Marine Reserves: from Beverton and Holt to the Present

The idea of using marine reserves, where all fishing is banned is not new to fisheries management. It was first formally considered by Beverton and Holt but rejected in favour of approaches such as fleet and gear control. Since that analysis, many fisheries have collapsed worldwide, illustrating the vulnerability of fishery resources and the ineffectiveness of these approaches. Empirical data and modelling suggest that marine reserves would generally increase yields, especially at the high fishing mortality that occurs in most fisheries. However, the most interesting feature of reserves is their ability to provide resilience to overexploitation, thereby reducing the risk of stock collapse. Benefits from reserves come from the increase in biomass and individual size within them, resulting in adult migration and/or larval dispersal that would replenish fishing grounds. The use of marine reserves in managing fisheries necessitates a thorough understanding of critical habitat requirements, fish movement, fish behaviour, the relations between subpopulations and the critical density effect for larval dispersal. When properly designed, and coupled with other management practices, reserves may provide a better insurance against uncertainties in stock assessment, fishing control and management by protecting a part of the population from exploitation. This strategy can be used for both sedentary and migratory species.     

Rapid Effects of Marine Reserves via Larval Dispersal

Marine reserves have been advocated worldwide as conservation and fishery management tools. It is argued that they can protect ecosystems and also benefit fisheries via density-dependent spillover of adults and enhanced larval dispersal into fishing areas. However, while evidence has shown that marine reserves can meet conservation targets, their effects on fisheries are less understood. In particular, the basic question of if and over what temporal and spatial scales reserves can benefit fished populations via larval dispersal remains unanswered. We tested predictions of a larval transport model for a marine reserve network in the Gulf of California, Mexico, via field oceanography and repeated density counts of recently settled juvenile commercial mollusks before and after reserve establishment. We show that local retention of larvae within a reserve network can take place with enhanced, but spatially-explicit, recruitment to local fisheries. Enhancement occurred rapidly (2 yrs), with up to a three-fold increase in density of juveniles found in fished areas at the downstream edge of the reserve network, but other fishing areas within the network were unaffected. These findings were consistent with our model predictions. Our findings underscore the potential benefits of protecting larval sources and show that enhancement in recruitment can be manifested rapidly. However, benefits can be markedly variable within a local seascape. Hence, effects of marine reserve networks, positive or negative, may be overlooked when only focusing on overall responses and not considering finer spatially-explicit responses within a reserve network and its adjacent fishing grounds. Our results therefore call for future research on marine reserves that addresses this variability in order to help frame appropriate scenarios for the spatial management scales of interest.     

When Is Spillover from Marine Reserves Likely to Benefit Fisheries?

The net movement of individuals from marine reserves (also known as no-take marine protected areas) to the remaining fishing grounds is known as spillover and is frequently used to promote reserves to fishers on the grounds that it will benefit fisheries. Here we consider how mismanaged a fishery must be before spillover from a reserve is able to provide a net benefit for a fishery. For our model fishery, density of the species being harvested becomes higher in the reserve than in the fished area but the reduction in the density and yield of the fished area was such that the net effect of the closure was negative, except when the fishery was mismanaged. The extent to which effort had to exceed traditional management targets before reserves led to a spillover benefit varied with rates of growth and movement of the model species. In general, for well-managed fisheries, the loss of yield from the use of reserves was less for species with greater movement and slower growth. The spillover benefit became more pronounced with increasing mis-management of the stocks remaining available to the fishery. This model-based result is consistent with the literature of field-based research where a spillover benefit from reserves has only been detected when the fishery is highly depleted, often where traditional fisheries management controls are absent. We conclude that reserves in jurisdictions with well-managed fisheries are unlikely to provide a net spillover benefit.     

Marine reserve effects on fishery profit

Some studies suggest that fishery yields can be higher with reserves than under conventional management. However, the economic performance of fisheries depends on economic profit, not fish yield. The predictions of higher yields with reserves rely on intensive fishing pressures between reserves; the exorbitant costs of harvesting low-density populations erode profits. We incorporated this effect into a bioeconomic model to evaluate the economic performance of reserve-based management. Our results indicate that reserves can still benefit fisheries, even those targeting species that are expensive to harvest. However, in contrast to studies focused on yield, only a moderate proportion of the coast in reserves (with moderate harvest pressures outside reserves) is required to maximize profit. Furthermore, reserve area and harvest intensity can be traded off with little impact on profits, allowing for management flexibility while still providing higher profit than attainable under conventional management.     

