Correction: Publication reform to safeguard wildlife from researcher harm.

[This corrects the article DOI: 10.1371/journal.pbio.3000193.].

Publication reform to safeguard wildlife from researcher harm.

Despite abundant focus on responsible care of laboratory animals, we argue that inattention to the maltreatment of wildlife constitutes an ethical blind spot in contemporary animal research. We begin by reviewing significant shortcomings in legal and institutional oversight, arguing for the relatively rapid and transformational potential of editorial oversight at journals in preventing harm to vertebrates studied in the field and outside the direct supervision of institutions. Straightforward changes to animal care policies in journals, which our analysis of 206 journals suggests are either absent (34%), weak, incoherent, or neglected by researchers, could provide a practical, effective, and rapidly imposed safeguard against unnecessary suffering. The Animals in Research: Reporting On Wildlife (ARROW) guidelines we propose here, coupled with strong enforcement, could result in significant changes to how animals involved in wildlife research are treated. The research process would also benefit. Sound science requires animal subjects to be physically, physiologically, and behaviorally unharmed. Accordingly, publication of methods that contravenes animal welfare principles risks perpetuating inhumane approaches and bad science.

Large carnivore hunting and the social license to hunt.

The social license to operate framework considers how society grants or withholds informal permission for resource extractors to exploit publicly owned resources. We developed a modified model, which we refer to as the social license to hunt (SLH). In it we similarly consider hunters as operators, given that wildlife are legally considered public resources in North America and Europe. We applied the SLH model to examine the controversial hunting of large carnivores, which are frequently killed for trophies. Killing for trophies is widespread, but undertaken by a minority of hunters, and can pose threats to the SLH for trophy-seeking carnivore hunters and potentially beyond. Societal opposition to large carnivore hunting relates not only to conservation concerns but also to misalignment between killing for trophies and dominant public values and attitudes concerning the treatment of animals. We summarized cases related to the killing of grizzly bears (Ursus arctos), wolves (Canis lupus), and other large carnivores in Canada, the United States, and Europe to illustrate how opposition to large carnivore hunting, now expressed primarily on social media, can exert rapid and significant pressure on policy makers and politicians. Evidence of the potential for transformative change to wildlife management and conservation includes proposed and realized changes to legislation, business practice, and wildlife policy, including the banning of some large carnivore hunts. Given that policy is ultimately shaped by societal values and attitudes, research gaps include developing increased insight into public support of various hunting policies beyond that derived from monitoring of social media and public polling. Informed by increased evidence, the SLH model can provide a conceptual foundation for predicting the likelihood of transient versus enduring changes to wildlife conservation policy and practice for a wide variety of taxa and contexts.

Cacería de Grandes Carnívoros y la Licencia Social para Cazar Resumen El marco de trabajo de la licencia social para operar considera cómo la sociedad otorga o restringe permisos informales para que los extractores de recursos puedan explotar los recursos públicos. Desarrollamos un modelo modificado, al cual nos referimos como la licencia social para cazar (LSC). En este modelo consideramos a los cazadores como símiles de los operadores puesto que en América del Norte y en Europa a la fauna se le considera legalmente como recurso público. Aplicamos el modelo de la LSC en un análisis de la cacería controversial de grandes carnívoros, a los cuales con frecuencia se les caza para convertirlos en trofeos. La cacería para trofeos es común pero sólo la realiza una minoría de los cazadores y puede presentar una amenaza para la LSC para los cazadores que cazan carnívoros para trofeos e incluso para otros tipos de cazadores. La oposición social a la cacería de grandes carnívoros se relaciona no sólo con el interés de conservación sino también con la discordancia entre la caza para trofeos y las actitudes y valores públicos dominantes con respecto al trato hacia los animales. Resumimos algunos casos relacionados con la muerte de osos pardos (Ursus arctos), lobos (Canis lupus) y otros grandes carnívoros en Canadá, los Estados Unidos y Europa para mostrar cómo la oposición a la cacería de grandes carnívoros, hoy en día expresada principalmente en las redes sociales, puede ejercer una presión rápida y significativa sobre los políticos y los formuladores de políticas. La evidencia de un potencial de cambio transformador en el manejo y conservación de fauna incluye los cambios propuestos y realizados a la legislación, la práctica comercial y las políticas para la fauna, incluyendo la prohibición de la caza de algunos grandes carnívoros. Ya que las políticas están finalmente moldeadas por las actitudes y los valores sociales, las lagunas en la investigación incluyen el desarrollo de un conocimiento mejorado del respaldo público para varias políticas de cacería más allá del conocimiento derivado del monitoreo de las redes sociales y las encuestas públicas. Si se informa con mucha más evidencia, el modelo de la LSC puede proporcionar una base conceptual para predecir la probabilidad de los cambios transitorios versus los duraderos en las políticas y las prácticas de conservación de fauna para una gama amplia de taxones y contextos.

