The battle between harvest and natural selection creates small and shy fish.

Harvest of fish and wildlife, both commercial and recreational, is a selective force that can induce evolutionary changes to life history and behavior. Naturally selective forces may create countering selection pressures. Assessing natural fitness represents a considerable challenge in broadcast spawners. Thus, our understanding about the relative strength of natural and fisheries selection is slim. In the field, we compared the strength and shape of harvest selection to natural selection on body size over four years and behavior over one year in a natural population of a freshwater top predator, the northern pike (Esox lucius). Natural selection was approximated by relative reproductive success via parent-offspring genetic assignments over four years. Harvest selection was measured by comparing individuals susceptible to recreational angling with individuals never captured by this gear type. Individual behavior was measured by high-resolution acoustic telemetry. Harvest and natural size selection operated with equal strength but opposing directions, and harvest size selection was consistently negative in all study years. Harvest selection also had a substantial behavioral component independent of body length, while natural behavioral selection was not documented, suggesting the potential for directional harvest selection favoring inactive, timid fish. Simulations of the outcomes of different fishing regulations showed that traditional minimum size-based harvest limits are unlikely to counteract harvest selection without being completely restrictive. Our study suggests harvest selection may be inevitable and recreational fisheries may thus favor small, inactive, shy, and difficult-to-capture fish. Increasing fractions of shy fish in angling-exploited stocks would have consequences for stock assessment and all fisheries operating with hook and line.

Reproductive hyperallometry and managing the world's fisheries.

Marine fisheries are an essential component of global food security, but many are close to their limits and some are overfished. The models that guide the management of these fisheries almost always assume reproduction is proportional to mass (isometry), when fecundity generally increases disproportionately to mass (hyperallometry). Judged against several management reference points, we show that assuming isometry overestimates the replenishment potential of exploited fish stocks by 22% (range: 2% to 78%) for 32 of the world's largest fisheries, risking systematic overharvesting. We calculate that target catches based on assumptions of isometry are more than double those based on assumptions of hyperallometry for most species, such that common reference points are set twice as high as they should be to maintain the target level of replenishment. We also show that hyperallometric reproduction provides opportunities for increasing the efficacy of tools that are underused in standard fisheries management, such as protected areas or harvest slot limits. Adopting management strategies that conserve large, hyperfecund fish may, in some instances, result in higher yields relative to traditional approaches. We recommend that future assessment of reference points and quotas include reproductive hyperallometry unless there is clear evidence that it does not occur in that species.

Expanding conservation culturomics and iEcology from terrestrial to aquatic realms.

The ongoing digital revolution in the age of big data is opening new research opportunities. Culturomics and iEcology, two emerging research areas based on the analysis of online data resources, can provide novel scientific insights and inform conservation and management efforts. To date, culturomics and iEcology have been applied primarily in the terrestrial realm. Here, we advocate for expanding such applications to the aquatic realm by providing a brief overview of these new approaches and outlining key areas in which culturomics and iEcology are likely to have the highest impact, including the management of protected areas; fisheries; flagship species identification; detection and distribution of threatened, rare, and alien species; assessment of ecosystem status and anthropogenic impacts; and social impact assessment. When deployed in the right context with awareness of potential biases, culturomics and iEcology are ripe for rapid development as low-cost research approaches based on data available from digital sources, with increasingly diverse applications for aquatic ecosystems.

Evolutionary Impact of Size-Selective Harvesting on Shoaling Behavior: Individual-Level Mechanisms and Possible Consequences for Natural and Fishing Mortality.

AbstractIntensive and size-selective harvesting is an evolutionary driver of life history as well as individual behavioral traits. Yet whether and to what degree harvesting modifies the collective behavior of exploited species are largely unknown. We present a multigeneration harvest selection experiment with zebrafish, Danio rerio, as a model species to understand the effects of size-selective harvesting on shoaling behavior. The experimental system is based on a large-harvested (typical of most wild-capture fisheries targeting larger size classes) and small-harvested (typical of specialized fisheries and gape-limited predators targeting smaller size classes) selection lines. By combining high-resolution tracking of fish behavior with computational agent-based modeling, we show that shoal cohesion changed in the direction expected by a trade-off between individual vigilance and the use of social cues. In particular, we document a decrease of individual vigilance in the small-harvested line, which was linked to an increase in the attention to social cues, favoring more cohesive shoals. Opposing outcomes were found for the large-harvested line, which formed less cohesive shoals. Using the agent-based model, we outline possible consequences of changes in shoaling behavior for both fishing and natural mortality. The changes in shoaling induced by large size-selective harvesting may decrease fishing mortality but increase mortality by natural predators. Our work suggests an insofar overlooked evolutionary mechanism by which size-selective harvesting can affect fishing and natural mortality of exploited fish.

