Watershed features shape spatial patterns of fish tissue mercury in a boreal river network.

Freshwater mercury (Hg) contamination is a widespread environmental concern but how proximate sources and downstream transport shape Hg spatial patterns in riverine food webs is poorly understood. We measured total Hg (THg) in slimy sculpin (Cottus cognatus) across the Kuskokwim River, a large boreal river in western Alaska and home to subsistence fishing communities which rely on fish for primary nutrition. We used spatial stream network models (SSNMs) to quantify watershed and instream conditions influencing sculpin THg. Spatial covariates for local watershed geology and slope accounted for 55 % of observed variation in sculpin THg and evidence for downstream transport of Hg in sculpins was weak. Empirical semivariograms indicated these spatial covariates accounted for most spatial autocorrelation in observed THg. Watershed geology and slope explained up to 70 % of sculpin THg variation when SSNMs accounted for instream spatial dependence. Our results provide network-wide predictions for fish tissue THg based largely on publicly available geospatial data and open-source software for SSNMs, and demonstrate how these emerging models can be used to understand contaminant behavior in spatially complex aquatic ecosystems.

Declines in body size of sockeye salmon associated with increased competition in the ocean.

Declining body sizes have been documented for several species of Pacific salmon; however, whether size declines are caused mainly by ocean warming or other ecological factors, and whether they result primarily from trends in age at maturation or changing growth rates remain poorly understood. We quantified changes in mean body size and contributions from shifting size-at-age and age structure of mature sockeye salmon returning to Bristol Bay, Alaska, over the past 60 years. Mean length declined by 3%, corresponding to a 10% decline in mean body mass, since the early 1960s, though much of this decline occurred since the early 2000s. Changes in size-at-age were the dominant cause of body size declines and were more consistent than trends in age structure among the major rivers that flow into Bristol Bay. Annual variation in size-at-age was largely explained by competition among Bristol Bay sockeye salmon and interspecific competition with other salmon in the North Pacific Ocean. Warm winters were associated with better growth of sockeye salmon, whereas warm summers were associated with reduced growth. Our findings point to competition at sea as the main driver of sockeye salmon size declines, and emphasize the trade-off between fish abundance and body size.

Getting ahead of climate change for ecological adaptation and resilience.

Changing the course of Earth's climate is increasingly urgent, but there is also a concurrent need for proactive stewardship of the adaptive capacity of the rapidly changing biosphere. Adaptation ultimately underpins the resilience of Earth's complex systems; species, communities, and ecosystems shift and evolve over time. Yet oncoming changes will seriously challenge current natural resource management and conservation efforts. We review forward-looking conservation approaches to enable adaptation and resilience. Key opportunities include expanding beyond preservationist approaches by including those that enable and facilitate ecological change. Conservation should not just focus on climate change losers but also on proactive management of emerging opportunities. Local efforts to conserve biodiversity and generate habitat complexity will also help to maintain a diversity of future options for an unpredictable future.

Assessing the potential for demographic restoration and assisted evolution to build climate resilience in coral reefs.

Interest is growing in developing conservation strategies to restore and maintain coral reef ecosystems in the face of mounting anthropogenic stressors, particularly climate warming and associated mass bleaching events. One such approach is to propagate coral colonies ex situ and transplant them to degraded reef areas to augment habitat for reef-dependent fauna, prevent colonization from spatial competitors, and enhance coral reproductive output. In addition to such "demographic restoration" efforts, manipulating the thermal tolerance of outplanted colonies through assisted relocation, selective breeding, or genetic engineering is being considered for enhancing rates of evolutionary adaptation to warming. Although research into such "assisted evolution" strategies has been growing, their expected performance remains unclear. We evaluated the potential outcomes of demographic restoration and assisted evolution in climate change scenarios using an eco-evolutionary simulation model. We found that supplementing reefs with pre-existing genotypes (demographic restoration) offers little climate resilience benefits unless input levels are large and maintained for centuries. Supplementation with thermally resistant colonies was successful at improving coral cover at lower input levels, but only if maintained for at least a century. Overall, we found that, although demographic restoration and assisted evolution have the potential to improve long-term coral cover, both approaches had a limited impact in preventing severe declines under climate change scenarios. Conversely, with sufficient natural genetic variance and time, corals could readily adapt to warming temperatures, suggesting that restoration approaches focused on building genetic variance may outperform those based solely on introducing heat-tolerant genotypes.

