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.     

Catastrophes,connectivity and Allee effects in the design of marine reserve networks

Catastrophic events, like oil spills and hurricanes, occur in many marine systems. One potential role of marine reserves is buffering populations against disturbances, including the potential for disturbance-driven population collapses under Allee effects. This buffering capacity depends on reserves in a network providing rescue effects, setting up a tradeoff where reserves need to be connected to facilitate rescue, but also distributed in space to prevent simultaneous extinction. We use a set of population models to examine how dispersal ability and the disturbance regime interact to determine the optimal reserve spacing. We incorporate fishing in a spatially-explicit model to understand the effect of objective choice (e.g. conservation versus fisheries yield) on the optimal reserve spacing. We show that the optimal spacing between reserves increases when accounting for catastrophes with larger spacing needed when Allee effects interact with catastrophes to increase the probability of extinction. We also show that classic tradeoffs between conservation and fishing objectives disappear in the presence of catastrophes. Specifically, we found that at intermediate levels of disturbance, it is optimal to spread out reserves in order to increase both population persistence and to maximize spillover into non-reserve areas.     

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.     

Spillover from marine reserves related to mechanisms of population regulation

Spillover of fish from marine reserves to adjacent harvested waters may be mediated by density-independent movement, density-dependent movement, or both. If dispersal is by random movement, populations within the reserve must be regulated by density-dependent population growth (DDG). Density-dependent movement (DDM) can also regulate the population if accelerated emigration from a reserve to the surrounding fishing grounds leads to substantially increased mortality. Using spatially explicit models, we show that stock per unit area is bounded for DDG and increases with size for DDM. With DDG, spillover rate per unit area of reserve is maximized with reserves around 50% larger in linear dimension than the minimum size for population persistence. With DDM, spillover per unit area of reserve increases with reserve size. The results highlight the need for the mechanism of population regulation to be incorporated into theoretical and empirical investigations of marine reserve ecology.     

Fiji’s largest marine reserve benefits reef sharks

To provide more information about whether sharks benefit from no-take marine reserves, we quantified the relative abundance and biomass of reef sharks inside and outside of Namena, Fiji’s largest reserve (60.6 km²). Using stereo baited remote underwater video systems (stereo-BRUVs), we found that the abundance and biomass of sharks was approximately two and four times greater in shallow and deep locations, respectively, within the Namena reserve compared to adjacent fished areas. The greater abundance and biomass of reef sharks inside Namena is likely a result of greater prey availability rather than protection from fishing. This study demonstrates that marine reserves can benefit sharks.     

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.     

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.     

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.     

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.     

Spatial population dynamics and the design of marine reserves

The failure of many fisheries world-wide, and the concern about marine biodiversity, has sparked a growing interest in the spatial aspects of harvested populations. If a population conforms to the Ideal Free Distribution and that one of the habitats is set aside as a reserve free from harvesting, the design of reserves may be problematic. If a substantial proportion of the unharvested population is to be preserved, then the reserve area must be unrealistically large, or have a much higher expected fitness than the unprotected area. Interestingly, the optimal harvest rate will be unaffected by both the size of the reserve and the quality of it relative to the harvested area. Even if the Ideal Free Distribution model is extended to include simple age-structure and "spillover" of recruits from the reserve, these conclusions largely remain intact. In a model that also includes spillover, the habitat quality of the reserve may also affect the catch.     

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.     

Reef sharks exhibit site-fidelity and higher relative abundance in marine reserves on the Mesoamerican Barrier Reef

Carcharhinid sharks can make up a large fraction of the top predators inhabiting tropical marine ecosystems and have declined in many regions due to intense fishing pressure. There is some support for the hypothesis that carcharhinid species that complete their life-cycle within coral reef ecosystems, hereafter referred to as “reef sharks”, are more abundant inside no-take marine reserves due to a reduction in fishing pressure (i.e., they benefit from marine reserves). Key predictions of this hypothesis are that (a) individual reef sharks exhibit high site-fidelity to these protected areas and (b) their relative abundance will generally be higher in these areas compared to fished reefs. To test this hypothesis for the first time in Caribbean coral reef ecosystems we combined acoustic monitoring and baited remote underwater video (BRUV) surveys to measure reef shark site-fidelity and relative abundance, respectively. We focused on the Caribbean reef shark (Carcharhinus perezi), the most common reef shark in the Western Atlantic, at Glover''s Reef Marine Reserve (GRMR), Belize. Acoustically tagged sharks (N = 34) were detected throughout the year at this location and exhibited strong site-fidelity. Shark presence or absence on 200 BRUVs deployed at GRMR and three other sites (another reserve site and two fished reefs) showed that the factor “marine reserve” had a significant positive effect on reef shark presence. We rejected environmental factors or site-environment interactions as predominant drivers of this pattern. These results are consistent with the hypothesis that marine reserves can benefit reef shark populations and we suggest new hypotheses to determine the underlying mechanism(s) involved: reduced fishing mortality or enhanced prey availability.     

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.     

