Individual quality overwrites carry-over effects across the annual cycle of a long-distance migrant.

In seasonal environments, the fitness of animals depends upon the successful integration of life-history stages throughout their annual cycle. Failing to do so can lead to negative carry-over effects where individuals are transitioning into the next season in different states, consequently affecting their future performance. However, carry-over effects can be masked by individual quality when individuals vary in their efficiency at acquiring resources year after year (i.e. 'quality'), leading to cross-seasonal consistency in individual performance. Here we investigated the relative importance of carry-over effects and individual quality in determining cross-seasonal interactions and consequences for breeding success over the full annual cycle of a migratory seabird (black-legged kittiwake Rissa tridactyla). We monitored the reproduction and annual movement of kittiwakes over 13 years using geolocators to estimate their breeding success, distribution and winter energy expenditure. We combined this with an experimental approach (clutch removal experiment, 2 years) to manipulate the reproductive effort irrespective of individual quality. Piecewise path analyses showed that successful breeders reproduced earlier and were more likely to breed successfully again the following year. This positive interaction among consecutive breeding stages disappeared after controlling for individual quality, suggesting that quality was dominant in determining seasonal interactions. Moreover, controlling experimentally for individual quality revealed underlying carry-over effects that were otherwise masked by quality, with breeding costs paid in higher energy expenditure and delayed onset of reproduction. We highlight the need to combine an experimental approach along with long-term data while assessing apparent carry-over effects in wild animals, and their potential impact on fitness and population demography.

Seabirds reveal mercury distribution across the North Atlantic.

Mercury (Hg) is a heterogeneously distributed toxicant affecting wildlife and human health. Yet, the spatial distribution of Hg remains poorly documented, especially in food webs, even though this knowledge is essential to assess large-scale risk of toxicity for the biota and human populations. Here, we used seabirds to assess, at an unprecedented population and geographic magnitude and high resolution, the spatial distribution of Hg in North Atlantic marine food webs. To this end, we combined tracking data of 837 seabirds from seven different species and 27 breeding colonies located across the North Atlantic and Atlantic Arctic together with Hg analyses in feathers representing individual seabird contamination based on their winter distribution. Our results highlight an east-west gradient in Hg concentrations with hot spots around southern Greenland and the east coast of Canada and a cold spot in the Barents and Kara Seas. We hypothesize that those gradients are influenced by eastern (Norwegian Atlantic Current and West Spitsbergen Current) and western (East Greenland Current) oceanic currents and melting of the Greenland Ice Sheet. By tracking spatial Hg contamination in marine ecosystems and through the identification of areas at risk of Hg toxicity, this study provides essential knowledge for international decisions about where the regulation of pollutants should be prioritized.

North Atlantic winter cyclones starve seabirds.

Each winter, the North Atlantic Ocean is the stage for numerous cyclones, the most severe ones leading to seabird mass-mortality events called "winter wrecks."1-3 During these, thousands of emaciated seabird carcasses are washed ashore along European and North American coasts. Winter cyclones can therefore shape seabird population dynamics4,5 by affecting survival rates as well as the body condition of surviving individuals and thus their future reproduction. However, most often the geographic origins of impacted seabirds and the causes of their deaths remain unclear.6 We performed the first ocean-basin scale assessment of cyclone exposure in a seabird community by coupling winter tracking data for ∼1,500 individuals of five key North Atlantic seabird species (Alle alle, Fratercula arctica, Uria aalge, Uria lomvia, and Rissa tridactyla) and cyclone locations. We then explored the energetic consequences of different cyclonic conditions using a mechanistic bioenergetics model7 and tested the hypothesis that cyclones dramatically increase seabird energy requirements. We demonstrated that cyclones of high intensity impacted birds from all studied species and breeding colonies during winter but especially those aggregating in the Labrador Sea, the Davis Strait, the surroundings of Iceland, and the Barents Sea. Our broad-scale analyses suggested that cyclonic conditions do not increase seabird energy requirements, implying that they die because of the unavailability of their prey and/or their inability to feed during cyclones. Our study provides essential information on seabird cyclone exposure in a context of marked cyclone regime changes due to global warming.8.