Glacier retreat creating new Pacific salmon habitat in western North America.

Glacier retreat poses risks and benefits for species of cultural and economic importance. One example is Pacific salmon (Oncorhynchus spp.), supporting subsistence harvests, and commercial and recreational fisheries worth billions of dollars annually. Although decreases in summer streamflow and warming freshwater is reducing salmon habitat quality in parts of their range, glacier retreat is creating new streams and lakes that salmon can colonize. However, potential gains in future salmon habitat associated with glacier loss have yet to be quantified across the range of Pacific salmon. Here we project future gains in Pacific salmon freshwater habitat by linking a model of glacier mass change for 315 glaciers, forced by five different Global Climate Models, with a simple model of salmon stream habitat potential throughout the Pacific Mountain ranges of western North America. We project that by the year 2100 glacier retreat will create 6,146 (±1,619) km of new streams accessible for colonization by Pacific salmon, of which 1,930 (±569) km have the potential to be used for spawning and juvenile rearing, representing 0 to 27% gains within the 18 sub-regions we studied. These findings can inform proactive management and conservation of Pacific salmon in this era of rapid climate change.

Evolution and connectivity influence the persistence and recovery of coral reefs under climate change in the Caribbean, Southwest Pacific, and Coral Triangle.

Corals are experiencing unprecedented decline from climate change-induced mass bleaching events. Dispersal not only contributes to coral reef persistence through demographic rescue but can also hinder or facilitate evolutionary adaptation. Locations of reefs that are likely to survive future warming therefore remain largely unknown, particularly within the context of both ecological and evolutionary processes across complex seascapes that differ in temperature range, strength of connectivity, network size, and other characteristics. Here, we used eco-evolutionary simulations to examine coral adaptation to warming across reef networks in the Caribbean, the Southwest Pacific, and the Coral Triangle. We assessed the factors associated with coral persistence in multiple reef systems to understand which results are general and which are sensitive to particular geographic contexts. We found that evolution can be critical in preventing extinction and facilitating the long-term recovery of coral communities in all regions. Furthermore, the strength of immigration to a reef (destination strength) and current sea surface temperature robustly predicted reef persistence across all reef networks and across temperature projections. However, we found higher initial coral cover, slower recovery, and more evolutionary lag in the Coral Triangle, which has a greater number of reefs and more larval settlement than the other regions. We also found the lowest projected future coral cover in the Caribbean. These findings suggest that coral reef persistence depends on ecology, evolution, and habitat network characteristics, and that, under an emissions stabilization scenario (RCP 4.5), recovery may be possible over multiple centuries.

Evolution reverses the effect of network structure on metapopulation persistence.

Global environmental change is challenging species with novel conditions, such that demographic and evolutionary trajectories of populations are often shaped by the exchange of organisms and alleles across landscapes. Current ecological theory predicts that random networks with dispersal shortcuts connecting distant sites can promote persistence when there is no capacity for evolution. Here, we show with an eco-evolutionary model that dispersal shortcuts across environmental gradients instead hinder persistence for populations that can evolve because long-distance migrants bring extreme trait values that are often maladaptive, short-circuiting the adaptive response of populations to directional change. Our results demonstrate that incorporating evolution and environmental heterogeneity fundamentally alters theoretical predictions regarding persistence in ecological networks.

Y-chromosome haplotypes are associated with variation in size and age at maturity in male Chinook salmon.

