Unintended consequences and trade‐offs of fish passage

We synthesized evidence for unintended consequences and trade‐offs associated with the passage of fishes. Provisioning of fish passageways at dams and dam removals are being carried out increasingly as resource managers seek ways to reduce fragmentation of migratory fish populations and restore biodiversity and nature‐like ecosystem services in tributaries altered by dams. The benefits of provisioning upstream passage are highlighted widely. Possible unwanted consequences and trade‐offs of upstream passage are coming to light, but remain poorly examined and underappreciated. Unintended consequences arise when passage of native and desirable introduced fishes is delayed, undone (fallback), results in patterns of movement and habitat use that reduce Darwinian fitness (e.g. ecological traps), or is highly selective taxonomically and numerically. Trade‐offs arise when passage decisions intended to benefit native species interfere with management decisions intended to control the unwanted spread of non‐native fishes and aquatic invertebrates, or genes, diseases and contaminants carried by hatchery and wild fishes. These consequences and trade‐offs will vary in importance from system to system and can result in large economic and environmental costs. For some river systems, decisions about how to manage fish passage involve substantial risks and could benefit from use of a formal, structured process that allows transparent, objective and, where possible, quantitative evaluation of these risks. Such a process can also facilitate the design of an adaptive framework that provides valuable insights into future decisions.     

Learning about danger: chemical alarm cues and local risk assessment in prey fishes

An individual's behaviour patterns can be conceptualized as a series of threat‐sensitive trade‐offs between ambient predation pressure and a suite of fitness‐related activities, such as resource defence, foraging and mating. Individuals that can reliably assess local predation risk could increase their fitness potential by exhibiting predator avoidance behaviours only at appropriate times. However, such learned risk assessment requires reliable information regarding current predation risks. A diverse range of prey fishes are known to possess chemical alarm cues, which when detected by conspecifics and some heterospecifics, elicit a variety of overt and covert responses. These chemical cues, either alone or as a part of a predator's dietary odour, provide reliable information regarding local predation risk. In this review, I describe recent works examining the role of chemosensory information in: (i) acquired predator recognition, (ii) predator inspection behaviour and (iii) the use of conspecific and heterospecific cues as social information sources.     

Foraging arena theory

There is a critical need for quantitative models that can help evaluate trade‐off decisions related to the impacts of harvesting and protection of aquatic ecosystems within an ecosystem context. Ecosystem models used to evaluate such trade‐offs need to have the capability of capturing the dynamic stability that can arise when predator‐prey interactions are restricted to spatial and temporal arenas. Foraging arenas appear common in aquatic systems and are created by a wide range of mechanisms, ranging from restrictions of predator distributions in response to predation risk caused by their own predators, to risk‐sensitive foraging behaviour by their prey. Foraging arenas partition the prey in each predator‐prey interaction in a food web into vulnerable and invulnerable states, with exchange between these states potentially limiting overall trophic flow. Inclusion of vulnerability exchange processes in models for recruitment processes and food web responses to disturbances like harvesting leads to very different predictions about dynamic stability, trophic cascades and maintenance of ecological diversity than do models based on large‐scale mass action (random mixing) interactions between prey and predators. Although a number of methods to estimate these critical exchange rates are presented, none are considered fully satisfactory. The most important challenge for the practical application of models that incorporate foraging arena theory today is not only developing new or improved methods for measuring exchange rates but also evaluating how such rates vary in responses to major fishery‐induced changes in abundances of predators.     

Forage fish fisheries management requires a tailored approach to balance trade‐offs