Real-time predictions of seabird distribution improve oil spill risk assessments.

Current knowledge of the distribution of sensitive seabirds is inadequate to safeguard seabird populations from impacts of oil spills in the Arctic. This gap is mainly driven by the fact that statistical models applied to survey data are coarse-scale and static with limited documentation of the distributional dynamics and patchiness of seabirds relevant to risk assessments related to oil spills. This paper describes a dynamic modelling framework solution for prediction of fine-scale densities and movements of seabirds in close-to-real time using fully integrated 3-D hydrodynamic models, dynamic habitat suitability models and agent-based models. The modelling framework has been developed and validated for the swimming migration of Brünnich's Guillemot Uria lomvia in the Barents Sea. The results document that the distributional dynamics of Brünnich's Guillemot and other seabird species to a large degree can be simulated with in-situ state variables and patterns reflecting the physical meteorology and oceanography and habitat suitability.

Blue justice: A survey for eliciting perceptions of environmental justice among coastal planners' and small-scale fishers in Northern-Norway.

Ocean-based economic development arising from an increasing interest in the 'blue economy' is placing ecosystems and small-scale fisheries under pressure. The dominant policy response for dealing with multiple uses is the allocation of coastal space through coastal zone planning (CZP). Recent studies have shown that the rush to develop the blue economy and regulate coastal activity can result in social injustices and the exclusion of less powerful and unrecognized groups (e.g., small-scale fishers, women, Indigenous peoples and youth). To achieve a primary goal of the 2030 sustainable development agenda to "leave no one behind", it is important to understand the implications of coastal planning and development for these groups. Here, we present a social survey protocol for examining perceptions of justice related to small-scale fisheries (SSF) in the context of the blue economy in coastal areas. Specifically, we designed the survey instrument and sampling protocol to assess whether decisions about the use of the coastal zone over the last five years have i) followed principles of good governance, ii) recognized fishers' knowledge, culture and rights and iii) been attentive to impacts of changed coastal zone use on fisheries. The survey will engage coastal planners (N = app. 120) and fishers (N = app. 4300) in all the coastal municipalities (N = 81) in Northern-Norway. The sampling protocol is designed to ensure representation of different sectors of society, including those defined by gender, age, ethnicity and occupation (e.g., small-scale fishers, large-scale fishers, coastal planners).

Meeting Paris agreement objectives will temper seabird winter distribution shifts in the North Atlantic Ocean.

We explored the implications of reaching the Paris Agreement Objective of limiting global warming to <2°C for the future winter distribution of the North Atlantic seabird community. We predicted and quantified current and future winter habitats of five North Atlantic Ocean seabird species (Alle alle, Fratercula arctica, Uria aalge, Uria lomvia and Rissa tridactyla) using tracking data for ~1500 individuals through resource selection functions based on mechanistic modeling of seabird energy requirements, and a dynamic bioclimate envelope model of seabird prey. Future winter distributions were predicted to shift with climate change, especially when global warming exceed 2°C under a "no mitigation" scenario, modifying seabird wintering hotspots in the North Atlantic Ocean. Our findings suggest that meeting Paris agreement objectives will limit changes in seabird selected habitat location and size in the North Atlantic Ocean during the 21st century. We thereby provide key information for the design of adaptive marine-protected areas in a changing ocean.

Stakeholders' perceptions of protected area management following a nationwide community-based conservation reform.

