Spearfishing Regulation Benefits Artisanal Fisheries: The ReGS Indicator and Its Application to a Multiple-Use Mediterranean Marine Protected Area

The development of fishing efficiency coupled with an increase of fishing effort led to the overexploitation of numerous natural marine resources. In addition to this commercial pressure, the impact of recreational activities on fish assemblages remains barely known. Here we examined the impact of spearfishing limitation on resources in a marine protected area (MPA) and the benefit it provides for the local artisanal fishery through the use of a novel indicator. We analysed trends in the fish assemblage composition using artisanal fisheries data collected in the Bonifacio Strait Natural Reserve (BSNR), a Mediterranean MPA where the spearfishing activity has been forbidden over 15% of its area. Fish species were pooled into three response groups according to their target level by spearfishing. We developed the new flexible ReGS indicator reflecting shifts in species assemblages according to the relative abundance of each response group facing external pressure. The catch per unit effort (CPUE) increased by ca. 60% in the BSNR between 2000 and 2007, while the MPA was established in 1999. The gain of CPUE strongly depended on the considered response group: for the highly targeted group, the CPUE doubled while the CPUE of the untargeted group increased by only 15.5%. The ReGS value significantly increased from 0.31 to 0.45 (on a scale between 0 and 1) in the general perimeter of this MPA while it has reached a threshold of 0.43, considered as a reference point, in the area protected from spearfishing since 1982. Our results demonstrated that limiting recreational fishing by appropriate zoning in multiple-use MPAs represents a real benefit for artisanal fisheries. More generally we showed how our new indicator may reveal a wide range of impacts on coastal ecosystems such as global change or habitat degradation.

Related papers:

title: Current issues with fish and fisheries: editor's overview and introduction
abstract: 1. By any measure, fishes are among the world's most important natural resources. Annual exploitation from wild populations exceeds 90 million tonnes, and fish supply over 15% of global protein needs as part of total annual trade exceeding $US 55 billion. Additionally, with over 25 000 known species, the biodiversity and ecological roles of fishes are being increasingly recognised in aquatic conservation, ecosystem management, restoration and aquatic environmental regulation. 
2. At the same time, substantial management problems now affect the production, exploitable stocks, global diversity, trophic structure, habitat quality and local composition of fish communities. 
3. In marine systems, key issues include the direct effects of exploitation on fish, habitats and other organisms, while habitat or water quality problems arise also from the atmospheric, terrestrial and coastal environments to which marine systems are linked. In freshwaters, flow regulation and obstruction by dams, fragmentation, catchment management, pollution, habitat alterations, exotic fish introductions and nursery-reared fish are widespread issues. 
4. Management responses to the problems of fish and fisheries include aquatic reserves in both marine and freshwater habitats, and their effectiveness is now being evaluated. Policies on marine exploitation increasingly emphasise fishes as integral components of aquatic ecosystems rather than individually exploitable stocks, but the rationalisation of fishing pressures presents many challenges. In Europe, North America and elsewhere, policies on freshwaters encourage habitat protection, integrated watershed management and restoration, but pressures on water resources will cause continued change. All these management approaches require development and evaluation, and will benefit from a perspective of ecological understanding with ecologists fully involved. 
5. Synthesis and applications. Although making a small contribution to the Journal of Applied Ecology in the past, leading work on aquatic problems and fish-related themes appear increasingly in this and other mainstream ecology journals. As this special profile of five papers shows, significant contributions arise on diverse issues that here include the benefit of aquatic reserves, river restoration for fish, the accumulation of contaminants, interactions with predators, and the fitness of salmonids from nurseries. This overview outlines the current context in which papers on the applied ecology of fish and fisheries are emerging, and it identifies scope for further contributions.

title: Limiting abuse: marine protected areas, a limited solution
abstract: Designation of marine protected areas (MPAs) is increasing as humans seek to combat overexploitation of marine resources and preserve the integrity of the ocean’s unique biodiversity. At present there are over 1300 MPAs. The primary legal responsibility for the designation of MPAs falls to individual countries, but protection of the marine environment at large scales is also critical because ocean circulation does not honor legal boundaries and often exceeds the influence of any one nation or group of nations. There are many reasons for establishing MPAs; the papers we surveyed principally referred to scientific, economic, cultural, and ethical factors. Two approaches predominated: fisheries management and habitat protection. Although the major threat to terrestrial systems is habitat loss, the major threats to the world’s oceans are fisheries overexploitation, coastal development, and chemical and biological pollution. MPAs may provide conservation of formerly exploited species as well as benefits to the fishery through leakage of ‘surplus’ adults (spillover) and larvae (larval replenishment) across reserve boundaries. Higher order effects, such as changes in species richness or changes in community structure and function, have only been superficially explored. Because many MPAs are along coastlines, within shipping lanes, and near human centers of activity, the chance of chemical and biological pollution is high. Use of MPAs to combat development and pollution is not appropriate, because MPAs do not have functional boundaries. The ocean is a living matrix carrying organisms as well as particles and therefore even relatively environmentally sensitive uses of coastal ecosystems can degrade ecosystem structure and function via increasing service demands (e.g. nutrient and toxics transformation) and visitation. Whether an MPA is effective is a function of the initial objectives, the level of enforcement, and its design. Single reserves need to be large and networked to accommodate bio-physical patterns of larval dispersal and recruitment. Some authors have suggested that reserve size needs to be extremely large — 50–90% of total habitat — to hedge against the uncertainties of overexploitation. On a local scale, marine protected areas can be effective conservation tools. On a global scale, MPAs can only be effective if they are substantively representative of all biogeographic zones, single reserves are networked within biogeographic zones, and the total amount of area reserved per zone is 20% or greater. The current size and placement of protected areas falls far short of comprehensive or even adequate conservation objectives.

title: Ecosystems Historical Overfishing and the Recent Collapse of Coastal
abstract: clicking here. colleagues, clients, or customers by , you can order high-quality copies for your If you wish to distribute this article to others here. following the guidelines can be obtained by Permission to republish or repurpose articles or portions of articles ): October 15, 2012 www.sciencemag.org (this information is current as of The following resources related to this article are available online at

title: Rapid worldwide depletion of predatory fish communities
abstract: Serious concerns have been raised about the ecological effects of industrialized fishing, spurring a United Nations resolution on restoring fisheries and marine ecosystems to healthy levels. However, a prerequisite for restoration is a general understanding of the composition and abundance of unexploited fish communities, relative to contemporary ones. We constructed trajectories of community biomass and composition of large predatory fishes in four continental shelf and nine oceanic systems, using all available data from the beginning of exploitation. Industrialized fisheries typically reduced community biomass by 80% within 15 years of exploitation. Compensatory increases in fast-growing species were observed, but often reversed within a decade. Using a meta-analytic approach, we estimate that large predatory fish biomass today is only about 10% of pre-industrial levels. We conclude that declines of large predators in coastal regions have extended throughout the global ocean, with potentially serious consequences for ecosystems. Our analysis suggests that management based on recent data alone may be misleading, and provides minimum estimates for unexploited communities, which could serve as the ‘missing baseline’ needed for future restoration efforts.

title: Uncertainty and sustainability in fisheries and the benefit of marine protected areas
abstract: Over-exploitation of fisheries is a serious and immediate global problem that current management policies struggle to solve. Scientists and managers accept that new strategies based on a long-term approach are required for future fisheries management policies. But to succeed, such strategies are likely to require more than just attempts by management to control annual levels of exploitation. Recent studies show that using MPAs (marine protected areas, marine reserves) can improve yield as well as protect stocks and sustain fishery viability. In this paper, we use a deliberately simple model, which describes an exploited fishery as a nonlinear dynamical system close to a point of bifurcation; small stochastic perturbations can therefore build up to cause fishery collapse. Extending the model shows that a MPA can buffer this stochasticity and alleviate the propensity to collapse. The model illustrates two main points: (i) evaluating fisheries management strategies such as MPAs in a deterministic framework is inherently misleading; at worst it leads to fishery extinction, at best it fails to maximise yield; (ii) management strategies that are designed explicitly to buffer uncertainty in the system can provide a sustainable and productive fishery. Compared with harvest control rules based on uncertain estimates of stock size, our simulations indicate that MPAs can substantially reduce the risk of fisheries collapse for only a very small cost to total yield.

title: Confronting the coral reef crisis
abstract: The worldwide decline of coral reefs calls for an urgent reassessment of current management practices. Confronting large-scale crises requires a major scaling-up of management efforts based on an improved understanding of the ecological processes that underlie reef resilience. Managing for improved resilience, incorporating the role of human activity in shaping ecosystems, provides a basis for coping with uncertainty, future changes and ecological surprises. Here we review the ecological roles of critical functional groups (for both corals and reef fishes) that are fundamental to understanding resilience and avoiding phase shifts from coral dominance to less desirable, degraded ecosystems. We identify striking biogeographic differences in the species richness and composition of functional groups, which highlight the vulnerability of Caribbean reef ecosystems. These findings have profound implications for restoration of degraded reefs, management of fisheries, and the focus on marine protected areas and biodiversity hotspots as priorities for conservation.

title: Depletion, Degradation, and Recovery Potential of Estuaries and Coastal Seas
abstract: Estuarine and coastal transformation is as old as civilization yet has dramatically accelerated over the past 150 to 300 years. Reconstructed time lines, causes, and consequences of change in 12 once diverse and productive estuaries and coastal seas worldwide show similar patterns: Human impacts have depleted >90% of formerly important species, destroyed >65% of seagrass and wetland habitat, degraded water quality, and accelerated species invasions. Twentieth-century conservation efforts achieved partial recovery of upper trophic levels but have so far failed to restore former ecosystem structure and function. Our results provide detailed historical baselines and quantitative targets for ecosystem-based management and marine conservation.