Size-selective mortality fosters ontogenetic changes in collective risk-taking behaviour in zebrafish, Danio rerio.

Size-selective mortality is common in fish populations and can operate either in a positive size-selective fashion by harvesting larger-than-average fish or be negatively size-selective by harvesting smaller-than-average fish. Through various mechanisms (like genetic correlations among behaviour and life-history traits or direct selection on behaviour co-varying with growth rate or size-at-maturation), size-selection can result in evolutionary changes in behavioural traits. Theory suggests that both positive and negative size-selection without additional selection on behaviour favours boldness, while evolution of shyness is possible if the largest fish are harvested. Here we examined the impact of size-selective mortality on collective boldness across ontogeny using three experimental lines of zebrafish (Danio rerio) generated through positive (large-harvested), negative (small-harvested) and random (control line) size-selective mortality for five generations and then relaxed selection for 10 generations to examine evolutionarily fixed outcomes. We measured collective risk-taking during feeding (boldness) under simulated aerial predation threat, and across four contexts in presence/absence of a cichlid. Boldness decreased across ontogeny under aerial predation threat, and the small-harvested line was consistently bolder than controls. The large and small-harvested lines showed higher behavioural plasticity as larvae and developed personality earlier compared to the controls. The large-harvested line showed increased variability and plasticity in boldness throughout ontogeny. In the presence of a live predator, fish did not differ in boldness in three contexts compared to the controls, but the large-harvested line showed reduced behavioural plasticity across contexts than controls. Our results confirmed theory by demonstrating that size-selective harvesting evolutionarily alters collective boldness and its variability and plasticity.

Ecological impacts of water-based recreational activities on freshwater ecosystems: a global meta-analysis.

Human presence at water bodies can have a range of ecological impacts, creating trade-offs between recreation as an ecosystem service and conservation. Conservation policies could be improved by relying on robust knowledge about the relative ecological impacts of water-based recreation. We present the first global synthesis on recreation ecology in aquatic ecosystems, differentiating the ecological impacts of shore use, (shoreline) angling, swimming and boating. Impacts were assessed at three levels of biological organization (individuals, populations and communities) for several taxa. We screened over 13 000 articles and identified 94 suitable studies that met the inclusion criteria, providing 701 effect sizes. Impacts of boating and shore use resulted in consistently negative, significant ecological impacts across all levels of biological organization. The results were less consistent for angling and swimming. The strongest negative effects were observed in invertebrates and plants. Recreational impacts on birds were most pronounced at the individual level, but not significant at the community level. Due to publication bias and knowledge gaps, generalizations of the ecological impacts of aquatic recreation are challenging. Impacts depend less on the form of recreation. Thus, selectively constraining specific types of recreation may have little conservation value, as long as other forms of water-based recreation continue.

Size-selective mortality induces evolutionary changes in group risk-taking behaviour and the circadian system in a fish.

Intensive and trait-selective mortality of fish and wildlife can cause evolutionary changes in a range of life-history and behavioural traits. These changes might in turn alter the circadian system due to co-evolutionary mechanisms or correlated selection responses both at behavioural and molecular levels, with knock-on effects on daily physiological processes and behavioural outputs. We examined the evolutionary impact of size-selective mortality on group risk-taking behaviour and the circadian system in a model fish species. We exposed zebrafish Danio rerio to either large or small size-selective harvesting relative to a control over five generations, followed by eight generations during which harvesting was halted to remove maternal effects. Size-selective mortality affected fine-scale timing of behaviours. In particular, small size-selective mortality, typical of specialized fisheries and gape-limited predators targeting smaller size classes, increased group risk-taking behaviuor during feeding and after simulated predator attacks. Moreover, small size-selective mortality increased early peaks of daily activity as well as extended self-feeding daily activity to the photophase compared to controls. By contrast large size-selective mortality, typical of most wild capture fisheries, only showed an almost significant effect of decreasing group risk-taking behaviour during the habituation phase and no clear changes in fine-scale timing of daily behavioural rhythms compared to controls. We also found changes in the molecular circadian core clockwork in response to both size-selective mortality treatments. These changes disappeared in the clock output pathway because both size-selected lines showed similar transcription profiles. This switch downstream to the molecular circadian core clockwork also resulted in similar overall behavioural rhythms (diurnal swimming and self-feeding in the last hours of darkness) independent of the underlying molecular clock. To conclude, our experimental harvest left an asymmetrical evolutionary legacy in group risk-taking behaviour and in fine-scale daily behavioural rhythms. Yet, the overall timing of activity showed evolutionary resistance probably maintained by a molecular switch. Our experimental findings suggest that size-selective mortality can have consequences for behaviour and physiological processes.

