Marine reserves and optimal harvesting

Advocates of no‐take marine reserves emphasize their conservation benefits. Critics counter that reserves would decrease fisheries yield. Analysis of a spatially explicit harvesting model, however, shows that no‐take marine reserves are always part of an optimal harvest designed to maximize yield. The optimal harvest generates a spatial source–sink structure with source populations placed in reserves. The sizes and locations of the optimal reserves depend on a dimensionless length parameter. For small values of this parameter, the maximum yield is obtained by placing a large reserve in the centre of the habitat. For large values of this parameter, the optimal harvesting strategy is a spatial ‘chattering control’ with infinite sequences of reserves alternating with areas of intense fishing. Such a chattering strategy would be impossible to actually implement, but in these cases an approximate yet practicable policy, utilizing a small number of reserves, can be constructed.     

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.     

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 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.     

Larval dispersal and movement patterns of coral reef fishes,and implications for marine reserve network design

Well‐designed and effectively managed networks of marine reserves can be effective tools for both fisheries management and biodiversity conservation. Connectivity, the demographic linking of local populations through the dispersal of individuals as larvae, juveniles or adults, is a key ecological factor to consider in marine reserve design, since it has important implications for the persistence of metapopulations and their recovery from disturbance. For marine reserves to protect biodiversity and enhance populations of species in fished areas, they must be able to sustain focal species (particularly fishery species) within their boundaries, and be spaced such that they can function as mutually replenishing networks whilst providing recruitment subsidies to fished areas. Thus the configuration (size, spacing and location) of individual reserves within a network should be informed by larval dispersal and movement patterns of the species for which protection is required. In the past, empirical data regarding larval dispersal and movement patterns of adults and juveniles of many tropical marine species have been unavailable or inaccessible to practitioners responsible for marine reserve design. Recent empirical studies using new technologies have also provided fresh insights into movement patterns of many species and redefined our understanding of connectivity among populations through larval dispersal. Our review of movement patterns of 34 families (210 species) of coral reef fishes demonstrates that movement patterns (home ranges, ontogenetic shifts and spawning migrations) vary among and within species, and are influenced by a range of factors (e.g. size, sex, behaviour, density, habitat characteristics, season, tide and time of day). Some species move <0.1–0.5 km (e.g. damselfishes, butterflyfishes and angelfishes), <0.5–3 km (e.g. most parrotfishes, goatfishes and surgeonfishes) or 3–10 km (e.g. large parrotfishes and wrasses), while others move tens to hundreds (e.g. some groupers, emperors, snappers and jacks) or thousands of kilometres (e.g. some sharks and tuna). Larval dispersal distances tend to be <5–15 km, and self‐recruitment is common. Synthesising this information allows us, for the first time, to provide species, specific advice on the size, spacing and location of marine reserves in tropical marine ecosystems to maximise benefits for conservation and fisheries management for a range of taxa. We recommend that: (i) marine reserves should be more than twice the size of the home range of focal species (in all directions), thus marine reserves of various sizes will be required depending on which species require protection, how far they move, and if other effective protection is in place outside reserves; (ii) reserve spacing should be <15 km, with smaller reserves spaced more closely; and (iii) marine reserves should include habitats that are critical to the life history of focal species (e.g. home ranges, nursery grounds, migration corridors and spawning aggregations), and be located to accommodate movement patterns among these. We also provide practical advice for practitioners on how to use this information to design, evaluate and monitor the effectiveness of marine reserve networks within broader ecological, socioeconomic and management contexts.     

Logic for designing nature reserves for multiple species

We examine the logic of designing nature reserves to understand better how to integrate the concepts of representativeness and persistence. Simple models of viability are used to evaluate how the expected number of species in the reserve system changes with variation in the risk of extinction among species, their rate of occurrence, and the distribution of species. The optimal size of individual reserves increased with the mean and variance of the probability of extinction among species and with the rate at which the risk of extinction declines with the cost of each reserve. In contrast, the rate of occurrence of species within reserves and their rate of accumulation with increasing reserve area had a relatively minor influence on the optimal size of reserves. Patterns of endemism were most important for the location of reserves. Including differences among species in the analysis reduced the optimal number of individual reserves (and increased the size of each) when operating under a fixed budget compared with reserve designs based on single species. A case study in the city of Melbourne, Australia, demonstrates the conservation value of small (approximately 1 ha) grassland reserves and the underrepresentation of Melbourne's volcanic plains in the region's conservation network.     

Optimized Floating Refugia: a new strategy for species conservation in production forest landscapes

Timber production forests can support diverse ecological communities, but existing conservation strategies fail to maximize this potential. While methods for limiting logging damage and locating biological reserves have been developed, strategies focused on the sequence and arrangement of harvest units are lacking, particularly for situations in which species-specific knowledge is limited. We present a new landscape-level approach to forest conservation that anticipates local extinctions and focuses on facilitating re-colonization via strategic spatiotemporal harvest plans (which are informed by species occurrence data only). As a proof of concept, we applied our framework to data from four tropical forest sites and found clear benefits of optimized spatiotemporal harvest plans relative to non-optimized harvest plans (random and three pattern-based plans). Our proposed approach, termed the Optimized Floating Refugia strategy, requires minimal species-specific knowledge and can be used to enhance existing conservation efforts (e.g. biological reserve establishment, reduced-impact logging). The approach effectively prioritizes logging-sensitive habitat specialists with restricted ranges and thus provides the largest benefits to the most extinction-prone species. This simple but novel method shows promise as a general strategy to improve biodiversity conservation in species-rich production forest landscapes.     

