Implementing the precautionary approach into fisheries management: Biomass reference points and uncertainty buffers

The precautionary approach to fisheries management advocates for risk-averse management strategies that include biological reference points and account for scientific uncertainty (i.e. process, model and observation uncertainty). In this regard, two approaches have been recommended: (a) biomass reference points to safeguard against low stock biomass, and (b) uncertainty buffers that reduce the catch limit as a function of the scientific uncertainty. This study compares the effectiveness of these two precautionary approaches in recovering over-exploited fish stocks. We evaluate the performance of more than 80 harvest control rules (HCRs) within a stochastic management strategy evaluation (MSE) framework for three stocks with contrasting life-history parameters and under various levels of scientific uncertainty. The results show that both approaches reduce the risk of overfishing at the expense of expected yield. This risk-yield trade-off strongly depends on the HCRs, life-history parameters of the species, as well as the level of the scientific uncertainty. Nevertheless, some combinations of biomass threshold and limit reference points as well as uncertainty buffers lead to a more favourable risk-yield trade-off than other rules. This study elucidates the multiple factors affecting the effectiveness of management strategies and highlights key features of HCRs for precautionary fisheries management.     

How consistent is the advice from stock assessments? Empirical estimates of inter-assessment bias and uncertainty for marine fish and invertebrate stocks

Fishery management frequently involves precautionary buffering for scientific uncertainty. For example, a precautionary buffer that scales with scientific uncertainty is used to calculate the acceptable biological catch downward from the overfishing limit in the US federal fishery management system. However, there is little empirical guidance to suggest how large buffers for scientific uncertainty should be. One important component of uncertainty is variation among different assessments of the same stock in estimates of management-relevant quantities. We analysed commercially exploited marine fish and invertebrate stocks around the world and developed Bayesian hierarchical models to quantify inter-assessment variation in terminal year biomass and fishing mortality estimates, reference points, relative biomass and fishing mortality estimates, and overfishing limits. There was little evidence of inter-assessment bias; stock assessment estimates in the terminal year of the assessment were not consistently higher or lower than estimates of the same quantities in future years. However, there was a tendency for extreme values from the terminal year to be pulled closer to the mean in future years. Inter-assessment variation in all estimates differed across regions, and a longer inter-assessment interval generally resulted in greater variation. Inter-assessment uncertainty was greatest for estimates of the overfishing limit, with coefficients of variation ranging from 17% in Europe (non-EU) to 107% for Pacific Ocean pelagic stocks. Because inter-assessment variation is only one component of scientific uncertainty, we suggest that these uncertainty estimates may provide a basis for determining the minimum size of precautionary buffers.     

大西洋中上层鲨鱼资源状况的初步探讨

据1994—2001年4个航次在金枪鱼延绳钓渔船对热带大洋性中上层鲨鱼资源进行调查，并分析国际大西洋金枪鱼资源保护委员会提供的关于中上层鲨鱼的渔业数据。结果表明：延绳钓渔业共兼捕13种鲨鱼，其中，尖吻鲭鲨和大青鲨是优势种类。尖吻鲭鲨在第1、第2和第4航次的CPUE分别为每千钩0．3502、0．1754和0．0642尾，呈下降趋势。大青鲨在第1～3航次的的CPUE达到每千钩5～7尾，而在第4航次下降为每千钩0．8尾。研究报告了中国金枪鱼船队自1993年开始在大西洋兼捕尖吻鲭鲨和大青鲨的渔获量，并根据捕捞死亡系数和大西洋总渔获量数据，初步探讨了尖吻鲭鲨和大青鲨年平均资源量，影响CPUE的因素和大西洋中上层资源状况。     

Abundance estimation of Northeast Atlantic mackerel based on tag recapture data—A useful tool for stock assessment?

