Rapid climate change increases diversity and homogenizes composition of coastal fish at high latitudes.

Rapid warming at high latitudes triggers poleward shifts of species' distributions that impact marine biodiversity. In the open sea, the documented redistributions of fish lead to a borealization of Arctic fauna. A climate-driven borealization and increased species diversity at high latitudes are also expected in coastal fish communities, but they have not been previously documented on a large, biogeographic scale. Here, we investigate the impact of temperature change over the last 25 years on fish communities along the coast of Norway. The study area, spanning different ecoclimatic zones between 62° and 71° N, harbors over 200 species of boreal and Arctic fish. Several of these fish species are harvested by coastal and indigenous communities, influencing settlement geography and livelihood. The long-term data on coastal water temperatures and fish species were obtained from monitoring stations and scientific surveys. Water temperature measured at three fixed sampling stations distributed along the coast show increased temperatures during the study period. The fish species distribution and abundance data were obtained from the annually standardized scientific bottom trawl survey program. Fish species richness, which was highest in the south, increased with warming first in the south and then, gradually, further north, eventually affecting biodiversity in the whole study area. Fish community composition showed a distinct latitudinal pattern early in the study, with Arctic fish species confined to the north and boreal species dominating the south. The poleward shifts eventually eroded this zoogeographic pattern, resulting in more boreal fish species in the north and an increased homogenization of species composition along the Norwegian coast. The climate-driven reorganization of fish communities affects coastal ecosystems that are exposed to fisheries, aquaculture, and other rapidly expanding human activities, stressing the urgent need for a climate adaptation of integrated coastal management.

Parasites alter food-web topology of a subarctic lake food web and its pelagic and benthic compartments.

We compared three sets of highly resolved food webs with and without parasites for a subarctic lake system corresponding to its pelagic and benthic compartments and the whole-lake food web. Key topological food-web metrics were calculated for each set of compartments to explore the role parasites play in food-web topology in these highly contrasting webs. After controlling for effects from differences in web size, we observed similar responses to the addition of parasites in both the pelagic and benthic compartments demonstrated by increases in trophic levels, linkage density, connectance, generality, and vulnerability despite the contrasting composition of free-living and parasitic species between the two compartments. Similar effects on food-web topology can be expected with the inclusion of parasites, regardless of the physical characteristics and taxonomic community compositions of contrasting environments. Additionally, similar increases in key topological metrics were found in the whole-lake food web that combines the pelagic and benthic webs, effects that are comparable to parasite food-web analyses from other systems. These changes in topological metrics are a result of the unique properties of parasites as infectious agents and the links they participate in. Trematodes were key contributors to these results, as these parasites have distinct characteristics in aquatic systems that introduce new link types and increase the food web's generality and vulnerability disproportionate to other parasites. Our analysis highlights the importance of incorporating parasites, especially trophically transmitted parasites, into food webs as they significantly alter key topological metrics and are thus essential for understanding an ecosystem's structure and functioning.

Successive extreme climatic events lead to immediate, large-scale, and diverse responses from fish in the Arctic.

The warming trend of the Arctic is punctuated by several record-breaking warm years with very low sea ice concentrations. The nature and reversibility of marine ecosystem responses to these multiple extreme climatic events (ECEs) are poorly understood. Here, we investigate the ecological signatures of three successive bottom temperature maxima concomitant with surface ECEs between 2004 and 2017 in the Barents Sea across spatial and organizational scales. We observed community-level redistributions of fish concurrent with ECEs at the scale of the whole Barents Sea. Three groups, characterized by different sets of traits describing their capacity to cope with short-term perturbations, reacted with different timing and intensity to each ECE. Arctic species co-occurred more frequently with large predators and incoming boreal taxa during ECEs, potentially affecting food web structures and functional diversity, accelerating the impacts of long-term climate change. On the species level, responses were highly diversified, with different ECEs impacting different species, and species responses (expansion, geographical shift) varying from one ECE to another, despite the environmental perturbations being similar. Past ECEs impacts, with potential legacy effects, lagged responses, thresholds, and interactions with the underlying warming pressure, could constantly set up new initial conditions that drive the unique ecological signature of each ECE. These results highlight the complexity of ecological reactions to multiple ECEs and give prominence to several sources of process uncertainty in the predictions of climate change impact and risk for ecosystem management. Long-term monitoring and studies to characterize the vertical extent of each ECE are necessary to statistically link demersal species and environmental spatial-temporal patterns. In the future, regular monitoring will be crucial to detect early signals of change and understand the determinism of ECEs, but we need to adapt our models and management to better integrate risk and stochasticity from the complex impacts of global change.

