Using marine reserves to estimate fishing mortality

The proportion of a fish stock that is killed by fishing activity is often calculated as the catch divided by the estimated stock biomass. However, stock biomass is notoriously difficult to estimate reliably, and moreover, the catch may be uncertain or misreported and does not include losses due to discarding. In all too many fisheries, these difficulties have lead to underestimates of total fishing mortality and the commercial demise of the fishery. No‐take marine reserves eliminate fishing mortality from within their boundaries and, for species that exhibit seasonal migratory behaviour, comparison of reserves with fished areas can provide direct estimates of the proportion killed by fishing. For an important exploited species in New Zealand, seasonal changes in density of sub‐legal fish at three marine reserves were similar in both reserve and adjacent non‐reserve areas. However, this result did not hold for legal‐size fish, and the difference in seasonal change between reserved and non‐reserved areas was used to obtain direct estimates of the total localized fishing mortality in the non‐reserve area over 6‐month periods. Estimates of the percentage of legal‐size fish killed by fishing ranged from 70 to 96%. These results demonstrate an unanticipated practical benefit from marine reserves that goes beyond their ecological role.     

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.     

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.     

Marine reserves with ecological uncertainty

To help manage the fluctuations inherent in fish populations scientists have argued for both an ecosystem approach to management and the greater use of marine reserves. Support for reserves includes empirical evidence that they can raise the spawning biomass and mean size of exploited populations, increase the abundance of species and, relative to reference sites, raise population density, biomass, fish size and diversity. By contrast, fishers often oppose the establishment and expansion of marine reserves and claim that reserves provide few, if any, economic payoffs. Using a stochastic optimal control model with two forms of ecological uncertainty we demonstrate that reserves create a resilience effect that allows for the population to recover faster, and can also raise the harvest immediately following a negative shock. The tradeoff of a larger reserve is a reduced harvest in the absence of a negative shock such that a reserve will never encompass the entire population if the goal is to maximize the economic returns from harvesting, and fishing is profitable. Under a wide range of parameter values with ecological uncertainty, and in the ‘worst case’ scenario for a reserve, we show that a marine reserve can increase the economic payoff to fishers even when the harvested population is not initially overexploited, harvesting is economically optimal and the population is persistent. Moreover, we show that the benefits of a reserve cannot be achieved by existing effort or output controls. Our results demonstrate that, in many cases, there is no tradeoff between the economic payoff of fishers and ecological benefits when a reserve is established at equal to, or less than, its optimum size.     

Effects of habitat,wave exposure,and marine protected area status on coral reef fish assemblages in the Hawaiian archipelago

The relationships between fish assemblages, their associated habitat, and degree of protection from fishing were evaluated over a broad spatial scale throughout the main Hawaiian islands. Most fish assemblage characteristics showed positive responses to protection whether it was physical (e.g. habitat complexity), biological (e.g. coral cover growth forms), or human-induced (e.g. marine reserves). Fish biomass was lowest in areas of direct wave exposure and highest in areas partially sheltered from swells. Higher values for fish species richness, number of individuals, biomass, and diversity were observed in locations with higher substrate complexity. Areas completely protected from fishing had distinct fish assemblages with higher standing stock and diversity than areas where fishing was permitted or areas that were partially protected from fishing. Locations influenced by customary stewardship harbored fish biomass that was equal to or greater than that of no-take protected areas. Marine protected areas in the main Hawaiian islands with high habitat complexity, moderate wave disturbance, a high percentage of branching and/or lobate coral coupled with legal protection from fishing pressure had higher values for most fish assemblage characteristics.     

