Weakened portfolio effects constrain management effectiveness for population aggregates.

Population diversity can reduce temporal variability in aggregate population abundances in a process known as the portfolio effect. Portfolio effects may weaken, however, due to greater synchrony among component populations. While weakened portfolio effects have been previously documented, the consequences of reduced stability on meeting conservation goals for population aggregates that are harvested (e.g., stock aggregates in fisheries) are rarely quantified. Here, we demonstrate how changes in variability within components, synchrony among components, and population productivity interact to influence the probability of achieving an array of management objectives for Fraser River sockeye salmon: a stock aggregate of high economic, ecological, and cultural value. We first present evidence that component variability and synchrony have increased over the last two decades, consistent with a weakening portfolio effect. We then parameterize a stochastic, closed-loop model that simulates the population dynamics of each stock, the fishery that harvests the stock aggregate, and the management framework used to establish mixed-stock exploitation rates. We find that while median aggregate abundance and catch through time were relatively insensitive to greater aggregate variability, catch stability and performance metrics associated with achieving management targets generally declined as component variability and synchrony increased. A notable exception we observed is that harvest control means that scale exploitation rates based on aggregate abundance may be more effective as synchrony increases. Reductions in productivity led to broad declines in performance, but also moderated the impacts of component variability and synchrony on the proportion of component stocks above management targets and catch stability. Our results suggest that even precautionary management strategies that account for declines in productivity may underestimate risk, particularly to socioeconomic objectives, if they fail to consider changes in aggregate variability. Adequately incorporating changes in portfolio effect strength may be particularly relevant when developing recovery strategies that are robust to climate change, which is likely to increase synchrony and component variability.

Individual variation, population-specific behaviours and stochastic processes shape marine migration phenologies.

The phenology of long-distance migrations can influence individual fitness, moderate population dynamics and regulate the availability of ecosystem services to other trophic levels. Phenology varies within and among populations, and can be influenced by conditions individuals experience both prior to departure and encounter en route. Assessing how intrinsic and extrinsic factors (e.g., individual physical condition vs. environmental conditions) interact to influence variation in migratory phenologies across ecological scales is often limited due to logistical constraints associated with tracking large numbers of individuals from multiple populations simultaneously. We used two natural tags, DNA and otolith microstructure analysis, to estimate the relative influence of individual traits (life-history strategy, body size at departure and growth during migration), population-specific behaviours and interannual variability on the phenology of marine migrations in juvenile sockeye salmon Oncorhynchus nerka. We show that the timing and duration of juvenile sockeye salmon migrations were correlated with both life-history strategy and body size, while migration duration was also correlated with departure timing and growth rates during migration. Even after accounting for the effect of individual traits, several populations exhibited distinct migration phenologies. Finally, we observed substantial interannual and residual variation, suggesting stochastic environmental conditions moderate the influence of carry-over effects that develop prior to departure, as well as population-specific strategies. Migratory phenologies are shaped by complex interactions between drivers acting at multiple ecological and temporal scales. Given evidence that intraspecific diversity can stabilize ecological systems, conservation efforts should seek to maintain migratory variation among populations and preserve locally adapted phenotypes; however, variation within populations, which may buffer systems from environmental stochasticity, should also be regularly assessed and preserved.

Repeated patterns of trait divergence between closely related dominant and subordinate bird species.