Ecosystem‐based fishery management requires considering the effects of actions on social, natural and economic systems. These considerations are important for forage fish fisheries, because these species provide ecosystem services as a key prey in food webs and support valuable commercial fisheries. Forage fish stocks fluctuate naturally, and fishing may make these fluctuations more pronounced, yet harvest strategies intended to ameliorate these effects might adversely affect fisheries and communities. Here, we evaluate trade‐offs among a diverse suite of management objectives by simulating outcomes from several harvest strategies on forage fish species. We demonstrate that some trade‐offs (like those between catches and minimizing collapse length) were universal among forage species and could not be eliminated by the use of different control rules. We also demonstrate that trade‐offs vary among forage fish species, with strong trade‐offs between stable, high catches and high‐biomass periods (“bonanzas”) for menhaden‐ and anchovy‐like fish, and counterintuitive trade‐offs for sardine‐like fish between shorter collapses and longer bonanzas. We find that harvest strategies designed to maintain stability in catches will result in more severe collapses. Finally, we show that the ability of assessments to detect rapid changes in population status greatly affects control rule performance and the degree and type of trade‐offs, increasing the risk and severity of collapses and reducing catches. Together, these results demonstrate that while default harvest strategies are useful in data‐poor situations, management strategy evaluations that are tailored to specific forage fish may better balance trade‐offs.     

A dynamic optimisation model for the behaviour of tunas at ocean fronts

We present a model that simulates the foraging behaviour of tunas in the vicinity of ocean fronts. Stochastic dynamic programming is used to determine optimal habitat choice and swimming speed in relation to environmental variables (water temperature and clarity) and prey characteristics (abundance and energy density). By incorporating submodels for obligate physiological processes (gastric evacuation, standard and active metabolic costs) and sensory systems (visual feeding efficiency), we have integrated into a single fitness-based model many of the factors that might explain the aggregation of tunas at ocean fronts. The modelling technique describes fitness landscapes for all combinations of states, and makes explicit, testable predictions about time- and state-dependent behaviour. Enhanced levels of searching activity when hungry and towards the end of the day are an important feature of the optimal behaviour predicted. We consider the model to be particularly representative of the behaviour of the warm-water tunas or Neothunnus (e.g. skipjack, Katsuwonus pelamis , and yellowfin, Thunnus albacares ) and for surface-dwelling temperate tunas (e.g. young albacore, Thunnus alalunga ), which are often observed to aggregate near fronts. For the bluefin group (i.e. older albacore; northern and southern bluefin, Thunnus thynnus and Thunnus maccoyii ), for which extended vertical migrations are a significant and as yet unexplained component of behaviour, the model is able to reproduce observed behaviour by adopting the lower optimal temperature and standard metabolic rate of albacore. The model cannot explain why physiological differences exist between and within the different tuna species, but it does show how differences in susceptibility to thermal stress will permit different behaviour.     

The mitigation hierarchy for sharks: A risk‐based framework for reconciling trade‐offs between shark conservation and fisheries objectives

Sharks and their cartilaginous relatives are one of the world's most threatened species groups. The primary cause is overfishing in targeted and bycatch fisheries. Reductions in fishing mortality are needed to halt shark population declines. However, this requires complex fisheries management decisions, which often entail trade‐offs between conservation objectives and fisheries objectives. We propose the mitigation hierarchy (MH)—a step‐wise precautionary approach for minimizing the impacts of human activity on biodiversity—as a novel framework for supporting these management decisions. We outline a holistic conceptual model for risks to sharks in fisheries, which includes biophysical, operational and socioeconomic considerations. We then demonstrate how this model, in conjunction with the MH, can support risk‐based least cost shark conservation. Through providing examples from real‐world fishery management problems, we illustrate how the MH can be applied to a range of species, fisheries and contexts, and explore some of the opportunities and challenges hereto. Finally, we outline next steps for research and implementation. This is important in the context of increasing international regulation of shark fishing and trade, which must lead to reductions in shark mortality, while managing trade‐offs between conservation objectives and the socioeconomic value of fisheries.     