Deep demersal fish communities respond rapidly to warming in a frontal region between Arctic and Atlantic waters.

The assessment of climate impact on marine communities dwelling deeper than the well-studied shelf seas has been hampered by the lack of long-term data. For a long time, the prevailing expectation has been that thermal stability in deep ocean layers will delay ecosystem responses to warming. Few observational studies have challenged this view and indicated that deep organisms can respond exceptionally fast to physical change at the sea surface. To address the depth-specific impact of climate change, we investigated spatio-temporal changes in fish community structure along a bathymetry gradient of 150-1500 m between 1998 and 2016 in East Greenland. Here, the Arctic East Greenland Current and the Atlantic Irminger Current meet and mix, representing a sub-Arctic transition zone. We found the strongest signals of community reorganizations at depths between 350 and 1000 m and only weak responses in the shallowest and deepest regions. Changes were in synchrony with atmospheric warming, loss in sea ice and variability in physical sea surface conditions both within our study region and North of the Denmark Strait. These results suggest that interannual variability and long-term climate trends of the larger ecoregion can rapidly affect fish communities down to 1000-m depth through atmospheric ocean coupling and food web interactions.

Medication adherence among persons with coronary heart disease and associations with blood pressure and low-density-lipoprotein-cholesterol.

PURPOSE: To describe medication adherence to lipid-lowering drugs (LLDs), antihypertensive drugs, and acetylsalicylic acid (ASA) among persons with coronary heart disease (CHD) and explore its association with low-density-lipoprotein (LDL)-cholesterol, and systolic and diastolic blood pressure. METHODS: Based on record linkage between the seventh wave of the Tromsø Study and the Norwegian Prescription Database, medication adherence was calculated as the proportion of days covered (PDC) for persistent prevalent users in the period of 365 days before the attendance date. Multivariable linear regression models were used to assess the association between systolic and diastolic blood pressure and medication nonadherence to antihypertensive drugs, age, sex, lifestyle, body mass index (BMI), current and previous diabetes, and between LDL-cholesterol and medication nonadherence to LLDs, age, sex, lifestyle, BMI, and current and previous diabetes. RESULTS: Mean PDC was 0.94 for LLDs and antihypertensive drugs and 0.97 for ASA. Among persons with PDC ≥ 0.80 for LLDs, 12.0% had an LDL-cholesterol < 1.8 mmol/L. Blood pressure < 140/90 mmHg (< 140/80 mmHg if diabetes patient) was reached by 55.1% of those with a PDC ≥ 0.80 for antihypertensive drugs. Adherence to LLDs was associated with lower LDL-cholesterol, while neither systolic nor diastolic blood pressure was associated with adherence to antihypertensive drugs. CONCLUSION: Adherence to antihypertensive drugs, LLDs, and ASA among persons with CHD were high despite low achievement of treatment goals for blood pressure and LDL-cholesterol. There was a statistically significant association between adherence to LLDs and LDL-cholesterol, but not between adherence to antihypertensive drugs and blood pressure.

Increased functional diversity warns of ecological transition in the Arctic.

As temperatures rise, motile species start to redistribute to more suitable areas, potentially affecting the persistence of several resident species and altering biodiversity and ecosystem functions. In the Barents Sea, a hotspot for global warming, marine fish from boreal regions have been increasingly found in the more exclusive Arctic region. Here, we show that this shift in species distribution is increasing species richness and evenness, and even more so, the functional diversity of the Arctic. Higher diversity is often interpreted as being positive for ecosystem health and is a target for conservation. However, the increasing trend observed here may be transitory as the traits involved threaten Arctic species via predation and competition. If the pressure from global warming continues to rise, the ensuing loss of Arctic species will result in a reduction in functional diversity.

