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Marine protected areas (MPAs) are globally important environmental management tools that provide protection from the effects of human exploitation and activities, supporting the conservation of marine biological diversity, habitats, ecosystems and the processes they host, as well as resources in a broad sense. Consequently, they are also expected to manage and enhance marine ecosystem services and material, non-material, consumptive and non-consumptive goods, and benefits for humans. There is however certain confusion on what constitutes an ecosystem service, and it is not always easy to distinguish between them and societal benefits. The main nuance is that an ecosystem service is the aptitude an ecosystem has or develops naturally or as consequence of a management action, and that manifests through its own properties (productivity, diversity, stability, quality of its key parameters, etc.), while a societal benefit is the economic or other profitability (emotional, educational, scientific, etc.) that humans obtain from said service or quality. In this work, 268 publications, together with our own experiences in the different investigations carried out in the MPAs that are part of the BiodivERsA3-2015-21 RESERVEBENEFIT European project, have been selected, reviewed and discussed to analyze the knowledge status of the expected ecosystem services of MPAs and the societal benefits derived from them, sometimes providing information on their evidence, when they exist. We define and classify the effects of protection, ecosystem services and societal benefits and elaborate a conceptual model of the cause-effect relationships between them.

Marine Protected Areas (MPAs) help replenish fish assemblages, though different trophic levels may show diverse recovery patterns. Long-term protection is required to achieve total recovery but poaching events may prevent the achievement of full carrying capacity. Here, we have analysed the effect of long-term protection on the entire reef fish community and the different trophic levels in the Cabo de Palos-Islas Hormigas MPA (SE Spain; SW Mediterranean Sea) in order to assess their recovery patterns after 23 years of protection. We compared the values for carrying capacity obtained with the maximum values achieved at regional scale, and we assessed the effect of a reduction in the surveillance over a few years, during which poaching events increased, on the recovery patterns. We found that, overall, biomass of fishes increased with time while density diminished. In particular, piscivorous and macro-invertivore fish increased while the other trophic groups remained constant or declined, suggesting top-down processes. For the entire study period, those trophic groups were approaching carrying capacity; however, when accounting only for the period in which enforcement was high and constant, they grew exponentially, indicating that full carrying capacity may have not been achieved yet. When compared to other Mediterranean MPAs, the Cabo de Palos-Islas Hormigas MPA showed values for biomass that were disproportionately higher, suggesting that local factors, such as habitat structure and associated oceanographic processes, may be responsible for the dynamics found. Our results help to understand the potential trajectories of fish assemblages over a consolidated MPA and highlight empirically how the reduction of surveillance in a period may change the recovery patterns.

Groupers species are extremely vulnerable to overfishing and many species are threatened worldwide. In recent decades, Mediterranean groupers experienced dramatic population declines. Marine protected areas (MPAs) can protect populations inside their boundaries and provide individuals to adjacent fishing areas through the process of spillover and larval export. This study aims to evaluate the effectiveness of six marine reserves in the Western Mediterranean Sea to protect the populations of three species of grouper, Epinephelus marginatus, Epinephelus costae and Mycteroperca rubra, and to understand in which circumstances MPAs are able to export biomass to neighbouring areas. All the studied MPAs, except one where no grouper was observed, were able to maintain high abundance, biomass and mean weight of groupers. Size classes were more evenly distributed inside than outside MPAs. In two reserves, biomass gradients could be detected through the boundaries of the reserve as an indication of spillover. In some cases, habitat structure appeared to exert a great influence on grouper abundance, biomass and mean individual weight, influencing the gradient shape. Because groupers are generally sedentary animals with a small home range, we suggest that biomass gradients could only occur where groupers attain sufficient abundance inside MPA limits, indicating a strongly density-dependent process.

Abstract Protection from direct human impacts can safeguard marine life, yet ocean warming crosses marine protected area boundaries. Here, we test whether protection offers resilience to marine heatwaves from local to network scales. We examine 71,269 timeseries of population abundances for 2269 reef fish species surveyed in 357 protected versus 747 open sites worldwide. We quantify the stability of reef fish abundance from populations to metacommunities, considering responses of species and functional diversity including thermal affinity of different trophic groups. Overall, protection mitigates adverse effects of marine heatwaves on fish abundance, community stability, asynchronous fluctuations and functional richness. We find that local stability is positively related to distance from centers of high human density only in protected areas. We provide evidence that networks of protected areas have persistent reef fish communities in warming oceans by maintaining large populations and promoting stability at different levels of biological organization.