A General Business Model for Marine Reserves

Marine reserves are an effective tool for protecting biodiversity locally, with potential economic benefits including enhancement of local fisheries, increased tourism, and maintenance of ecosystem services. However, fishing communities often fear short-term income losses associated with closures, and thus may oppose marine reserves. Here we review empirical data and develop bioeconomic models to show that the value of marine reserves (enhanced adjacent fishing + tourism) may often exceed the pre-reserve value, and that economic benefits can offset the costs in as little as five years. These results suggest the need for a new business model for creating and managing reserves, which could pay for themselves and turn a profit for stakeholder groups. Our model could be expanded to include ecosystem services and other benefits, and it provides a general framework to estimate costs and benefits of reserves and to develop such business models.     

Density dependence and the economic efficacy of marine reserves

Predictions on the efficacy of marine reserves for benefiting fisheries differ in large part due to considerations of models of either intra- or inter-cohort population density regulating fish recruitment. Here, I consider both processes acting on recruitment and show using a bioeconomic model how for many fisheries density dependent recruitment dynamics interact with harvest costs to influence fishery profit with reserves. Reserves consolidate fishing effort, favoring fisheries that can profitably harvest low-density stocks of species where adult density mediates recruitment. Conversely, proportion coastline in reserves that maximizes profit, and relative improvement in profit from reserves over conventional management, decline with increasing harvest costs and the relative importance of intra-cohort density dependence. Reserves never increase profit when harvest cost is high, regardless of density dependent recruitment dynamics. I quantitatively synthesize diverse results in the literature, show disproportionate effects on the economic performance of reserves from considering only inter- or intra-cohort density dependence, and highlight fish population and fishery dynamics predicted to be complementary to reserve management. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The Role of Pre-Existing Disturbances in the Effect of Marine Reserves on Coastal Ecosystems: A Modelling Approach

We have used an end-to-end ecosystem model to explore responses over 30 years to coastal no-take reserves covering up to 6% of the fifty thousand square kilometres of continental shelf and slope off the coast of New South Wales (Australia). The model is based on the Atlantis framework, which includes a deterministic, spatially resolved three-dimensional biophysical model that tracks nutrient flows through key biological groups, as well as extraction by a range of fisheries. The model results support previous empirical studies in finding clear benefits of reserves to top predators such as sharks and rays throughout the region, while also showing how many of their major prey groups (including commercial species) experienced significant declines. It was found that the net impact of marine reserves was dependent on the pre-existing levels of disturbance (i.e. fishing pressure), and to a lesser extent on the size of the marine reserves. The high fishing scenario resulted in a strongly perturbed system, where the introduction of marine reserves had clear and mostly direct effects on biomass and functional biodiversity. However, under the lower fishing pressure scenario, the introduction of marine reserves caused both direct positive effects, mainly on shark groups, and indirect negative effects through trophic cascades. Our study illustrates the need to carefully align the design and implementation of marine reserves with policy and management objectives. Trade-offs may exist not only between fisheries and conservation objectives, but also among conservation objectives.     

Spillover of fish naïveté from marine reserves

Spillover of adult fish biomass is an expected benefit from no‐take marine reserves to adjacent fisheries. Here, we show fisher‐naïve behaviour in reef fishes also spills over from marine reserves, potentially increasing access to fishery benefits by making fishes more susceptible to spearguns. The distance at which two targeted families of fishes began to flee a potential fisher [flight initiation distance (FID)] was lower inside reserves than in fished areas, and this reduction extended outside reserve boundaries. Reduced FID persisted further outside reserves than increases in fish biomass. This finding could help increase stakeholder support for marine reserves and improve current models of spillover by informing estimates for spatial changes in catchability. Behavioural changes of fish could help explain differences between underwater visual census and catch data in quantifying the spatial extent of spillover from marine reserves, and should be considered in the management of adjacent fisheries.     