In the present study we utilize tag recapture data to estimate year class abundance and spawning stock biomass of mackerel (Scomber scombrus L.) in the Northeast Atlantic for the period 1986–2008. On average 20,000 jigged mackerel have been tagged annually with internal steel tags in the spawning area west of Ireland and the British Isles, and the tags have been recaptured in commercial catches screened through metal detectors. The spawning stock biomass estimates derived from two different tag-based models were highly variable but were on average 2 and 2.3 times higher than the ICES official estimate. The official estimate is considered uncertain and most likely an underestimate of the actual biomass, due to unregistered mortality in the fisheries and lack of fishery-independent, age-disaggregated data. Hence, tag-based estimates could potentially improve the current assessment if included in the ICES stock assessment on a regular basis. These estimates also involve some uncertainty that needs consideration, especially related to variable tagging mortality, detector efficiency and migrations of the stock.     

Measuring uncertainty in fisheries stock assessment: the delta method,bootstrap, and MCMC

Fisheries management depends on reliable quantification of uncertainty for decision‐making. We evaluate which uncertainty method can be expected to perform best for fisheries stock assessment. The method should generate confidence intervals that are neither too narrow nor too wide, in order to cover the true value of estimated quantities with a probability matching the claimed confidence level. This simulation study compares the performance of the delta method, the bootstrap, and Markov chain Monte Carlo (MCMC). A statistical catch‐at‐age model is fitted to 1000 simulated datasets, with varying recruitment and observation noise. Six reference points are estimated, and confidence intervals are constructed across a range of significance levels. Overall, the delta method and MCMC performed considerably better than the bootstrap, and MCMC was the most reliable method in terms of worst‐case performance, for our relatively data‐rich scenario and catch‐at‐age model, which was not subject to substantial model misspecification. All three methods generated too narrow confidence intervals, underestimating the true uncertainty. Bias correction improved the bootstrap performance, but not enough to match the performance of the delta method and MCMC. We recommend using MCMC as the default method for quantifying uncertainty in fisheries stock assessment, although the delta method is the fastest to apply, and the bootstrap is useful to diagnose estimator bias.     

Variation in abundance of Norwegian spring-spawning herring (Clupea harengus, Clupeidae) throughout the 20th century and the influence of climatic fluctuations

A long-term (1907–98) virtual population analysis (VPA) was made for Norwegian spring-spawning herring (NSSH), which is a huge pelagic fish stock in the north-east Atlantic. It shows that this herring stock has had large fluctuations during the last century; these fluctuations have mainly been determined by variations in the temperature of the inflowing water masses to the region. The spawning stock biomass (SSB) increased from a rather low level in the early years of this century and reached a high level of around 14 million tons by 1930. The spawning stock biomass then decreased to a level of around 10 million tons by 1940, but increased again to a record high level of 16 million tons by 1945. The stock then started to decrease and during the next 20-year period fell to a level of less than 50 000 tons by the late 1960s. Through the 1970s and 1980s, the stock slowly recovered and after the recruitment of strong year classes in 1983 and 1990–1992 the stock recovered to a spawning stock biomass of about 10 million tons. The long-term fluctuation in spawning stock biomass is caused by variations in the survival of recruits. It is found that the long-term changes in spawning stock abundance are highly correlated with the long-term variations in the mean annual temperature of the inflowing Atlantic water masses (through the Kola section) into the north-east Atlantic region. The recruitment is positively correlated with the average temperature in the Kola section in the winter months, January–April, which indicates that environmental factors govern the large-scale fluctuations in production for this herring stock.     

A stochastic surplus production model in continuous time

Surplus production modelling has a long history as a method for managing data‐limited fish stocks. Recent advancements have cast surplus production models as state‐space models that separate random variability of stock dynamics from error in observed indices of biomass. We present a stochastic surplus production model in continuous time (SPiCT), which in addition to stock dynamics also models the dynamics of the fisheries. This enables error in the catch process to be reflected in the uncertainty of estimated model parameters and management quantities. Benefits of the continuous‐time state‐space model formulation include the ability to provide estimates of exploitable biomass and fishing mortality at any point in time from data sampled at arbitrary and possibly irregular intervals. We show in a simulation that the ability to analyse subannual data can increase the effective sample size and improve estimation of reference points relative to discrete‐time analysis of aggregated annual data. Finally, subannual data from five North Sea stocks are analysed with particular focus on using residual analysis to diagnose model insufficiencies and identify necessary model extensions such as robust estimation and incorporation of seasonality. We argue that including all known sources of uncertainty, propagation of that uncertainty to reference points and checking of model assumptions using residuals are critical prerequisites to rigorous fish stock management based on surplus production models.     