title: Impacts of Biodiversity Loss on Ocean Ecosystem Services
abstract: Human-dominated marine ecosystems are experiencing accelerating loss of populations and species, with largely unknown consequences. We analyzed local experiments, long-term regional time series, and global fisheries data to test how biodiversity loss affects marine ecosystem services across temporal and spatial scales. Overall, rates of resource collapse increased and recovery potential, stability, and water quality decreased exponentially with declining diversity. Restoration of biodiversity, in contrast, increased productivity fourfold and decreased variability by 21%, on average. We conclude that marine biodiversity loss is increasingly impairing the ocean's capacity to provide food, maintain water quality, and recover from perturbations. Yet available data suggest that at this point, these trends are still reversible.

title: The recreational fishery off Majorca Island (western Mediterranean): some implications for coastal resource management
abstract: The sociology and habits of recreational anglers on the Island of Majorca (western Mediterranean) were evaluated using telephone and on-site surveys, as well as fishing logbooks and recreational fishing competitions. The recreational fishery is one of the island's main leisure activities, 5.14% of the population (37 265 people) participating. Enthusiasts tend to be mainly middle class (most anglers own boats moored at marinas), middle-aged males (90% male, mean age 46 ± 2 years). The most popular fishing method is from a boat (62.9%), followed by fishing from shore (32.4%), and spearfishing (3.6%). The mean time spent fishing is 3.86 ± 0.03 h d −1 , and more than one type of gear (mean 1.27 ± 0.21) is used simultaneously by a single angler. The frequency of fishing is 4-6 times per month, mainly on holidays and weekends, increasing in summer. The activity has a sizeable impact on the coastal fauna, with diverse catches of at least 1209.25 t year −1 (about 615 000 fishing outings year −1 ). Thus, the amount of carbon extracted annually is at least 137.34 kg C km −2 year −1 , and the recreational fishery removes about 31% of production at trophic level 4. Although these are gross estimates and more detailed study of the effect on trophic level and local production is needed, the values do highlight the pressure the recreational fishery exerts on coastal fish communities. Assuming that this level of exploitation is common to north-shore Mediterranean countries, there may be cause for concern about sustainable exploitation in the recreational fishery, and the effectiveness of current protection measures is discussed. Despite the limitations of the study, recreational fisheries clearly should be taken into account when considering measures for fisheries management. Moreover, fishing has considerable social import, and the benefits must be taken into account when investments to protect exploited resources are being contemplated.

title: The Role of Recreational Fishing in Global Fish Crises
abstract: Exploitation of fishery resources has become a major conservation issue on a global scale. Commercial fisheries have been repeatedly blamed for the worldwide declines in fish populations. However, we contend that the recreational fishing sector also has the potential to negatively affect fish and fisheries. Here we present evidence to show that both recreational and commercial fishing sectors deserve consideration as contributors to the exploitation of fish in marine and inland waters. The lack of global monitoring and compiling of statistics on recreational fishing participation, harvest, and catch-and-release has retarded our ability to understand the magnitude of this fishing sector. Using data from Canada, we estimate that the potential contribution of recreational fish harvest around the world may represent approximately 12 percent of the global fish harvest. Failure to recognize the potential contribution of recreational fishing to fishery declines, environmental degradation, and ecosystem alterations places ecologically and economically important resources at risk. Elevating recreational fishing to a global conservation concern would facilitate the development of strategies to increase the sustainability of this activity.

title: Spearfishing pressure on fish communities in rocky coastal habitats in a Mediterranean marine protected area
abstract: This study constitutes the first empirical investigation in the Mediterranean Sea quantifying in situ the pressure of spearfishing on fish communities in rocky coastal habitats in the Mediterranean. The investigation was carried out in the Cape Creus Marine Protected Area (MPA). Overall, 20 fish species and 1 invertebrate species were identified (three were coastal pelagic and the rest rocky benthic or demersal). In terms of abundance, two omnivorous species made up the 44% of the catch but in terms of weight, the main part (77%) of the catches relied on seven piscivorous (i.e. top-predator) fish (with trophic level values greater than 3.80). With the exception of three fish, the specie's intrinsic vulnerability values ranged between 40 and 90 (out of 100), implying a moderate to very high vulnerability to fishing. Spearfishing mainly targets large individuals measuring more than 30 cm in length. It is estimated that the total annual biomass extracted by spearfishing in the MPA is equivalent to ca. 40% of the total biomass extracted annually by artisanal (commercial) fishing. Overall, results highlight the pressure spearfishing exerts on the reproductive potential of fish species in rocky habitats along the Mediterranean coast, particularly those that are vulnerable (i.e. long lived and slow growing species with low reproductive potential) and have a higher trophic level. Results support the assumption that spearfishing can induce changes in both the trophic structure and the intrinsic vulnerability of taxa in the catch. Considering the biological implications of spearfishing, management should be supported by additional regulations in order to protect species that are vulnerable and have a higher trophic level.

title: The Decline of the Artisanal Fisheries in Mediterranean Coastal Areas: The Case of Cap de Creus (Cape Creus)
abstract: This article examines the sociocultural, economic, and environmental causes of the decline of the artisanal fisheries in the rocky coast of Cape Creus (NW Mediterranean) over the past decades. The changes that have occurred over time have favored trawlers, purse-seiners, shellfish fisheries, and tourism activities, to the detriment of artisanal fisheries. This article shows that the establishment of a marine reserve in 1998 could not reverse the observed decline in the artisanal fisheries. This raises the necessity of implementing a fishery management plan integrated into a coastal management plan, which should accompany the habitat protection. These plans could assist in the maintenance and the sustainable development of the artisanal fishing sector in Cape Creus as well as in other Mediterranean coastal areas where artisanal fisheries are also declining.

title: Dangerous targets? Unresolved issues and ideological clashes around marine protected areas
abstract: While conservationists, resource managers, scientists and coastal planners have recognized the broad applicability of marine protected areas (MPAs), they are often implemented without a firm understanding of the conservation science } both ecological and socio-economic } underlying marine protection. The rush to implement MPAs has set the stage for paradoxical differences of opinions in the marine conservation community. 2. The enthusiastic prescription of simplistic solutions to marine conservation problems risks polarization of interests and ultimately threatens bona fide progress in marine conservation. The blanket assignment and advocacy of empirically unsubstantiated rules of thumb in marine protection creates potentially dangerous targets for conservation science. 3. Clarity of definition, systematic testing of assumptions, and adaptive application of diverse MPA management approaches are needed so that the appropriate mix of various management tools can be utilized, depending upon specific goals and conditions. Scientists have a professional and

title: Marine protected areas and artificial reefs: A review of the interactions between management and scientific studies
abstract: Inshore marine areas host key habitats for the biological cycle of many marine populations, but are subject to strong anthropogenic pressure. Uses of inshore areas are multiple and often give rise to conflicting interests. In this context, marine protected areas (MPAs) and artificial reefs (ARs) are increasingly regarded as interesting management measures, in that they contribute to ecosystem conservation, fisheries sustainability, and because they can be helpful in zoning coastal areas in order to reduce conflicts between users. Beyond these expected benefits, it is important to appraise whether MPAs and ARs meet managers' objectives. This review focuses on the interaction between ecological sciences and management in the purpose of establishing or monitoring MPAs and ARs. We first study how objectives have to be formalized through management plans, and which kinds of information need to be collected in this respect. We then report management objectives related to MPAs and ARs, and list corresponding observed effects. Finally, we discuss the present role of monitoring and scientific research in the management process and propose a few perspectives for scientific research.

title: Benefits beyond boundaries: the fishery effects of marine reserves
abstract: Marine reserves are areas of the sea where fishing is not allowed. They provide refuges where populations of exploited species can recover and habitats modified by fishing can regenerate. In some places, closed areas have been used for fisheries management for centuries  and, until recently, natural refugia also existed, inaccessible through depth, distance or adverse conditions. Developments in technology have left few areas of fishing interest beyond our reach. Recently, the idea of marine reserves as fisheries management tools has re-emerged with developing interest in ecosystembased management, and observations of incidental fisheries benefits from reserves established for conservation. In light of new evidence, we argue that, by integrating large-scale networks of marine reserves into fishery management, we could reverse global fishery declines and provide urgently needed protection for

title: Biological responses in marine no-take reserves versus partially protected areas
abstract: Marine Protected Areas (MPAs) are a common tool for conserving and managing marine and coastal ecosystems. MPAs encompass a range of protection levels, from fully protected no-take reserves to restriction of only particular activities, gear types, user groups, target species, or extrac- tion periods. There is a growing body of scientific evidence supporting the ecological benefits of full reserve protection, but it is more difficult to generalize about the effects of other types of MPAs, in part because they include a range of actual protection levels. However, it is critical to determine whether partial protection and no-take reserves provide similar ecological benefits given potential economic costs of lost fishing grounds in no-take areas, common sociopolitical opposition to full pro- tection, and promotion of partially protected areas as a compromise solution in ocean zoning dis- putes. Here we synthesize all empirical studies comparing biological measures (biomass, density, species richness, and size of organisms) in no-take marine reserves and adjacent partially protected and unprotected areas across a range of geographic locations worldwide. We demonstrate that while partially protected areas may confer some benefits over open access areas, no-take reserves gener- ally show greater benefits and yield significantly higher densities of organisms within their bound- aries relative to partially protected sites nearby.