Lack of Cross-Scale Linkages Reduces Robustness of Community-Based Fisheries Management

Community-based management and the establishment of marine reserves have been advocated worldwide as means to overcome overexploitation of fisheries. Yet, researchers and managers are divided regarding the effectiveness of these measures. The “tragedy of the commons” model is often accepted as a universal paradigm, which assumes that unless managed by the State or privatized, common-pool resources are inevitably overexploited due to conflicts between the self-interest of individuals and the goals of a group as a whole. Under this paradigm, the emergence and maintenance of effective community-based efforts that include cooperative risky decisions as the establishment of marine reserves could not occur. In this paper, we question these assumptions and show that outcomes of commons dilemmas can be complex and scale-dependent. We studied the evolution and effectiveness of a community-based management effort to establish, monitor, and enforce a marine reserve network in the Gulf of California, Mexico. Our findings build on social and ecological research before (1997–2001), during (2002) and after (2003–2004) the establishment of marine reserves, which included participant observation in >100 fishing trips and meetings, interviews, as well as fishery dependent and independent monitoring. We found that locally crafted and enforced harvesting rules led to a rapid increase in resource abundance. Nevertheless, news about this increase spread quickly at a regional scale, resulting in poaching from outsiders and a subsequent rapid cascading effect on fishing resources and locally-designed rule compliance. We show that cooperation for management of common-pool fisheries, in which marine reserves form a core component of the system, can emerge, evolve rapidly, and be effective at a local scale even in recently organized fisheries. Stakeholder participation in monitoring, where there is a rapid feedback of the systems response, can play a key role in reinforcing cooperation. However, without cross-scale linkages with higher levels of governance, increase of local fishery stocks may attract outsiders who, if not restricted, will overharvest and threaten local governance. Fishers and fishing communities require incentives to maintain their management efforts. Rewarding local effective management with formal cross-scale governance recognition and support can generate these incentives.     

Ecological guidelines for designing networks of marine reserves in the unique biophysical environment of the Gulf of California

No-take marine reserves can be powerful management tools, but only if they are well designed and effectively managed. We review how ecological guidelines for improving marine reserve design can be adapted based on an area’s unique evolutionary, oceanic, and ecological characteristics in the Gulf of California, Mexico. We provide ecological guidelines to maximize benefits for fisheries management, biodiversity conservation and climate change adaptation. These guidelines include: representing 30% of each major habitat (and multiple examples of each) in marine reserves within each of three biogeographic subregions; protecting critical areas in the life cycle of focal species (spawning and nursery areas) and sites with unique biodiversity; and establishing reserves in areas where local threats can be managed effectively. Given that strong, asymmetric oceanic currents reverse direction twice a year, to maximize connectivity on an ecological time scale, reserves should be spaced less than 50–200 km apart depending on the planktonic larval duration of target species; and reserves should be located upstream of fishing sites, taking the reproductive timing of focal species in consideration. Reserves should be established for the long term, preferably permanently, since full recovery of all fisheries species is likely to take?>?25 years. Reserve size should be based on movement patterns of focal species, although marine reserves?>?10 km long are likely to protect?~?80% of fish species. Since climate change will affect species’ geographic range, larval duration, growth, reproduction, abundance, and distribution of key recruitment habitats, these guidelines may require further modifications to maintain ecosystem function in the future.     

Marine reserve effects on fishery profits: a comment on White et al. (2008)

A recent study ( White et al. 2008 ) claimed that fishery profits will often be higher with management that employs no‐take marine reserves than conventional fisheries management alone. However, this conclusion was based on the erroneous assumption that all landed fish have equal value regardless of size, and questionable assumptions regarding density‐dependence. Examination of an age‐structured version of the White et al. (2008) model demonstrates that their results are not robust to these assumptions. Models with more realistic assumptions generally do not indicate increased fishery yield or profits from marine reserves except for overfished stocks.     

On the Optimal Size of Marine Reserves

The excessive and unsustainable exploitation of our marine resources has led to the promotion of marine reserves as a fisheries management tool. Marine reserves, areas in which fishing is restricted or prohibited, can offer opportunities for the recovery of exploited stock and fishery enhancement. This study examines the impact of the creation of marine protected areas, from both economic and biological perspectives. The consequences of reserve establishment on the long-run equilibrium fish biomass and fishery catch levels are evaluated. We include reserve size as control variable to maximize catch at equilibrium. A continuous time model is used to simulate the effects of reserve size on fishing catch. Fish movements between the sites is assumed to take place at a faster time scale than the variation of the stock and the change of the fleet size. We take advantage of these two time scales to derive a reduced model governing the dynamics of the total fish stock and the fishing effort. Simulation results suggest that the establishment of a protected marine reserve will always lead to an increase in total fish biomass, an optimal size of a marine reserve can achieve to maximize the catch at equilibrium.     