Ecologically similar species often compete aggressively for shared resources. These interactions are frequently asymmetric, with one species behaviorally dominant to another and excluding it from preferred resources. Despite the potential importance of this type of interference competition as a source of selection, we know little about patterns of trait divergence between dominant and subordinate species. We compiled published data on phylogenetically independent, closely related species of North American birds where one species was consistently dominant in aggressive interactions with a congeneric species. We then compared the body size, breeding phenology, life history, ecological breadth, and biogeography of these species. After accounting for body size and phylogeny, we found repeated patterns of trait divergence between subordinate and dominant species within genera. Subordinate species that migrated seasonally arrived 4-7 days later than dominants on their sympatric breeding grounds, and both resident and migratory subordinates initiated breeding 7-8 days later than their dominant, sympatric congeners. Subordinate species had a 5.2% higher annual adult mortality rate and laid eggs that were 0.02 g heavier for their body mass. Dominant and subordinate species used a similar number of different foods, foraging behaviors, nest sites, and habitats, but subordinates were more specialized in their foraging behaviors compared with closely related dominant species. The breeding and wintering ranges of subordinate species were 571 km farther apart than the ranges of dominant species, suggesting that subordinate species migrate greater distances. Range sizes and latitudinal distributions did not consistently differ, although subordinate species tended to breed farther north or winter farther south. These results are consistent with dominant species directly influencing the ecological strategies of subordinate species (via plastic or genetically based changes), either by restricting their access to resources or simply through aggression. Alternatively, these ecological traits may covary with patterns of behavioral dominance, with no direct consequences of interactions. Regardless of the mechanism, recognizing that the relative position of a species within a dominance hierarchy is correlated with a suite of other ecological and fitness related traits has far-reaching implications for the mechanisms underlying species distributions and the structure of biological assemblages.

Chinook salmon depth distributions on the continental shelf are shaped by interactions between location, season, and individual condition.

BACKGROUND: Ecological and physical conditions vary with depth in aquatic ecosystems, resulting in gradients of habitat suitability. Although variation in vertical distributions among individuals provides evidence of habitat selection, it has been challenging to disentangle how processes at multiple spatio-temporal scales shape behaviour. METHODS: We collected thousands of observations of depth from > 300 acoustically tagged adult Chinook salmon Oncorhynchus tshawytscha, spanning multiple seasons and years. We used these data to parameterize a machine-learning model to disentangle the influence of spatial, temporal, and dynamic oceanographic variables while accounting for differences in individual condition and maturation stage. RESULTS: The top performing machine learning model used bathymetric depth ratio (i.e., individual depth relative to seafloor depth) as a response. We found that bathymetry, season, maturation stage, and spatial location most strongly influenced Chinook salmon depth. Chinook salmon bathymetric depth ratios were deepest in shallow water, during winter, and for immature individuals. We also identified non-linear interactions among covariates, resulting in spatially-varying effects of zooplankton concentration, lunar cycle, temperature and oxygen concentration. CONCLUSIONS: Our results suggest Chinook salmon vertical habitat use is a function of ecological interactions, not physiological constraints. Temporal and spatial variation in depth distributions could be used to guide management decisions intended to reduce fishery impacts on Chinook salmon. More generally, our findings demonstrate how complex interactions among bathymetry, seasonality, location, and life history stage regulate vertical habitat selection.

An integrated model of seasonal changes in stock composition and abundance with an application to Chinook salmon.

Population-specific spatial and temporal distribution data are necessary to identify mechanisms regulating abundance and to manage anthropogenic impacts. However the distributions of highly migratory species are often difficult to resolve, particularly when multiple populations' movements overlap. Here we present an integrated model to estimate spatially-stratified, seasonal trends in abundance and population composition, using data from extensive genetic sampling of commercial and recreational Chinook salmon (Oncorhynchus tshawytscha) fisheries in southern British Columbia. We use the model to estimate seasonal changes in population-specific standardized catch per unit effort (a proxy for abundance) across six marine regions, while accounting for annual variability in sampling effort and uncertain genetic stock assignment. We also share this model as an R package stockseasonr for application to other regions and species. Even at the relatively small spatial scales considered here, we found that patterns in seasonal abundance differed among regions and stocks. While certain locations were clearly migratory corridors, regions within the Salish Sea exhibited diverse, and often weak, seasonal patterns in abundance, emphasizing that they are important, year-round foraging habitats. Furthermore, we found evidence that stocks with similar freshwater life histories and adult run timing, as well as relatively proximate spawning locations, exhibited divergent distributions. Our findings highlight subtle, but important differences in how adult Chinook salmon use marine habitats. Down-scaled model outputs could be used to inform ecosystem-based management efforts by resolving the degree to which salmon overlap with other species of concern, as well as specific fisheries. More broadly, variation in stock-specific abundance among regions indicates efforts to identify mechanisms driving changes in size-at-maturity and natural mortality should account for distinct marine distributions.