Novel feeding interactions amplify the impact of species redistribution on an Arctic food web.

Species are redistributing globally in response to climate warming, impacting ecosystem functions and services. In the Barents Sea, poleward expansion of boreal species and a decreased abundance of Arctic species are causing a rapid borealization of the Arctic communities. This borealization might have profound consequences on the Arctic food web by creating novel feeding interactions between previously non co-occurring species. An early identification of new feeding links is crucial to predict their ecological impact. However, detection by traditional approaches, including stomach content and isotope analyses, although fundamental, cannot cope with the speed of change observed in the region, nor with the urgency of understanding the consequences of species redistribution for the marine ecosystem. In this study, we used an extensive food web (metaweb) with nearly 2,500 documented feeding links between 239 taxa coupled with a trait data set to predict novel feeding interactions and to quantify their potential impact on Arctic food web structure. We found that feeding interactions are largely determined by the body size of interacting species, although species foraging habitat and metabolic type are also important predictors. Further, we found that all boreal species will have at least one potential resource in the Arctic region should they redistribute therein. During 2014-2017, 11 boreal species were observed in the Arctic region of the Barents Sea. These incoming species, which are all generalists, change the structural properties of the Arctic food web by increasing connectance and decreasing modularity. In addition, these boreal species are predicted to initiate novel feeding interactions with the Arctic residents, which might amplify their impact on Arctic food web structure affecting ecosystem functioning and vulnerability. Under the ongoing species redistribution caused by environmental change, we propose merging a trait-based approach with ecological network analysis to efficiently predict the impacts of range-shifting species on food webs.

Arctic coastal benthos long-term responses to perturbations under climate warming.

Climate warming influences structure and function of Arctic benthic ecosystems. Assessing the response of these systems to perturbations requires long-term studies addressing key ecological processes related to recolonization and succession of species. Based on unique time-series (1980-2017), this study addresses successional patterns of hard-bottom benthos in two fjords in NW Svalbard after a pulse perturbation in 1980 and during a period of rapid climate warming. Analysis of seafloor photographs revealed different return rates of taxa, and variability in species densities, through time. It took 13 and 24 years for the community compositions of cleared and control transects to converge in the two fjords. Nearly two decades after the study initiation, an increase in filamentous and foliose macroalgae was observed with a subsequent reorganization in the invertebrate community. Trait analyses showed a decrease in body size and longevity of taxa in response to the pulse perturbation and a shift towards small/medium size and intermediate longevity following the macroalgae takeover. The observed slow recovery rates and abrupt shifts in community structure document the vulnerability of Arctic coastal ecosystems to perturbations and continued effects of climate warming. This article is part of the theme issue 'The changing Arctic Ocean: consequences for biological communities, biogeochemical processes and ecosystem functioning'.

Climate-driven changes in functional biogeography of Arctic marine fish communities.

Climate change triggers poleward shifts in species distribution leading to changes in biogeography. In the marine environment, fish respond quickly to warming, causing community-wide reorganizations, which result in profound changes in ecosystem functioning. Functional biogeography provides a framework to address how ecosystem functioning may be affected by climate change over large spatial scales. However, there are few studies on functional biogeography in the marine environment, and none in the Arctic, where climate-driven changes are most rapid and extensive. We investigated the impact of climate warming on the functional biogeography of the Barents Sea, which is characterized by a sharp zoogeographic divide separating boreal from Arctic species. Our unique dataset covered 52 fish species, 15 functional traits, and 3,660 stations sampled during the recent warming period. We found that the functional traits characterizing Arctic fish communities, mainly composed of small-sized bottom-dwelling benthivores, are being rapidly replaced by traits of incoming boreal species, particularly the larger, longer lived, and more piscivorous species. The changes in functional traits detected in the Arctic can be predicted based on the characteristics of species expected to undergo quick poleward shifts in response to warming. These are the large, generalist, motile species, such as cod and haddock. We show how functional biogeography can provide important insights into the relationship between species composition, diversity, ecosystem functioning, and environmental drivers. This represents invaluable knowledge in a period when communities and ecosystems experience rapid climate-driven changes across biogeographical regions.

