Abstract. Marine reserves are assumed to protect a wide range of species from deleterious effects stemming from exploitation. However, some species, due to their ecological characteristics, may not respond positively to protection. Very little is known about the effects of life history and ecological traits (e.g., mobility, growth, and habitat) on responses of fish species to marine reserves. Using 40 data sets from 12 European marine reserves, we show that there is significant variation in the response of different species of fish to protection and that this heterogeneity can be explained, in part, by differences in their traits. Densities of targeted sizeclasses of commercial species were greater in protected than unprotected areas. This effect of protection increased as the maximum body size of the targeted species increased, and it was greater for species that were not obligate schoolers. However, contrary to previous theoretical findings, even mobile species with wide home ranges benefited from protection: the effect of protection was at least as strong for mobile species as it was for sedentary ones. Noncommercial bycatch and unexploited species rarely responded to protection, and when they did (in the case of unexploited bentho-pelagic species), they exhibited the opposite response: their densities were lower inside reserves. The use of marine reserves for marine conservation and fisheries management implies that they should ensure protection for a wide range of species with different life-history and ecological traits. Our results suggest this is not the case, and instead that effects vary with economic value, body size, habitat, depth range, and schooling behavior.

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SummaryThe success of MPAs in conserving fishing resources and protecting marine biodiversity relies strongly on how well they meet their planned (or implicit) management goals. From a review of empirical studies aiming at assessing the ecological effects of Mediterranean and Macaronesian MPAs, we conclude that establishing an MPA is successful for (i) increasing the abundance/biomass, (ii) increasing the proportion of larger/older individuals, and (iii) enhancing the fecundity of commercially harvested populations; also, MPAs demonstrated to be Meta-analysis; Mediterranean; Macaronesia effective for (iv) augmenting local fishery yields through biomass exportation from the protected area, and (v) inducing shifts in fish assemblage structure by increasing the dominance of large predator species. However, the attraction for tourism and diving due to ecological benefits of protection can cause damages likely to reverse some of the MPA effects. Other expected effects are more subject to uncertainty, and hence need more research, such as (vi) causing density-dependent changes in life history traits and (vii) protecting the recruitment of commercially important species, (viii) protecting marine biodiversity (including genetic diversity), (ix) causing ecosystem-wide effects such as trophic cascades, and (x) increasing community and ecosystem stability, thus promoting resilience and faster recovery from disturbance. Meta-analysis of data arising from these case studies are used to establish the overall effect of MPAs, and its relationship to MPA features, such as size of no-take area or time since protection. Based on the review and the metaanalyses, specific recommendations are provided for MPA management, regarding the establishment of goals and objectives, site selection, MPA design and zoning, planning, and monitoring. Finally, a series of recommendations for MPA research are offered to drive future research in MPA issues in the Mediterranean and Macaronesia.

Marine protected areas (MPAs) are often promoted as tools for biodiversity conservation as well as for fisheries management. Despite increasing evidence of their usefulness, questions remain regarding the optimal design of MPAs, in particular concerning their function as fisheries management tools, for which empirical studies are still lacking. Using 28 data sets from seven MPAs in Southern Europe, we developed a meta-analytical approach to investigate the effects of protection on adjacent fisheries and asking how these effects are influenced by MPA size and age. Southern European MPAs showed clear effects on the surrounding fisheries, on the 'catch per unit effort' (CPUE) of target species, but especially on the CPUE of the marketable catch. These effects depended on the time of protection and on the size of the no-take area. CPUE of both target species and the marketable catch increased gradually by 2-4% per year over a long time period (at least 30 years). The influence of the size of the no-take area appeared to be more complex. The catch rates of the entire fishery in and around the MPA were higher when the no-take areas were smaller. Conversely, catch rates of selected fisheries that were expected to benefit most from protection increased when the no-take area was larger. Our results emphasize the importance of MPA size on its export functions and suggest that an adequate, often extended, time frame be used for the management and the evaluation of effectiveness of MPAs.

Although site-attached fish can be expected to increase in abundance in marine protected areas (MPAs), there is little known about recruitment effects. The present work reviews concepts and field evidence for enhancement of recruitment of species in MPAs, focusing geographically on the Mediterranean littoral because of its long history of intensive fishery exploitation, but drawing on evidence from studies on recruitment processes in general on MPAs both in the Mediterranean and elsewhere. We considered recruitment as the process of a fish being added to the local population. The general questions of interest are whether the increase in biomass of species protected in MPAs has an effect on recruitment in the MPAs or in neighbouring areas, and, on competition and predation effects on new recruits. A flow diagram of the effects of MPA status on recruitment is developed and employed to identify the relevant processes. The diagram incorporates three levels of factors: (1) characteristics of MPAs (location, size, habitat type, oceanography and level of protection); (2) life stages of species protected in MPAs relevant to recruitment (eggs, larvae, settlers and juveniles); and (3) fundamental processes of dispersal/movement, predation and competition. From this conceptual diagram, the following main components of the recruitment process were identified and used to structure the review: (1) relationship between the ecology of pelagic stages and the design, location and oceanographic regime of MPAs; (2) effects of protection in MPAs from fishery exploitation of nursery habitats on settlement success; and (3) effects of protection on survival of settlers and juveniles from competition and predation. We found an exceptionally low number of studies specifically addressing recruitment processes in MPAs. This was particularly the case in what concerns the relationship between larval ecology and the characteristics and oceanographic regime of MPAs. The effectiveness of MPAs in promoting recruitment mainly depends on the locations and on sizes of the MPAs in relation to the reproductive biology and larval ecology of the species concerned. The locations and sizes of MPAs in turn depend on MPA objectives, whether the purpose is to protect entire life cycles, the juveniles, or to increase egg production and larval export. The assessment of the relationship between the protection of nursery habitats and settlement success indicates that the magnitude of the effects of protection depends on whether the recruitment of the species involved is restricted to a narrowly-defined set of environmental conditions or, on the contrary, can occur in diverse environments, including areas beyond the influence of the MPAs. Thus, the locations of MPAs determine the habitats which are protected and, consequently, the species, the settlement of which will be favoured. For Mediterranean shallow-water species, the near-shore zone encompasses most of the essential nursery habitats for protection. Recruitment studies conducted in MPAs in the north-western Mediterranean have showed no differences in survival of newly-settled littoral fish between MPAs and the areas outside of them. Conversely, for older recruits, mortality was found to be higher inside MPAs, probably due to the increased abundance and size of large predators. This study highlights the almost total absence of studies addressing even the most elementary questions of recruitment in the specific context of MPAs.

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.

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a b s t r a c tThe brown meagre Sciaena umbra, an iconic demersal fish species in Mediterranean coastal habitats, is particularly vulnerable to fishing pressure and presents a worrying population decline. Fish numbers and sizes were surveyed by visual census at Scandola (Corsica) in and outside reserve zones subject to increasing levels of protection, including unprotected zones (UP) where all fishing activities are permitted, buffer zones (BZ) subject to partial protection and a totally protected no-take integral reserve zone (IR). The numerical abundance, individual size and biomass of the brown meagre were found to increase with levels of reserve protection. The abundance of the larger size classes and the numbers of fish per shoal were significantly lower in unprotected zones. A comparison with similar censuses performed in 1983 showed a significant increase of S. umbra abundance in IR, but no difference in UP. That increasing levels of protection resulted in increased abundance and biomass of the brown meagre suggested a prominent role of fishing, particularly spearfishing, activities in the persistence of its low abundance in the unprotected zones. As a consequence, protective action for the brown meagre (including a ban on both spearfishing and recreational hook-and-line fishing) has been introduced in France since January 2014.