Spillover Effects of a Community-Managed Marine Reserve

The value of no-take marine reserves as fisheries-management tools is controversial, particularly in high-poverty areas where human populations depend heavily on fish as a source of protein. Spillover, the net export of adult fish, is one mechanism by which no-take marine reserves may have a positive influence on adjacent fisheries. Spillover can contribute to poverty alleviation, although its effect is modulated by the number of fishermen and fishing intensity. In this study, we quantify the effects of a community-managed marine reserve in a high poverty area of Northern Mozambique. For this purpose, underwater visual censuses of reef fish were undertaken at three different times: 3 years before (2003), at the time of establishment (2006) and 6 years after the marine reserve establishment (2012). The survey locations were chosen inside, outside and on the border of the marine reserve. Benthic cover composition was quantified at the same sites in 2006 and 2012. After the reserve establishment, fish sizes were also estimated. Regression tree models show that the distance from the border and the time after reserve establishment were the variables with the strongest effect on fish abundance. The extent and direction of the spillover depends on trophic group and fish size. Poisson Generalized Linear Models show that, prior to the reserve establishment, the survey sites did not differ but, after 6 years, the abundance of all fish inside the reserve has increased and caused spillover of herbivorous fish. Spillover was detected 1km beyond the limit of the reserve for small herbivorous fishes. Six years after the establishment of a community-managed reserve, the fish assemblages have changed dramatically inside the reserve, and spillover is benefitting fish assemblages outside the reserve.     

Effect of Small Versus Large Clusters of Fish School on the Yield of a Purse-Seine Small Pelagic Fishery Including a Marine Protected Area

We consider a fishery model with two sites: (1) a marine protected area (MPA) where fishing is prohibited and (2) an area where the fish population is harvested. We assume that fish can migrate from MPA to fishing area at a very fast time scale and fish spatial organisation can change from small to large clusters of school at a fast time scale. The growth of the fish population and the catch are assumed to occur at a slow time scale. The complete model is a system of five ordinary differential equations with three time scales. We take advantage of the time scales using aggregation of variables methods to derive a reduced model governing the total fish density and fishing effort at the slow time scale. We analyze this aggregated model and show that under some conditions, there exists an equilibrium corresponding to a sustainable fishery. Our results suggest that in small pelagic fisheries the yield is maximum for a fish population distributed among both small and large clusters of school.     

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.     

Changes in Fish Assemblages following the Establishment of a Network of No-Take Marine Reserves and Partially-Protected Areas

Networks of no-take marine reserves and partially-protected areas (with limited fishing) are being increasingly promoted as a means of conserving biodiversity. We examined changes in fish assemblages across a network of marine reserves and two different types of partially-protected areas within a marine park over the first 5 years of its establishment. We used Baited Remote Underwater Video (BRUV) to quantify fish communities on rocky reefs at 20–40 m depth between 2008–2011. Each year, we sampled 12 sites in 6 no-take marine reserves and 12 sites in two types of partially-protected areas with contrasting levels of protection (n = 4 BRUV stations per site). Fish abundances were 38% greater across the network of marine reserves compared to the partially-protected areas, although not all individual reserves performed equally. Compliance actions were positively associated with marine reserve responses, while reserve size had no apparent relationship with reserve performance after 5 years. The richness and abundance of fishes did not consistently differ between the two types of partially-protected areas. There was, therefore, no evidence that the more regulated partially-protected areas had additional conservation benefits for reef fish assemblages. Overall, our results demonstrate conservation benefits to fish assemblages from a newly established network of temperate marine reserves. They also show that ecological monitoring can contribute to adaptive management of newly established marine reserve networks, but the extent of this contribution is limited by the rate of change in marine communities in response to protection.     

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.     

Effects of marine reserves versus nursery habitat availability on structure of reef fish communities

No-take marine fishery reserves sustain commercial stocks by acting as buffers against overexploitation and enhancing fishery catches in adjacent areas through spillover. Likewise, nursery habitats such as mangroves enhance populations of some species in adjacent habitats. However, there is lack of understanding of the magnitude of stock enhancement and the effects on community structure when both protection from fishing and access to nurseries concurrently act as drivers of fish population dynamics. In this study we test the separate as well as interactive effects of marine reserves and nursery habitat proximity on structure and abundance of coral reef fish communities. Reserves had no effect on fish community composition, while proximity to nursery habitat only had a significant effect on community structure of species that use mangroves or seagrass beds as nurseries. In terms of reef fish biomass, proximity to nursery habitat by far outweighed (biomass 249% higher than that in areas with no nursery access) the effects of protection from fishing in reserves (biomass 21% lower than non-reserve areas) for small nursery fish (≤ 25 cm total length). For large-bodied individuals of nursery species (>25 cm total length), an additive effect was present for these two factors, although fish benefited more from fishing protection (203% higher biomass) than from proximity to nurseries (139% higher). The magnitude of elevated biomass for small fish on coral reefs due to proximity to nurseries was such that nursery habitats seem able to overrule the usually positive effects on fish biomass by reef reserves. As a result, conservation of nursery habitats gains importance and more consideration should be given to the ecological processes that occur along nursery-reef boundaries that connect neighboring ecosystems.     

