrient loads. The study results highlight that river discharge plays a significant role in temporal differences in sediment and water column nutrient concentrations. The mean sediment nutrient concentration was high for the dry season, with high values being observed for the urban river system. Nutrient loads were high above the sewage treatment works outflow (i.e. urban sites), as such, a decreasing trend was observed with increasing distance from the urban environment. Nutrient loads were generally high for the dry season in comparison to the wet season indicating organic matter retention (i.e. accumulation from burst sewage pipes) most likely due to low flows. While it was evident that the ageing wastewater infrastructure contributed to the observed state of the Bloukrans River, the high natural nutrient retention capacity seemed to mitigate eutrophication of downstream aquatic ecosystems. As such, the nutrient retention capacity and management of the system is central to the entire Bloukrans River catchment management practices. Therefore, the study contributes to our understanding of water and sediment nutrient pollution dynamics in an arid temperate river landscape where vast spatio-temporal differences in base flow characterise the riverscape., (Copyright © 2019 Elsevier B.V. All rights reserved.)

This study explores diatom community dynamics in a highly modified semi-arid temperate region river system characterised by inconsistent river flow. Various water and sediment environmental variables were assessed using a multi-faceted analysis approach to determine the spatio-temporal drivers of benthic diatom communities in the river system. Overall, the diatom community was generally dominated by pollution tolerant species, reflecting the anthropogenic intensity and activities on the river system. Diatom community composition was found to be largely determined by water column chemistry variables particularly nutrient concentrations in comparison to sediment chemistry and physical variables. Strong seasonal diatom species composition was also observed and this was driven by strong seasonal variations in nutrient loads and metal concentrations, a result of the variable water flow across the two seasons. However, the greater temporal variation in communities was observed in the smaller systems with the mainstream river system being more homogenous over time. In addition, diatom community composition and environmental variables were found to be different and more pronounced between streams and mainstream sites, than between canals and streams. The study highlights the complex interaction between water column, sediment and physical variables in determining the diatom species composition in small river systems. It also highlights the importance of river flow inconsistency as an indirect variable that alters primary drivers such as nutrient concentrations in the water column and heavy metal levels in the sediment., (Copyright © 2017 Elsevier B.V. All rights reserved.)

Trophic variation in food web structure occurs among and within ecosystems. The magnitude of variation, however, differs from system to system. In ephemeral pond ecosystems, temporal dynamics are relatively more important than in many systems given that hydroperiod is the ultimate factor determining the presence of an aquatic state. Here, using stable isotopes we tested for changes in trophic chain length and shape over time in these dynamic aquatic ecosystems. We found that lower and intermediate trophic level structure increased over time. We discuss these findings within the context of temporal environmental stability. The dynamic nature of these ephemeral systems seems to be conducive to greater levels of intermediate and lower trophic level diversity, with omnivorous traits likely being advantageous.

Plastic pollution is a major anthropogenic contaminant effecting the marine environment and is often associated with high human population densities and industrial activities. The microplastic (63 to 5000μm) burden of beach sediment and surf-zone water was investigated at selected sites along the entire length of the South African coastline. It was predicted that samples collected in areas of high population density, would contain a higher microplastic burden than those along coasts that demonstrate very low population densities. With the exception of water column microplastics within Richard's Bay Harbour (413.3±77.53particles·m -3 ) and Durban Harbour (1200±133.2particles·m -3 ), there were no significant spatial differences in microplastic loads. This supports the theory that harbours act as a source of microplastics for the surrounding marine environment. Additionally, the absence of any spatial variation highlights the possible long range distribution of microplastic pollutants by large scale ocean currents., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

The extent of microplastic pollution (<5mm) in the southern hemisphere, particularly southern Africa, is largely unknown. This study aimed to evaluate microplastic pollution along the south-eastern coastline of South Africa, looking at whether bays are characterised by higher microplastic densities than open stretches of coastline in both beach sediment and surf-zone water. Microplastic (mean ± standard error) densities in the beach sediment ranged between 688.9 ± 348.2 and 3308 ± 1449 particles · m(-2), while those in the water column varied between 257.9 ± 53.36 and 1215 ± 276.7 particles · m(-3). With few exceptions there were no significant spatial patterns in either the sediment or water column microplastic densities; with little differences in density between bays and the open coast (P>0.05). These data indicate that the presence of microplastics were not associated with proximity to land-based sources or population density, but rather is governed by water circulation., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

As part of a larger investigation, the effect of apex predation on estuarine bacterial community structure, through trophic cascading, was investigated using experimental in situ mesocosms. Through either the removal (filtration) or addition of specific size classes of planktonic groups, four different trophic scenarios were established using estuarine water and its associated plankton. One such treatment represented a "natural" scenario in which stable apex predatory pressure was qualified. Water samples were collected over time from each of the treatments for bacterial community evaluation. These samples were assessed through pyrosequencing of the variable regions 4 and 5 of the bacterial 16S rRNA gene and analysed at the species operational taxonomic unit (OTU) level using a community procedure. The blue-green group dominated the samples, followed by Proteobacteria and Bacteroidetes. Samples were the most similar among treatments at the commencement of the experiment. While the bacterial communities sampled within each treatment changed over time, the deviation from initial appeared to be linked to the treatment trophic scenarios. The least temporal deviation-from-initial in bacterial community was found within the stable apex predatory pressure treatment. These findings are consistent with trophic cascade theory, whereby predators mediate interactions at multiple lower trophic levels with consequent repercussions for diversity.