Marine subsidies mediate patterns in avian island biogeography.

The classical theory of island biogeography, which predicts species richness using island area and isolation, has been expanded to include contributions from marine subsidies, i.e. subsidized island biogeography (SIB) theory. We tested the effects of marine subsidies on species diversity and population density on productive temperate islands, evaluating SIB predictions previously untested at comparable scales and subsidy levels. We found that the diversity of terrestrial breeding bird communities on 91 small islands (approx. 0.0001-3 km2) along the Central Coast of British Columbia, Canada were correlated most strongly with island area, but also with marine subsidies. Species richness increased and population density decreased with island area, but isolation had no measurable influence. Species richness was negatively correlated with marine subsidy, measured as forest-edge soil δ15N. Density, however, was higher on islands with higher marine subsidy, and a negative interaction between area and subsidy indicates that this effect is stronger on smaller islands, offering some support for SIB. Our study emphasizes how subsidies from the sea can shape diversity patterns on islands and can even exceed the importance of isolation in determining species richness and densities of terrestrial biota.

Natural re-colonization and admixture of wolves (Canis lupus) in the US Pacific Northwest: challenges for the protection and management of rare and endangered taxa.

Admixture resulting from natural dispersal processes can potentially generate novel phenotypic variation that may facilitate persistence in changing environments or result in the loss of population-specific adaptations. Yet, under the US Endangered Species Act, policy is limited for management of individuals whose ancestry includes a protected taxon; therefore, they are generally not protected under the Act. This issue is exemplified by the recently re-established grey wolves of the Pacific Northwest states of Washington and Oregon, USA. This population was likely founded by two phenotypically and genetically distinct wolf ecotypes: Northern Rocky Mountain (NRM) forest and coastal rainforest. The latter is considered potentially threatened in southeast Alaska and thus the source of migrants may affect plans for their protection. To assess the genetic source of the re-established population, we sequenced a ~ 300 bp portion of the mitochondrial control region and ~ 5 Mbp of the nuclear genome. Genetic analysis revealed that the Washington wolves share ancestry with both wolf ecotypes, whereas the Oregon population shares ancestry with NRM forest wolves only. Using ecological niche modelling, we found that the Pacific Northwest states contain environments suitable for each ecotype, with wolf packs established in both environmental types. Continued migration from coastal rainforest and NRM forest source populations may increase the genetic diversity of the Pacific Northwest population. However, this admixed population challenges traditional management regimes given that admixture occurs between an adaptively distinct ecotype and a more abundant reintroduced interior form. Our results emphasize the need for a more precise US policy to address the general problem of admixture in the management of endangered species, subspecies, and distinct population segments.

Salmonid species diversity predicts salmon consumption by terrestrial wildlife.

