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 reassessment of equivalence in yield from marine reserves and traditional fisheries managament

Lively debate continues over whether marine reserves can lead to increased fishery yields when compared to conventional, quota‐based management, apparently driven by differences in the complexity and biological richness of the models being used. In an influential article, Hastings and Botsford used an analytically tractable, spatially implicit, non‐age‐structured model to assert that reserves are typically incapable of increasing yields relative to conventional management, regardless of the type (pre‐ or post‐dispersal, involving adults and/or larvae) or functional form (Ricker or Beverton‐Holt) of density dependence present. A recent numerical (simulation) model by Gaylord et al. concludes that reserves can enhance yield compared to conventional management, a result the authors attribute to their spatially‐explicit evaluation of stage‐structured adult growth, survivability and fecundity; and intercohort (adult‐on‐larvae) post‐dispersal density dependent population dynamics. Here we demonstrate that the increased model complexity is not responsible for the different conclusions. We analyze a spatially‐implicit model without stage structure that incorporates intercohort post‐dispersal density dependence. In this simple model we still find annual extirpation of adult populations outside reserves due to fishing to enhance larval recruitment there, allowing for increased yields compared to those achieved when harvest is evenly spread across the entire domain under conventional management. Consideration of neither spatially‐explicit dispersal dynamics nor stage‐structure in adult demographics is required for reserves to substantially improve yield beyond that attainable under conventional management. In contrast, consideration of within cohort post‐dispersal density dependence among larva during settlement in an otherwise identical model generates equivalence in yield between the two management strategies. These results recast a common message characterizing the relative benefit of reserve versus non‐reserve management from “equivalence at best” to “potentially improved”.     

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 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.     

A general production model with dependence between data from multiple surveys

A fishery‐independent survey for stock assessments is made sometimes more than once per year to detect a difference in relative sizes of fish populations (e.g., catch‐per‐unit‐effort [CPUE]) in response to a seasonal change in fish spatial distributions. Many managers tended to treat such data independently instead of systematically synthesizing them. A primary objective of this study was to synthesize all survey data via a simple hierarchical structure. I used the general (Pella‐Tomlinson) surplus production model for the illustration, because the purpose of this study was not a stock assessment, and the model was simpler than an age‐structured model. The surplus production model has about an eight decade history (since Graham's paper in 1935) and has been prominent in fish population dynamics. The logistic (Graham‐Schaefer) version was useful in the sense of simplifying the dynamics of a fish population in relation to its intrinsic growth, natural mortality, recruitment, density‐dependence, and fishery catch, but it was criticized because of its unrealistic limitations. Subsequently, the general version was suggested to accommodate flexibility and be realistic. In this study, I inferred parameters in the general surplus production model, simultaneously synthesizing all available data even from different temporal ranges. I used Georges Bank yellowtail flounder (Limanda ferruginea) data for demonstration.     

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.     

Marine reserves and reproductive biomass: a case study of a heavily targeted reef fish

Recruitment overfishing (the reduction of a spawning stock past a point at which the stock can no longer replenish itself) is a common problem which can lead to a rapid and irreversible fishery collapse. Averting this disaster requires maintaining a sufficient spawning population to buffer stochastic fluctuations in recruitment of heavily harvested stocks. Optimal strategies for managing spawner biomass are well developed for temperate systems, yet remain uncertain for tropical fisheries, where the danger of collapse from recruitment overfishing looms largest. In this study, we explored empirically and through modeling, the role of marine reserves in maximizing spawner biomass of a heavily exploited reef fish, Lethrinus harak around Guam, Micronesia. On average, spawner biomass was 16 times higher inside the reserves compared with adjacent fished sites. Adult density and habitat-specific mean fish size were also significantly greater. We used these data in an age-structured population model to explore the effect of several management scenarios on L. harak demography. Under minimum-size limits, unlimited extraction and all rotational-closure scenarios, the model predicts that preferential mortality of larger and older fish prompt dramatic declines in spawner biomass and the proportion of male fish, as well as considerable declines in total abundance. For rotational closures this occurred because of the mismatch between the scales of recovery and extraction. Our results highlight how alternative management scenarios fall short in comparison to marine reserves in preserving reproductively viable fish populations on coral reefs.     