Antibiotics to outpatients in Norway-Assessing effect of latitude and municipality population size using quantile regression in a cross-sectional study.

High antibiotic consumption rates are associated to high prevalence of antimicrobial resistance. Geographical differences in dispensing rates of antibiotics are frequently analysed using statistical methods addressing the central tendency of the data. Yet, examining extreme quantiles may be of equal or greater interest if the problem relates to the extremes of consumption rates, as is the case for antimicrobial resistance. The objective of this study was to investigate how geographic location (latitude) and municipality population size affect antibiotic consumption in Norway. We analysed all outpatient antibiotic prescriptions (n > 14 000 000) in Norway between 2004 and 2010 using quantile regression. Data were stratified by year, and we aggregated individual data to municipality, county, or latitudinal range. We specified the quantile regression models using directed acyclic graphs and selected the model based on Akaike information criteria. Yearly outpatient antibiotic consumption in Norway varied up to 10-fold at municipality level. We found geographical variation to depend on the number of inhabitants in a municipality and on latitude. These variables interacted, so that consumption declined with increasing latitude when municipality population sizes were small, but the effect of latitude diminished as the number of inhabitants increased. Aggregation to different levels of spatial resolution did not significantly affect our results. In Norway, outpatient antibiotic dispensing rates decreases with latitude at a rate contingent on municipality population size. Quantile regression analysis provides a flexible and powerful tool to address problems related to high, or low, dispensing rates.

Costs and benefits of natural transformation in Acinetobacter baylyi.

BACKGROUND: Natural transformation enables acquisition of adaptive traits and drives genome evolution in prokaryotes. Yet, the selective forces responsible for the evolution and maintenance of natural transformation remain elusive since taken-up DNA has also been hypothesized to provide benefits such as nutrients or templates for DNA repair to individual cells. RESULTS: We investigated the immediate effects of DNA uptake and recombination on the naturally competent bacterium Acinetobacter baylyi in both benign and genotoxic conditions. In head-to-head competition experiments between DNA uptake-proficient and -deficient strains, we observed a fitness benefit of DNA uptake independent of UV stress. This benefit was found with both homologous and heterologous DNA and was independent of recombination. Recombination with taken-up DNA reduced survival of transformed cells with increasing levels of UV-stress through interference with nucleotide excision repair, suggesting that DNA strand breaks occur during recombination attempts with taken-up DNA. Consistent with this, we show that absence of RecBCD and RecFOR recombinational DNA repair pathways strongly decrease natural transformation. CONCLUSIONS: Our data show a physiological benefit of DNA uptake unrelated to recombination. In contrast, recombination during transformation is a strand break inducing process that represents a previously unrecognized cost of natural transformation.

Climate change alters the structure of arctic marine food webs due to poleward shifts of boreal generalists.

Climate-driven poleward shifts, leading to changes in species composition and relative abundances, have been recently documented in the Arctic. Among the fastest moving species are boreal generalist fish which are expected to affect arctic marine food web structure and ecosystem functioning substantially. Here, we address structural changes at the food web level induced by poleward shifts via topological network analysis of highly resolved boreal and arctic food webs of the Barents Sea. We detected considerable differences in structural properties and link configuration between the boreal and the arctic food webs, the latter being more modular and less connected. We found that a main characteristic of the boreal fish moving poleward into the arctic region of the Barents Sea is high generalism, a property that increases connectance and reduces modularity in the arctic marine food web. Our results reveal that habitats form natural boundaries for food web modules, and that generalists play an important functional role in coupling pelagic and benthic modules. We posit that these habitat couplers have the potential to promote the transfer of energy and matter between habitats, but also the spread of pertubations, thereby changing arctic marine food web structure considerably with implications for ecosystem dynamics and functioning.