Misspecification in stock assessments: Common uncertainties and asymmetric risks

Common uncertainties in stock assessment relate to parameters or assumptions that strongly determine both the estimates of quantities of management interest (e.g. stock depletion) and related reference points (e.g. biomass at maximum sustainable yield). The risks associated with these uncertainties are often presented to managers in the form of decision tables. However, a formal evaluation of the risks from misspecifying an assessment model over time‐horizons spanning multiple assessment cycles requires closed‐loop simulation. There were two aims of this study: (a) develop an approach to identify and evaluate asymmetries in risk to yields and spawning biomass due to biases in key parameters and data sources in a stock assessment model, (b) quantify the relative importance of correctly specifying the various assessment attributes. A computationally efficient stock reduction analysis was evaluated using closed‐loop simulation to identify risks associated with a stock assessment with persistent positive and negative biases in the key parameters and inaccurate assumptions regarding data sources. Six types of assessment misspecification were examined, namely the assumed natural mortality rate, the assumed recruitment compensation ratio, the assumed age of maturity, a hyper‐stable or hyper‐deplete index of abundance, over‐ or under‐reporting of historical catch, and misspecification of the assumed shape of the selectivity curve. This study reveals large asymmetries in risk associated with common uncertainties in stock assessment processes. We highlight the value of reproducible and computationally efficient stock assessment models that can be investigated by closed‐loop simulation before being used for fisheries management.     

Evaluation of a spatially sex‐specific assessment method incorporating a habitat preference model for blue marlin (Makaira nigricans) in the Pacific Ocean

Blue marlin, widely distributed throughout the Pacific Ocean, are sexually dimorphic, have certain preferred habitats, and migrate seasonally. These characteristics have been ignored in previous stock assessment models. A population dynamics model that includes spatial structure, and sex and age structure was therefore constructed and fitted to fisheries data for blue marlin, along with information on the relative density of the population over space derived from a habitat preference model that uses the oceanographic and biological variables sea‐surface temperature, mixed layer depth, sea‐surface height anomaly, and chlorophyll‐a concentration. Monte Carlo simulation was then used to examine the estimation performance of the stock assessment method. Estimates of management‐related quantities including current spawning stock biomass are substantially biased when the assessment method ignores seasonal movement and sexual dimorphism. We also found that (i) uncertainty about the relationship between catch rate and abundance influences estimation performance to a larger extent than uncertainty in catches, (ii) the outcomes of the assessment are sensitive to the values assumed for natural mortality and stock‐recruitment steepness, and (iii) the ratio of current spawning stock biomass to that at pre‐exploitation equilibrium appears to be the most robust among the quantities considered. We conclude that assessment methods for blue marlin in the Pacific Ocean need to take account of seasonal migration and sex structure to improve stock assessments.     

Re‐evaluating the effect of wind on recruitment in Gulf of Maine Atlantic Cod (Gadus morhua) using an environmentally‐explicit stock recruitment model

A previous study documented a correlation between Atlantic Cod (Gadus morhua) recruitment in the Gulf of Maine and an annual index of the north component of May winds. This correlation was supported by modeling studies that indicated strong recruitment of Gulf of Maine Atlantic Cod results from high retention of spring‐spawned larvae in years when winds were predominately out of the north, which favor downwelling. We re‐evaluated this relationship using updated recruitment estimates and found that the correlation decreased between recruitment and wind. The original relationship was largely driven by two recruitment estimates, one of which (2005 year‐class) was highly uncertain because it was near the terminal year of the assessment. With additional data, the updated assessment estimated lower recruitment for the 2005 year‐class, which consequently lowered the correlation between recruitment and wind. We then investigated whether an environmentally‐explicit stock recruit function that incorporated an annual wind index was supported by either the original or updated assessment output. Although incorporation of the annual wind index produced a better fitting model, the uncertainty in the estimated parameters and the implied unexploited conditions were not appropriate for providing management advice. These results suggest the need for caution in the development of environmentally‐explicit stock recruitment relationships, in particular when basing relationships and hypotheses on recruitment estimates from the terminal years of stock assessment models. More broadly, this study highlights a number of sources of uncertainty that should be considered when analyzes are performed on the output of stock assessment models.     