People's perceptions can point to reasons for conservation support or lack thereof. In this study, we surveyed the perceptions of conservation among local stakeholders who participated in protected area (PA) management following a reform towards community-based conservation in Norway. We analyzed the link between perceptions of threats to conservation values, prioritized management actions and trust in PA governance, and assessed how these perceptions aligned with the stakeholders' preferred overarching conservation approach and their support for PAs. Conservation perceptions differed mostly between property owners and representatives of nature conservation (such as NGOs). Property owners perceived modern farming, grazing and hay making, and securing the interests of rights holders as a priority. They had a lower support for PAs and favored a conservation approach focusing on "people and nature". Representatives from nature conservation prioritized management actions to increase biodiversity and reduce land development, had higher trust in environmental authorities and identified motorized vehicle use as a threat to conservation values. They had a high support for PAs and favored a conservation approach that mitigates threats from human activity (i.e. "nature despite people"). The nationwide reform aimed at increasing support for PAs, but 31% of the members of the stakeholder advisory councils were willing to downgrade or degazette PAs for the benefit of economic development, which is much more than general population surveys. However, the level of trust in local governance was less polarized among the members of stakeholder advisory councils compared with the former state governance, which suggests that that the community-based conservation reform has the potential to improve collaboration and conflict mitigation.

Local land use associated with socio-economic development in six arctic regions.

The socioeconomic causes of land use change are complex. They are highly context dependent, but most often studied through case studies. Here, we use a quasi-experimental paired block design to investigate whether better access to wage income leads to more visible land use around 28 settlements in six regions of the circumpolar Arctic. We mapped visible land use on high-resolution satellite images taken both close to the settlements, and in a more remote area of extensive land use, and payed special attention to tracks of off-road vehicles (ORV). Despite considerable differences among regions, there was an overall positive relationship between better access to wage income and land use. Reindeer herding was also associated with more visible use, in particular ORV tracks. These results suggest that access to wage income in the mixed subsistence-cash communities of the Arctic could lead to more local use related to harvesting and reindeer herding.

Arctic greening from warming promotes declines in caribou populations.

The migratory tundra caribou herds in North America follow decadal population cycles, and browsing from abundant caribou could be expected to counteract the current climate-driven expansion of shrubs in the circumpolar tundra biome. We demonstrate that the sea ice cover in the Arctic Ocean has provided a strong signal for climate-induced changes on the adjacent caribou summer ranges, outperforming other climate indices in explaining the caribou-plant dynamics. We found no evidence of a negative effect of caribou abundance on vegetation biomass. On the contrary, we found a strong bottom-up effect in which a warmer climate related to diminishing sea ice has increased the plant biomass on the summer pastures, along with a paradoxical decline in caribou populations. This result suggests that this climate-induced greening has been accompanied by a deterioration of pasture quality. The shrub expansion in Arctic North America involves plant species with strong antibrowsing defenses. Our results might therefore be an early signal of a climate-driven shift in the caribou-plant interaction from a system with low plant biomass modulated by cyclic caribou populations to a system dominated by nonedible shrubs and diminishing herds of migratory caribou.

Phenology and cover of plant growth forms predict herbivore habitat selection in a high latitude ecosystem.

The spatial and temporal distribution of forage quality is among the most central factors affecting herbivore habitat selection. Yet, for high latitude areas, forage quantity has been found to be more important than quality. Studies on large ungulate foraging patterns are faced with methodological challenges in both assessing animal movements at the scale of forage distribution, and in assessing forage quality with relevant metrics. Here we use first-passage time analyses to assess how reindeer movements relate to forage quality and quantity measured as the phenology and cover of growth forms along reindeer tracks. The study was conducted in a high latitude ecosystem dominated by low-palatable growth forms. We found that the scale of reindeer movement was season dependent, with more extensive area use as the summer season advanced. Small-scale movement in the early season was related to selection for younger stages of phenology and for higher abundances of generally phenologically advanced palatable growth forms (grasses and deciduous shrubs). Also there was a clear selection for later phenological stages of the most dominant, yet generally phenologically slow and low-palatable growth form (evergreen shrubs). As the summer season advanced only quantity was important, with selection for higher quantities of one palatable growth form and avoidance of a low palatable growth form. We conclude that both forage quality and quantity are significant predictors to habitat selection by a large herbivore at high latitude. The early season selectivity reflected that among dominating low palatability growth forms there were palatable phenological stages and palatable growth forms available, causing herbivores to be selective in their habitat use. The diminishing selectivity and the increasing scale of movement as the season developed suggest a response by reindeer to homogenized forage availability of low quality.