Bull sharks (Carcharhinus leucas) are globally distributed top predators that play an important ecological role within coastal marine communities. However, little is known about the spatial and temporal scales of their habitat use and associated ecological role. In this study, we employed passive acoustic telemetry to investigate the residency patterns and migration dynamics of 18 adult bull sharks (195-283 cm total length) tagged in southern Mozambique for a period of between 10 and 22 months. The majority of sharks (n = 16) exhibited temporally and spatially variable residency patterns interspersed with migration events. Ten individuals undertook coastal migrations that ranged between 433 and 709 km (mean  = 533 km) with eight of these sharks returning to the study site. During migration, individuals exhibited rates of movement between 2 and 59 km x d(-1) (mean = 17.58 km x d(-1)) and were recorded travelling annual distances of between 450 and 3760 km (mean = 1163 km). Migration towards lower latitudes primarily took place in austral spring and winter and there was a significant negative correlation between residency and mean monthly sea temperature at the study site. This suggested that seasonal change is the primary driver behind migration events but further investigation is required to assess how foraging and reproductive activity may influence residency patterns and migration. Results from this study highlight the need for further understanding of bull shark migration dynamics and suggest that effective conservation strategies for this vulnerable species necessitate the incorporation of congruent trans-boundary policies over large spatial scales.

As apex predators, sharks play an important role shaping their respective marine communities through predation and associated risk effects. Understanding the predatory dynamics of sharks within communities is, therefore, necessary to establish effective ecologically based conservation strategies. We employed non-lethal sampling methods to investigate the feeding ecology of bull sharks (Carcharhinus leucas) using stable isotope analysis within a subtropical marine community in the southwest Indian Ocean. The main objectives of this study were to investigate and compare the predatory role that sub-adult and adult bull sharks play within a top predatory teleost fish community. Bull sharks had significantly broader niche widths compared to top predatory teleost assemblages with a wide and relatively enriched range of δ(13)C values relative to the local marine community. This suggests that bull sharks forage from a more diverse range of δ(13)C sources over a wider geographical range than the predatory teleost community. Adult bull sharks appeared to exhibit a shift towards consistently higher trophic level prey from an expanded foraging range compared to sub-adults, possibly due to increased mobility linked with size. Although predatory teleost fish are also capable of substantial migrations, bull sharks may have the ability to exploit a more diverse range of habitats and appeared to prey on a wider diversity of larger prey. This suggests that bull sharks play an important predatory role within their respective marine communities and adult sharks in particular may shape and link ecological processes of a variety of marine communities over a broad range.

Background: Classically, estuarine planktonic research has focussed largely on the physico-chemical drivers of community assemblages leaving a paucity of information on important biological interactions., Methodology/principal Findings: Within the context of trophic cascades, various treatments using in situ mesocosms were established in a closed estuary to highlight the importance of predation in stabilizing estuarine plankton abundances. Through either the removal (filtration) or addition of certain planktonic groups, five different trophic systems were established. These treatments contained varied numbers of trophic levels and thus different "predators" at the top of the food chain. The abundances of zooplankton (copepod and polychaete), ciliate, micro-flagellate, nano-flagellate and bacteria were investigated in each treatment, over time. The reference treatment containing apex zooplanktivores (early juvenile mullet) and plankton at natural densities mimicked a natural, stable state of an estuary. Proportional variability (PV) and coefficient of variation (CV) of temporal abundances were calculated for each taxon and showed that apex predators in this experimental ecosystem, when compared to the other systems, induced stability. The presence of these predators therefore had consequences for multiple trophic levels, consistent with trophic cascade theory., Conclusions/significance: PV and CV proved useful indices for comparing stability. Apex predators exerted a stabilizing pressure through feeding on copepods and polychaetes which cascaded through the ciliates, micro-flagellates, nano-flagellates and bacteria. When compared with treatments without apex predators, the role of predation in structuring planktonic communities in closed estuaries was highlighted.

Mutualistic relations between plants and animals are well documented on land but have received less attention in marine systems. This study examined the relationship between the territorial intertidal limpet Patella longicosta and the crustose brown alga Ralfsia verrucosa. Adult Patella are found exclusively in association with Ralfsia, on which they feed, while Ralfsia occurs primarily, but not exclusively, in Patella territories. Ralfsia benefits directly from both the presence and the territorial behaviour of Patella. Algal productivity was assessed by measuring oxygen evolution and utilization in situ and deriving photosynthesis/irradiance curves. Productivity was increased by about 30% by the presence of Patella in both summer (P max of grazed algae 0.0098; ungrazed algae 0.0063 mg C · cm -2 · h -1 ) and winter (P max grazed algae 0.0081; ungrazed algae 0.0053 mg -2 · C · h -1 ). Algal growth rates were not significantly increased by the application of limpet mucus in the laboratory. We did not examine nutrient regeneration by the limpet, but the increase in photosynthetic rate may depend on the limpet's grazing pattern which creates secondary sites for growth. Ralfsia also benefited from the territorial behaviour of Patella. The effects of different grazing regimes were investigated in different seasons by removing territorial limpets and either excluding all limpets using copper-based antifouling paint, or allowing access to non-territorial limpets (mostly P. oculus) using partial paint barriers. Exclusion of all limpets resulted in rapid overgrowth of Ralfsia plants by the foliose green alga Ulva sp.. Where non-territorial limpets had access to the plants overgrowth was reduced but Ralfsia plants were entirely removed by destructive grazing. Non-territorial grazers removed 90% of Ralfsia plants within 4 weeks in summer and 60% in winter. In control treatments P. longicosta prevented overgrowth by Ulva and actively excluded vagrant grazers, preventing overgrazing. Based on these findings, the association between the limpet and alga can be regarded as a nonobligate mutualism.