Climate-driven regime shifts in Arctic marine benthos.

Climate warming can trigger abrupt ecosystem changes in the Arctic. Despite the considerable interest in characterizing and understanding the ecological impact of rapid climate warming in the Arctic, few long time series exist that allow addressing these research goals. During a 30-y period (1980-2010) of gradually increasing seawater temperature and decreasing sea ice cover in Svalbard, we document rapid and extensive structural changes in the rocky-bottom communities of two Arctic fjords. The most striking component of the benthic reorganization was an abrupt fivefold increase in macroalgal cover in 1995 in Kongsfjord and an eightfold increase in 2000 in Smeerenburgfjord. Simultaneous changes in the abundance of benthic invertebrates suggest that the macroalgae played a key structuring role in these communities. The abrupt, substantial, and persistent nature of the changes observed is indicative of a climate-driven ecological regime shift. The ecological processes thought to drive the observed regime shifts are likely to promote the borealization of these Arctic marine communities in the coming years.

Parasite communities of two three-spined stickleback populations in subarctic Norway--effects of a small spatial-scale host introduction.

Co-introduction and colonization of parasites with the introduction of new host species into aquatic habitats may depend on the host specificity and dispersal capabilities of the parasites. We compared the metazoan parasite community of an introduced three-spined stickleback (Gasterosteus aculeatus) population with that of the nearby source population in subarctic Norway. As expected from a small spatial scale (5 km), the parasite component communities in the two lakes were highly similar. All identifiable allogenic parasite taxa (Diphyllobothrium dendriticum, Diphyllobothrium ditremum, Diphyllobothrium spp., Schistocephalus solidus, Apatemon sp. and Diplostomum spp.) were also observed in both lakes while inter-lake differences were driven by autogenic parasite taxa (Eubothrium spp., Crepidostomum spp., Nematoda spp., Proteocephalus sp. and Gyrodactylus arcuatus). Contrary to expectation, the total number of parasite taxa was higher in the introduced stickleback population (12) compared to that found in the source population (9) with three parasite taxa (Eubothrium spp., Crepidostomum spp., Nematoda spp.) only occurring in the introduced population. These parasites were uncommon however and normally restricted to salmonids. Sticklebacks from both populations were heavily infected, particularly with eye-infecting metacercariae. Sequences from the DNA barcode region of cytochrome oxidase 1 indicated that these include Diplostomum lineage 6, a member of the Diplostomum baeri complex and a member of the Strigeinae. Despite high similarity between the two component communities, quantitative inter-lake differences were found at the infracommunity level. At this scale, parasite intensity was significantly higher in the source population for the two autogenic stickleback specialists: G. arcuatus and Proteocephalus sp., assumed to be the autogenic stickleback specialist Proteocephalus filicollis. Parasite infracommunities within each lake also resembled each other significantly more than infracommunities between lakes, primarily driven by the allogenic cestode D. ditremum, as well as G. arcuatus and Proteocephalus sp. Overall, quantitative dissimilarities between the two parasite communities were possibly explained by inter-lake differences in the density of sticklebacks and intermediate hosts.

A trade-off between the fitness cost of functional integrases and long-term stability of integrons.

Horizontal gene transfer (HGT) plays a major role in bacterial microevolution as evident from the rapid emergence and spread of antimicrobial drug resistance. Few studies have however addressed the population dynamics of newly imported genetic elements after HGT. Here, we show that newly acquired class-1 integrons from Salmonella enterica serovar Typhimurium and Acinetobacter baumannii, free of associated transposable elements, strongly reduce host fitness in Acinetobacter baylyi. Insertional inactivation of the integron intI1 restored fitness, demonstrating that the observed fitness costs were due to the presence of an active integrase. The biological cost of harboring class-1 integrons was rapidly reduced during serial transfers due to intI1 frameshift mutations leading to inactivated integrases. We use a mathematical model to explore the conditions where integrons with functional integrases are maintained and conclude that environmental fluctuations and episodic selection is necessary for the maintenance of functional integrases. Taken together, the presented data suggest a trade-off between the ability to capture gene cassettes and long-term stability of integrons and provide an explanation for the frequent observation of inactive integron-integrases in bacterial populations.