Constant proportion harvest policies: dynamic implications in the Pacific halibut and Atlantic cod fisheries

Overfishing, pollution and other environmental factors have greatly reduced commercially valuable stocks of fish. In a 2006 Science article, a group of ecologists and economists warned that the world may run out of seafood from natural stocks if overfishing continues at current rates. In this paper, we explore the interaction between a constant proportion harvest policy and recruitment dynamics. We examine the discrete-time constant proportion harvest policy discussed in Ang et al. (2009) and then expand the framework to include stock-recruitment functions that are compensatory and overcompensatory, both with and without the Allee effect.We focus on constant proportion policies (CPPs). CPPs have the potential to stabilize complex overcompensatory stock dynamics, with or without the Allee effect, provided the rates of harvest stay below a threshold. If that threshold is exceeded, CPPs are known to result in the sudden collapse of a fish stock when stock recruitment exhibits the Allee effect. In case studies, we analyze CPPs as they might be applied to Gulf of Alaska Pacific halibut fishery and the Georges Bank Atlantic cod fishery based on harvest rates from 1975 to 2007. The best fit models suggest that, under high fishing mortalities, the halibut fishery is vulnerable to sudden population collapse while the cod fishery is vulnerable to steady decline to zero. The models also suggest that CPP with mean harvesting levels from the last 30 years can be effective at preventing collapse in the halibut fishery, but these same policies would lead to steady decline to zero in the Atlantic cod fishery. We observe that the likelihood of collapse in both fisheries increases with increased stochasticity (for example, weather variability) as predicted by models of global climate change.     

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.     

Age is not just a number—Mathematical model suggests senescence affects how fish populations respond to different fishing regimes

Senescence is often described as an age‐dependent increase in natural mortality (known as actuarial senescence) and an age‐dependent decrease in fecundity (known as reproductive senescence), and its role in nature is still poorly understood. Based on empirical estimates of reproductive and actuarial senescence, we used mathematical simulations to explore how senescence affects the population dynamics of Coregonus albula, a small, schooling salmonid fish. Using an empirically based eco‐evolutionary model, we investigated how the presence or absence of senescence affects the eco‐evolutionary dynamics of a fish population during pristine, intensive harvest, and recovery phases. Our simulation results showed that the presence or absence of senescence affected how the population responded to the selection regime. At an individual level, gillnetting caused a larger decline in asymptotic length when senescence was present, compared to the nonsenescent population, and the opposite occurred when fishing was done by trawling. This change was accompanied by evolution toward younger age at maturity. At the population level, the change in biomass and number of fish in response to different fishery size‐selection patterns depended on the presence or absence of senescence. Since most life‐history and fisheries models ignore senescence, they may be over‐estimating reproductive capacity and under‐estimating natural mortality. Our results highlight the need to understand the combined effects of life‐history characters such as senescence and fisheries selection regime to ensure the successful management of our natural resources.     

Monitoring demographics of a commercially exploited population of shovelnose sturgeon in the Wabash River,Illinois/Indiana,USA

Shovelnose sturgeon (Scaphirhynchus platorynchus, Rafinesque, 1820) in the Wabash River, Illinois/Indiana, USA, provide an important recreational sport and commercial caviar fishery. In fact, it is one of the last commercially viable populations for sturgeon roe harvest. Due to increased demand in the caviar trade and endangered species legislation that protect shovelnose sturgeon in only a portion of their range, efforts of the roe harvest market may continue to divert toward unprotected populations like the shovelnose sturgeon in the Wabash River. Previous studies have shown that increased harvest pressure in this species can affect the age‐at‐maturation and result in recruitment overfishing. Therefore, it is important to closely and continuously monitor commercially exploited populations. Over the past decade (2007–2016), 13,170 shovelnose sturgeon were sampled with boat electroshocking, hoop nets, drift nets, trotlines, and benthic electrified trawls. Captured fish ranged from 61 to 910 mm fork length (FL; mean = 668 mm), with very few fish less than 550 mm FL. Although fish were found to be in a healthy condition (mean relative weight = 87), there was a decrease in the mean condition over time. In addition, we saw declines in mean FL, weight of roe‐per‐fish, and size‐at‐maturity for female fish directly impacted by harvest. The decline of these population parameters, coupled with an increase in total annual mortality and a truncated age frequency distribution, suggest that harvest is negatively impacting the demographics and recruitment of shovelnose sturgeon in the Wabash River. Considering the downward trajectory of population dynamics and high estimates of mortality, their resiliency to continued harvest and environmental changes will be limited.     