Resource waves-spatial variation in resource phenology that extends feeding opportunities for mobile consumers-can affect the behaviour and productivity of recipient populations. Interspecific diversity among Pacific salmon species (Oncorhynchus spp.) creates staggered spawning events across space and time, thereby prolonging availability to terrestrial wildlife. We sought to understand how such variation might influence consumption by terrestrial predators compared with resource abundance and intra- and interspecific competition. Using stable isotope analysis, we investigated how the proportion of salmon in the annual diet of male black bears (Ursus americanus; n = 405) varies with species diversity and density of spawning salmon biomass, while also accounting for competition with sympatric black and grizzly bears (U. arctos horribilis), in coastal British Columbia, Canada. We found that the proportion of salmon in the annual diet of black bears was ≈40% higher in the absence of grizzly bears, but detected little effect of relative black bear density and salmon biomass density. Rather, salmon diversity had the largest positive effect on consumption. On average, increasing diversity from one salmon species to ~four (with equal biomass contributions) approximately triples the proportion of salmon in diet. Given the importance of salmon to bear life histories, this work provides early empirical support for how resource waves may increase the productivity of consumers at population and landscape scales. Accordingly, terrestrial wildlife management might consider maintaining not only salmon abundance but also diversity.

BMC ecology image competition 2017: the winning images.

For the fifth year, BMC Ecology is proud to present the winning images from our annual image competition. The 2017 edition received entries by talented shutterbug-ecologists from across the world, showcasing research that is increasing our understanding of ecosystems worldwide and the beauty and diversity of life on our planet. In this editorial we showcase the winning images, as chosen by our Editorial Board and guest judge Chris Darimont, as well as our selection of highly commended images. Enjoy!

Hunting as a management tool? Cougar-human conflict is positively related to trophy hunting.

BACKGROUND: Overexploitation and persecution of large carnivores resulting from conflict with humans comprise major causes of declines worldwide. Although little is known about the interplay between these mortality types, hunting of predators remains a common management strategy aimed at reducing predator-human conflict. Emerging theory and data, however, caution that such policy can alter the age structure of populations, triggering increased conflict in which conflict-prone juveniles are involved. RESULTS: Using a 30-year dataset on human-caused cougar (Puma concolor) kills in British Columbia (BC), Canada, we examined relationships between hunter-caused and conflict-associated mortality. Individuals that were killed via conflict with humans were younger than hunted cougars. Accounting for human density and habitat productivity, human hunting pressure during or before the year of conflict comprised the most important variables. Both were associated with increased male cougar-human conflict. Moreover, in each of five regions assessed, conflict was higher with increased human hunting pressure for at least one cougar sex. CONCLUSION: Although only providing correlative evidence, such patterns over large geographic and temporal scales suggest that alternative approaches to conflict mitigation might yield more effective outcomes for humans as well as cougar populations and the individuals within populations.

Using grizzly bears to assess harvest-ecosystem tradeoffs in salmon fisheries.

Implementation of ecosystem-based fisheries management (EBFM) requires a clear conceptual and quantitative framework for assessing how different harvest options can modify benefits to ecosystem and human beneficiaries. We address this social-ecological need for Pacific salmon fisheries, which are economically valuable but intercept much of the annual pulse of nutrient subsidies that salmon provide to terrestrial and aquatic food webs. We used grizzly bears, vectors of salmon nutrients and animals with densities strongly coupled to salmon abundance, as surrogates for "salmon ecosystem" function. Combining salmon biomass and stock-recruitment data with stable isotope analysis, we assess potential tradeoffs between fishery yields and bear population densities for six sockeye salmon stocks in Bristol Bay, Alaska, and British Columbia (BC), Canada. For the coastal stocks, we find that both bear densities and fishery yields would increase substantially if ecosystem allocations of salmon increase from currently applied lower to upper goals and beyond. This aligning of benefits comes at a potential cost, however, with the possibility of forgoing harvests in low productivity years. In contrast, we detect acute tradeoffs between bear densities and fishery yields in interior stocks within the Fraser River, BC, where biomass from other salmon species is low. There, increasing salmon allocations to ecosystems would benefit threatened bear populations at the cost of reduced long-term yields. To resolve this conflict, we propose an EBFM goal that values fisheries and bears (and by extension, the ecosystem) equally. At such targets, ecosystem benefits are unexpectedly large compared with losses in fishery yields. To explore other management options, we generate tradeoff curves that provide stock-specific accounting of the expected loss to fishers and gain to bears as more salmon escape the fishery. Our approach, modified to suit multiple scenarios, provides a generalizable method to resolve conflicts over shared resources in other systems.