Evidence of artisanal fishing impacts and depth refuge in assemblages of Fijian reef fish

Protection from fishing generally results in an increase in the abundance and biomass of species targeted by fisheries within marine reserve boundaries. Natural refuges such as depth may also protect such species, yet few studies in the Indo Pacific have investigated the effects of depth concomitant with marine reserves. We studied the effects of artisanal fishing and depth on reef fish assemblages in the Kubulau District of Vanua Levu Island, Fiji, using baited remote underwater stereo-video systems. Video samples were collected from shallow (5–8 m) and deep (25–30 m) sites inside and outside of a large old marine reserve (60.6 km², 13 years old) and a small new marine reserve (4.25 km², 4 years old). Species richness tended to be greater in the shallow waters of the large old reserve when compared to fished areas. In the deeper waters, species richness appeared to be comparable. The difference in shallow waters was driven by species targeted by fisheries, indicative of a depth refuge effect. In contrast, differences in the abundance composition of the fish assemblage existed between protected and fished areas for deep sites, but not shallow. Fish species targeted by local fisheries were 89% more abundant inside the large old reserve than surrounding fished areas, while non-targeted species were comparable. We observed no difference in the species richness or abundance of species targeted by fisheries inside and outside of the small new reserve. This study suggests that artisanal fishing impacts on the abundance and species richness of coral reef fish assemblages and effects of protection are more apparent with large reserves that have been established for a long period of time. Observed effects of protection also vary with depth, highlighting the importance of explicitly incorporating multiple depth strata in studies of marine reserves.     

An ecological perspective on marine reserves in prey–predator dynamics

This paper describes a prey–predator type fishery model with prey dispersal in a two-patch environment, one of which is a free fishing zone and other is a protected zone. The existence of possible steady states, along with their local stability, is discussed. A geometric approach is used to derive the sufficient conditions for global stability of the system at the positive equilibrium. Relative size of the reserve is considered as control in order to study optimal sustainable yield policy. Subsequently, the optimal system is derived and then solved numerically using an iterative method with Runge–Kutta fourth-order scheme. Numerical simulations are carried out to illustrate the importance of marine reserve in fisheries management. It is noted that the marine protected area enables us to protect and restore multi-species ecosystem. The results illustrate that dynamics of the system is extremely interesting if simultaneous effects of a regulatory mechanism like marine reserve is coupled with harvesting effort. It is observed that the migration of the resource, from protected area to unprotected area and vice versa, is playing an important role towards the standing stock assessment in both the areas which ultimately control the harvesting efficiency and enhance the fishing stock up to some extent.     

Spatial and temporal variation of abundance,biomass and diversity within marine reserves in the Philippines

Aim The objective of this study was to investigate the influence of protection duration (years of fishing closure) and location (distance from shore) on reef fish diversity. Location Danajon Double Barrier Reef, Bohol, Philippines. Methods Reef fish abundance and size structure, by species, were obtained monthly using replicated underwater visual belt transects (n = 8; 70 × 5‐m belt transects) over 3 years (2002–2005) at eight sites that included six marine reserves and two unprotected reef areas. We analysed species accumulation curves, diversity indices and abundance–biomass comparison (ABC) curves within and across the study sites to assess the influence of protection duration and location. Results Analyses showed that longer protection duration impacted reef fish diversity at both inshore and offshore sites by shifting ABC curves from higher abundance than biomass curves at fished sites to higher biomass than abundance curves at most of the protected sites. Protection duration did not significantly influence either the rate of species accumulation within sites or the 12 diversity indices measured across the study sites. The offshore sites consistently showed higher rates of species accumulation and diversity indices values than inshore sites with similar protection duration. One protected offshore young marine reserve site that has been assessed as the least well‐managed showed patterns more consistent with the fished sites. Main conclusions Analyses showed that protection duration mainly impacted diversity by increasing the dominance of large‐bodied species and enhancing total biomass. Besides protection duration, reserve location influenced species accumulation curves and diversity indices.     