Behavioural and fitness effects of translocation to a novel environment: Whole-lake experiments in two aquatic top predators.

Translocation into a novel environment through common fisheries management practices, such as fish stocking, provides opportunities to study behavioural and fitness impacts of translocations at realistic ecological scales. The process of stocking, as well as the unfamiliarity with novel ecological conditions and the interactions with resident fish may affect translocated individuals, leading to alterations of behaviours and causing fitness impacts. Our objectives were to investigate how aquatic top predators behaviourally establish themselves and compete with resident individuals following introduction in a novel lake environment and to investigate the resulting fitness consequences. Using high-resolution acoustic telemetry, we conducted whole-lake experiments and compared the activity, activity-space size and fate of translocated and resident individuals in two model top predators, the northern pike Esox lucius (n = 160) and European catfish Silurus glanis (n = 33). Additionally, we compared the reproductive success of translocated and resident northern pike. The experiment was conducted with large (adult) individuals of different origins, resilient to predation, but subject to agonistic interactions and competition with resident fish. Over a period of several months, the translocated catfish exhibited consistently larger activity-space sizes than resident catfish, but did not differ from residents in activity and survival. The pike from one of the two translocated origins we tested also showed elevated space-use, and both translocated origins revealed higher mortality rates than their resident conspecifics, indicating maladjustment to their novel environment. When non-resident pike reproduced, they overwhelmingly produced hybrid offspring with resident fish, indicating that introductions fostered gene flow of non-native genes. Our study indicates that fish introductions result in behavioural and fitness impacts even in large-bodied top predators that experience low levels of natural predation risk.

Size-selective harvesting fosters adaptations in mating behaviour and reproductive allocation, affecting sexual selection in fish.

The role of sexual selection in the context of harvest-induced evolution is poorly understood. However, elevated and trait-selective harvesting of wild populations may change sexually selected traits, which in turn can affect mate choice and reproduction. We experimentally evaluated the potential for fisheries-induced evolution of mating behaviour and reproductive allocation in fish. We used an experimental system of zebrafish (Danio rerio) lines exposed to large, small or random (i.e. control) size-selective mortality. The large-harvested line represented a treatment simulating the typical case in fisheries where the largest individuals are preferentially harvested. We used a full factorial design of spawning trials with size-matched individuals to control for the systematic impact of body size during reproduction, thereby singling out possible changes in mating behaviour and reproductive allocation. Both small size-selective mortality and large size-selective mortality left a legacy on male mating behaviour by elevating intersexual aggression. However, there was no evidence for line-assortative reproductive allocation. Females of all lines preferentially allocated eggs to the generally less aggressive males of the random-harvested control line. Females of the large-harvested line showed enhanced reproductive performance, and males of the large-harvested line had the highest egg fertilization rate among all males. These findings can be explained as an evolutionary adaptation by which individuals of the large-harvested line display an enhanced reproductive performance early in life to offset the increased probability of adult mortality due to harvest. Our results suggest that the large-harvested line evolved behaviourally mediated reproductive adaptations that could increase the rate of recovery when populations adapted to high fishing pressure come into secondary contact with other populations.

Public perception of river fish biodiversity in four European countries.

Public support for biodiversity conservation is shaped by people's values and their knowledge, beliefs, and attitudes toward the environment. We conducted the first multinational representative survey of the general public's perceptions of river fish biodiversity in France, Germany, Norway, and Sweden. For the online survey, 1000 respondents per country were randomly selected from large panels following country-specific quotas set on age, gender, and educational level. Questions covered people's level of knowledge, beliefs, values, and attitudes toward river fish, environmental threats, and conservation measures. We found that the public had limited knowledge of freshwater fishes. Two non-native species, rainbow trout (Oncorhynchus mykiss) and brook trout (Salvelinus fontinalis), were widely perceived as native, whereas native Atlantic salmon (Salmo salar) was mostly classified as native in Scandinavia and largely as non-native in central Europe. These results suggest an extinction of experience paralleling the extirpation or decline of salmon stocks in countries such as Germany and France. Respondents thought pollution was the dominant threat to riverine fish biodiversity. In reality, habitat loss, dams, and the spread of non-native fishes are equally important. Despite limited biological knowledge, respondents from all countries held an overwhelmingly proecological worldview, supported conservation stocking, and appreciated native fishes, although only a minority interacted with them directly. Differences among the 4 countries related to several conservation issues. For example, threats to biodiversity stemming from aquaculture were perceived as more prevalent in Norway compared with the other 3 countries. Promoting fish conservation based on charismatic species and use values of fishes may work well in countries with a strong economic and cultural link to the freshwater environment, such as Norway. In countries where people rather abstractly care for nature, focusing conservation messaging on broader ecosystem traits and non-use values of fishes is likely to win more support.