title: When can marine reserves improve fisheries management
abstract: Marine reserves are a promising tool for fisheries management and conservation of biodiversity, but they are not a panacea for fisheries management problems. For fisheries that target highly mobile single species with little or no by-catch or habitat impact, marine reserves provide few benefits compared to conventional fishery management tools. For fisheries that are multi-species or on more sedentary stocks, or for which broader ecological impacts of fishing are an issue, marine reserves have some potential advantages. Their successful use requires a case-by-case understanding of the spatial structure of impacted fisheries, ecosystems and human communities. Marine reserves, together with other fishery management tools, can help achieve broad fishery and biodiversity objectives, but their use will require careful planning and evaluation. Mistakes will be made, and without planning, monitoring and evaluation, we will not learn what worked, what did not, and why. If marine reserves are implemented without case by case evaluation and appropriate monitoring programs, there is a risk of unfulfilled expectations, the creation of disincentives, and a loss of credibility of what potentially is a valuable management tool.

title: Marine reserves: size and age do matter.
abstract: 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.

title: TRAJECTORIES AND CORRELATES OF COMMUNITY CHANGE IN NO-TAKE MARINE RESERVES
abstract: Marine reserves are a spatial approach to marine management and conservation aimed at protecting and restoring multispecies assemblages and the structure and function of marine ecosystems. We used meta-analyses of published data to address the questions of how and over what time frames marine assemblages change within no-take marine reserves as they recover from fishing and other human uses. We used 20 studies of coastal fish assemblages from 31 temperate and tropical locations, reporting abundances, and in some cases biomass, of 10–134 species from reserve and reference conditions (i.e., conditions in nearby fished sites or before reserve establishment) spanning 1–25 years of protection. Synthesis of data from these diverse sets of assemblages showed that: (1) a species' level of exploitation, trophic level, and the duration of protection through the no-take reserve explain small but significant amounts of variation in individual species responses to protection, with only species that are targeted by fishing or by aquarium trade showing overall enhanced abundances in protected areas, and increasing positive effects of protection on abundances at top trophic levels through time; (2) up to a third of species in different studies (19% on average) appeared to be negatively affected by protection, indicating that indirect effects of protection through competitive or predatory interactions may be common; and (3) variation and lags in species responses to protection resulted in protected assemblages diverging from reference conditions, with greater proportions of total fish biomass at top trophic levels in protected compared to fished assemblages. These results indicate that marine reserves are effective in enhancing local abundances of exploited species and restoring the structure of whole communities, though these changes occur through a series of transient states and, for some communities, over long time frames (decades). In contrast with the more predictable increases of aggregate community variables such as total abundance and biomass, individual species and community structure exhibited broad variation in their responses to protection. Marine protected areas represent multiple human-exclusion “experiments,” replicated in a variety of ecosystem types and geographic locations, providing key insights on community-wide impacts of the removal of human extraction. Long-term monitoring of community trajectories in marine protected areas and modeling studies scaling up local effects to relevant spatial and temporal scales are needed to increase our ability to protect and restore whole marine systems and to set realistic targets for the conservation and restoration of specific assemblages.

title: Temperate marine reserves enhance targeted but not untargeted fishes in multiple no-take MPAs.
abstract: Although many papers report the effects of no-take marine protected areas (MPAs or reserves), scientifically rigorous empirical studies are rare, particularly for temperate reef fishes. We evaluated the responses of fish populations to protection from fishing in reserves by comparing densities and sizes inside and outside of five no-take reserves in southern California, USA. Our results are robust because we compared responses across multiple rocky-reef reserves in two different years and controlled for possible site differences by (a) ensuring that habitat characteristics were the same inside and outside reserves, and (b) sampling species that are not targeted, which would not be expected to have a direct response to fishing. We compared fish density and size and calculated biomass and egg production across all five sites. Fishes targeted by recreational and/or commercial fisheries consistently exhibited increases in mean density (150%), size (30%), biomass (440%), and egg production (730%) inside reserves. Reserve effects were greatest for legal-sized targeted fishes: significantly greater densities were found exclusively inside reserves for targeted species (580%), the largest size classes existed only inside reserves, and mean biomass was 1000% higher. These responses were unlikely to have been caused by habitat differences because there were no significant differences in habitat characteristics between reserve and control locations. Densities of non-targeted species did not differ between reserve and non-reserve locations, further supporting the conclusions that differences in targeted species between reserve and control locations were due to harvesting rather than site-specific effects. Although MPAs cannot replace traditional fisheries management, the concentration of increased biomass and egg production is a unique MPA benefit that serves both reserves and fisheries. Scientifically rigorous studies that include multiple reserves, such as this study, are needed to inform management and policy decisions.

title: Effects of habitat on spillover from marine protected areas to artisanal fisheries
abstract: Marine protected areas (MPAs) potentially enhance the long-term sustainability of coastal fish resources that have been overexploited. The types and quality of habitats, both inside and outside the MPAs, may determine the likelihood of migration by fish to surrounding unprotected areas where spillover to fisheries occurs. We assessed whether MPAs enhanced catches of artisanal fisheries, using an experimental fishing study with the same fishing gear as that used by local fishers. This approach allowed us to test the hypothesis of increased catches along the borders of MPAs in comparison with those in other fishing grounds located at medium and far distances from 3 Medi- terranean MPAs: Tabarca Marine Reserve, Carry-le-Rouet Marine Reserve and Cerbere-Banyuls Marine Reserve. Surveys were done over 2 homogeneous habitats (Posidonia oceanica meadow and sand), in 3 different seasons. Catches were significantly higher for some species near the borders of the MPAs when fishing on P. oceanica meadows, but not when fishing on sandy bottoms. The spillover effect appears to be limited by a lack of continuous suitable habitat through the boundaries of the MPA. Some of the species that showed a significant response to protection and concurrent higher catches near the MPA borders, such as Dentex dentex, Mullus surmuletus, Phycis phycis, Sciaena umbra and Scorpaena porcus, are target species of artisanal fisheries. Although we found that the spatial scale of the spillover-induced density gradient was localized, it was sufficient to provide local benefits to artisanal fisheries. We conclude that spillover effects are not a universal consequence of siting MPAs in temperate waters and that they are related to the distribution of habi- tats inside and around MPAs.

title: Marine reserve benefits local fisheries
abstract: The utility of no-take marine reserves as fisheries-management tools is controversial. It is hypothesized that marine reserves will help to sustain fisheries external to them by becoming net exporters of adults (the “spillover effect”) and net exporters of propagules (the “recruitment effect”). Local fishery benefits from spillover will likely generate support from fishing communities for marine reserves. We used underwater visual census to show that biomass of Acanthuridae (surgeonfish) and Carangidae (jacks), two families of reef fish that account for 40–75% of the fishery yield from Apo Island, Philippines, tripled in a well-protected no-take reserve over 18 years (1983–2001). Biomass of these families did not change significantly over the same period at a site open to fishing. The reserve protected 10% of the total reef fishing area at the island. Outside the reserve, biomass of these families increased significantly closer to (200–250 m) than farther away from (250–500 m) the reserve boundary over time. We used published estimates of fishery catch and effort, and fisher interviews (creel surveys) to show that the total catch of Carangidae and Acanthuridae combined at Apo Island was significantly higher after (1985–2001) than before (1981) reserve establishment. Hook-and-line catch per unit effort (CPUE) at the island was 50% higher during 1998–2001 (reserve protected 16–19 years) than during 1981–1986 (pre-reserve and early phases of reserve protection). Total hook-and-line effort declined by 46% between 1986 and 1998–2001. Hook-and-line CPUE of Acanthuridae was significantly higher close to (within 200 m) than far from the reserve. CPUE of Carangidae was significantly higher away from the reserve, possibly reflecting a local oceanographic effect. The benefits of the reserve to local fisheries at the island were higher catch, increased catch rate, and a reduction in fishing effort. The fishery and tourism benefits generated by the reserve have enhanced the living standard of the fishing community.

title: Spatial assessment of benefits of a coastal Mediterranean Marine Protected Area
abstract: Marine reserves are increasingly advocated not only as conservation but also as fisheries management tools to safeguard the decline of coastal fishing resources. Still, conclusive evidence of their functioning is lacking, amongst others due to the influence of spatio-temporal variations in fish populations and habitat heterogeneity which could hamper a sound data interpretation. We conducted a spatial analysis of the benefits of the Medes Island Marine Reserve by combining geostatistical and Geographic Information System (GIS) tools. Concurrently, we analysed effects of trends reflecting habitat heterogeneity and spatial structuring of data on spatial predictions of fish catch per unit effort (CPUE) and length. Predicted spatial patterns showed the complexity and simultaneous action of trend factors leading to mostly non-linear gradients in CPUE and length data. CPUE of total fish and CPUE and length of common pandora (Pagellus erythrinus) increased close to the Integral Reserve due to direct and indirect reserve effects. CPUE and length of striped red mullet (Mullus surmuletus) slightly increased also near the Integral Reserve, but distinct reserve effects could not be identified due to the strong influence of artificial reefs. We conclude that the spatial dimension of the Buffer Zone, where artisanal fisheries are allowed, permits in general protection only for target species, favouring a habitat with no discontinuities from the reserve outwards. Our spatial approach to assess reserve benefits provides major insights into complex systems like coastal marine reserves in the northwestern Mediterranean. In addition, it contributes to a crucial aspect of marine conservation, viz. the decision on the spatial dimension of protected areas.