Density dependence,prey accessibility and prey depletion by fisheries drive Peruvian seabird population dynamics

In marine ecosystems top predator populations are shaped by environmental factors affecting their prey abundance. Coupling top predators’ population studies with independent records of prey abundance suggests that prey fluctuations affect fecundity parameters and abundance of their predators. However, prey may be abundant but inaccessible to their predators and a major challenge is to determine the relative importance of prey accessibility in shaping seabird populations. In addition, disentangling the effects of prey abundance and accessibility from the effects of prey removal by fisheries, while accounting for density dependence, remains challenging for marine top predators. Here, we investigate how climate, population density, and the accessibility and removal of prey (the Peruvian anchovy Engraulis ringens) by fisheries influence the population dynamics of the largest sedentary seabird community (≈ 4 million individuals belonging to guanay cormorant Phalacrocorax bougainvillii, Peruvian booby Sula variegata and Peruvian pelican Pelecanus thagus) of the northern Humboldt Current System over the past half‐century. Using Gompertz state–space models we found strong evidence for density dependence in abundance for the three seabird species. After accounting for density dependence, sea surface temperature, prey accessibility (defined by the depth of the upper limit of the subsurface oxygen minimum zone) and prey removal by fisheries were retained as the best predictors of annual population size across species. These factors affected seabird abundance the current year and with year lags, suggesting effects on several demographic parameters including breeding propensity and adult survival. These findings highlight the effects of prey accessibility and fishery removals on seabird populations in marine ecosystems. This will help refine management objectives of marine ecosystems in order to ensure sufficient biomass of forage fish to avoid constraining seabird population dynamics, while taking into account of the effects of environmental variability.     

Population dynamics and potential of fisheries stock enhancement: practical theory for assessment and policy analysis

The population dynamics of fisheries stock enhancement, and its potential for generating benefits over and above those obtainable from optimal exploitation of wild stocks alone are poorly understood and highly controversial. I review pertinent knowledge of fish population biology, and extend the dynamic pool theory of fishing to stock enhancement by unpacking recruitment, incorporating regulation in the recruited stock, and accounting for biological differences between wild and hatchery fish. I then analyse the dynamics of stock enhancement and its potential role in fisheries management, using the candidate stock of North Sea sole as an example and considering economic as well as biological criteria. Enhancement through release of recruits or advanced juveniles is predicted to increase total yield and stock abundance, but reduce abundance of the naturally recruited stock component through compensatory responses or overfishing. Economic feasibility of enhancement is subject to strong constraints, including trade-offs between the costs of fishing and hatchery releases. Costs of hatchery fish strongly influence optimal policy, which may range from no enhancement at high cost to high levels of stocking and fishing effort at low cost. Release of genetically maladapted fish reduces the effectiveness of enhancement, and is most detrimental overall if fitness of hatchery fish is only moderately compromised. As a temporary measure for the rebuilding of depleted stocks, enhancement cannot substitute for effort limitation, and is advantageous as an auxiliary measure only if the population has been reduced to a very low proportion of its unexploited biomass. Quantitative analysis of population dynamics is central to the responsible use of stock enhancement in fisheries management, and the necessary tools are available.     

Ecological drivers of guanaco recruitment: variable carrying capacity and density dependence

Ungulates living in predator-free reserves offer the opportunity to study the influence of food limitation on population dynamics without the potentially confounding effects of top-down regulation or livestock competition. We assessed the influence of relative forage availability and population density on guanaco recruitment in two predator-free reserves in eastern Patagonia, with contrasting scenarios of population density. We also explored the relative contribution of the observed recruitment to population growth using a deterministic linear model to test the assumption that the studied populations were closed units. The observed densities increased twice as fast as our theoretical populations, indicating that marked immigration has taken place during the recovery phase experienced by both populations, thus we rejected the closed-population assumption. Regarding the factors driving variation in recruitment, in the low- to medium-density setting, we found a positive linear relationship between recruitment and surrogates of annual primary production, whereas no density dependence was detected. In contrast, in the high-density scenario, both annual primary production and population density showed marked effects, indicating a positive relationship between recruitment and per capita food availability above a food-limitation threshold. Our results support the idea that environmental carrying capacity fluctuates in response to climatic variation, and that these fluctuations have relevant consequences for herbivore dynamics, such as amplifying density dependence in drier years. We conclude that including the coupling between environmental variability in resources and density dependence is crucial to model ungulate population dynamics; to overlook temporal changes in carrying capacity may even mask density dependence as well as other important processes.     