Effect of recreational-fisheries management on fish biodiversity in gravel pit lakes, with contrasts to unmanaged lakes.

Gravel pit lakes are novel ecosystems that can be colonized by fish through natural or anthropogenic pathways. In central Europe, many of them are managed by recreational anglers and thus experience regular fish stocking. However, also unmanaged gravel pits may be affected by stocking, either through illegal fish introductions or, occasionally, by immigration from connected water bodies. We sampled 23 small (< 20 ha) gravel pit lakes (16 managed and 7 unmanaged) in north-western Germany using littoral electrofishing and multimesh gillnets. Our objective was to compare the fish biodiversity in gravel pit lakes in the presence or absence of recreational fisheries. Given the size of the sampled lakes, we expected species poor communities and elevated fish diversity in the managed systems due to regular stocking of game fish species. Our study lakes were primarily mesotrophic and did not differ in key abiotic and biotic environmental characteristics. Lakes of both management types hosted similar fish abundances and biomasses, but were substantially different in terms of fish community structure and species richness. Fish were present in all lakes, with a minimum of three species. Higher α-diversity and lower ß-diversity was discovered in managed gravel pit lakes compared to unmanaged lakes. Consequently, recreational-fisheries management fostered homogenization of fish communities, by stocking a similar set of fish species desired by anglers such as piscivorous fish and large bodied cyprinids. However, unmanaged gravel pit lakes were also affected by human-mediated colonization, presumably by illegal fish releases. Hardly any non-native species were detected, suggesting that recreational-fisheries management did not foster the spread of exotic species in our study region.

Searching for responsible and sustainable recreational fisheries in the Anthropocene.

Recreational fisheries that use rod and reel (i.e., angling) operate around the globe in diverse freshwater and marine habitats, targeting many different gamefish species and engaging at least 220 million participants. The motivations for fishing vary extensively; whether anglers engage in catch-and-release or are harvest-oriented, there is strong potential for recreational fisheries to be conducted in a manner that is both responsible and sustainable. There are many examples of recreational fisheries that are well-managed where anglers, the angling industry and managers engage in responsible behaviours that both contribute to long-term sustainability of fish populations and the sector. Yet, recreational fisheries do not operate in a vacuum; fish populations face threats and stressors including harvest from other sectors as well as environmental change, a defining characteristic of the Anthropocene. We argue that the future of recreational fisheries and indeed many wild fish populations and aquatic ecosystems depends on having responsible and sustainable (R&S) recreational fisheries whilst, where possible, addressing, or at least lobbying for increased awareness about the threats to recreational fisheries emanating from outside the sector (e.g., climate change). Here, we first consider how the concepts of R&S intersect in the recreational fishing sector in an increasingly complex socio-cultural context. Next, we explore the role of the angler, angling industry and decision-makers in achieving R&S fisheries. We extend this idea further by considering the consequences of a future without recreational fisheries (either because of failures related to R&S) and explore a pertinent case study situated in Uttarakahand, India. Unlike other fisheries sectors where the number of participants is relatively small, recreational angling participants are numerous and widespread, such that if their actions are responsible, they have the potential to be a key voice for conservation and serve as a major force for good in the Anthropocene. What remains to be seen is whether this will be achieved, or if failure will occur to the point that recreational fisheries face increasing pressure to cease, as a result of external environmental threats, the environmental effects of recreational fishing and emerging ethical concerns about the welfare of angled fish.

The value artificial lake ecosystems provide to recreational anglers: Implications for management of biodiversity and outdoor recreation.