title: Spillover from six western Mediterranean marine protected areas: evidence from artisanal fisheries
abstract: This study investigated spillover (biomass export) around 6 marine protected areas (MPAs) in the western Mediterranean based on catch and effort data from artisanal fisheries. The selected MPAs were Cerbere-Banyuls and Carry-le-Rouet in France, and Medes, Cabrera, Tabarca, and Cabo de Palos in Spain. These MPAs had been functional for more than 8 yr and incorporate areas of fisheries closure and restricted use where fishing is limited. We based our study on the hypotheses that, in the presence of biomass export, (1) fishing effort would concentrate close to MPA boundaries, and (2) fishery production, expressed as catch per unit area (CPUA), would be highest near MPA boundaries and decrease with dis- tance. We selected data from 14 'fishing tactics' using gill nets, trammel nets and bottom long-lines target- ing sparids, mullids, serranids, scorpaenids and palinurids. We analyzed the spatial distribution of effort, fishery production and revenues per unit area, using generalized additive models (GAMs), and we tested regression slopes of effort density and CPUA with distance to closure boundaries, using generalized lin- ear models (GLMs). GAMs allowed us to recognize habitat discontinuities or 'hot spots' of high production in the vicinity of the MPAs, and to identify the extent of potential spillover effects in order to implement GLMs. We found evidence of effort concentration and high fishery production near fisheries closures for all fishing tactics analyzed and significant negative slopes for most. Revenues generally followed trends similar to CPUA. Significant negative slopes from GLM of effort density and CPUA with distance from fisheries closures were indicative of biomass export where habitats across closure boundaries had some degree of continuity. The spatial extent of spillover was consistent with species mobility and fisheries effi- ciency and extended 700 to 2500 m from fishery closure boundaries. Our results suggest that coastal MPAs can be an effective management tool for artisanal fisheries in the region and can be extended to the rest of the western Mediterranean, as the fishing tactics studied are typical of the region.

title: Indicator species analysis as a test of non-random distribution of species in the context of marine protected areas
abstract: Species are identified as bioindicators when their presence or abundance within sites of a particular habitat or a conservation area are high. Dufrêne and Legendre (1997), Ecological Monographs 67: 345–366, proposed the use of a flexible asymmetrical approach to detect indicator species and species assemblages characterizing groups of sites, the indicator value for each species being subsequently tested by randomization. Although the advantages of these methods in comparison to other multivariate or two-way indicator species analyses have been demonstrated, relatively few ecological studies have made use of them. The aim of the present study was to demonstrate the advantages of the indicator species test using a data set generated by traditional fisheries activities at four areas in the Bonifacio Strait Marine Reserve, Corsica, France. ANOVA, Kruskal-Wallis and another randomization test were also performed on the same data set. Of the 54 species caught in the fishing nets, the indicator species analyses identified 19 species for which the indicator value was significant for a given area, whereas the a posteriori Tukey test for ANOVA detected only 12 species for which the mass capture per unit effort was significantly greater for a given area. The efficiency of this new approach within the framework of conservation and marine reserve studies is demonstrated by focusing on three species (Sciaena umbra, Palinurus elephas and Maja verrucosa).

title: Paradox of estuarine quality: benthic indicators and indices, consensus or debate for the future.
abstract: The European Water Framework Directive will have instituted the concept of Ecological Quality Status (EQS) as a way to assess the biological quality of water masses. The EQS will be based mainly upon the composition of the different biological compartments in the ecosystem specially the benthos as compared to certain reference sites. Such management tools are already well established for freshwater (i.e. biotic indices), but not for coastal and estuarine (i.e. transitional) waters. In the framework of the Seine-Aval programme a workshop on benthic indicators was organized at Wimereux (France) in June 2005. The aim of this workshop and this paper is (1) to present the experiences of the Seine Aval researchers, and the French scientific approaches to benthic indicators, with those international experiences and approaches that have been published or are under development; and (2) to examine the existing benthic tools and their possible use in the characterization of the state of estuarine ecosystems. The debate during the workshop and the numerous recently published on the WFD are discussed in term of the implementation of the WFD in transitional water bodies using benthic indicators and indices. Some proposals for the future underline the needs to re-examine and adapt the different index thresholds, to take into account physical disturbances, to inventory the existing conditions vs reference conditions and to be as pragmatic as possible in using the WFD in transitional waters.

title: Assessing the effects of marine protected area (MPA) on a reef fish assemblage in a northwestern Mediterranean marine reserve: Identifying community-based indicators
abstract: Marine protected areas (MPAs) are increasingly envisaged as a tool to manage coastal ecosystems and fisheries. Assessment of their performance with respect to management objectives is therefore important. A number of WAS provided conservation benefits for fished species. Observed benefits do not apply to all species at all times, and responses to protection are also highly variable among fish taxa. Among the many empirical studies on marine reserves, only a few designs considered 'before and after data' and spatial variation. In this paper, we are interested in assessing the effect of a no-take reserve on the reef fish assemblage in a northwestern Mediterranean example. Data were obtained from a three-year survey using underwater visual censuses (UVC), before and after MPA establishment. Permutational multivariate analysis of variance (PERMANOVA) and multivariate regression trees (MRT) were used to evaluate the effects of reserve protection on the reef fish assemblage, while accounting for habitat. Modelled biological responses were abundances and diversity indices calculated at different levels of the assemblage. Significant effects were found for many of these metrics. in addition to PERMANOVA, univariate models provided more insight into the magnitude and direction of effects. The most sensitive metrics were related to large species and species targeted by fishing. These results may be used to choose the metrics that are more suitable as community-based indicators of MPA impact in the perspective of monitoring programs.

title: Designing indicators for assessing the effects of marine protected areas on coral reef ecosystems: A multidisciplinary standpoint
abstract: The present paper aims at identifying and assessing indicators of the effects of Marine Protected Areas (MPAs) in coral reef regions, based on a bibliography review in ecology, economics and social sciences. First the various effects studied within each of these domains and the variables used to measure them were censused. Poten- tial ecological indicators were assessed through their link with the question used (here termed "relevance") and their "effectiveness" which encompasses the issues of precision, accuracy and statistical power. Relevance and effectiveness were respectively measured by the frequency of use of each indicator and the proportion of significant results in the reviewed articles. For social and economic effects, the approach was not possible due to the low number of references; we thus discussed the issue of finding appropriate indicators for those fields.  1- the unbalance in liter- ature between disciplines; 2- the need for protocols and methodologies which include controls in order to assess MPA effects; 3- an important proportion of ecological indicators with low effectiveness; 4- the large number of ecological effects still not studied or not demonstrated at present.

title: A Marine Biotic Index to Establish the Ecological Quality of Soft-Bottom Benthos Within European Estuarine and Coastal Environments
abstract: In this paper, a marine Biotic Index (BI) for soft-bottom benthos of European estuarine and coastal environments is proposed. This is derived from the proportions of individual abundance in five ecological groups, which are related to the degree of sensitivity/tolerance to an environmental stress gradient. The main diAerence with previously published indices is the use of a simple formula that produces a continuous Biotic CoeAcient (BC) ‐ which makes it more suitable for statistical analysis, in opposition with previous discreet biotic indices ‐ not affected by subjectivity. Relationships between this coeAcient and a complementary BI with several environmental variables are discussed. Finally, a validation of the proposed index is made with data from systems aAected by recent human disturbances, showing that diAerent anthropogenic changes in the environment can be detected through the use of this BI. ” 2000 Elsevier Science Ltd. All rights reserved.

title: Standardizing catch and effort data: a review of recent approaches
abstract: The primary indices of abundance for many of the world's most valuable species (e.g. tunas) and vulnerable species (e.g. sharks) are based on catch and effort data collected from commercial and recreational fishers. These indices can, however, be misleading because changes over time in catch rates can occur because of factors other than changes in abundance. Catch-effort standardization is used to attempt to remove the impact of these factors. This paper reviews the current state of the art in the methods for standardizing catch and effort data. It outlines the major estimation approaches being applied, the methods for dealing with zero observations, how to identify and select appropriate explanatory variables, and how standardized catch rate data can be used when conducting stock assessments.

title: R graphics
abstract: Extensively updated to reflect the evolution of statistics and computing, the second edition of the bestselling R Graphics comes complete with with new packages and new examples. Paul Murrell, widely known as the leading expert on R graphics, has developed an in-depth resource that helps both neophyte and seasoned users master the intricacies of R graphics. New in the Second Edition Updated information on the core graphics engine, the traditional graphics system, the grid graphics system, and the lattice package A new chapter on the ggplot2 package New chapters on applications and extensions of R Graphics, including geographic maps, dynamic and interactive graphics, and node-and-edge graphs Organized into five parts, R Graphics covers both "traditional" and newer, R-specific graphics systems. The book reviews the graphics facilities of the R language and describes Rs powerful grid graphics system. The book then covers the graphics engine, which represents a common set of fundamental graphics facilities, and provides a series of brief overviews of some of the major areas of application for R graphics, and some of the major extensions of R graphics.

title: Computational aspects of nonparametric smoothing with illustrations from the sm library
abstract: Smoothing techniques such as density estimation and nonparametric regression are widely used in applied work and the basic estimation procedures can be implemented relatively easily in standard statistical computing environments. However, computationally efficient procedures quickly become necessary with large datasets, many evaluation points or more than one covariate. Further computational issues arise in the use of smoothing techniques for inferential, rather than simply descriptive, purposes. These issues are addressed in two ways by (i) deriving efficient matrix formulations of nonparametric smoothing methods and (ii) by describing further simple modifications to these for the use of 'binned' data when sample sizes are large. The implications for other graphical and inferential aspects of the estimators are also discussed. These issues are dealt with in an algorithmic manner, to allow implementation in any programming environment, but particularly those which are geared towards vector and matrix representations of data. Specific examples of S-Plus code from the sm library of Bowman and Azzalini (Applied Smoothing Techniques for Data Analysis: the Kernel Approach With S-Plus Illustrations,  1997) are given in an appendix as illustrations.

title: An R And S Plus Companion To Applied Regression
abstract: an r and s plus companion to applied regression is available in our digital library an online access to it is set as public so you can download it instantly. Our book servers saves in multiple countries, allowing you to get the most less latency time to download any of our books like this one. Kindly say, the an r and s plus companion to applied regression is universally compatible with any devices to read.