Human demography and reserve size predict wildlife extinction in West Africa.

Species-area models have become the primary tool used to predict baseline extinction rates for species in isolated habitats, and have influenced conservation and land-use planning worldwide. In particular, these models have been used to predict extinction rates following the loss or fragmentation of natural habitats in the absence of direct human influence on species persistence. Thus, where direct human influences, such as hunting, put added pressure on species in remnant habitat patches, we should expect to observe extinction rates higher than those predicted by simple species-area models. Here, we show that extinction rates for 41 species of large mammals in six nature reserves in West Africa are 14-307 times higher than those predicted by models based on reserve size alone. Human population and reserve size accounted for 98% of the observed variation in extinction rates between reserves. Extinction occurred at higher rates than predicted by species-area models for carnivores, primates and ungulates, and at the highest rates overall near reserve borders. Our results indicate that, where the harvest of wildlife is common, conservation plans should focus on increasing the size of reserves and reducing the rate of hunting.     

Marine reserve design theory for species with ontogenetic migration

Models for marine reserve design have been developed primarily with ‘reef fish’ life histories in mind: sedentary adults in patches connected by larval dispersal. However, many fished species undertake ontogenetic migrations, such as from nursery grounds to adult spawning habitats, and current theory does not fully address the range of reserve options posed by that situation. I modelled a generic species with ontogenetic migration to investigate the possible benefits of reserves under three alternative scenarios. First, the fishery targets adult habitat, and reserves can sustain yields under high exploitation, unless habitat patches are well connected. Second, the fishery targets the nursery, and reserves are highly effective, regardless of connectivity patterns. Third, the fishery targets both habitats, and reserves only succeed if paired on adjacent, well-connected nursery and adult patches. In all cases, reserves can buffer populations against overexploitation but would not enhance fishery yield beyond that achievable by management without reserves. These results summarize the general situations in which management using reserves could be useful for ontogenetically migrating species, and the type of connectivity data needed to inform reserve design.     

Design of Nature Reserve System for Red-Crowned Crane in China

A system of nature reserves is a necessary component of an integrated conservation strategy. The basic problem in the design of a nature reserve system is to select the smallest number of sites from a region for some conservation objectives. There are 33 nature reserves intended for the conservation of Red-Crowned Crane (Grus japonensis), totalling 3.1 million ha. Other habitats of Red-Crowned Crane are facing severe problems due to economic development and other human activities. GIS, iterative methods and the integer programming approach were employed to design the nature reserve system for Red-Crowned Crane, with conservation goals to protect 70 and 60% areas of highly and moderately suitable wetlands, respectively. The results indicated the need to designate six new nature reserves in Wulagai marsh, Duluhe River, Tumen River, Rizhao, Gaoyou Lake and Hongze Lake, and showed a protection zone in Songnen Plain (between Heilongjiang Province and Jilin Province) and three clusters of nature reserves in Sanjing Plain, which, will be used as reference for the designation of new nature reserves, or enlargement and adjustment of existing nature reserves. The iterative method and integer programming approaches were feasible for design of nature reserve system.     

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.     

Beyond opportunism: Key principles for systematic reserve selection

The intention and practice of conservation reserve selection are different. A major reason for systems of reserves is to sustain biological diversity. This involves protecting examples of as many natural features, e.g. species, communities or environments, as possible. In reality, however, new reserves have rarely been dedicated for their representation of features. Furthermore, the opportunism that has characterized the development of reserve systems can actually jeopardize the representation of all features in reserves through the inefficient allocation of limited resources. More systematic approaches are essential if reserves are to play their role in protecting biodiversity. Some basic principles for conservation planning are emerging from recent systematic procedures for reserve selection. These principles will help to link intention and practice.     

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.     

自然保护区生态安全设计的方法研究

由于生态破坏和自然栖息地的丧失,造成野生动植物种群的破碎化．自然保护区已成为孤立的生境岛屿,目前以单个、孤立保护区为主的生物多样性保护模式是远远不够的,应在广泛的时空尺度上保护生态过程和生物多样性各组成成份,建立一个整体的保护网络．根据国内外生物多样性保护的要求和发展趋势,提出了自然保护区生态安全设计的概念,它是综合考虑了生态、社会、经济的一种协调设计战略,首先从区域层次研究保护区网络的优化设计;其次,在网络的每个节点(保护区),研究保护区的面积、形状和内部功能分区;最后,研究网络与节点的连接(廊道)．自然保护区网络设计应维持生态系统的地域完整性和生态过程完整性,采用迭代法、整数规划方法和地理途径方法,为一个或多个保护目标勾画出多种保护规划蓝图。     