Inclusion of discards in stock assessment models

A large portion of the catch in many stocks may comprise discards which need to be accounted for in assessments in order to avoid bias in estimates of fishing mortality, stock biomass and reference points. In age‐structured assessment models, discards are sometimes treated as a separate fleet or are added to the landings before fitting so that information about discard behaviour and sampling error is lost. In this paper, an assessment model is developed to describe the discard process with size as a covariate while retaining age‐structured population dynamics. Discard size selection, high grading and bulk dumping of fish at sea are modelled so that the temporal dynamics of the process can be quantified within the assessment. The model is used to show that discarding practices have changed over time in a range of Northeast Atlantic demersal fish. In some stocks, there is a substantial increase in high grading and evidence for bulk discarding which can be related to regulatory measures. The model offers a means of identifying transient effects in the discard process that should be removed from both short‐term forecasts and equilibrium reference point calculations.     

Risk of exceeding bioeconomic limit reference points in shrimp aquaculture systems

Abstract

Designing intelligent management strategies for shrimp aquaculture systems require recognising the uncertainty and risks associated with different processes of aquaculture production. To account for natural variability and other sources of uncertainty, estimates of appropriate aquaculture bio‐economic indicators are needed to re‐evaluate periodically the production system and establish new reference points and corresponding management strategies. This paper concentrates on this aspect of the management process and presents a simple classification of indicators and reference points for aquaculture production systems. The uncertainty causing variability in the estimated values of bio‐economic parameters is incorporated through the use of Monte Carlo analysis to estimate the probability of exceeding limit reference points. To illustrate this process, the risks (i.e. probabilities of exceeding bioeconomic limit reference points) associated to alternative timing of harvesting decisions were estimated by randomly generating uncertain prices and natural mortalities with the appropriate probability density functions and corresponding variance. Alternative approaches to deal with risk and uncertainty in data limited management contexts are discussed.     

Assessing the diet of North American Atlantic salmon (Salmo salar L.) off the West Greenland coast using gut content and stable isotope analyses

Investigations on the marine feeding of Atlantic salmon (Salmo salar L.) in the Northwest Atlantic are limited compared with the Northeast Atlantic. Climate‐induced changes to food webs in Atlantic salmon feeding areas have been noted, alongside increased mortality despite a cessation of most marine fisheries. As forage efficiency may be hampering survival, it was important to address this knowledge gap. Atlantic salmon were sampled at three sites on the West Greenland coast (Sisimiut, Nuuk and Qaqortoq) between 2009 and 2011. Gut content and stable isotope analyses were combined to assess spatial and temporal differences in feeding. Capelin (Mallotus villosus) dominated the diet at Nuuk and Qaqortoq, whereas boreoatlantic armhook squid (Gonatus fabricii) was the dominant prey at Sisimiut. Hyperiid amphipods (Themisto spp.) and sand lance (Ammodytes spp.) were also important. Significant differences were found among sites for both gut contents and stable isotope analyses, with fewer differences evident temporally. Dietary differences were also evident across larger scales, with little overlap demonstrated with Northeast Atlantic diets and the emergence of boreoatlantic armhook squid as an important prey item over time. Atlantic salmon diets are frequently anchored on one or two prey items, on which they appear to specialize, but they will diversify to consume other available pelagic prey. Thus, Atlantic salmon are an opportunistic, generalist predator within the pelagic food web. The variability evident in diet suggests that the limited data available are insufficient to appropriately understand potential vulnerabilities that the species may have to ecosystem changes, and suggest further research is needed.     