title: Niche overlap estimates based on quantitative functional traits: a new family of non-parametric indices
abstract: The concept of niche overlap appears in studies of the mechanisms of the maintenance of species diversity, in searches for assembly rules, and in estimation of within-community species redundancy. For plant traits measured on a continuous scale, existing indices are inadequate because they split the scale into a number of categories thus losing information. An index is easy to construct if we assume a normal distribution for each trait within a species, but this assumption is rarely true. We extend and apply an index, NOK, which is based on kernel density functions, and can therefore work with distributions of any shape without prior assumptions. For cases where the ecologist wishes to downweight traits that are inter-correlated, we offer a variant that does this: NOKw. From either of these indices, an index of the mean niche overlap in a community can be calculated: NOK,community and NOKw,community. For all these indices, the variance can be calculated and formulae for this are given. To give examples of the new indices in use, we apply them to a coastal fish dataset and a sand-dune plant dataset. The former exhibits considerable non-normality, emphasising the need for kernel-based indices. Accordingly, there was a considerable difference in index values, with those for an index based on a normal distribution being significantly higher than those from an index which, being based on kernel fitting, is not biased by an assumption for the distribution. The NOK values were ecologically consistent for the fish species concerned, varying from 0.02 to 0.53. The sand-dune plant data also showed a wide range of overlap values. Interestingly, the least overlap was between two graminoids, which would have been placed in the same functional group in the coarse classification often used in functional-type/ecosystem-function work.

title: Impacts of atypical data on Bayesian inference and robust Bayesian approach in fisheries
abstract: Bayesian inference is increasingly used in fisheries. In formulating likelihood functions in Bayesian inference, data have been analyzed as if they are normally, identically, and independently distributed. It has come to be believed that the first two of the assumptions are frequently inappropriate in fisheries studies. In fact, data distributions are likely to be leptokurtic and (or) contaminated by occasional bad values giving rise to outliers in many fisheries studies. Despite the likelihood of having outliers in fisheries studies, the impacts of outliers on Bayesian inference have received little attention. In this study, using a simple growth model as an example, we evaluate the impacts of outliers on the derivation of posterior distributions in Bayesian analyses. Posterior distributions derived from the Bayesian method commonly used in fisheries are found to be sensitive to outliers. The distributions are severely biased in the presence of atypical values. The sensitivity of normality-based Bayesian analyses on atypical data may result from small "tails" of normal distribution so that the probability of occurrence of an event drops off quickly as one moves away from the mean a distance of a few standard deviations. A robust Bayesian method can be derived by including a mixture distribution that increases the size of tail so that the probability of occurrence of an event does not drop off too quickly as one moves away from the mean. The posterior distributions derived from this proposed approach are found to be robust to atypical data in this study. The proposed approach offers a potentially useful addition to Bayesian methods used in fisheries.

title: Gradients of abundance and biomass across reserve boundaries in six Mediterranean marine protected areas : Evidence of fish spillover?
abstract: Marine protected areas (MPAs) are considered as an effective tool in marine coastal management, and considered able to enhance local fisheries through adult fish spillover. Indirect evidence of fish spillover could be obtained by horizontal gradients in fish abundance. To address this question, the existence of gradients of fish abundance and biomass across marine reserve boundaries was assessed in six Mediterranean MPAs using underwater visual censuses performed at various distances from the core of the MPA, in integral reserve (IR), to buffer zone (BZ) and fished areas. A reserve effect was evidenced with higher values of fish species richness (×1.1), abundance (×1.3), and biomass (×4.7) recorded inside MPAs compared to adjacent fished areas. Linear correlations revealed significant negative gradients in mean fish biomass in all the reserves studied after the effect of habitat had been removed, whereas negative gradients in abundance were less conspicuous. Generalized additive models suggested two main patterns of biomass gradients, with a sharp decrease at the IR–BZ boundary or at the BZ–fished area boundary. It was estimated that fish spillover beneficial to local fisheries occurred mostly at a small spatial scale (100s of metres). The existence of regular patterns of negative fish biomass gradients from within MPAs to fished areas was consistent with the hypothesis of adult fish biomass spillover processes from marine reserves and could be considered as a general pattern in this Mediterranean region.

title: The influence of bottom type and shelf position on biodiversity of tropical fish inside a recently enlarged marine reserve
abstract: 1. 
A necessary component of implementing a successful marine reserve is the quantification of the biological resources that fall under its protection. Without such an initial assessment, the future effects of the reserve on the local habitat and biotic community cannot be quantified and will remain the subject of debate. 
 
2. 
This study provides such a baseline assessment of fish diversity and habitat types within a recently enlarged marine reserve. Buck Island Reef National Monument, US Virgin Islands, was recently enlarged from approximately 4 km2 to over 76 km2. Areas of sand, seagrass, and hard-bottom under protection were increased from 0.29 km2, 0.47 km2, and 1.96 km2 to 2.70 km2, 2.89 km2, and 18.30 km2 respectively when the Monument was expanded. A 53 km2 area of pelagic/deep-water habitat with unknown bottom type is now also protected by the Monument. 
 
3. 
Visual counts of fish within 25×4 m2 transects conducted during the day were used to assess fish community structure and habitat utilization patterns. Species richness, diversity, assemblage structure, and fish density were evaluated and compared among sand, seagrass, and hard-bottom habitats. Hard-bottom sites had over twice the mean species richness and diversity as sand and seagrass sites, and several times greater mean fish density. 
 
4. 
Quantification of the fish community in pelagic and deep-water habitats within the reserve is recommended to provide a more comprehensive assessment of the offshore areas of the reserve. Fish numbers, size, and diversity outside the reserve boundaries must also be evaluated to allow quantification of the effects of the marine reserve on the adjacent fish communities.

title: Differences in recreationally targeted fishes between protected and fished areas of a coral reef marine park
abstract: Many comparisons have been made between sanctuary (no-fishing) and fished areas, where fishing pressure is exerted by artisanal or commercial fishers, but few have examined the effect of recreational fishing on fish assemblages in coral reef habitats. In this study, we compared assemblages of targeted fish from coral reef habitats in sanctuary (no-fishing) and recreationally fished zones of a marine protected area (MPA). Surface visual census (SVC) transects were conducted two times, at three regions, to compare the composition of predatory fish assemblages and the abundance, biomass, and size of the most commonly targeted fish. Baited remote underwater video (BRUV) was used to make relative counts of fish between zones. We also measured benthic cover and rugosity, which may influence fish assemblages. Analysis of similarity (ANOSIM) revealed significant differences in the composition of fish families/genera targeted by fishers (Lethrinidae, Lutjanidae, Haemulidae, Serranidae, and the genus Choerodon of the family Labridae) in terms of biomass (P<0.01) and abundance (P<0.05). The most consistent trends were recorded for biomass and this was supported by clustering of replicates in nonmetric multidimensional scaling (nMDS) ordinations. Similarity percentages (SIMPER) analysis indicated that the family Lethrinidae accounted for 73% (as abundance), and up to 69% (as biomass), of the dissimilarity between zones. Three-factor ANOVA highlighted significantly greater biomass, size, and abundance of legal-sized lethrinids (the most targeted family in the region) in sanctuary zones, but no differences in other families/genera. Results of BRUV supported SVC with greater relative counts of lethrinids (P<0.01) in sanctuaries, but no significant differences for other families. Cover of Acropora coral and hard substrate differed between zones at some regions but differences were inconsistent. There were no significant differences in algal cover or rugosity between zones. Given the inconsistency in benthic cover, the similarity of rugosity between zones, the consistently greater biomass of lethrinids in sanctuaries, and the abundance of large lethrinids in sanctuaries, the cessation of fishing in sanctuary zones appears responsible for observed differences in the populations of these fish. These results demonstrate that recreational

title: The Cap Roux MPA (Saint-Raphaël, French Mediterranean): changes in fish assemblages within four years of protection
abstract: Seytre, C., and Francour, P. 2009. The Cap Roux MPA (Saint-Raphael, French Mediterranean): changes in fish assemblages within four years of protection. - ICES Journal of Marine Science, 66: 180-187.In recent decades, marine reserves have been established to protect ecosystem structure and biological diversity, or as management tools to combat the overexploitation of fish stocks. The Cap Roux Marine Protected Area (MPA) was established by professional fishers in December 2003, in the French Mediterranean between Cannes and Saint-Raphael. It was implemented to enhance target fish stocks for local fisheries. The objective of this 3-year study was to investigate the initial responses of fish assemblages, using complementary methods: experimental net fishing performed by a professional fisher and underwater visual census. Within 3 years, this study detected early changes in the fish assemblages. The methods also detected an increase in abundance and diversity of fish, but also a decrease of seasonal fluctuations of the assemblage structure, which was characterized by winter values close to summer values in the protected zone but not outside of the MPA. These results helped clarify the dynamic by which fish assemblages respond to fishing prohibition in a newly created protected area.

title: SPILLOVER OF EXPLOITABLE FISHES FROM A MARINE PARK AND ITS EFFECT ON THE ADJACENT FISHERY
abstract: The role of a marine protected area in enhancing local fisheries, through the emigration or spillover of exploitable fishes, was studied in a coral reef park (Mombasa Marine Park, Kenya) and fishery over a seven-year period during a time when the park's border changed and pull seines were eliminated. We measured catches before and after the park's establishment and during the management changes and compared these catches with the unmanaged side of the park. Additionally, we placed baited traps on both sides of the park over a full tidal cycle which allowed us to measure the spillover from the park compared to the deeper, rougher, and less fished reef edge. The total wet mass of catches per trap, the mean size of the trapped fish, and the number of fish species caught per trap declined as a function of the distance away from the park edge on both the southern and northern sides. However, this relationship was truncated on the unmanaged side which also had smaller catches, smaller fish, and fewer species th...