Risk-sensitive reproductive allocation: fitness consequences of body mass losses in two contrasting environments.

For long-lived organisms, the fitness value of survival is greater than that of current reproduction. Asymmetric fitness rewards suggest that organisms inhabiting unpredictable environments should adopt a risk-sensitive life history, predicting that it is adaptive to allocate resources to increase their own body reserves at the expense of reproduction. We tested this using data from reindeer populations inhabiting contrasting environments and using winter body mass development as a proxy for the combined effect of winter severity and density dependence. Individuals in good and harsh environments responded similarly: Females who lost large amounts of winter body mass gained more body mass the coming summer compared with females losing less mass during winter. Additionally, females experienced a cost of reproduction: On average, barren females gained more body mass than lactating females. Winter body mass development positively affected both the females' reproductive success and offspring body mass. Finally, we discuss the relevance of our findings with respect to scenarios for future climate change.

Population densities, vegetation green-up, and plant productivity: impacts on reproductive success and juvenile body mass in reindeer.

Global warming is expected to cause earlier springs and increased primary productivity in the Arctic. These changes may improve food availability for Arctic herbivores, but may also have negative effects by generating a mismatch between the surge of high quality food in the spring and the timing of reproduction. We analyzed a 10 year dataset of satellite derived measures of vegetation green-up, population densities, calf body masses and female reproductive success in 19 reindeer (Rangifer tarandus) populations in Northern Norway. An early onset of spring and high peak plant productivity had positive effects on calf autumn body masses and female reproductive success. In addition, body masses and reproductive success were both negatively related to population density. The quantity of food available, as determined by the onset of vegetation green-up and plant productivity over the summer were the main drivers of body mass growth and reproductive success. We found no evidence for an effect of the speed of spring green-up. Nor did we detect a negative mismatch between early springs and subsequent recruitment. Effects of global warming on plant productivity and onset of spring is likely to positively affect sub-Arctic reindeer.

Community-based management: under what conditions do Sámi pastoralists manage pastures sustainably?

Community-based management (CBM) has been implemented in socio-ecological systems (SES) worldwide. CBM has also been the prevailing policy in Sámi pastoral SES in Norway, but the outcomes tend to vary extensively among resource groups ("siidas"). We asked why do some siidas self-organize to manage common pool resources sustainably and others do not? To answer this question we used a mixed methods approach. First, in the statistical analyses we analyzed the relationship between sustainability indicators and structural variables. We found that small winter pastures that are shared by few siidas were managed more sustainably than larger pastures. Seasonal siida stability, i.e., a low turnover of pastoralists working together throughout the year, and equality among herders, also contributed to more sustainable outcomes. Second, interviews were conducted in the five largest pastures to explain the relationships between the structural variables and sustainability. The pastoralists expressed a high level of agreement with respect to sustainable policies, but reported a low level of trust and cooperation among the siidas. The pastoralists requested siida tenures or clear rules and sanctioning mechanisms by an impartial authority rather than flexible organization or more autonomy for the siidas. The lack of nestedness in self-organization for managing pastures on larger scales, combined with the past economic policies, could explain why CBM is less sustainable on the largest winter pastures. We conclude that the scale mis-match between self-organization and the formal governance is a key condition for sustainability.

Wasp-waist interactions in the North Sea ecosystem.