On the fraction of habitat allocated to marine reserves

The case for marine reserves is strengthening, and both deterministic and stochastic calculations show that fisheries management using reserves may achieve harvests comparable with management without reserves. Thus, depending upon the metric used, reserves need not disadvantage harvest. Reserves provide a buffer that increases the chances of sustainability of the stock, and thus the fishery. In this paper, I develop methods (deterministic and stochastic) that allow one to determine how much habitat needs to be set aside as reserve, once societal decisions concerning the goals of reserves are made. The answer to the question: "how much habitat needs to be allocated to reserves" is not a simple single number. Rather, it is a procedure that can be employed once biological, operational and social information are provided. The methods also apply to reserves used to aid stock recovery.     

Large recovery of fish biomass in a no-take marine reserve

No-take marine reserves are effective management tools used to restore fish biomass and community structure in areas depleted by overfishing. Cabo Pulmo National Park (CPNP) was created in 1995 and is the only well enforced no-take area in the Gulf of California, Mexico, mostly because of widespread support from the local community. In 1999, four years after the establishment of the reserve, there were no significant differences in fish biomass between CPNP (0.75 t ha(-1) on average) and other marine protected areas or open access areas in the Gulf of California. By 2009, total fish biomass at CPNP had increased to 4.24 t ha(-1) (absolute biomass increase of 3.49 t ha(-1), or 463%), and the biomass of top predators and carnivores increased by 11 and 4 times, respectively. However, fish biomass did not change significantly in other marine protected areas or open access areas over the same time period. The absolute increase in fish biomass at CPNP within a decade is the largest measured in a marine reserve worldwide, and it is likely due to a combination of social (strong community leadership, social cohesion, effective enforcement) and ecological factors. The recovery of fish biomass inside CPNP has resulted in significant economic benefits, indicating that community-managed marine reserves are a viable solution to unsustainable coastal development and fisheries collapse in the Gulf of California and elsewhere.     

Effects of marine reserve characteristics on the protection of fish populations: a meta-analysis

Meta-analyses of published data for 19 marine reserves reveal that marine protected areas enhance species richness consistently, but their effect on fish abundance is more variable. Overall, there was a slight (11%) but significant increase in fish species number inside marine reserves, with all reserves sharing a common effect. There was a substantial but non-significant increase in overall fish abundance inside marine reserves compared to adjacent, non-reserve areas. When only species that are the target of fisheries were considered, fish abundance was significantly higher (by 28%) within reserve boundaries. Marine reserves vary significantly in the extent and direction of their response. This variability in relative abundance was not attributable to differences in survey methodology among studies, nor correlated with reserve characteristics such as reserve area, years since protection, latitude nor species diversity. The effectiveness of marine reserves in enhancing fish abundance may be largely related to the intensity of exploitation outside reserve boundaries and to the composition of the fish community within boundaries. It is recommended that studies of marine reserve effectiveness should routinely report fishing intensity, effectiveness of enforcement and habitat characteristics.     

Rapid increase in fish numbers follows creation of world's largest marine reserve network

No-take marine reserves (NTMRs) are much advocated as a solution to managing marine ecosystems, protecting exploited species and restoring natural states of biodiversity [1,2]. Increasingly, it is becoming clear that effective marine conservation and management at ecosystem and regional scales requires extensive networks of NTMRs [1,2]. The world's largest network of such reserves was established on Australia's Great Barrier Reef (GBR) in 2004. Closing such a large area to all fishing has been socially and politically controversial, making it imperative that the effectiveness of this new reserve network be assessed. Here we report evidence, first, that the densities of the major target species of the GBR reef line fisheries were significantly higher in the new NTMRs, compared with fished sites, in just two years; and second, that the positive differences were consistent for multiple marine reserves over an unprecedented spatial scale (>1,000 km).     