title: Do partial marine reserves protect reef fish assemblages
abstract: Fish assemblages in the Mimiwhangata Marine Park, an area closed to commercial fishing but open to most forms of recreational fishing, were compared with adjacent fished areas. Two survey methodologies were used; baited underwater video and underwater visual census. Snapper (Pagrus auratus), the most heavily targeted fish species in the region, showed no difference in abundance or size between the Marine Park and adjacent control areas. When compared to the fully no-take Poor Knights Island Marine Reserve and two other reference areas open to all kinds of fishing (Cape Brett and the Mokohinau Islands), the abundance and size of snapper at the Marine Park were most similar to fished reference areas. In fact, the Marine Park had the lowest mean numbers and sizes of snapper of all areas, no-take or open to fishing. Baited underwater video found that pigfish (Bodianus unimaculatus), leatherjackets (Parika scaber) and trevally (Pseudocaranx dentex) were significantly more common in the Marine Park, than in the adjacent control areas. However, none of these species are heavily targeted by fishers. Underwater visual census found similar results with five species significantly more abundant in the Marine Park and five species more abundant outside the Marine Park. The lack of any recovery by snapper within the Marine Park, despite the exclusion of commercial fishers and restrictions on recreational fishing, indicates that partial closures are ineffective as conservation tools. The data suggest fishing pressure within the Marine Park is at least as high as at other ‘fished’ sites.

title: Alternatives to taxonomic-based approaches to assess changes in transitional water communities
abstract: 1. 
Transitional waters, described as critical transition zones because of their position at terrestrial, freshwater and marine interfaces, provide essential goods and services to the biosphere including human populations. These ecotones face increasing human influence mainly due to population density increase in coastal areas. 
 
2. 
Transitional water bodies have, to date, received little attention in the development of ecological status indicators; this is a critical deficiency when trying to meet the Water Framework Directive objective of all significant water bodies achieving good ecological status by the year 2015. 
 
3. 
In order to assess changes in transitional water communities many taxonomic-based indicators have already been proposed but there are a number of concerns for their use such as taxonomic classification difficulties, their unsuitability for multi-site comparisons and their inconsistent relationships with disturbance or stress. 
 
4. 
Alternative methods based on body size, abundance distribution among functional groups, functional diversity and productivity descriptors are proposed. These methods offer the opportunity to compare sites with different taxonomic compositions and allow derivation of indicators related to ecological status of communities under scrutiny. 
 
5. 
Finally, the suitability of these taxonomic-free descriptors to provide relevant information for each of the four main biotic compartments in coastal lagoons is discussed. The use of biomass distribution among functional groups for fish, benthos and macrophyte and to use body-size distribution for benthos and plankton is proposed.

title: Long-term and spillover effects of a marine protected area on an exploited fish community
abstract: We assessed the development of the exploited fish community inside and around the Columbretes Islands Marine Reserve (CIMR), a marine protected area (MPA), 8 to 16 yr after fishing ceased in the reserve. Sampling was by annual lobster trammel net fishing, an experimental tech- nique used inside the CIMR, and on-board commercial operations in adjacent fishing grounds. We examined trends in combined fish abundance and biomass (catch per unit effort), species richness and diversity, size structure, trophic level and species composition of the community. Our results showed the CIMR fish community continued to change throughout the study period as (1) abundance and biomass increased, (2) mean body size and trophic level increased and (3) species composition changed according to a linear model. Relative to nearby fished areas the CIMR fish community had (1) higher abundance and biomass, (2) lower species diversity and higher taxonomic distinctness, (3) larger relative body size and (4) no difference in mean trophic level. We found clear evidence of spillover of fish from the CIMR to the adjacent fishery as commercial fish yields at the MPA border (<0.5 km from the boundary) increased continuously during the study period, despite being locally depleted due to fishing effort concentration (fishing the line). Furthermore, fish size and diversity at the border were intermediate between the CIMR and other fished zones, suggesting that this is a transitional zone influenced by this MPA. Our results show that changes in community abundance, biomass, size structure and species composition provide a clear and interpretable view of MPA recov- ery. Diversity indices are also useful; however, their interpretation is more difficult. We conclude that the creation of the CIMR has had a positive effect on the exploitable fish community and that there is evidence of exportation of biomass to the surrounding fishery. We highlight the advantage of using multiple community metrics to study changes in fish communities, yet recommend the need for caution when interpreting them.

title: Structural change in an exploited fish community: a consequence of differential fishing effects on species with contrasting life histories
abstract: 1. An understanding of the links between life histories and responses to exploitation could provide the basis for predicting shifts in community structure by identifying susceptible species and linking life-history tactics with population dynamics. 
 
 
 
2. We examined long-term trends in the abundance of species in the North Sea bottom-dwelling (demersal) fish community. Between 1925 & 1996 changes in species composition led to an increase in mean growth rate, while mean maximum size, age at maturity and size at maturity decreased. The demersal fish community was increasingly heavily fished during this period. 
 
 
 
3. Trends in mean life-history characteristics of the community were linked to trends in abundance of component species. An approach based on phylogenetic comparisons was used to examine the differential effects of fishing on individual species with contrasting life histories. 
 
 
 
4. Those species that decreased in abundance relative to their nearest relative, matured later at a greater size, grew more slowly towards a greater maximum size and had lower rates of potential population increase. The phylogenetically based analyses demonstrated that trends in community structure could be predicted from the differential responses of related species to fishing. 
 
 
 
5. This is the first study to link exploitation responses of an entire community to the life histories of individual species. The results demonstrate that fishing has greater effects on slower growing, larger species with later maturity and lower rates of potential population increase. The comparative approach provides a basis for predicting structural change in other exploited communities.

title: Short-term effects of fishing on life history traits of fishes
abstract: An important aspect of species susceptibility to fishing are the changes in demographic characteristics of populations that fishing might induce. The purpose of this study is to show the short-term effects of fishing on growth and reproduction patterns. This assessment is made through a comparative study of key parameters (mortality, size, age and size at maturity, fecundity) among stocks subject to various levels of exploitation. Data have been assembled for 77 separate (primarily commercial) fish stocks.Trait variation is partitioned into effects attributable to size, phylogeny, and population. High adult mortality appears to lead to short-term apparently plastic changes in age and size at maturity: exploited populations are characterized by earlier age and increased size at maturity. This compensatory response to exploitation may conceal longer term selection effects, and may be worth considering in stock assessments.

title: Maladaptive changes in multiple traits caused by fishing: impediments to population recovery.
abstract: Some overharvested fish populations fail to recover even after considerable reductions in fishing pressure. The reasons are unclear but may involve genetic changes in life history traits that are detrimental to population growth when natural environmental factors prevail. We empirically modelled this process by subjecting populations of a harvested marine fish, the Atlantic silverside, to experimental size-biased fishing regimes over five generations and then measured correlated responses across multiple traits. Populations where large fish were selectively harvested (as in most fisheries) displayed substantial declines in fecundity, egg volume, larval size at hatch, larval viability, larval growth rates, food consumption rate and conversion efficiency, vertebral number, and willingness to forage. These genetically based changes in numerous traits generally reduce the capacity for population recovery.

title: What about recreational catch?: Potential impact on stock assessment for Hawaii's bottomfish fisheries
abstract: The bottom-fish stocks of the Main Hawaiian Islands (MHI) are intensively fished, both commercially and recreationally. Recent assessments of the Bottomfish Management Unit Species (BMUS) complex suggested overfishing, and expressed concerns about missing non-commercial data. We used reported commercial time-series data and estimation ratios to indirectly estimate non-commercial catches for non-pelagic species (i.e., excluding tuna and billfishes) for 1950–2005. Using adjustment ratios, we also accounted for commercial under-reporting, which suggested that total commercial non-pelagic catches were 28–128% higher than reported commercial catches for any given year. Estimated non-commercial catches for 1950–2005 were 2.1 times higher than reported commercial catches. Reported catches underestimated likely total catches (reported and un-reported commercial plus non-commercial) of non-pelagic species and BMUS components for 1950–2005 by a factor of 3.9 and 2.9, respectively. We incorporated the reconstructed BMUS non-commercial catches into stock assessments of the officially reported commercial BMUS catches via a Schaefer production model. Total catch increased by 2.5–3.5 times with the addition of non-commercial BMUS catch estimates, which in turn increased model estimates of MSY and carrying capacity (k) by approximately four times compared to analyses with reported commercial data alone. As the CPUE data lacked information to resolve the confounding between large, unproductive and small, productive stocks, an informative prior was used for fishing mortality rate to attain MSY (FMSY). To address uncertainty in key management parameters, independent estimates of exploitation rate, or fisheries independent estimates of abundance, and informative trends in recreational effort or catches are required.

title: Some effects of marine reserve protection on the population structure of two spearfishing target-fish species, Dicentrarchus labrax (Moronidae) and Sparus aurata (Sparidae), in shallow inshore waters, along a rocky coast in the northwestern Mediterranean Sea
abstract: 1. The relative abundances of two highly sought after spearfishing target-fish species, Dicentrarchus labrax (Linnaeus 1758) and Sparus aurata Linnaeus 1758, were compared using underwater visual censuses in shallow (<5 m) inshore waters along rocky shores inside and outside a marine reserve area in the northwestern Mediterranean Sea (Banyuls-sur-Mer region, France). 
 
 
 
2. The relative abundances of these two species were consistently lower outside the reserve area, which comprised around 40% of the 25.6 km of inshore coastal waters surveyed. 
 
 
 
3. The demographic structure of D. labrax was also different inside and outside the protected area, the proportion of larger fish being significantly higher in the marine reserve. 
 
 
 
4. The direct and indirect impacts of spearfishing are postulated to explain the distributional and population structural characteristics of these two species in the shallow waters off this rocky coast. 
 
