An important principle of environmental science is that changes in single components of systems are likely to have consequences elsewhere in the same systems. In the sea, food web data are one of the few foundations for predicting such indirect effects, whether of fishery exploitation or following recovery in marine protected areas (MPAs). We review the available literature on one type of indirect interaction in benthic marine ecosystems, namely trophic cascades, which involve three or more trophic levels connected by predation. Because many indirect effects have been revealed through fishery exploitation, in some cases we include humans as trophic levels. Our purpose is to establish how widespread cascades might be, and infer how likely they are to affect the properties of communities following the implementation of MPAs or intensive resource exploitation. We review 39 documented cascades (eight of which include humans as a trophic level) from 21 locations around the world; all but two of the cascades are from shallow systems underlain by hard substrata (kelp forests, rocky subtidal, coral reefs and rocky intertidal). We argue that these systems are well represented because they are accessible and also amenable to the type of work that is necessary. Nineteen examples come from the central-eastern and north-eastern Pacific, while no well-substantiated benthic cascades have been reported from the NE, CE or SW Atlantic, the Southern Oceans, E Indian Ocean or NW Pacific. The absence of examples from those zones is probably due to lack of study. Sea urchins are very prominent in the subtidal examples, and gastropods, especially limpets, in the intertidal examples; we suggest that this may reflect their predation by fewer specialist predators than is the case with fishes, but also their conspicuousness to investigators. The variation in ecological resolution amongst studies, and in intensity of study amongst systems and regions, indicates that more cascades will likely be identified in due course. Broadening the concept of cascades to include pathogenic interactions would immediately increase the number of examples. The existing evidence is that cascade effects are to be expected when hard-substratum systems are subject to artisanal resource exploitation, but that the particular problems of macroalgal overgrowth on Caribbean reefs and the expansion of coralline barrens in the Mediterranean rocky-sublittoral will not be readily reversed in MPAs, probably because factors other than predation-based cascades have contributed to them in the first place. More cascade effects are likely to be found in the soft-substratum systems that are crucial to so many large-scale fisheries, when opportunities such as those of MPAs and fishing gradients become available for study of such systems, and the search is widened to less conspicuous focal organisms such as polychaetes and crustaceans.

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Marine bioconstructions are biodiversity-rich, three-dimensional biogenic structures, regulating key ecological functions of benthic ecosystems worldwide. Tropical coral reefs are outstanding for their beauty, diversity and complexity, but analogous types of bioconstructions are also present in temperate seas. The main bioconstructions in the Mediterranean Sea are represented by coralligenous formations, vermetid reefs, deep-sea cold-water corals, Lithophyllum byssoides trottoirs, coral banks formed by the shallow-water corals Cladocora caespitosa or Astroides calycularis, and sabellariid or serpulid worm reefs. Bioconstructions change the morphological and chemicophysical features of primary substrates and create new habitats for a large variety of organisms, playing pivotal roles in ecosystem functioning. In spite of their importance, Mediterranean bioconstructions have not received the same attention that tropical coral reefs have, and the knowledge of their biology, ecology and distribution is still fragmentary. All existing data about the spatial distribution of Italian bioconstructions have been collected, together with information about their growth patterns, dynamics and connectivity. The degradation of these habitats as a consequence of anthropogenic pressures (pollution, organic enrichment, fishery, coastal development, direct physical disturbance), climate change and the spread of invasive species was also investigated. The study of bioconstructions requires a holistic approach leading to a better understanding of their ecology and the application of more insightful management and conservation measures at basin scale, within ecologically coherent units based on connectivity: the cells of ecosystem functioning.

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.

Vermetids form reefs in sub-tropical and warm-temperate waters that protect coasts from erosion, regulate sediment transport and accumulation, serve as carbon sinks and provide habitat for other species. The gastropods that form these reefs brood encapsulated larvae; they are threatened by rapid environmental changes since their ability to disperse is very limited. We used transplant experiments along a natural CO2 gradient to assess ocean acidification effects on the reef-building gastropod Dendropoma petraeum. We found that although D. petraeum were able to reproduce and brood at elevated levels of CO2, recruitment success was adversely affected. Long-term exposure to acidified conditions predicted for the year 2100 and beyond caused shell dissolution and a significant increase in shell Mg content. Unless CO2 emissions are reduced and conservation measures taken, our results suggest these reefs are in danger of extinction within this century, with significant ecological and socioeconomic ramifications for coastal systems.

The aesthetic appeal of marine reserves and the facilities provided, together with the increased public awareness of nature, all contribute to creating massive tourism in MPAs. Human activities are being changed inside MPAs in two ways: humans as top predators are generally being removed, but in turn they could come back at great numbers as visitors. Many authors have studied the impact of visitors, and the results highlight that the consequences can be very substantial and may represent a severe threat to the overall diversity of marine communities. To date, the documented effects of human recreational activities on natural communities are restricted to assessing the consequences of trampling over intertidal and upper infralittoral areas, boat anchoring in seagrass meadows and tropical reefs, or SCUBA-diving. In this paper we review the available literature world-wide on the effects of human recreational activities in marine communities. The objective is to address the extent of these impacts and to highlight the gaps of knowledge to be filled in order to optimise decision making on research, monitoring, and management of Mediterranean MPAs. A specific plan for managing tourism use in each Mediterranean MPA should be designated. These strategies should be implemented through education, training, and changes in legislation and policy.

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