Snow avalanches are a primary climate-linked driver of mountain ungulate populations.

Snow is a major, climate-sensitive feature of the Earth's surface and catalyst of fundamentally important ecosystem processes. Understanding how snow influences sentinel species in rapidly changing mountain ecosystems is particularly critical. Whereas effects of snow on food availability, energy expenditure, and predation are well documented, we report how avalanches exert major impacts on an ecologically significant mountain ungulate - the coastal Alaskan mountain goat (Oreamnos americanus). Using long-term GPS data and field observations across four populations (421 individuals over 17 years), we show that avalanches caused 23-65% of all mortality, depending on area. Deaths varied seasonally and were directly linked to spatial movement patterns and avalanche terrain use. Population-level avalanche mortality, 61% of which comprised reproductively important prime-aged individuals, averaged 8% annually and exceeded 22% when avalanche conditions were severe. Our findings reveal a widespread but previously undescribed pathway by which snow can elicit major population-level impacts and shape demographic characteristics of slow-growing populations of mountain-adapted animals.

Human disturbance in riparian areas disrupts predator-prey interactions between grizzly bears and salmon.

Wildlife must increasingly balance trade-offs between the need to access important foods and the mortality risks associated with human-dominated landscapes. Human disturbance can profoundly influence wildlife behavior, but managers know little about the relationship between disturbance-behavior dynamics and associated consequences for foraging. We address this gap by empirically investigating the consequences of human activity on a keystone predator-prey interaction in a region with limited but varied industrial disturbance. Using stable isotope data from 226 hair samples of grizzly bears (Ursus arctos horribilis) collected from 1995 to 2014 across 22 salmon-bearing watersheds (88,000 km2) in British Columbia, Canada, we examined how human activity influenced their consumption of spawning salmon (Oncorhynchus spp.), a fitness-related food. Accounting for the abundance of salmon and other foods, salmon consumption strongly decreased (up to 59% for females) with increasing human disturbance (as measured by the human footprint index) in riparian zones of salmon-bearing rivers. Declines in salmon consumption occurred with disturbance even in watersheds with low footprints. In a region currently among the least influenced by industrial activity, intensification of disturbance in river valleys is predicted to increasingly decouple bears from salmon, possibly driving associated reductions in population productivity and provisioning of salmon nutrients to terrestrial ecosystems. Accordingly, we draw on our results to make landscape-scale and access-related management recommendations beyond current streamside protection buffers. This work illustrates the interaction between habitat modification and food security for wildlife, highlighting the potential for unacknowledged interactions and cumulative effects in increasingly modified landscapes.

Ecology of fear alters behavior of grizzly bears exposed to bear-viewing ecotourism.

Humans are perceived as predators by many species and may generate landscapes of fear, influencing spatiotemporal activity of wildlife. Additionally, wildlife might seek out human activity when faced with predation risks (human shield hypothesis). We used the anthropause, a decrease in human activity resulting from the COVID-19 pandemic, to test ecology of fear and human shield hypotheses and quantify the effects of bear-viewing ecotourism on grizzly bear (Ursus arctos) activity. We deployed camera traps in the Khutze watershed in Kitasoo Xai'xais Territory in the absence of humans in 2020 and with experimental treatments of variable human activity when ecotourism resumed in 2021. Daily bear detection rates decreased with more people present and increased with days since people were present. Human activity was also associated with more bear detections at forested sheltered sites and less at exposed sites, likely due to the influence of habitat on bear perception of safety. The number of people negatively influenced adult male detection rates, but we found no influence on female with young detections, providing no evidence that females responded behaviorally to a human shield effect from reduced male activity. We also observed apparent trade-offs of risk avoidance and foraging. When salmon levels were moderate to high, detected bears were more likely to be females with young than adult males on days with more people present. Should managers want to minimize human impacts on bear activity and maintain baseline age-sex class composition at ecotourism sites, multiday closures and daily occupancy limits may be effective. More broadly, this work revealed that antipredator responses can vary with intensity of risk cues, habitat structure, and forage trade-offs and manifest as altered age-sex class composition of individuals using human-influenced areas, highlighting that wildlife avoid people across multiple spatiotemporal scales.