 
 
5. Prohibition of spearfishing has favoured the persistence of these species during summer (the main spearfishing season) in the shallower inshore waters inside the area of the Cerbere/Banyuls-sur-Mer marine reserve. 
 
 
 
6. These preliminary results indicate the need for additional work in this area and further surveys to ascertain spearfishing impacts on target-fish species in the vicinity of other marine protected areas.

title: Fish diversity patterns in the Mediterranean Sea: deviations from a mid-domain model
abstract: Many ecological drivers have been proposed to explain the spatial distribution of species richness (productivity, energy, climate), but historical and paleogeographical contingencies may also be important. All these hypotheses make the implicit assumption that species richness would be similar everywhere in absence of any direct environmental variations; this is known as the mid-domain effect (MDE). The Mediterranean Sea is an 'archetypal situation' where we would expect this effect, so we used the MDE as a null model for fish richness and analyzed the deviations. Temperature, primary production and distance from both the Gibraltar Strait and Suez Canal have been included in our models as predictors. To obtain our objective we (1) built a database containing the geographic distributions of 619 Mediterranean fishes, (2) implemented 1- and 2-dimensional MDE models, and (3) carried out spatial autoregressive models on MDE deviations. It appeared that in 1 dimension, the only true MDE was displayed longitudinally by the large-ranged endemic species. In 2 dimensions, the MDE accounted for 21% of variation in fish richness distribution on the continental shelf. However, we showed that neglecting the MDE when relating fish richness to envi- ronmental predictors is misleading. In fact, minimum temperature and primary production negatively influenced fish species richness per se, but had a positive effect on deviations from MDE predictions. Finally, our study suggested that by analyzing the deviations from MDE predictions and taking into account spatial autocorrelation we developed a more rigorous assessment of the factors influencing diversity patterns.

title: Evaluating the ecological effects of Mediterranean marine protected areas: habitat, scale and the natural variability of ecosystems
abstract: The capability to detect and predict the responses of marine populations and communities to the establishment of marine protected areas (MPAs) depends on the ability to distinguish between the influences of management and natural variability due to the effects of factors other than protection. Thus, it is important to understand and quantify the magnitude and range of this natural variability at each scale of observation. Here we review the scale of responses of target populations and communities to protection within Mediterranean MPAs, against their ‘normal’ spatio-temporal heterogeneity, and compare those with documented cases from other temperate and tropical marine ecosystems. Additionally, we approach the problem of the relative importance of habitat structure, considered as a set of biological and physical elements of the seascape hierarchically arranged in space at multiple scales, to drive natural variability. We conclude that much more effort has to be made to characterize heterogeneity in relation to Mediterranean MPAs, and to quantify and explain relationships between target species and their habitats as sources of such variability. These studies should be based on sound sampling designs, which (1) generate long-term data sets, and would ideally (2) be based on a Mediterranean-wide comparison of a number of protected and unprotected localities, (3) be designed from a multi-scaled perspective, and (4) control for factors other than protection, in order to avoid their confounding effects. The need for appropriate spatial and temporal replication, nested designs and power analysis is advocated.

title: Considering multiple-species attributes to understand better the effects of successive changes in protection status on a coral reef fish assemblage
abstract: The response of fish assemblages to changes in protection status is a major issue for both biodiversity conservation and fishery management. In New Caledonia, the Abore reef marine reserve harbours more than 500 fish species, and has been subjected to changes in protection status since 1988. The present study investigates the impact of these changes on a wide subset of species (213), based on underwater visual counts collected before the opening and after the closure to fishing of this marine protected area (MPA). We analysed the spatial and temporal variability in fish assemblage attributable to protection status, explicitly considering habitat. To understand the successive responses of fish assemblage to fishing and protection, the assessment models included four criteria defining species groups that partition the fish assemblage: trophic regime, adult size, mobility, and interest for fishing. We could therefore identify the negative impact of opening the MPA to fishing on piscivores and highly mobile species. Surprisingly, target species were not affected more than non-target species. Model results were used to identify species groups that respond to fishing and protection. These results utilize fisheries-related criteria to provide new insight into the response of fish assemblages to protection from the perspective of MPA monitoring.

title: Modelling spatial and temporal scales for spill-over and biomass exportation from MPAs and their potential for fisheries enhancement
abstract: Marine protected areas (MPAs) are considered as a tool for marine conservation and sustainable fishery resource management. Improvements in fishery yields should take place via the spill-over of individuals from the reserve. In general, it has been demonstrated that MPAs affect the density and biomass of the organisms within them, however, little evidence has been found in order to assess the exportation of individuals across their boundaries. In this study, a simple model involving population growth, harvest, and the diffusion coefficient for individuals was used to explore the effects of protection on populations inside the reserve and the spill-over of individuals to the fished area. The model showed that biological responses inside marine reserves appear to develop quickly, reaching mean levels within a short (1–5 year) time period. Mean population abundance is always higher inside the reserve and highlights the effectiveness of protection, particularly when there is strong fishing pressure outside the reserve. However, reserves smaller than 2000 m radius show significantly lower levels of abundance inside than larger sites. Large MPAs (i.e. about 2000 m in radius) offer nearly the maximum capacity for recovery (close to 100% of the system carrying capacity) and nearly the maximum flux of individuals per unit boundary length. Very large MPAs (i.e. larger than 6000 m in radius) could be a guaranteed means of providing resilience in order to prevent population crises, with the added advantage that the flux of individuals is slightly higher at larger distances from the boundary. However, in practice they provide no further advantage towards increasing the density of individuals or the exportation of biomass, and a network of smaller MPAs could be more beneficial, both from the point of view of conservation and of benefits to fisheries.

title: Incorporating ecosystem objectives into management of sustainable marine fisheries, including 'best practice' reference points and use of marine protected areas.
abstract: The broadening of fisheries management to include ecosystem-related objectives raises a potentially confusing range of possible issues for consideration in management decisions, in reporting, and in assessing management performance. However, there are methods available and approaches to addressing the issues that are practical, accessible to stakeholder participation, and scientifically assessable. Three broad and interrelated elements are described that allow ecosystem objectives to be practically and operationally incorporated into marine fisheries management systems. Reporting and assessment of the whole management system against sustainability objectives. Three major points are developed and emphasized: (i) Indicators and reference points – and consequently performance measures – must explicitly relate to the high-level objectives of management. (ii) The structure and focus of reports on sustainability must be transparently derived from the high-level objectives. A methodology for this is described that can be used in meetings with stakeholders to elucidate the issues, indicators and reference points, management response and the justification for decisions. It can include risk-based methods to help identify the relative importance of different issues. (iii) Performance assessment must be of the management system as a whole, rather than solely on the merits of particular parts in isolation. An established methodology (Management Strategy Evaluation) is described that can be used to quantitatively test the likely performance of different management strategies in achieving ecosystem objectives. A management strategy in this context is a combination of monitoring, use of the monitoring data for assessment against reference points, identification of appropriate management measures, and implementation of these measures. This methodology can be used to test any aspect of the strategy in the “common currency” of the management objectives, and to identify the circumstances in which particular strategies are likely to perform well or fail. It has already been used in fisheries in relation to target species, important by-catch species, predator-prey dependencies, and sea-bed habitats. Keith SAINSBURY Ussif Rashid SUMALIA Keith.Sainsbury@marine.csiro.au R.Sumaila@fisheries.ubc.ca 2 Incorporating ecosystem objectives into management of sustainable marine fisheries K. Sainsbury and U.R. Sumalia Indicators, reference points and performance measures for fisheries ecosystem objectives. There are many options available and some recent summaries are identified. A set of targetand limit-reference points for fisheries ecosystem objectives are provided. These are based broadly on experience to date, and could be practically implemented in the short term. It is not claimed that these reference points are necessary or adequate to achieve sustainability for fisheries and marine ecosystems. Rather, they represent a practical and emerging “best practice” means of operationally accommodating ecosystem-related objectives in fisheries management. Use of marine protected areas to achieve ecosystem objectives in fisheries management Fisheries have long used some forms of spatial management, such as closure of nursery areas to protect juvenile fish, but more recently there has been a focus on use of marine protected areas (MPAs) to achieve fishery objectives for the target species and for the ecosystem more generally. MPAs hold promise as a rational and practical way of managing ocean resources to achieve fishery ecosystem objectives, although this promise should not be overstated. MPAs are best seen as part of a collection of management tools and measures, with a combination of on-reserve and off-reserve measures being used together to achieve sustainable fisheries and marine ecosystems. Several new technological developments are making their design and management more practical. These recent developments are reviewed.

title: Avoidance of fisheries-induced evolution: management implications for catch selectivity and limit reference points
abstract: I examined how the fitness (r) associated with early‐ and late‐maturing genotypes varies with fishing mortality (F) and age‐/size‐specific probability of capture. Life‐history data on Newfoundland’s northern Atlantic cod (Gadus morhua) allowed for the estimation of r for individuals maturing at 4 and 7 year in the absence of fishing. Catch selectivity data associated with four types of fishing gear (trap, gillnet, handline, otter trawl) were then incorporated to examine how r varied with gear type and with F. The resulting fitness functions were then used to estimate the F above which selection would favour early (4 year) rather than delayed (7 year) maturity. This evolutionarily‐sensitive threshold, Fevol, identifies a limit reference point somewhat similar to those used to define overfishing (e.g., Fmsy, F0.1). Over‐exploitation of northern cod resulted in fishing mortalities considerably greater than those required to effect evolutionary change. Selection for early maturity is reduced by the dome‐shaped selectivities characteristic of fixed gears such as handlines (the greater the leptokurtosis, the lower the probability of a selection response) and enhanced by the knife‐edged selectivities of bottom trawls. Strategies to minimize genetic change are consistent with traditional management objectives (e.g., yield maximization, population increase). Compliance with harvest control rules guided by evolutionarily‐sensitive limit reference points, which may be achieved by adherence to traditional reference points such as Fmsy and F0.1, should be sufficient to minimize the probability of fisheries‐induced evolution for commercially exploited species.