Efficiency of fisheries is increasing at the ecosystem level

Managing fisheries presents trade‐offs between objectives, for example yields, profits, minimizing ecosystem impact, that have to be weighed against one another. These trade‐offs are compounded by interacting species and fisheries at the ecosystem level. Weighing objectives becomes increasingly challenging when managers have to consider opposing objectives from different stakeholders. An alternative to weighing incomparable and conflicting objectives is to focus on win–wins until Pareto efficiency is achieved: a state from which it is impossible to improve with respect to any objective without regressing at least one other. We investigate the ecosystem‐level efficiency of fisheries in five large marine ecosystems (LMEs) with respect to yield and an aggregate measure of ecosystem impact using a novel calibration of size‐based ecosystem models. We estimate that fishing patterns in three LMEs (North Sea, Barents Sea and Benguela Current) are nearly efficient with respect to long‐term yield and ecosystem impact and that efficiency has improved over the last 30 years. In two LMEs (Baltic Sea and North East US Continental Shelf), fishing is inefficient and win–wins remain available. We additionally examine the efficiency of North Sea and Baltic Sea fisheries with respect to economic rent and ecosystem impact, finding both to be inefficient but steadily improving. Our results suggest the following: (i) a broad and encouraging trend towards ecosystem‐level efficiency of fisheries; (ii) that ecosystem‐scale win–wins, especially with respect to conservation and profits, may still be common; and (iii) single‐species assessment approaches may overestimate the availability of win–wins by failing to account for trade‐offs across interacting species.     

Diel distribution,behaviour and consumption of juvenile Chinook salmon (Oncorhynchus tshawytscha) in a Wilderness stream

Abstract – Understanding population regulation in juvenile salmonids requires distinguishing the effects of intrinsic (size, behaviour) and extrinsic (food, competition) factors. To examine the relative influence of these variables on juvenile Chinook salmon (Oncorhynchus tshawytscha) in the Salmon River drainage (ID, USA), we examined diel differences in foraging microhabitats, behaviour and consumption in two areas with consistent differences in parr‐to‐smolt survival. In lower Big Creek (LBC, high‐survival area) and upper Big Creek (UBC, low‐survival area), we observed fish by snorkelling, recording length, behaviour (foraging rate and aggression) and physical (depth, velocity, cover, temperature) and biotic (density, size and species of neighbouring fish) microhabitat features. Stomach contents were extracted to estimate consumption. Depth and temperature were greater in LBC, where Chinook salmon were significantly larger and present at lower densities. Fish in LBC exhibited higher foraging activity during the day than night, but there were no size differences between day and night foragers. In UBC, a higher density area, foraging behaviour did not change between day and night, although the smallest size classes did not forage nocturnally. Regression models that integrated physical and biotic variables suggested that physical factors influenced foraging in both areas, but competition also affected foraging in UBC. Our results demonstrate that fish from low‐ and high‐survival populations in Big Creek are exposed to different physical and biological influences during their first growth season, which are reflected in different diel foraging behaviours.     

Diverse juvenile life‐history behaviours contribute to the spawning stock of an anadromous fish population

Habitat quality often varies substantially across space and time, producing a shifting mosaic of growth and mortality trade‐offs across watersheds. Traditional studies of juvenile habitat use have emphasised the evolution of single optimal strategies that maximise recruitment to adulthood and eventual fitness. However, linking the distribution of individual behaviours that contribute to recruitment at the population level has been elusive, particularly for highly fecund aquatic organisms. We examined juvenile habitat use within a population of sockeye salmon (Oncorhynchus nerka) that spawn in a watershed consisting of two interconnected lakes and a marine lagoon. Otolith microchemical analysis revealed that the productive headwater lake accounted for about half of juvenile growth for those individuals surviving to spawn in a single river in the upper watershed. However, 47% of adults had achieved more than half of their juvenile growth in the downstream less productive lake, and 3% of individuals migrated to the estuarine environment during their first summer and returned to freshwater to overwinter before migrating back to sea. These results describe a diversity of viable habitat‐use strategies by juvenile sockeye salmon that may buffer the population against poor conditions in any single rearing environment, reduce density‐dependent mortality and have implications for the designation of critical habitat for conservation purposes. A network of accessible alternative habitats providing trade‐offs in growth and survival may be important for long‐term viability of populations.     