When the enemy of an enemy is no friend.

Some wildlife species mistakenly seek human-inhabited areas to avoid predators.

The diversity of animals identified as keystone species.

Although the keystone species concept was conceived of over 50 years ago, contemporary efforts to synthesize related literature have been limited. Our objective was to create a list of keystone animal species identified in the literature and to examine the variation in the traits of species and the ecosystem influences they elicit. We documented 230 species considered keystones. A clustering analysis classified them into five archetypes based on combinations of their taxonomic class, body size, trophic level, and role (consumers, modifiers, or prey). Although conservation and public perception of keystones primarily focuses on large vertebrate consumers, our analysis reveals that researchers have defined a wide diversity of keystone species, with large variation in associated ecosystem processes. Future research may confront ambiguity in the definition of keystone status, as well as clarify the type, abundance, and quality of data required to assign the term. Identifying keystones with increased rigor would not only enrich the literature but also inform intervention to safeguard threatened keystones and their associated influences on ecosystems.

Humanity's diverse predatory niche and its ecological consequences.

Although humans have long been predators with enduring nutritive and cultural relationships with their prey, seldom have conservation ecologists considered the divergent predatory behavior of contemporary, industrialized humans. Recognizing that the number, strength and diversity of predator-prey relationships can profoundly influence biodiversity, here we analyze humanity's modern day predatory interactions with vertebrates and estimate their ecological consequences. Analysing IUCN 'use and trade' data for ~47,000 species, we show that fishers, hunters and other animal collectors prey on more than a third (~15,000 species) of Earth's vertebrates. Assessed over equivalent ranges, humans exploit up to 300 times more species than comparable non-human predators. Exploitation for the pet trade, medicine, and other uses now affects almost as many species as those targeted for food consumption, and almost 40% of exploited species are threatened by human use. Trait space analyses show that birds and mammals threatened by exploitation occupy a disproportionally large and unique region of ecological trait space, now at risk of loss. These patterns suggest far more species are subject to human-imposed ecological (e.g., landscapes of fear) and evolutionary (e.g., harvest selection) processes than previously considered. Moreover, continued overexploitation will likely bear profound consequences for biodiversity and ecosystem function.

Seasonal and biogeographical patterns of gastrointestinal parasites in large carnivores: wolves in a coastal archipelago.

Parasites are increasingly recognized for their profound influences on individual, population and ecosystem health. We provide the first report of gastrointestinal parasites in gray wolves from the central and north coasts of British Columbia, Canada. Across 60 000 km(2), wolf feces were collected from 34 packs in 2005-2008. At a smaller spatial scale (3300 km(2)), 8 packs were sampled in spring and autumn. Parasite eggs, larvae, and cysts were identified using standard flotation techniques and morphology. A subset of samples was analysed by PCR and sequencing to identify tapeworm eggs (n=9) and Giardia cysts (n=14). We detected ≥14 parasite taxa in 1558 fecal samples. Sarcocystis sporocysts occurred most frequently in feces (43·7%), followed by taeniid eggs (23·9%), Diphyllobothrium eggs (9·1%), Giardia cysts (6·8%), Toxocara canis eggs (2·1%), and Cryptosporidium oocysts (1·7%). Other parasites occurred in ≤1% of feces. Genetic analyses revealed Echinococcus canadensis strains G8 and G10, Taenia ovis krabbei, Diphyllobothrium nehonkaiense, and Giardia duodenalis assemblages A and B. Parasite prevalence differed between seasons and island/mainland sites. Patterns in parasite prevalence reflect seasonal and spatial resource use by wolves and wolf-salmon associations. These data provide a unique, extensive and solid baseline for monitoring parasite community structure in relation to environmental change.