title: Indicators to support an ecosystem approach to fisheries
abstract: Indicators are needed to support the implementation of an ecosystem approach to fisheries (EAF), by providing information on the state of the ecosystem, the extent and intensity of effort or mortality and the progress of management in relation to objectives. Here, I review recent work on the development, selection and application of indicators and consider how indicators might support an EAF. Indicators should guide the management of fishing activities that have led to, or are most likely to lead to, unsustainable impacts on ecosystem components or attributes. The numbers and types of indicators used to support an EAF will vary among management regions, depending on resources available for monitoring and enforcement, and actual and potential fishing impacts. State indicators provide feedback on the state of ecosystem components or attributes and the extent to which management objectives, which usually relate to state, are met. State can only be managed if the relationships with fishing (pressure) and management (response) are known. Predicting such relationships is fundamental to developing a management system that supports the achievement of objectives. In a management framework supported by pressure, state and response indicators, the relationship between the value of an indicator and a target or limit reference point, reference trajectory or direction provides guidance on the management action to take. Values of pressure, state and response indicators may be affected by measurement, process, model and estimation error and thus different indicators, and the same indicators measured at different scales and in different ways, will detect true trends on different timescales. Managers can use several methods to estimate the effects of error on the probability of detecting true trends and/or to account for error when setting reference points, trajectories and directions. Given the high noise to signal ratio in many state indicators, pressure and response indicators would often guide short-term management decision making more effectively, with state indicators providing longer-term policy-focused feedback on the effects of management action.

title: A review of approaches for classifying benthic habitats and evaluating habitat quality.
abstract: We have assessed the current state of knowledge relative to methods used in assessing sub-tidal benthic habitat quality and the classification of benthic habitats. While our main focus is on marine habitat, we extensively draw on knowledge gained in freshwater systems where benthic assessment procedures are at an advanced stage of maturity. We found a broad range of sophistication/complication in terms of the methods applied in assessing and mapping benthic habitats. The simplest index or metric involved some assessment of species richness, while the most complicated required utilizing multi-variate analysis. The simplest mapping attempts equated physical substrate with benthic habitat while the most sophisticated relied on extensive environmental preference and groundtruth data for species of concern. The leading edge of methods for benthic habitat mapping involves combining the advances in optical and acoustic methods that allow for routine classifying and mapping of the seafloor with biological and habitat data for species of concern. The objective of this melding of dispirit methods is to produce benthic habitat maps with broad system wide coverage and sound biological underpinning. It is clear that the disparity in information density between the physical and biological sides of the equation currently hinder applicability and acceptability of benthic habitat mapping efforts. In addition to the lack of basic information on the biological and environmental tolerances of targeted species, the proliferation of metrics for characterizing and assessing biological conditions further clouds the usefulness of any broad scale mapping attempt. The problem of data density mismatch between physical and biological methods will likely not be solved until acoustic methods can routinely resolve the elusive biological components that make a physical substrate a habitat.

title: Do climate and fishing influence size-based indicators of Celtic Sea fish community structure?
abstract: Ecosystem-based management requires the development of indicators that allow anthropogenic impacts to be detected against the background of natural variation. Size-based community metrics are potentially useful indicators because of their theoretical foundation and practical utility. Temporal and spatial patterns in size-based community metrics for Celtic Sea fish are described and calculated using data from the English groundfish survey of the area (1987-2003). The results reveal that the size structure of the community has changed over time, and that a decrease in the relative abundance of larger fish was accompanied by an increase in smaller fish (4-25 g). Temporal analyses of the effects of fishing and climate variation suggest that fishing generally has had a stronger effect on size structure than changes in temperature. Therefore, size-based metrics respond clearly to the effects of fishing even in variable environments, reflecting the ubiquity of size-based processes in defining community structure and responses to mortality. Spatial analyses were inconclusive, probably owing to the limited area for which fishing effort, temperature, and survey data were all available.

title: The Mediterranean Sea as a 'cul-de-sac' for endemic fishes facing climate change.
abstract: The Mediterranean Sea is a hotspot of biodiversity, and climate warming is expected to have a significant influence on its endemic fish species. However, no previous studies have predicted whether fish species will experience geographic range extensions or contractions as a consequence of warming. Here, we projected the potential future climatic niches of 75 Mediterranean Sea endemic fish species based on a global warming scenario implemented with the Mediterranean model OPAMED8 and a multimodel inference, which included uncertainty. By 2070-2099, the average surface temperature of the Mediterranean Sea was projected to warm by 3.1 °C. Projections for 2041-2060 are that 25 species would qualify for the International Union for the Conservation of Nature and Natural Resources (IUCN) Red List, and six species would become extinct. By 2070-2099, 45 species were expected to qualify for the IUCN Red List whereas 14 were expected to become extinct. By the middle of the 21st century, the coldest areas of the Mediterranean Sea (Adriatic Sea and Gulf of Lion) would act as a refuge for cold-water species, but by the end of the century, those areas were projected to become a 'cul-de-sac' that would drive those species towards extinction. In addition, the range size of endemic species was projected to undergo extensive fragmentation, which is a potentially aggravating factor. Since a majority of endemic fishes are specialists, regarding substratum and diet, we may expect a reduced ability to track projected climatic niches. As a whole, 25% of the Mediterranean Sea continental shelf was predicted to experience a total modification of endemic species assemblages by the end of the 21st century. This expected turnover rate could be mitigated by marine protected areas or accelerated by fishing pressure or competition from exotic fishes. It remains a challenge to predict how these assemblage modifications might affect ecosystem function.

title: Predicting indirect effects of fishing in Mediterranean rocky littoral communities using a dynamic simulation model
abstract: Modelling may significantly enhance our understanding of the potential impacts of fisheries at larger spatial scales and on groups that would otherwise be very difficult to study. An aggregated biomass-based simulation model of a Mediterranean infralittoral zone was developed and used to carry out fishing ‘experiments’ where fishing intensity and catch selection were varied. The model was constructed for the Bay of Calvi, Corsica, using the Ecopath with Ecosim software, and was composed of 27 compartments, including seabirds, 11 groups of fish, 12 groups of invertebrates, 2 primary producers, bacteria and detritus. Several instances of indirect fishing effects (‘trophic cascades’ and ‘keystone predation’) have been proposed from anecdotal evidence in the western Mediterranean. Model outcomes provided little support for the widely accepted paradigm that fishing, by removing invertebrate-feeding fish, allows increases in the biomass of sea urchins and as a consequence the formation of overgrazed ‘barrens’ of bare substrate. Simulated harvesting of sea urchins by humans did, however, results in an increase of macroalgal biomass as reported previously. Intensified fishing pressure on ‘macrocarnivorous’ fish resulted in a ‘release’ of small fish species (e.g. blennies), and as a consequence a decline in the biomass of some small invertebrates on which they feed (e.g. amphipods). Increased fishing on large ‘piscivores’ resulted in increases in other small fish groups and consequential effects on other benthic invertebrate groups (e.g. polychaetes). Depletion of piscivorous fish resulted in a dramatic increase in the biomass of seabirds, which apparently compete with piscivores for small demersal and pelagic fish. An intensification of fishing pressure overall resulted in an increase in cephalopod biomass. Responses of target species to increased fishing pressure were most marked within the first 5 years of the new fishing regime. Indirect responses exhibited varying degrees of inertia, and biomasses of many groups did not assume a new equilibrium within the first 20 years of the simulation. The Mediterranean infralittoral rocky-bottom ecosystem was predicted to be relatively resilient to pulses of increased fishing and exhibited a high degree of detritus recycling. However, the speed and magnitude of ecosystem responses was shown to depend greatly on the extent of ‘top-down’ or ‘bottom-up’ control assumed for components within the system.

title: Marine reserves demonstrate top-down control of community structure on temperate reefs
abstract: Replicated ecological studies in marine reserves and associated unprotected areas are valuable in examining top-down impacts on communities and the ecosystem-level effects of fishing. We carried out experimental studies in two temperate marine reserves to examine these top-down influences on shallow subtidal reef communities in northeastern New Zealand. Both reserves examined are known to support high densities of predators and tethering experiments showed that the chance of predation on the dominant sea urchin, Evechinus chloroticus, within both reserves was approximately 7 times higher relative to outside. Predation was most intense on the smallest size class (30–40 mm) of tethered urchins, the size at which urchins cease to exhibit cryptic behaviour. A high proportion of predation on large urchins could be attributed to the spiny lobster, Jasus edwardsii. Predation on the smaller classes was probably by both lobsters and predatory fish, predominantly the sparid Pagrus auratus. The density of adult Evechinus actively grazing the substratum in the urchin barrens habitat was found to be significantly lower at marine reserve sites (2.2±0.3 m–2) relative to non-reserve sites (5.5±0.4 m–2). There was no difference in the density of cryptic juveniles between reserve and non-reserve sites. Reserve populations were more bimodal, with urchins between 40 and 55 mm occurring at very low numbers. Experimental removal of Evechinus from the urchin barrens habitat over 12 months lead to a change from a crustose coralline algal habitat to a macroalgal dominated habitat. Such macroalgal habitats were found to be more extensive in both reserves, where urchin densities were lower, relative to the adjacent unprotected areas that were dominated by urchin barrens. The patterns observed provide evidence for a top-down role of predators in structuring shallow reef communities in northeastern New Zealand and demonstrate how marine reserves can reverse the indirect effects of fishing and re-establish community-level trophic cascades.