Variation in size and age at maturity is an important component of life history that is influenced by both environmental and genetic factors. In salmonids, large size confers a direct reproductive advantage through increased fecundity and egg quality in females, while larger males gain a reproductive advantage by monopolizing access to females. In addition, variation in size and age at maturity in males can be associated with different reproductive strategies; younger smaller males may gain reproductive success by sneaking among mating pairs. In both sexes, there is a trade-off between older age and increased reproductive success and increased risk of mortality by delaying reproduction. We identified four Y-chromosome haplogroups that showed regional- and population-specific variation in frequency using RADseq data for 21 populations of Alaska Chinook salmon. We then characterized the range-wide distribution of these haplogroups using GT-seq assays. These haplogroups exhibited associations with size at maturity in multiple populations, suggesting that lack of recombination between X and Y-chromosomes has allowed Y-chromosome haplogroups to capture different alleles that influence size at maturity. Ultimately, conservation of life history diversity in Chinook salmon may require conservation of Y-chromosome haplotype diversity.

Deeper waters are changing less consistently than surface waters in a global analysis of 102 lakes.

Globally, lake surface water temperatures have warmed rapidly relative to air temperatures, but changes in deepwater temperatures and vertical thermal structure are still largely unknown. We have compiled the most comprehensive data set to date of long-term (1970-2009) summertime vertical temperature profiles in lakes across the world to examine trends and drivers of whole-lake vertical thermal structure. We found significant increases in surface water temperatures across lakes at an average rate of + 0.37 °C decade-1, comparable to changes reported previously for other lakes, and similarly consistent trends of increasing water column stability (+ 0.08 kg m-3 decade-1). In contrast, however, deepwater temperature trends showed little change on average (+ 0.06 °C decade-1), but had high variability across lakes, with trends in individual lakes ranging from - 0.68 °C decade-1 to + 0.65 °C decade-1. The variability in deepwater temperature trends was not explained by trends in either surface water temperatures or thermal stability within lakes, and only 8.4% was explained by lake thermal region or local lake characteristics in a random forest analysis. These findings suggest that external drivers beyond our tested lake characteristics are important in explaining long-term trends in thermal structure, such as local to regional climate patterns or additional external anthropogenic influences.

Individual behavior drives ecosystem function and the impacts of harvest.

Current approaches for biodiversity conservation and management focus on sustaining high levels of diversity among species to maintain ecosystem function. We show that the diversity among individuals within a single population drives function at the ecosystem scale. Specifically, nutrient supply from individual fish differs from the population average >80% of the time, and accounting for this individual variation nearly doubles estimates of nutrients supplied to the ecosystem. We test how management (i.e., selective harvest regimes) can alter ecosystem function and find that strategies targeting more active individuals reduce nutrient supply to the ecosystem up to 69%, a greater effect than body size-selective or nonselective harvest. Findings show that movement behavior at the scale of the individual can have crucial repercussions for the functioning of an entire ecosystem, proving an important challenge to the species-centric definition of biodiversity if the conservation and management of ecosystem function is a primary goal.

Glacier Retreat and Pacific Salmon.

Glaciers have shaped past and present habitats for Pacific salmon (Oncorhynchus spp.) in North America. During the last glacial maximum, approximately 45% of the current North American range of Pacific salmon was covered in ice. Currently, most salmon habitat occurs in watersheds in which glacier ice is present and retreating. This synthesis examines the multiple ways that glacier retreat can influence aquatic ecosystems through the lens of Pacific salmon life cycles. We predict that the coming decades will result in areas in which salmon populations will be challenged by diminished water flows and elevated water temperatures, areas in which salmon productivity will be enhanced as downstream habitat suitability increases, and areas in which new river and lake habitat will be formed that can be colonized by anadromous salmon. Effective conservation and management of salmon habitat and populations should consider the impacts of glacier retreat and other sources of ecosystem change.

Resurgence of an apex marine predator and the decline in prey body size.