Towards ecosystem-based fisheries management

Performance measures and reference points for the management of target species are now widely used in the fisheries of the developed world. To move us closer to an ecosystem‐based fisheries management framework, we look at the prospects for expanding current single‐species approaches for target species, by considering nontarget species. We also examine the development of performance measures and reference points for emergent ecosystem‐level properties. We conclude that the expansion of single‐species reference points to take account of the nontarget species of a fishery is tractable and desirable. In contrast, the use of performance measures for a single or a small selection of ecosystem metrics is not possible at present, owing to the absence of a clear understanding of their dynamics and a lack of underlying theory to explain their behaviour. However, recent methods that aggregate a wide range of metrics to provide an overall picture of the ecosystem status show promise and have a particular attraction because they have the potential to provide a framework for establishing a dialogue on management issues between all interested parties.     

Genetic structure,origin, and connectivity between nesting and foraging areas of hawksbill turtles of the Yucatan Peninsula: A study for conservation and management

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Integrating ecosystem services into conservation strategies for freshwater and marine habitats: a review

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Heavy loads of parasitic freshwater pearl mussel (Margaritifera margaritifera L.) larvae impair foraging,activity and dominance performance in juvenile brown trout (Salmo trutta L.)

The life cycle of the endangered freshwater pearl mussel (Margaritifera margaritifera) includes a parasitic larval phase (glochidia) on the gills of a salmonid host. Glochidia encystment has been shown to affect both swimming ability and prey capture success of brown trout (Salmo trutta), which suggests possible fitness consequences for host fish. To further investigate the relationship between glochidia encystment and behavioural parameters in brown trout, pairs (n = 14) of wild‐caught trout (infested vs. uninfested) were allowed to drift feed in large stream aquaria and foraging success, activity, agonistic behaviour and fish coloration were observed. No differences were found between infested and uninfested fish except for in coloration, where infested fish were significantly darker than uninfested fish. Glochidia load per fish varied from one to several hundred glochidia, however, and high loads had significant effects on foraging, activity and behaviour. Trout with high glochidia loads captured less prey, were less active and showed more subordinate behaviour than did fish with lower loads. Heavy glochidia loads therefore may negatively influence host fitness due to reduced competitive ability. These findings have implications not only for management of mussel populations in the streams, but also for captive breeding programmes which perhaps should avoid high infestation rates. Thus, low levels of infestation on host fish which do not affect trout behaviour but maintains mussel populations may be optimal in these cases.     

Behaviour of adult Pacific lamprey in near‐field flow and fishway design experiments

Abstract An experimental fishway was used to examine adult Pacific lamprey, Lampetra tridentata (Gairdner), behaviour in a series of attraction and passage‐performance tests. Among all experiments, lamprey oriented to the fishway floor and walls and were attracted to both ambient and concentrated flow. When confronted with high‐velocity areas (vertical‐slot and submerged‐orifice weirs), many lamprey failed to pass upstream. However, lamprey were able to find and take advantage of low‐velocity refuges when they were provided. Lamprey climbed shallow‐ and steep‐angled ramps when attraction cues were sufficient and other passage routes were restricted. The combined results demonstrated the passage challenges that fishways designed and operated for salmonids present to non‐salmonid species. They also highlight the importance of evaluating trade‐offs between fishway attraction and passage efficiency. The experiments were integrated with tagging studies and development of lamprey‐specific passage structures, a research combination that provided an effective template for fishway performance evaluations.     

Attending to spatial social–ecological sensitivities to improve trade‐off analysis in natural resource management

Balancing trade‐offs amongst social–ecological objectives is a central aim of natural resource management. However, objectives and resources often have spatial dimensions, which are usually ignored in trade‐off analyses. We examine how simultaneously integrating social–ecological benefits and their spatial complexities can improve trade‐off analysis. We use Pacific herring (Clupea pallasii, Clupeidae)—an ecologically important forage fish with social, cultural and economic value to communities and commercial fisheries—as a case study. By combining spatial management strategy evaluation with social benefits analysis, we illustrate when policies aimed at aggregate stocks versus spatially segregated substocks of fish fail to balance trade‐offs amongst social–ecological objectives. Spatial measures (e.g. area‐based closures) may achieve some objectives but produce alternative trade‐offs that are sensitive to assumptions about fish population dynamics and social complexities. Our analyses identify policies that are inefficient (e.g. yielding economic costs without producing social or ecological gains), highlight management strategies that generate trade‐offs and indicate when costs are distributed unequally for different user groups. We also point to strategies with outcomes that are robust to spatial uncertainties and reveal research priorities by identifying which performance metrics exhibit sensitivity to spatial ecological assumptions. Collectively, our analyses demonstrate how incorporating social objectives and spatial dynamics into management strategy evaluation can reveal trade‐offs and the implications of management decisions.     