Under ice plankton and lipid dynamics in a subarctic lake.

Climate warming causes shorter winters and changes in ice and snow cover in subarctic lakes, highlighting the need to better understand under-ice ecosystem functioning. The plankton community in a subarctic, oligotrophic lake was studied throughout the ice-covered season, focusing on lipid dynamics and life history traits in two actively overwintering copepods, Cyclops scutifer and Eudiaptomus graciloides. Whereas C. scutifer was overwintering in C-IV to C-V stage, E. graciloides reproduced under ice cover. Both species had accumulated lipids prior to ice-on and showed a substantial decrease in total lipid content throughout the ice-covered period: E. graciloides (60%-38% dw) and C. scutifer (73%-33% dw). Polyunsaturated fatty acids of algal origin were highest in E. graciloides and declined strongly in both species. Stearidonic acid (18:4n-3) content in E. graciloides was particularly high and decreased rapidly during the study period by 50%, probably due to reproduction. The copepods differed in feeding behavior, with the omnivore C. scutifer continuing to accumulate lipids until January, whereas the herbivorous E. graciloides accumulated lipids from under-ice primary production during the last months of ice-cover. Our findings emphasize the importance of lipid accumulation and utilization for actively overwintering copepods irrespective of the timing of their reproduction.

New parasites and predators follow the introduction of two fish species to a subarctic lake: implications for food-web structure and functioning.

Introduced species can alter the topology of food webs. For instance, an introduction can aid the arrival of free-living consumers using the new species as a resource, while new parasites may also arrive with the introduced species. Food-web responses to species additions can thus be far more complex than anticipated. In a subarctic pelagic food web with free-living and parasitic species, two fish species (arctic charr Salvelinus alpinus and three-spined stickleback Gasterosteus aculeatus) have known histories as deliberate introductions. The effects of these introductions on the food web were explored by comparing the current pelagic web with a heuristic reconstruction of the pre-introduction web. Extinctions caused by these introductions could not be evaluated by this approach. The introduced fish species have become important hubs in the trophic network, interacting with numerous parasites, predators and prey. In particular, five parasite species and four predatory bird species depend on the two introduced species as obligate trophic resources in the pelagic web and could therefore not have been present in the pre-introduction network. The presence of the two introduced fish species and the arrival of their associated parasites and predators increased biodiversity, mean trophic level, linkage density, and nestedness; altering both the network structure and functioning of the pelagic web. Parasites, in particular trophically transmitted species, had a prominent role in the network alterations that followed the introductions.

Climate warming accelerates somatic growth of an Arctic fish species in high-latitude lakes.

High-latitude aquatic ecosystems are responding to rapid climate warming. A longer ice-free season with higher water temperatures may accelerate somatic growth in lake ectotherms, leading to widespread ecological implications. In fish, rising temperatures are expected to boost rates of food intake and conversion, and predictions based on empirical relationships between temperature and growth suggest a substantial increase in fish growth rates during the last decades. Fish abundance negatively affects growth by limiting food availability. This field study addresses the effects of climate warming on growth of a subarctic population of Arctic charr (Salvelinus alpinus (L.) over nearly 40 years. Juvenile growth of 680 individuals of Arctic charr, was reconstructed by sclerochronological analysis using sagittal otoliths sampled annually from the early 1980s to 2016. Statistical modelling revealed a positive effect of water temperature, and a negative effect of abundance on somatic growth in juvenile individuals. Temperature dependence in growth was significant for average and fast-growing individuals across all investigated age classes. These findings suggest that, as temperatures rise, somatic growth of Arctic charr will increase in high latitude lakes. Climate warming will thus influence cold water fish life history and size-structured interactions, with important consequences for their populations and ecosystems.