In light of recent recoveries of marine mammal populations worldwide and heightened concern about their impacts on marine food webs and global fisheries, it has become increasingly important to understand the potential impacts of large marine mammal predators on prey populations and their life-history traits. In coastal waters of the northeast Pacific Ocean, marine mammals have increased in abundance over the past 40 to 50 y, including fish-eating killer whales that feed primarily on Chinook salmon. Chinook salmon, a species of high cultural and economic value, have exhibited marked declines in average size and age throughout most of their North American range. This raises the question of whether size-selective predation by marine mammals is generating these trends in life-history characteristics. Here we show that increased predation since the 1970s, but not fishery selection alone, can explain the changes in age and size structure observed for Chinook salmon populations along the west coast of North America. Simulations suggest that the decline in mean size results from the selective removal of large fish and an evolutionary shift toward faster growth and earlier maturation caused by selection. Our conclusion that intensifying predation by fish-eating killer whales contributes to the continuing decline in Chinook salmon body size points to conflicting management and conservation objectives for these two iconic species.

Does lipid-correction introduce biases into isotopic mixing models? Implications for diet reconstruction studies.

Carbon isotopes are commonly used in trophic ecology to estimate consumer diet composition. This estimation is complicated by the fact that lipids exhibit a more depleted carbon signature (δ13C) than other macromolecules, and are often found at different concentrations among individual organisms. Some researchers argue that lipids bias diet reconstructions using stable isotopes and should be accounted for prior to analysis in food web mixing models, whereas others contend that removing lipids may result in erroneous interpretations of the trophic interactions under study. To highlight this disagreement on best practices for applying δ13C in food web studies, we sampled the recent literature to determine the frequency and method of lipid-correction. We then quantified the potential magnitude and source of bias in mixing model results from a theoretical example and case study of diet reconstruction. The literature was split nearly evenly as to whether lipid-correction was applied to δ13C data in mixing model estimates of diet composition. Comparative mixing model scenarios demonstrated that lipid-correction can substantially alter the estimated diet composition and interpretation of consumer foraging habits. Given the lack of consensus on whether or not to lipid-correct prey and/or consumers, and the associated variation in mixing model results, we call for the establishment of a unified framework that will guide diet reconstruction in stable isotope ecology. Uncertainty in the prevalence of direct routing versus de novo synthesis of lipids across ecosystems, taxa, and trophic levels must be resolved to better guide treatment of lipids in isotope studies using carbon.

Shifting habitat mosaics and fish production across river basins.

Watersheds are complex mosaics of habitats whose conditions vary across space and time as landscape features filter overriding climate forcing, yet the extent to which the reliability of ecosystem services depends on these dynamics remains unknown. We quantified how shifting habitat mosaics are expressed across a range of spatial scales within a large, free-flowing river, and how they stabilize the production of Pacific salmon that support valuable fisheries. The strontium isotope records of ear stones (otoliths) show that the relative productivity of locations across the river network, as both natal- and juvenile-rearing habitat, varies widely among years and that this variability is expressed across a broad range of spatial scales, ultimately stabilizing the interannual production of fish at the scale of the entire basin.

Effects of warming climate and competition in the ocean for life-histories of Pacific salmon.

The life-histories of exploited fish species, such as Pacific salmon, are vulnerable to a wide variety of anthropogenic stressors including climate change, selective exploitation and competition with hatchery releases for finite foraging resources. However, these stressors may generate unexpected changes in life-histories due to developmental linkages when species complete their migratory life cycle in different habitats. We used multivariate time-series models to quantify changes in the prevalence of different life-history strategies of sockeye salmon from Bristol Bay, Alaska, over the past half-century-specifically, how they partition their lives between freshwater habitats and the ocean. Climate warming has decreased the time spent by salmon in their natal freshwater habitat, as climate-enhanced growth opportunities have enabled earlier migration to the ocean. Migration from freshwater at a younger age, and increasing competition from wild and hatchery-released salmon, have tended to delay maturation toward the salmon spending an additional year feeding in the ocean. Models evaluating the effects of size-selective fishing on these patterns had only small support. These stressors combine to reduce the size-at-age of fish vulnerable to commercial fisheries and have increasingly favoured a single-age class, potentially affecting the age class complexity that stabilizes this highly reliable resource.

Isotopes in teeth and a cryptic population of coastal freshwater seals.

