Marine protected areas promote stability of reef fish communities under climate warming.

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.

Coral reefs at Sir Bu Nair Island: An offshore refuge of Acropora in the southern Arabian Gulf.

Coral reefs across the southern Arabian Gulf have declined in the past two decades, with extensive loss of formerly Acropora table corals, which are now functionally extinct in nearshore reefs. This study documents the coral community at Sir Bu Nair (SBN), an offshore island buffered by less extreme environmental conditions, which contains the last remaining large stands of Acropora in the southern Gulf. We found that Acropora is a major reef-building coral throughout SBN. Mean coral cover was 27% (range: 6%-49%) across all sites and depths, of which more than half was comprised by Acropora. This varied around the island, with the highest densities to the south and southwest in shallow waters. Our study provides essential information for the management and conservation of these highly valuable and vulnerable corals.

Quotas regulation is necessary but not sufficient to mitigate the impact of SCUBA diving in a highly visited marine protected area.

When effectively managed, Marine Protected Areas (MPAs) can produce wide ecosystem benefits that can foster, directly and indirectly, local economies. Tourism is one of the sectors mainly benefited by the effect of conservation. SCUBA diving represents an important tourism activity, especially in the context of MPAs, where it is one of the few activities often fostered rather than limited, for its capacity to integrate environmental and socio-economic sustainability. However, SCUBA diving can also produce negative impacts on the environment when tourism frequentation exceeds a sustainable threshold, these potentially generating negative effects on the sector itself. In this study, we (1) investigated the impact of SCUBA diving in one of the most frequented diving areas of the Mediterranean Sea (Cabo de Palos - Islas Hormigas marine reserve), and (2) assessed the potential benefits over time related to the adoption of a regulation change for the diving activity (i.e., formally adoption of diving quotas). Specifically, we compared demographic (density of alive and dead colonies) and morphometric (height, width and complexity) characteristics of the false coral (Myriapora truncata) between dived and fully protected (non-dived) locations over four diving seasons (one before and three after the change in diving quotas). The density of alive colonies of the false coral was, on average, six times lower in dived locations compared to controls, highlighting a clear impact of SCUBA diving (consistent over time). Colonies were also significantly smaller in dived locations. The diving quotas produced a significant reduction of the ratio dead/total colonies in the dived locations soon after their adoption, but these benefits disappeared over the following years, possibly due to a gradual decline in operators' and divers' observance and concern, rather than an increasing number of dives. This suggests that the adoption of effective regulations is crucial for the environmental sustainability of diving tourism in protected areas and can provide positive effects, but an effort is needed to ensure that compliance is consistent over time, and that low-impact diving practices are adopted by this important recreational sector.

Exceptionally high but still growing predatory reef fish biomass after 23 years of protection in a Marine Protected Area.

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.

Use of Lagrangian simulations to hindcast the geographical position of propagule release zones in a Mediterranean coastal fish.

The study of organism dispersal is fundamental for elucidating patterns of connectivity between populations, thus crucial for the design of effective protection and management strategies. This is especially challenging in the case of coastal fish, for which information on egg release zones (i.e. spawning grounds) is often lacking. Here we assessed the putative location of egg release zones of the saddled sea bream (Oblada melanura) along the south-eastern coast of Spain in 2013. To this aim, we hindcasted propagule (egg and larva) dispersal using Lagrangian simulations, fed with species-specific information on early life history traits (ELTs), with two approaches: 1) back-tracking and 2) comparing settler distribution obtained from simulations to the analogous distribution resulting from otolith chemical analysis. Simulations were also used to assess which factors contributed the most to dispersal distances. Back-tracking simulations indicated that both the northern sector of the Murcia region and some traits of the North-African coast were hydrodynamically suitable to generate and drive the supply of larvae recorded along the coast of Murcia in 2013. With the second approach, based on the correlation between simulation outputs and field results (otolith chemical analysis), we found that the oceanographic characteristics of the study area could have determined the pattern of settler distribution recorded with otolith analysis in 2013 and inferred the geographical position of main O. melanura spawning grounds along the coast. Dispersal distance was found to be significantly affected by the geographical position of propagule release zones. The combination of methods used was the first attempt to assess the geographical position of propagule release zones in the Mediterranean Sea for O. melanura, and can represent a valuable approach for elucidating dispersal and connectivity patterns in other coastal species.

Response of rocky reef top predators (Serranidae: Epinephelinae) in and around marine protected areas in the Western Mediterranean Sea.

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.

Influence of habitat structure on fish assemblage of an artificial reef in southern Brazil.

Habitat complexity strongly influences reef fish community composition. An understanding of the underlying reasons for this relationship is important for evaluating the suitability of artificial reef (AR) habitats as a marine resource management tool. We studied the influence of AR habitat structure on fish assemblage composition off the southern coast of Brazil. We found that reef blocks with greater area and number of holes possessed the greatest fish species richness and abundance. Reef blocks with greater complexity had higher abundance of almost 30% of fish species present. Natural reef (NR) and AR were different in their fish species composition, trophic structure and categories of water column occupancy by fish (spatial categories). Although NR was more diverse and harboured more trophic levels, AR presented the higher abundances and the presence of distinct fish species that underlined their importance at a regional scale. The greater availability of sheltering habitat where hard substrate is scarce, together with their frequent use by economically important species, make AR a useful tool for coastal management when certain ecological conditions are met.

Combined effects of levels of protection and environmental variables at different spatial resolutions on fish assemblages in a marine protected area.

The links between species-environment relations and species' responses to protection are unclear, but the objectives of marine protected areas (MPAs) are most likely to be achieved when those relations are known and inform MPA design. The components of a species' habitat vary with the spatial resolution of the area considered. We characterized areas at two resolutions: 250 m(2) (transect) and approximately 30,000 m(2) (seascape). We considered three categories of environmental variables: substrate type, bottom complexity, and depth. We sought to determine at which resolution habitat characteristics were a better predictor of abundance and species composition of fishes and whether the relations with environmental variables at either resolution affected species' responses to protection. Habitat features accounted for a larger proportion of spatial variation in species composition and abundances than differences in protection status. This spatial variation was explained best by habitat characteristics at the seascape level than at the transect level. Species' responses to protected areas were specific to particular seascape characteristics, primarily depth, and bottom complexity. Our method may be useful for prioritizing marine areas for protection, designing MPAs, and monitoring their effectiveness. It identified areas that provided natural shelter, areas acting as buffer zones, and areas where fish species were most responsive to protection. The identification of such areas is necessary for cost-effective establishment and monitoring of MPAs.

Marine reserves: size and age do matter.

Marine reserves are widely used throughout the world to prevent overfishing and conserve biodiversity, but uncertainties remain about their optimal design. The effects of marine reserves are heterogeneous. Despite theoretical findings, empirical studies have previously found no effect of size on the effectiveness of marine reserves in protecting commercial fish stocks. Using 58 datasets from 19 European marine reserves, we show that reserve size and age do matter: Increasing the size of the no-take zone increases the density of commercial fishes within the reserve compared with outside; whereas the size of the buffer zone has the opposite effect. Moreover, positive effects of marine reserve on commercial fish species and species richness are linked to the time elapsed since the establishment of the protection scheme. The reserve size-dependency of the response to protection has strong implications for the spatial management of coastal areas because marine reserves are used for spatial zoning.