Small lakes largely outnumber large lakes among the world's lentic ecosystems. Despite being common landscape elements, however, little is known about the value small lakes provide to recreationists. This paper presents results of an economic valuation study concerned with small gravel pits in Lower Saxony, Germany. Gravel pits are artificially created lake ecosystems that, particularly in Europe, are regularly used and managed by privately organized recreational anglers in an angling club context. A stated choice experiment provided insights into anglers' preferences for the abundance of target fish species, biodiversity of taxa other than fishes in the lakes as well as in the lake surroundings, and about the use of gravel pits by other recreationists for walking, swimming or boating. Latent class analysis identified five segments that varied in preferences. For the majority of anglers, the value of angling at gravel pit lakes was improved by an increase in the abundance of predatory fishes. Additionally, the presence of aquatic and terrestrial endangered species at the lakes increased the value of the ecosystems as perceived by the majority of anglers. By contrast, the presence of other recreational uses reduced the value of angling, with swimming being considered the most disturbing, while the degree of shoreline development had the least impact on the recorded choices. The results suggest that managing the gravel pit lakes for high biodiversity and the presence of desired game fish species maximizes the value for anglers. However, also two smaller segments with anglers revealing lexicographic preferences were identified. These anglers expressed either strong preferences against swimming in the lakes or strongly preferred the opportunity to use boats. Lake management may address the preference heterogeneity and the aversion against other recreational uses, such as swimming, by spatial zoning.

Managing River Fish Biodiversity Generates Substantial Economic Benefits in Four European Countries.

Ecosystems and biodiversity produce benefits to society, but many of them are hard to quantify. For example, it is unclear whether European societies gain benefits from experiencing rivers that host high native biodiversity. Without such knowledge, monetary investments into ecologically oriented river management plans are difficult to justify. The objective of this study was to reveal how the public in four European countries values ecological characteristics of domestic rivers and the outcomes of hypothetical river basin management plans designed to improve river ecosystems, particularly fish biodiversity. We conducted a choice experiment among the populations in Norway, Sweden, Germany, and France. We found similar preference structures in all countries with high marginal willingness-to-pay for improvements of abiotic river attributes (increased accessiblity of the river banks, improved bathing water quality, decreased river fragmentation). Citizens also benefited from certain fish species occurring in a river with native salmonid species being more valued than nonnatives, particularly in Norway, and from the degree of a river's native biodiversity. Welfare measures calculated for selected river basin management plans (policy scenarios) revealed societal benefits that were primarily derived from ecological river management whereas a scenario focusing on hydroelectricity production generated the lowest utility. We conclude that ecological river management may produce high nonmarket economic benefits in all study countries, particularly through the management of abiotic river attributes and the restoration of declining or extinct fish species. Our results help to inform decisions on restoration efforts by showcasing the benefits that these measures have for the public.

How ecology shapes exploitation: a framework to predict the behavioural response of human and animal foragers along exploration-exploitation trade-offs.

Understanding how humans and other animals behave in response to changes in their environments is vital for predicting population dynamics and the trajectory of coupled social-ecological systems. Here, we present a novel framework for identifying emergent social behaviours in foragers (including humans engaged in fishing or hunting) in predator-prey contexts based on the exploration difficulty and exploitation potential of a renewable natural resource. A qualitative framework is introduced that predicts when foragers should behave territorially, search collectively, act independently or switch among these states. To validate it, we derived quantitative predictions from two models of different structure: a generic mathematical model, and a lattice-based evolutionary model emphasising exploitation and exclusion costs. These models independently identified that the exploration difficulty and exploitation potential of the natural resource controls the social behaviour of resource exploiters. Our theoretical predictions were finally compared to a diverse set of empirical cases focusing on fisheries and aquatic organisms across a range of taxa, substantiating the framework's predictions. Understanding social behaviour for given social-ecological characteristics has important implications, particularly for the design of governance structures and regulations to move exploited systems, such as fisheries, towards sustainability. Our framework provides concrete steps in this direction.

How ecological processes shape the outcomes of stock enhancement and harvest regulations in recreational fisheries.