Human activities threaten the biodiversity of aquatic mammals across the globe. Conservation of these species hinges on the ability to delineate movements and foraging behaviors of animals, but gaining such insights is hampered by difficulties in tracing individuals over their lives. We determined isotope ratios in teeth (87 Sr/86 Sr, 13 C/12 C, and 18 O/16 O) to examine lifelong movement and resource-use patterns of a unique freshwater population of a wide-ranging pinniped species (harbor seal [Phoca vitulina]) that resides in Iliamna Lake, Alaska (U.S.A.). This population's potentially unique migratory behavior and use of different trophic resources are unknown. The isotope ratios we measured in teeth showed that seals were born in the lake, remained lifelong residents, and relied principally on resources produced from in the lake, even when seasonally abundant and nutrient-dense spawning anadromous fish (i.e., sockeye salmon [Oncorhynchus nerka]) were available in the lake. Our results illustrate how serial isotope records in teeth, particularly 87 Sr/86 Sr ratios, can be used to quantify how coastal mammal populations exploit both freshwater and marine ecosystems. Understanding lifelong patterns of habitat and resource use is essential information when designing effective conservation plans for threatened coastal mammals. We present the Iliamna Lake harbor seals as a unique case study into how isotope records within teeth can help reveal the cryptic ecology of such a population residing in an intact ecosystem. The results also provide critical baseline information for the Kvichak River system, which is facing an uncertain future due to proposed large-scale industrial development and a rapidly changing climate.

Isotopos Dentales y una Población Críptica de Focas Costeras de Agua Dulce Resumen Las actividades humanas amenazan a la diversidad de mamíferos acuáticos en todo el mundo. La conservación de estas especies depende de la habilidad para delinear los movimientos y los comportamientos de búsqueda de alimento de los animales, pero la obtención de dicha información está obstaculizada por las dificultades en el rastreo de individuos a lo largo del transcurso de sus vidas. Determinamos la proporción de isotopos dentales (87 Sr/86 Sr, 13 C/12 C y 18 O/16 O) para examinar el movimiento a lo largo de la vida y los patrones de uso de recursos de una población única de una especie de pinnípedos de agua dulce con una distribución amplia (foca común [Phoca vitulina]), la cual reside en el lago Iliamna, Alaska (E.U.A.). Se desconocen el comportamiento migratorio potencialmente único de esta población y el uso que le dan a los diferentes recursos tróficos. La proporción de isotopos que medimos en los dientes mostró que las focas nacieron en el lago, permanecieron como residentes de toda la vida y dependieron principalmente de los recursos producidos en el lago, incluso cuando estaban disponibles en aquel lugar por razones reproductivas los peces anádromos abundantes estacionalmente y con densidad de nutrientes (es decir, el salmón rojo [Oncorhynchus nerka]). Nuestros resultados ilustran cómo los registros seriales de isotopos dentales, particularmente la proporción 87 Sr/86 Sr, pueden usarse para cuantificar cómo las poblaciones de mamíferos costeros explotan tanto los ecosistemas marinos como los de agua dulce. El entendimiento de los patrones ontogénicos del uso de recursos y de hábitat es esencial cuando se diseñan planes efectivos de conservación para los mamíferos costeros en peligro. Presentamos a las focas comunes del lago Iliamna como un estudio de caso único sobre cómo los registros de isotopos dentales pueden ayudar a revelar la ecología críptica de dicha población que reside en un ecosistema intacto. Los resultados también proporcionan información importante de línea base para el sistema el río Kvichak, el cual está enfrentando un futuro incierto debido a la propuesta de un desarrollo industrial de gran escala y al rápido clima cambiante.

Recruitment variation disrupts the stability of alternative life histories in an exploited salmon population.