Marine reserves help coastal ecosystems cope with extreme weather

Natural ecosystems have experienced widespread degradation due to human activities. Consequently, enhancing resilience has become a primary objective for conservation. Nature reserves are a favored management tool, but we need clearer empirical tests of whether they can impart resilience. Catastrophic flooding in early 2011 impacted coastal ecosystems across eastern Australia. We demonstrate that marine reserves enhanced the capacity of coral reefs to withstand flood impacts. Reserve reefs resisted the impact of perturbation, whilst fished reefs did not. Changes on fished reefs were correlated with the magnitude of flood impact, whereas variation on reserve reefs was related to ecological variables. Herbivory and coral recruitment are critical ecological processes that underpin reef resilience, and were greater in reserves and further enhanced on reserve reefs near mangroves. The capacity of reserves to mitigate external disturbances and promote ecological resilience will be critical to resisting an increased frequency of climate‐related disturbance.     

Implications of fish home range size and relocation for marine reserve function

Reserves are being used increasingly to conserve fish communities and populations under threat from overfishing, but little consideration has been given to how fish behavior might affect reserve function. This review examines the implications of how fish use space, in particular the occurrence and size of home ranges and the frequency and direction of home range relocations. Examples are drawn primarily from the literature on coral reef fishes, but the principles apply to other habitats. Reserves can protect fish species only if individuals restrict their movements to a localized home range during at least part of the life cycle. Home range sizes increase with body size. In small reserves, a significant proportion of fish whose home ranges are centered within the reserve can be exposed to fishing mortality because their home ranges include non-reserve areas. Relocation of home ranges following initial settlement increases exposure to the fishery, especially if habitat selection is frequency-dependent. Distance, barriers, and costs of movement counter such redistribution. These considerations lead to predictions that population density and mean fish size (1) will form gradients across reserve boundaries with maxima in the center of the reserve and minima outside the reserve away from the boundary; (2) will increase rapidly in newly established reserves, only later providing spillover to adjacent fisheries as density-dependent emigration begins to take effect; and (3) will be higher in reserves that are larger and have higher area:edge ratios, more habitat types, natural barriers between reserve and non-reserve areas, and higher habitat quality inside than outside the reserve. (4) Species with low mobility and weak density-dependence of space use will show the greatest increase in reserves and the strongest benefit for population reproductive capacity, but those with intermediate levels of these traits will provide the greatest spillover benefit to nearby fisheries.     

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.     

Designing connected marine reserves in the face of global warming

Marine reserves are widely used to protect species important for conservation and fisheries and to help maintain ecological processes that sustain their populations, including recruitment and dispersal. Achieving these goals requires well‐connected networks of marine reserves that maximize larval connectivity, thus allowing exchanges between populations and recolonization after local disturbances. However, global warming can disrupt connectivity by shortening potential dispersal pathways through changes in larval physiology. These changes can compromise the performance of marine reserve networks, thus requiring adjusting their design to account for ocean warming. To date, empirical approaches to marine prioritization have not considered larval connectivity as affected by global warming. Here, we develop a framework for designing marine reserve networks that integrates graph theory and changes in larval connectivity due to potential reductions in planktonic larval duration (PLD) associated with ocean warming, given current socioeconomic constraints. Using the Gulf of California as case study, we assess the benefits and costs of adjusting networks to account for connectivity, with and without ocean warming. We compare reserve networks designed to achieve representation of species and ecosystems with networks designed to also maximize connectivity under current and future ocean‐warming scenarios. Our results indicate that current larval connectivity could be reduced significantly under ocean warming because of shortened PLDs. Given the potential changes in connectivity, we show that our graph‐theoretical approach based on centrality (eigenvector and distance‐weighted fragmentation) of habitat patches can help design better‐connected marine reserve networks for the future with equivalent costs. We found that maintaining dispersal connectivity incidentally through representation‐only reserve design is unlikely, particularly in regions with strong asymmetric patterns of dispersal connectivity. Our results support previous studies suggesting that, given potential reductions in PLD due to ocean warming, future marine reserve networks would require more and/or larger reserves in closer proximity to maintain larval connectivity.     

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.     

A theory for optimal monitoring of marine reserves

Monitoring of marine reserves has traditionally focused on the task of rejecting the null hypothesis that marine reserves have no impact on the population and community structure of harvested populations. We consider the role of monitoring of marine reserves to gain information needed for management decisions. In particular we use a decision theoretic framework to answer the question: how long should we monitor the recovery of an over‐fished stock to determine the fraction of that stock to reserve? This exposes a natural tension between the cost (in terms of time and money) of additional monitoring, and the benefit of more accurately parameterizing a population model for the stock, that in turn leads to a better decision about the optimal size for the reserve with respect to harvesting. We found that the optimal monitoring time frame is rarely more than 5 years. A higher economic discount rate decreased the optimal monitoring time frame, making the expected benefit of more certainty about parameters in the system negligible compared with the expected gain from earlier exploitation.