BACKGROUND: In a "wasp-waist" ecosystem, an intermediate trophic level is expected to control the abundance of predators through a bottom-up interaction and the abundance of prey through a top-down interaction. Previous studies suggest that the North Sea is mainly governed by bottom-up interactions driven by climate perturbations. However, few studies have investigated the importance of the intermediate trophic level occupied by small pelagic fishes. METHODOLOGY/PRINCIPAL FINDINGS: We investigated the numeric interactions among 10 species of seabirds, two species of pelagic fish and four groups of zooplankton in the North Sea using decadal-scale databases. Linear models were used to relate the time series of zooplankton and seabirds to the time series of pelagic fish. Seabirds were positively related to herring (Clupea harengus), suggesting a bottom-up interaction. Two groups of zooplankton; Calanus helgolandicus and krill were negatively related to sprat (Sprattus sprattus) and herring respectively, suggesting top-down interactions. In addition, we found positive relationships among the zooplankton groups. Para/pseudocalanus was positively related to C. helgolandicus and C. finmarchicus was positively related to krill. CONCLUSION/SIGNIFICANCE: Our results indicate that herring was important in regulating the abundance of seabirds through a bottom-up interaction and that herring and sprat were important in regulating zooplankton through top-down interactions. We suggest that the positive relationships among zooplankton groups were due to selective foraging and switching in the two clupeid fishes. Our results suggest that "wasp-waist" interactions might be more important in the North Sea than previously anticipated. Fluctuations in the populations of pelagic fish due to harvesting and depletion of their predators might accordingly have profound consequences for ecosystem dynamics through trophic cascades.

Scale-dependent response diversity of seabirds to prey in the North Sea.

Functional response diversity is defined as the diversity of responses to environmental change among species that contribute to the same ecosystem function. Because different ecological processes dominate on different spatial and temporal scales, response diversity is likely to be scale dependent. Using three extensive data sets on seabirds, pelagic fish, and zooplankton, we investigate the strength and diversity in the response of seabirds to prey in the North Sea over three scales of ecological organization. Two-stage analyses were used to partition the variance in the abundance of predators and prey among the different scales of investigation: variation from year to year, variation among habitats, and variation on the local patch scale. On the year-to-year scale, we found a strong and synchronous response of seabirds to the abundance of prey, resulting in low response diversity. Conversely, as different seabird species were found in habitats dominated by different prey species, we found a high diversity in the response of seabirds to prey on the habitat scale. Finally, on the local patch scale, seabirds were organized in multispecies patches. These patches were weakly associated with patches of prey, resulting in a weak response strength and a low response diversity. We suggest that ecological similarities among seabird species resulted in low response diversity on the year-to-year scale. On the habitat scale, we suggest that high response diversity was due to interspecific competition and niche segregation among seabird species. On the local patch scale, we suggest that facilitation with respect to the detection and accessibility of prey patches resulted in overlapping distribution of seabirds but weak associations with prey. The observed scale dependencies in response strength and diversity have implications for how the seabird community will respond to different environmental disturbances.

One- and five-year follow-ups on blood pressure and renal function in kidney donors.

It is considered safe to donate a kidney if internationally accepted medical criteria are fulfilled. However, some donors have encountered hypertension, proteinuria and impaired renal function after donation. The study was based on retrospective data on 908 donors, donating in the period 1997-2007. Preoperative and follow-up data were collected from patient files and the Norwegian Living Donor Registry. Follow-up data were available for 665 donors at 1 year after donation, and 256 donors at 5 years after donation. We calculated the estimated glomerular filtration rate (eGFR) using the four variable Modification of Diet in Renal Disease equation. At 1 and 5 years after donation, the prevalence of hypertension was 11.7% and 27.1% respectively compared to 2.6% before donation. Proteinuria was present in 3.3% and 1.6% at 1 and 5 years. Mean eGFR was 56.1 ± 10.8 ml/min/1.73 m² at 1 year and 61.0 ± 11.8 ml/min/1.73 m² at 5 years. Mean blood pressure was 122.5 ± 10.6/76.2 ± 7.5 mmHg at donation (n = 908), 124.3 ± 14.2/77.9 ± 8.2 mmHg at 1-year (n = 649) and 127.2 ± 15.4/78.8 ± 8.3 mmHg at 5-year follow-ups (n = 247). We found no evidence of further decline in renal function beyond the initial decrement following nephrectomy.