53 A Seaweed's perspective on marine reserves

In response to major changes in coastal ecosystems in recent decades, a number of governmental agencies around the world are establishing marine reserves – areas where removal of animals or plants is prohibited. Although marine reserves are touted as an ecosystem based approach to management of marine resources, the vast majority of attention on reserve design and impact focuses solely on fish. Although a few species of algae are commercially harvested, most are not. As a result, they will receive little direct benefit from protection by reserves aside from habitat protection. From the perspective of a seaweed, the primary impacts of marine reserves will therefore be indirect through species interactions. We examine the rapidly growing theoretical and empirical literature on marine reserves to anticipate the likely responses of seaweeds to exclusion of fishing. The key issues that emerge are: the trophic level of prior fishing and the dispersal scales of seaweeds relative to their competitors and consumers. The latter issue is poorly understood and poses a key challenge to phycologists if we are to effectively incorporate seaweeds into future marine reserve design.     

Increased Disease Calls for a Cost-Benefits Review of Marine Reserves

Marine reserves (or No-Take Zones) are implemented to protect species and habitats, with the aim of restoring a balanced ecosystem. Although the benefits of marine reserves are commonly monitored, there is a lack of insight into the potential detriments of such highly protected waters. High population densities attained within reserves may induce negative impacts such as unfavourable trophic cascades and disease outbreaks. Hence, we investigated the health of lobster populations in the UK’s Marine Conservation Zone (MCZ) at Lundy Island. Comparisons were made between the fished, Refuge Zone (RZ) and the un-fished, No-Take Zone (NTZ; marine reserve). We show ostensibly positive effects such as increased lobster abundance and size within the NTZ; however, we also demonstrate apparent negative effects such as increased injury and shell disease. Our findings suggest that robust cost-benefit analyses of marine reserves could improve marine reserve efficacy and subsequent management strategies.     

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.     

Potential of a no‐take marine reserve to protect home ranges of anadromous brown trout (Salmo trutta)

The extent to which no‐take marine reserves can benefit anadromous species requires examination. Here, we used acoustic telemetry to investigate the spatial behavior of anadromous brown trout (sea trout, Salmo trutta) in relation to a small marine reserve (~1.5 km²) located inside a fjord on the Norwegian Skagerrak coast. On average, sea trout spent 42.3 % (±5.0% SE) of their time in the fjord within the reserve, a proportion similar to the area of the reserve relative to that of the fjord. On average, sea trout tagged inside the reserve received the most protection, although the level of protection decreased marginally with increasing home range size. Furthermore, individuals tagged outside the reserve received more protection with increasing home range size, potentially opposing selection toward smaller home range sizes inflicted on fish residing within reserves, or through selective fishing methods like angling. Monthly sea trout home ranges in the marine environment were on average smaller than the reserve, with a mean of 0.430 (±0.0265 SE) km². Hence, the reserve is large enough to protect the full home range of some individuals residing in the reserve. Synthesis and applications: In general, the reserve protects sea trout to a varying degree depending on their individual behavior. These findings highlight evolutionary implications of spatial protection and can guide managers in the design of marine reserves and networks that preserve variation in target species' home range size and movement behavior.     

Very Slow Search and Reach: Failure to Maximize Expected Gain in an Eye-Hand Coordination Task

We examined an eye-hand coordination task where optimal visual search and hand movement strategies were inter-related. Observers were asked to find and touch a target among five distractors on a touch screen. Their reward for touching the target was reduced by an amount proportional to how long they took to locate and reach to it. Coordinating the eye and the hand appropriately would markedly reduce the search-reach time. Using statistical decision theory we derived the sequence of interrelated eye and hand movements that would maximize expected gain and we predicted how hand movements should change as the eye gathered further information about target location. We recorded human observers'' eye movements and hand movements and compared them with the optimal strategy that would have maximized expected gain. We found that most observers failed to adopt the optimal search-reach strategy. We analyze and describe the strategies they did adopt.     

How Well Will Brazil's System of Atlantic Forest Reserves Maintain Viable Bird Populations?

It is crucial for biodiversity conservation that protected areas are large and effective enough to support viable populations of their original species. We used a point count distance sampling method to estimate population sizes of a range of bird species in three Atlantic forest protected areas of size 5600, 22,500, and 46,050 ha. Population sizes were generally related to reserve area, although in the mid-sized reserve, there were many rare species reflecting a high degree of habitat heterogeneity. The proportions of forest species having estimated populations >500 ranged from 55% of 210 species in the largest reserve to just 25% of 140 species in the smallest reserve. All forest species in the largest reserves had expected populations >100, but in the small reserve, 28% (38 species) had populations <100 individuals. Atlantic forest endemics were no more or less likely to have small populations than widespread species. There are 79 reserves (>1000 ha) in the Atlantic forest lowlands. However, all but three reserves in the north of the region (Espírito Santo and states north) are smaller than 10,000 ha, and we predict serious levels of local extinction from these reserves. Habitat heterogeneity within reserves may promote species richness within them, but it may also be important in determining species loss over time by suppressing populations of individual species. We suggest that most reserves in the region are so small that homogeneity in the habitat/altitude within them is beneficial for maintenance of their (comparatively small) original species compliment. A lack of protection in the north, continued detrimental human activity inside reserves, and our poor knowledge of how well the reserve system protects individual taxa, are crucial considerations in biodiversity management in the region.