Learning of foraging skills by fish

This chapter outlines the relationships between a number of key factors that influence learning and memory, and illustrates them by reference to studies on the foraging behaviour of fish. Learning can lead to significant improvements in foraging performance in only a few exposures, and at least some fish species are capable of adjusting their foraging strategy as patterns of patch profitability change. There is also evidence that the memory window for prey varies between fish species, and that this may be a function of environmental predictability. Convergence between behavioural ecology and comparative psychology offers promise in terms of developing more mechanistically realistic foraging models and explaining apparently ‘suboptimal’ patterns of behaviour. Foraging decisions involve the interplay between several distinct systems of learning and memory, including those that relate to habitat, food patches, prey types, conspecifics and predators. Fish biologists, therefore, face an interesting challenge in developing integrated accounts of fish foraging that explain how cognitive sophistication can help individual animals to deal with the complexity of the ecological context.     

Catchment zoning to enhance co‐benefits and minimize trade‐offs between ecosystem services and freshwater biodiversity conservation

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Expanding the concept of sustainable seafood using Life Cycle Assessment

Fisheries management and sustainability assessment of fisheries more generally have recently expanded their scope from single‐species stock assessment to ecosystem‐based approaches, aiming to incorporate economic, social and local environmental impacts, while still excluding global‐scale environmental impacts. In parallel, Life Cycle Assessment (LCA) has emerged as a widely used and recommended framework to assess environmental impacts of products, including global‐scale impacts. For over a decade, LCA has been applied to seafood supply chains, leading to new insights into the environmental impact of seafood products. We present insights from seafood LCA research with particular focus on evaluating fisheries management, which strongly influences the environmental impact of seafood products. Further, we suggest tangible ways in which LCA could be taken up in management. By identifying trade‐offs, LCA can be a useful decision support tool and avoids problem shifting from one concern (or activity) to another. The integrated, product‐based and quantitative perspective brought by LCA could complement existing tools. One example is to follow up fuel use of fishing, as the production and combustion of fuel used dominates overall results for various types of environmental impacts of seafood products, and is also often linked to biological impacts of fishing. Reducing the fuel use of fisheries is therefore effective to reduce overall impacts. Allocating fishing rights based on environmental performance could likewise facilitate the transition to low‐impact fisheries. Taking these steps in an open dialogue between fishers, managers, industry, NGOs and consumers would enable more targeted progress towards sustainable fisheries.     

Ecosystem models for fisheries management: finding the sweet spot

The advent of an ecosystem‐based approach dramatically expanded the scope of fisheries management, creating a critical need for new kinds of data and quantitative approaches that could be integrated into the management system. Ecosystem models are needed to codify the relationships among drivers, pressures and resulting states, and to quantify the trade‐offs between conflicting objectives. Incorporating ecosystem considerations requires moving from the single‐species models used in stock assessments, to more complex models that include species interactions, environmental drivers and human consequences. With this increasing model complexity, model fit can improve, but parameter uncertainty increases. At intermediate levels of complexity, there is a ‘sweet spot’ at which the uncertainty in policy indicators is at a minimum. Finding the sweet spot in models requires compromises: for example, to include additional component species, the models of each species have in some cases been simplified from age‐structured to logistic or bioenergetic models. In this paper, we illuminate the characteristics, capabilities and short‐comings of the various modelling approaches being proposed for ecosystem‐based fisheries management. We identify key ecosystem needs in fisheries management and indicate which types of models can meet these needs. Ecosystem models have been playing strategic roles by providing an ecosystem context for single‐species management decisions. However, conventional stock assessments are being increasingly challenged by changing natural mortality rates and environmentally driven changes in productivity that are observed in many fish stocks. Thus, there is a need for more tactical ecosystem models that can respond dynamically to changing ecological and environmental conditions.