Males of many fish species exhibit alternative reproductive tactics, which can influence the maturation schedules, fishery productivity, and resilience to harvest of exploited populations. While alternative mating phenotypes can persist in stable equilibria through frequency-dependent selection, shifts in tactic frequencies have been observed and can have substantial consequences for fisheries. Here, we examine the dynamics of precocious sneaker males called "jacks" in a population of sockeye salmon (Oncorhynchus nerka) from Frazer Lake, Alaska. Jacks, which are of little commercial value due to their small body sizes, have recently been observed at unusually high levels in this stock, degrading the value of regional fisheries. To inform future strategies for managing the prevalence of jacks, we used long-term monitoring data to identify what regulates the frequencies of alternative male phenotypes in the population over time. Expression of the jack life history could not be explained by environmental factors expected to influence juvenile body condition and maturation probability. Instead, we found a strong positive association between the proportion of individuals maturing as jacks within a cohort and the prevalence of jacks among the males that sired that cohort. Moreover, due to differences in age-at-maturity between male phenotypes, and large interannual variability in recruitment strength, jacks from strong year-classes often spawn among older males from the weaker recruitments of earlier cohorts. Through such "cohort mismatches," which are amplified by size-selective harvest on older males, jacks frequently achieve substantial representation in the breeding population, and likely high total fertilizations. The repeated occurrence of these cohort mismatches appears to disrupt the stabilizing influence of frequency-dependent selection, allowing the prevalence of jacks to exceed what might be expected under equilibrium conditions. These results emphasize that the dynamics of alternative life histories can profoundly influence fishery performance and should be explicitly considered in the management of exploited populations.

The phenology of migration in an unpredictable world.

In Focus: Freshwater, C., Trudel, M., Beacham, T. D., Gauthier, S., Johnson, S. C., Neville, C. & Juanes, F. (2016) Individual variation, population-specific migration behaviours and stochastic processes shape marine migration phenologies. Journal of Animal Ecology, 88, 67-78. https://doi.org/10.1111/1365-2656.12852 Pacific salmon undertake arduous and risky migrations from their freshwater nursery grounds to the coastal ocean, northwards to their feeding grounds, and then back to their freshwater natal habitats to spawn. Understanding the phenology of such migrations has largely been viewed through the lens of microevolution producing optimal strategies that reflect local selection pressures; less emphasis has been placed on quantifying how variation in migration patterns can spread the risks associated with life in variable and unpredictable ecosystems. In this issue, Freshwater et al. use the information contained in ear stones (otoliths) and DNA of migrating juvenile sockeye salmon from the Fraser River of western Canada to quantify variation in the timing of their marine migrations. Not only were there population-specific differences in migration phenology of fish from the same river, but there was substantial variation among individuals from specific populations. These patterns also varied from year to year. Data like these emphasize the risks involved in such migrations and suggest that variation in key migration traits are maintained because of the inherent unpredictability of ecosystems. Management and conservation efforts would be well-served to consider actions that maintain such ecological variation to facilitate meta-population persistence in a rapidly changing world.

Spatial heterogeneity contributes more to portfolio effects than species variability in bottom-associated marine fishes.

Variance of community abundance will be reduced relative to its theoretical maximum whenever population densities fluctuate asynchronously. Fishing communities and mobile predators can switch among fish species and/or fishing locations with asynchronous dynamics, thereby buffering against variable resource densities (termed 'portfolio effects', PEs). However, whether variation among species or locations represent the dominant contributor to PE remains relatively unexplored. Here, we apply a spatio-temporal model to multidecadal time series (1982-2015) for 20 bottom-associated fishes in seven marine ecosystems. For each ecosystem, we compute the reduction in variance over time in total biomass relative to its theoretical maximum if species and locations were perfectly correlated (total PE). We also compute the reduction in variance due to asynchrony among species at each location (species PE) or the reduction due to asynchrony among locations for each species (spatial PE). We specifically compute total, species and spatial PE in 10-year moving windows to detect changes over time. Our analyses revealed that spatial PE are stronger than species PE in six of seven ecosystems, and that ecosystems where species PE is constant over time can exhibit shifts in locations that strongly contribute to PE. We therefore recommend that spatial and total PE be monitored as ecosystem indicators representing risk exposure for human and natural consumers.