Fish stocking and harvest regulations are frequently used to maintain or enhance freshwater recreational fisheries and contribute to fish conservation. However, their relative effectiveness has rarely been systematically evaluated using quantitative models that account for key size- and density-dependent ecological processes and adaptive responses of anglers. We present an integrated model of freshwater recreational fisheries where the population dynamics of two model species affect the effort dynamics of recreational anglers. With this model, we examined how stocking various fish densities and sizes (fry, fingerlings, and adults) performed relative to minimum-length limits using a variety of biological, social, and economic performance measures, while evaluating trade-offs. Four key findings are highlighted. First, stocking often augmented the exploited fish population, but size- and density-dependent bottlenecks limited the number of fry and fingerlings surviving to a catchable size in self-sustaining populations. The greatest enhancement of the catchable fish population occurred when large fish that escaped early bottlenecks were stocked, but this came at the cost of wild-stock replacement, thereby demonstrating a fundamental trade-off between fisheries benefits and conservation. Second, the relative performance of stocking naturally reproducing populations was largely independent of habitat quality and was generally low. Third, stocking was only economically advisable when natural reproduction was impaired or absent, stocking rates were low, and enough anglers benefitted from stocking to offset the associated costs. Fourth, in self-sustaining fish populations, minimum-length limits generally outperformed stocking when judged against a range of biological, social and economic objectives. By contrast, stocking in culture-based fisheries often generated substantial benefits. Collectively, our study demonstrates that size- and density-dependent processes, and broadly the degree of natural recruitment, drive the biological, social, and economic outcomes of popular management actions in recreational fisheries. To evaluate these outcomes and the resulting trade-offs, integrated fisheries-management models that explicitly consider the feedbacks among ecological and social processes are needed.

Rapid, broad-scale gene expression evolution in experimentally harvested fish populations.

Gene expression changes potentially play an important role in adaptive evolution under human-induced selection pressures, but this has been challenging to demonstrate in natural populations. Fishing exhibits strong selection pressure against large body size, thus potentially inducing evolutionary changes in life history and other traits that may be slowly reversible once fishing ceases. However, there is a lack of convincing examples regarding the speed and magnitude of fisheries-induced evolution, and thus, the relevant underlying molecular-level effects remain elusive. We use wild-origin zebrafish (Danio rerio) as a model for harvest-induced evolution. We experimentally demonstrate broad-scale gene expression changes induced by just five generations of size-selective harvesting, and limited genetic convergence following the cessation of harvesting. We also demonstrate significant allele frequency changes in genes that were differentially expressed after five generations of size-selective harvesting. We further show that nine generations of captive breeding induced substantial gene expression changes in control stocks likely due to inadvertent selection in the captive environment. The large extent and rapid pace of the gene expression changes caused by both harvest-induced selection and captive breeding emphasizes the need for evolutionary enlightened management towards sustainable fisheries.

Fast-slow life history is correlated with individual differences in movements and prey selection in an aquatic predator in the wild.

Fast and slow life histories are proposed to covary with consistent individual differences in behaviour, but little is known whether it holds in the wild, where individuals experience natural fluctuations of the environment. We investigated whether individual differences in behaviour, such as movement traits and prey selection, are linked to variation in life-history traits in Eurasian perch (Perca fluviatilis) in the wild. Using high-resolution acoustic telemetry, we collected the positional data of fish in a whole natural lake and estimated individual movement traits by fitting a two-state correlated random walk model. Prey selection was inferred from stable isotope analysis using scale samples. Life-history traits were estimated by fitting a biphasic growth model to an individual growth trajectory back-calculated from scale samples. Life-history traits were correlated with behavioural traits such as movements and prey selection. Individuals with higher reproductive effort were found to switch more frequently between active and inactive modes and show greater reliance on prey from pelagic pathways (indicated by lower δ13 C). Further, individuals with faster juvenile growth were found to stay active for a longer time during the adult stage. Our results demonstrate the link between individual behavioural differences and fast-slow life-history traits under ecologically relevant conditions.

Why social values cannot be changed for the sake of conservation.

The hope for creating widespread change in social values has endured among conservation professionals since early calls by Aldo Leopold for a "land ethic." However, there has been little serious attention in conservation to the fields of investigation that address values, how they are formed, and how they change. We introduce a social-ecological systems conceptual approach in which values are seen not only as motivational goals people hold but also as ideas that are deeply embedded in society's material culture, collective behaviors, traditions, and institutions. Values define and bind groups, organizations, and societies; serve an adaptive role; and are typically stable across generations. When abrupt value changes occur, they are in response to substantial alterations in the social-ecological context. Such changes build on prior value structures and do not result in complete replacement. Given this understanding of values, we conclude that deliberate efforts to orchestrate value shifts for conservation are unlikely to be effective. Instead, there is an urgent need for research on values with a multilevel and dynamic view that can inform innovative conservation strategies for working within existing value structures. New directions facilitated by a systems approach will enhance understanding of the role values play in shaping conservation challenges and improve management of the human component of conservation.