Is spawning stock biomass a robust proxy for reproductive potential?

Reference points based on fishing mortality (F) and spawning stock biomass (SSB) are a requirement of many fisheries management frameworks. SSB is assumed to be a proxy for stock reproductive potential (SRP). Limit reference points based on SSB are used to indicate the level of biomass below which productivity is affected. SSB fails to account for changes in fecundity, egg viability and sex ratio, and it has been argued that total egg production (TEP) provides a better reflection of SRP. We explore how accounting for TEP impacts limit reference points and evidence for a relationship between stock and recruit. Time series of SSB and TEP are compared for three North Sea stocks: cod (Gadus morhua), herring (Clupea harengus) and plaice (Pleuronectes platessa). Dynamics based on TEP are different from those based on SSB for cod and plaice, but the stock–recruit relationships were not ‘improved’ using TEP. Shifts in productivity (spawner per recruit) occur in all three time series and SSB underestimated uncertainty. Yet again, it was shown that assumptions of stationarity about fish population productivity are incorrect. We argue that the use of TEP does improve the realism in our understanding of stock dynamics, and demographically, more complex management strategy evaluation is required to develop management procedures that are robust to uncertainty and integrate F and the demographic health of a stock. Empirical feedback control systems based on fisheries independent indices including surveys of eggs, larvae, recruits, juveniles or spawning adults should be evaluated and compared to traditional approaches.     

Do albacore exert top‐down pressure on northern anchovy? Estimating anchovy mortality as a result of predation by juvenile north pacific albacore in the California current system

Quantifying the mortality of marine fishes is important for understanding spawner–recruit relationships, predicting year‐class strength, and improving fishery stock assessment models. There is increasing evidence that pelagic predators can exert a top‐down influence on prey, especially during critical early life‐history stages. The objective of this study was to quantify predation by North Pacific albacore on Northern anchovy in the California current system (CCS). I estimated the abundance of juvenile albacore in the CCS from 1966–2005 using stock assessment models and spatially explicit catch‐per‐unit‐effort time series. Anchovy abundance (1966–93), both recruitment and total biomass, was obtained from a stock assessment model. Annual rates of anchovy consumption by albacore were calculated using diet studies of albacore in the CCS, an age‐structured bioenergetics model, and regional estimates of albacore abundance. The range of estimates was large: albacore may remove from less than 1% to over 17% of anchovy pre‐recruitment biomass annually. Relationships between predation and recruitment biomass were consistent with expectations from top‐down effects, but further study is required. This is the first attempt to quantify a specific source of mortality on anchovy recruits and to demonstrate potential top‐down effects of predation on anchovy.     

Stock–environment–recruitment models for North Atlantic albacore (Thunnus alalunga)

Different stock–recruitment models were fitted to North Atlantic albacore (Thunnus alalunga) recruitment and spawning stock biomass data. A classical density dependence hypothesis, a recent environmental‐dependence hypothesis and a combination of both were considered. For the latter case, four stock–environment–recruitment models were used: Ricker, Beverton‐Holt, Deriso's General Model (modified to take into account environmental effects) and conditioned Neural Networks. Cross‐validation analysis showed that the modified Deriso model had the best predictive capability. It detected an inverse effect of the North Atlantic Oscillation (NAO) on recruitment, a Ricker‐type behaviour with density dependent overcompensation when environmental conditions are unfavourable and a Beverton–Holt‐type behaviour towards an asymptotic recruitment carrying capacity with favourable environmental conditions. The Neural Network model also detected that under favourable environmental conditions high spawning stock biomass does not necessarily have a depensatory effect on recruitment. Moreover, they suggest that under extremely favourable environmental conditions, albacore recruitment could increase well above the asymptotic carrying capacity predicted by Beverton–Holt‐type models. However, the general decrease in spawning stock biomass in recent years and increasing NAO trends suggest that there is low probability of exceptionally large recruitment in the future and instead there is a danger of recruitment overfishing.     