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.     

Management strategies for sustainable invertebrate fisheries in coastal ecosystems of Galicia (NW Spain)

Artisanal coastal invertebrate fisheries in Galicia are socio-economically important and ecologically relevant. Their management, however, has been based on models of fish population dynamics appropriate for highly mobile demersal or pelagic resources and for industrial fisheries. These management systems focus on regulating fishing effort, but in coastal ecosystems activities that change or destruct key habitats may have a greater effect on population abundance than does fishing mortality. The Golfo Artabro was analysed as a representative example of a coastal ecosystem in Galicia, and the spider crab Maja squinado used as a model of an exploited coastal invertebrate, for which shallow coastal areas are key habitats for juvenile stages. The commercial legal gillnet fishery for the spider crab harvests adults during their reproductive migrations to deep waters and in their wintering habitats. Illegal fisheries operate in shallow waters. The annual rate of exploitation is >90%, and <10% of the primiparous females reproduce effectively at least once. A simple spatially-explicit cohort model was constructed to simulate the population dynamics of spider crab females. Yield- and egg-per-recruit analyses corresponding to different exploitation regimes were performed to compare management policies directed to control the fishing effort or to protect key habitats. It was found that the protection of juvenile habitats could allow increases in yield and reproductive effort higher than in the present system, with such protection based in the control of the fishing effort of the legal fishery. Additionally, there is an urgent need for alternative research and management strategies in artisanal coastal fisheries based on the implementation of a system of territorial use rights for fishers, the integration of the fishers into assessment and management processes, and the protection of key habitats (marine reserves) as a basic tool for the regulation of the fisheries.     

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.     

Regime, phase and paradigm shifts: making community ecology the basic science for fisheries

Modern fishery science, which began in 1957 with Beverton and Holt, is ca. 50 years old. At its inception, fishery science was limited by a nineteenth century mechanistic worldview and by computational technology; thus, the relatively simple equations of population ecology became the fundamental ecological science underlying fisheries. The time has come for this to change and for community ecology to become the fundamental ecological science underlying fisheries. This point will be illustrated with two examples. First, when viewed from a community perspective, excess production must be considered in the context of biomass left for predators. We argue that this is a better measure of the effects of fisheries than spawning biomass per recruit. Second, we shall analyse a simple, but still multi-species, model for fishery management that considers the alternatives of harvest regulations, inshore marine protected areas and offshore marine protected areas. Population or community perspectives lead to very different predictions about the efficacy of reserves.     

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.     

Genetic recruitment patterns are patchy and spatiotemporally unpredictable in a deep-water snapper (Lutjanus vivanus) sampled in fished and protected areas of western Puerto Rico

Marine protected areas (MPAs) have the potential to conserve biodiversity and improve fishery sustainability, but their efficacy depends on sound design and implementation, which requires an understanding of connectivity among reserves and between reserves and fished areas. Most studies of connectivity involving reserves focus on fishes with characteristics atypical for exploited species, making the results less applicable to fisheries management. Here, patterns of genomic diversity were assessed within and among geographic samples of juvenile of silk snapper, Lutjanus vivanus, collected in protected and fished areas on the western coast of Puerto Rico. The results indicate significant variation in spatiotemporal genetic recruitment patterns, with the two MPAs located off the shelf having partially decoupled recruitment processes from sites on the shelf. Spatial autocorrelation was found at distances less than 20 km within years, but the degree and pattern of spatial structure differed across years, suggesting that recruitment along the west coast of Puerto Rico originates from semi-independent units of spawners whose contribution varies in space and time. The results suggest that while MPAs may work to supplement fisheries where recruitment is spatiotemporally predictable, in species for which adult contribution is variable in space and time, other management strategies should be explored as well.