Predator-prey reversal: a possible mechanism for ecosystem hysteresis in the North Sea?

Removal of large predatory fishes from marine ecosystems has resulted in persistent ecosystem shifts, with collapsed predator populations and super-abundant prey populations. One explanation for these shifts is reversals of predator-prey roles that generate internal feedbacks in the ecosystems. Pelagic forage fish are often predators and competitors to the young life stages of their larger fish predators. I show that cod recruitment in the North Sea has been negatively related to the spawning-stock biomass of herring for the last 44 years. Herring, together with the abundance of Calanus finmarchicus, the major food for cod larvae, were the main predictors of cod recruitment. These predictors were of equivalent importance, worked additively, and explained different parts of the dynamics in cod recruitment. I suggest that intensive harvesting of cod has released herring from predator control, and that a large population of herring suppresses cod recruitment through predation on eggs and larvae. This feedback mechanism can promote alternative stable states and therefore cause hysteresis to occur under changing conditions; however, harvesting of herring might at present prevent a shift in the ecosystem to a herring-dominated state.

Observational evidence of risk-sensitive reproductive allocation in a long-lived mammal.

Organisms should adopt a risk-sensitive reproductive allocation when summer reproductive allocation competes with survival in the coming winter. This trade off is shown through autumn female body mass, which acts as an insurance against unpredictable winter environmental conditions. We tested this hypothesis on female reindeer in a population that has experienced a time period of dramatic increase in abundance. Environmental conditions during winter were fairly stable (with the exception of 1 year). We conclude that increased population abundance (perhaps in interaction with winter environmental conditions) could have represented a worsening of winter environmental conditions as both autumn offspring and spring female body mass decreased during the course of the study. Moreover, we found that the cost of reproduction was related to environmental conditions as: (1) autumn body mass was larger for barren than for lactating females, and this difference was temporally highly variable; (2) lactating females produced smaller offspring than barren ones in the following year; and (3) reproductive output (offspring size) decreased over time. We also found evidence of quality effects as lactating females had a higher reproductive success in the following year. In sum, a worsening of winter conditions lead to: (1) decreased reproductive output; (2) lowered autumn body mass for lactating females; and (3) increased body mass for barren females. Since females reduce their reproductive allocation as winter conditions becomes more severe, we conclude that reindeer have adopted a risk-sensitive reproductive allocation.

Short- and long-term consequences of reproductive decisions: an experimental study in the puffin.

The purpose of the present study was to inspect the response of the Atlantic Puffin (Fratercula arctica) to an experimental manipulation of the investment needed to successfully raise an offspring. We achieved this by replacing an old offspring with a younger chick, and vice versa, thereby prolonging and shortening the chick-rearing period. To examine any costs of reproduction we then followed the breeding success, the recruitment of young to the population, and the survival of parents for 11 years following the manipulation. Parents in the prolonged and shortened category had a lower breeding success than controls mainly because parents deserted their chick shortly after swapping. Among those that raised their chick, the age and body mass of foster chicks at fledging were the same in all three categories even though the parents had raised chicks for different lengths of time. The recruitment of young to the breeding population was high and independent of treatment. Likewise, the survival of adults was independent of treatment. For the 11 years after the experiment, however, the resighting rate of those that deserted their chick was clearly lower than among those that accepted their foster chick. For parents that raised their foster chick, the survival to the following year was positively related to their body mass. The results support the hypothesis that puffins have a highly flexible parental investment, which they adjust according to their own individual quality and the survival prospects of the chick.