Multi‐decadal variations in spawning ground use in Northeast Arctic haddock (Melanogrammus aeglefinus)

Despite the importance both in an ecological and management context, much uncertainty remains about the underlying factors controlling spawning ground use in marine fish. Here, we investigate how spawning ground use of Northeast Arctic (NEA) haddock (Melanogrammus aeglefinus) has changed over time. By combining data from a Soviet‐Russian egg survey conducted over a 35 year period (1959–1993) with a biophysical drift model, we estimate annual median spawning latitudes of NEA haddock. Furthermore, we test for statistical association between the median latitude of spawning and covariates that are candidates for shaping the spawning distribution of haddock. The results indicate no association with temperature, as measured at the Kola transect, or demographic structure, as measured by the mean weight of spawners. However, we find a positive and significant association between spawning latitude and spawning stock biomass, indicating that density‐dependence may be more important in shaping spawning location than other factors such as temperature and demographic structure. A potential mechanism for the reported effect of spawning stock biomass may act through the distribution at the feeding grounds, which is reported to expand north‐ and eastwards at high population density.     

Importance of the Straits of Florida spawning ground to Atlantic sailfish (Istiophorus platypterus) and blue marlin (Makaira nigricans)

Much of the uncertainty in managing highly migratory pelagic species results from the scarcity of fisheries-independent data relevant to determining long-term trends in abundance, migratory movements, and the relative importance of different spawning grounds. To address these issues, we used an ichthyoplankton-based method to quantify the overall level of spawning of sailfish ( Istiophorus platypterus ) and blue marlin ( Makaira nigricans ) in the Straits of Florida (SF). We estimated that during the 2 years (2003–2004) of the study, 4.60 × 10¹¹ sailfish eggs and 4.49 × 10¹¹ blue marlin eggs were produced on an annual basis in this region. These egg production values, when combined with estimates of annual fecundity for each species and the most recent stock assessment estimate of total biomass, indicate that about 2.1% of Western Atlantic sailfish spawning and 1.6% of Atlantic-wide blue marlin spawning occurs in the SF. Additionally, pop-up satellite tags deployed on sailfish at the start of the spawning season revealed their short residency times in the SF, suggesting that a large (≈13%) transient portion of the sailfish population is responsible for the SF egg production. Overall, this study provides a critically needed fisheries-independent method of quantifying spatial and temporal trends in the abundance of highly migratory species. The application of this methodology in the SF indicated that above-average levels of sailfish and blue marlin spawning occur in this area and, possibly more importantly, that the SF is a migratory bottleneck for these species.     

Conservation of endemic landlocked salmonids in regulated rivers: a case‐study from Lake Vänern,Sweden

Conservation of migratory salmonids requires understanding their ecology at multiple scales, combined with assessing anthropogenic impacts. We present a case‐study from over 100 years of data for the endemic landlocked Atlantic salmon (Salmo salar, Salmonidae) and brown trout (Salmo trutta, Salmonidae) in Lake Vänern, Sweden. We use this case‐study to develop life history‐based research and monitoring priorities for migratory salmonids. In Vänern, small wild populations of salmon and trout remain only in the heavily regulated Rivers Klar (Klarälven) and Gullspång (Gullspångsälven), and commercial and sport fisheries are maintained by hatchery stocking. These populations represent some of the last remaining large‐bodied (up to 20 kg) landlocked salmon stocks worldwide. We found that one of four stocks of wild fish has increased since 1996; the other three remain critically low. Hatchery return rates for three of four stocks appear stable at roughly 1% and annual fisheries catch is roughly 75 metric tons, with an estimated 7.5% of hatchery smolts being recruited to the fishery; this also appears relatively stable since 1990. Our analysis reveals much uncertainty in key data requirements, including both river return and fisheries catch rates, estimates of wild smolt production and survival, and hatchery breeding and genetics protocols. These uncertainties, coupled with a lack of information on their riverine and lacustrine ecology, preclude effective management of these unique populations. We conclude with a framework for a life history‐based approach to research and monitoring for Vänern salmon and trout, which should be applicable for all endemic, migratory salmonid populations.