Diversification insulates fisher catch and revenue in heavily exploited tropical fisheries.

Declines in commercial landings and increases in fishing fleet power have raised concerns over the continued provisioning of nutritional and economic services by tropical wild fisheries. Yet, because tropical fisheries are often data-poor, mechanisms that might buffer fishers to declines are not understood. This data scarcity undermines fisheries management, making tropical fishing livelihoods particularly vulnerable to changes in marine resources. We use high-resolution fisheries data from Seychelles to understand how fishing strategy (catch diversification) influences catch rates and revenues of individual fishing vessels. We show that average catch weight decreased by 65% over 27 years, with declines in all nine species groups coinciding with increases in fishing effort. However, for individual vessels, catch diversity was associated with larger catches and higher fishing revenues and with slower catch declines from 1990 to 2016. Management strategies should maximize catch diversity in data-poor tropical fisheries to help secure nutritional security while protecting fishing livelihoods.

The outcomes of most aggressive interactions among closely related bird species are asymmetric.

Aggressive interactions among closely related species are common, and can play an important role as a selective pressure shaping species traits and assemblages. The nature of this selective pressure depends on whether the outcomes of aggressive contests are asymmetric between species (i.e., one species is consistently dominant), yet few studies have estimated the prevalence of asymmetric versus symmetric outcomes to aggressive contests. Here we use previously published data involving 26,212 interactions between 270 species pairs of birds from 26 taxonomic families to address the question: How often are aggressive interactions among closely related bird species asymmetric? We define asymmetry using (i) the proportion of contests won by one species, and (ii) statistical tests for asymmetric outcomes of aggressive contests. We calculate these asymmetries using data summed across different sites for each species pair, and compare results to asymmetries calculated using data separated by location. We find that 80% of species pairs had aggressive outcomes where one species won 80% or more of aggressive contests. We also find that the majority of aggressive interactions among closely related species show statistically significant asymmetries, and above a sample size of 52 interactions, all outcomes are asymmetric following binomial tests. Species pairs with dominance data from multiple sites showed the same dominance relationship across locations in 93% of the species pairs. Overall, our results suggest that the outcome of aggressive interactions among closely related species are usually consistent and asymmetric, and should thus favor ecological and evolutionary strategies specific to the position of a species within a dominance hierarchy.

Trends in extinction risk for imperiled species in Canada.

Protecting and promoting recovery of species at risk of extinction is a critical component of biodiversity conservation. In Canada, the Committee on the Status of Endangered Wildlife in Canada (COSEWIC) determines whether species are at risk of extinction or extirpation, and has conducted these assessments since 1977. We examined trends in COSEWIC assessments to identify whether at-risk species that have been assessed more than once tended to improve, remain constant, or deteriorate in status, as a way of assessing the effectiveness of biodiversity conservation in Canada. Of 369 species that met our criteria for examination, 115 deteriorated, 202 remained unchanged, and 52 improved in status. Only 20 species (5.4%) improved to the point where they were 'not at risk', and five of those were due to increased sampling efforts rather than an increase in population size. Species outcomes were also dependent on the severity of their initial assessment; for example, 47% of species that were initially listed as special concern deteriorated between assessments. After receiving an at-risk assessment by COSEWIC, a species is considered for listing under the federal Species at Risk Act (SARA), which is the primary national tool that mandates protection for at-risk species. We examined whether SARA-listing was associated with improved COSEWIC assessment outcomes relative to unlisted species. Of 305 species that had multiple assessments and were SARA-listed, 221 were listed at a level that required identification and protection of critical habitat; however, critical habitat was fully identified for only 56 of these species. We suggest that the Canadian government should formally identify and protect critical habitat, as is required by existing legislation. In addition, our finding that at-risk species in Canada rarely recover leads us to recommend that every effort be made to actively prevent species from becoming at-risk in the first place.