Marine reserves: size and age do matter

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

Trajectories and magnitude of change in coral reef fish populations in Philippine marine reserves: a meta-analysis

Marine reserves are widely implemented worldwide to meet both conservation and fisheries management goals. This study examines the efficacy of Philippine marine reserves using meta-analysis by comparing variations in fish density (1) between reserves and adjacent fished reefs (spatial comparison), (2) within reserves before establishment relative to years following the establishment (temporal comparison), and (3) among reserves classified based on size, age, and enforcement capacity. A grand (total) mean of nineteen 22.3 ha coral reef reserves, protected for a mean duration of 8.2 years, were included in the meta-analyses. The overall density of fishes was higher in the reserves compared with the fished reefs and this difference was largely accounted for by exploited fishes. However, the overall density of fishes within the same reserves remained similar from the period before its establishment to several years following its establishment. Only the density of nonexploited fishes increased significantly during years subsequent to the establishment of the reserves. Neither age nor size of reserves correlated with pattern of change in fish density following the establishment of the reserves; however, fish density was consistently higher in larger and older reserves relative to smaller and younger reserves in the spatial comparison. Furthermore, well-enforced reserves had higher density of exploited fishes relative to less-enforced reserves in both spatial and temporal comparisons. In general, the magnitude and trajectory of change in fish density following the establishment of Philippine marine reserves are influenced by (1) functional groups of fishes under consideration, (2) size and age of the reserve, and (3) level of enforcement of the regulatory mechanisms necessary to sustain a marine reserve.     

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.     

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.     

Marine reserves can enhance ecological resilience

The goals of ecosystem‐based management (EBM) include protecting ecological resilience, the magnitude of a perturbation that a community can withstand and remain in a given state. As a tool to achieve this goal, no‐take marine reserves may enhance resilience by protecting source populations or reduce it by concentrating fishing in harvested areas. Here, we test whether spatial management with marine reserves can increase ecological resilience compared to non‐spatial (conventional) management using a dynamic model of a simplified fish community with structured predation and competition that causes alternative stable states. Relative to non‐spatial management, reserves increase the resilience of the desired (predator‐dominated) equilibrium state in both stochastic and deterministic environments, especially under intensive fishing. As a result, spatial management also increases the feasibility of restoring degraded (competitor‐dominated) systems, particularly if combined with culling of competitors or stock enhancement of adult predators.     

Modelling the Impact of Marine Reserves on a Population with Depensatory Dynamics

In this study, we use a spatially implicit, stage-structured model to evaluate marine reserve effectiveness for a fish population exhibiting depensatory (strong Allee) effects in its dynamics. We examine the stability and sensitivity of the equilibria of the modelled system with regards to key system parameters and find that for a reasonable set of parameters, populations can be protected from a collapse if a small percentage of the total area is set aside in reserves. Furthermore, the overall abundance of the population is predicted to achieve a maximum at a certain ratio \(A\) of reserve area to fished area, which depends heavily on the other system parameters such as the net export rate of fish from the marine reserves to the fished areas. This finding runs contrary to the contested “equivalence at best” result when comparing fishery management through traditional catch or effort control and management through marine reserves. Lastly, we analyse the problem from a bioeconomics perspective by computing the optimal harvesting policy using Pontryagin’s Maximum Principle, which suggests that the value for \(A\) which maximizes the optimal equilibrium fishery yield also maximizes population abundance when the cost per unit harvest is constant, but can increase substantially when the cost per unit harvest increases with the area being harvested.