Concepts and issues in marine ecosystem management

Ecosystem management means different things to different people, but the underlying concept is similar to that of the long-standing ethic of conservation. Current interest in marine ecosystem management stems from concerns about overexploitation of world fisheries and the perceived need for broader perspectives in fisheries management. A central scientific question is whether the effects of harvesting (top down) or changes in the physical environment (bottom up) are responsible for major changes in abundance.Historically, ecology, fisheries biology, oceanography, fisheries management and the fishing industry have gone somewhat separate ways. Since the 1980s, increasing attention has been given to multispecies aspects of fisheries, the linkages between oceanography and fish abundance and more holistic approaches to fisheries management.Sorting out the causes and effects of fluctuations in fish abundance is complicated by the lack of reliability of fisheries statistics. Discards, dishonesty and the inherent logistic difficulties of collecting statistics all combine to confuse interpretation. The overcapacity of fishing fleets and their unrestricted use are widely recognized as a contributing cause to overfishing and declines in fish stocks in many parts of the world.Ecosystem management, as shorthand for more holistic approaches to resource management, is, from a fisheries management perspective, centred on multispecies interactions in the context of a variable physical and chemical environment. Broader perspectives include social, economic and political elements which are best considered pragmatically as a part of the context of fisheries management.Objectives in marine ecosystem management are varied. From a biological perspective, an underlying principle of management is commonly assumed to be a sustained yield of products for human consumption. Whether that should be taken to mean that the yield should always be of the same products is less certain. Fishing commonly changes the relative abundance of species of fishes. Thus, a biological objective should specify the species mix that is desired.Concern for the maintenance of global diversity has generated a substantial literature on threatened and endangered species. In general, it has not been considered likely that marine fish species could be rendered extinct and greatest attention has been given to marine mammals, sea birds and sea turtles. The provision of marine parks and sanctuary areas are obvious first steps in providing a measure of protection, at least for the less widely ranging species.Related to the current concepts of ecosystem management are expressions such as ecosystem health and ecosystem integrity which are given a wide range of different meanings, none of which are readily translated into operational language for resource management. These and similar expressions are best assessed as rhetorical devices. The essential components of ecosystem management are sustainable yield, maintenance of biodiversity and protection from the effects of pollution and habitat degradation.Theory for marine ecosystem management has a long history in fisheries and ecological literature. Ecological models such as Lotka-Volterra equations, ECOPATH, trophic cascades and chaos theory do not give practical guidance for management. Fleet interaction and multispecies virtual population analysis models hold more promise for fisheries managers.Alaska provides particular opportunities for developing new concepts in fisheries management. Statistics of catch are good, stock assessments are at the state-of-the-art level and management has been prudent. Debate is active on the causes of substantial changes in abundance of many species including marine mammals, because substantial changes in the fisheries have been accompanied by major changes in oceanographic conditions.As elsewhere, the resultant changes may be a consequence of top-down and bottom-up effects. The bottom part is beyond human control, and ecosystem management is centred on managing the top-down or fisheries component in the context of special measures of protection for particular species.Whether that is a realistic goal depends in part on how much special protection is to be afforded to which species. Marine mammals, for example, are given high priority for special protection, but like fisheries they too may have significant roles in shaping the structure of marine ecosystems. Eventually, ecosystem management must come to grips with the question of how much protection of particular species is desirable in achieving optimal use of living marine resources.     

Climate impacts and oceanic top predators: moving from impacts to adaptation in oceanic systems

Climate impacts are now widely reported from coastal marine systems, but less is known for the open ocean. Here we review progress in understanding impacts on large pelagic species presented at an international workshop for the Climate Impacts on Oceanic Top Predators programme, and discuss the future with regard to the next phase of adaptation-focused research. Recent highlights include a plan to map the distribution of key species in the foodweb using both acoustics and biochemical techniques, and development of a new data sharing and access tool for fisheries and associated data, including socio-economic information. A common research focus in pelagic ecosystems is on understanding climate variability and climate change impacts on marine species, but a greater emphasis on developing future scenarios and adaptation options is needed. Workshop participants also concluded that engagement with and provision of science support to regional fisheries management organisations are critical elements for ensuring successful uptake of research. This uptake will be required for future management of fisheries as global warming continues such that some open ocean top predators can be sustainably harvested, impacts on conservation-dependent species can be avoided, and ecosystem function is not compromised.     

Ocean zoning within a sparing versus sharing framework

The land-sparing versus land-sharing debate centers around how different intensities of habitat use can be coordinated to satisfy competing demands for biodiversity persistence and food production in agricultural landscapes. We apply the broad concepts from this debate to the sea and propose it as a framework to inform marine zoning based on three possible management strategies, establishing: no-take marine reserves, regulated fishing zones, and unregulated open-access areas. We develop a general model that maximizes standing fish biomass, given a fixed management budget while maintaining a minimum harvest level. We find that when management budgets are small, sea-sparing is the optimal management strategy because for all parameters tested, reserves are more cost-effective at increasing standing biomass than traditional fisheries management. For larger budgets, the optimal strategy switches to sea-sharing because, at a certain point, further investing to grow the no-take marine reserves reduces catch below the minimum harvest constraint. Our intention is to illustrate how general rules of thumb derived from plausible, single-purpose models can help guide marine protected area policy under our novel sparing and sharing framework. This work is the beginning of a basic theory for optimal zoning allocations and should be considered complementary to the more specific spatial planning literature for marine reserve as nations expand their marine protected area estates.