The role of the circumarctic forest–tundra ecotone for Arctic biodiversity

Accepted VersionThe arctic forest–tundra ecotone (FTE), which links species communities of the boreal forest with those of the arctic tundra, is expected to respond swiftly to climate change with a profound reduction of tundra as the dominating scenario. With its circumarctic expanse and up to several hundred kilometres in width, the zone occupies a large part of the vegetated surface at high latitudes. Relocation and structural changes of the ecotone vegetation will affect not only plant but also animal and other biological diversity. A large number of arctic species are dependent on the forest–tundra ecotone in terms of food and habitat during parts of their life cycle or annual migration. In the ‘Arctic Species Trend Index’, developed to provide trends in arctic vertebrates, more than half of the species and data are from the forest–tundra ecotone. However, in assessments of arctic biodiversity, only the northernmost tundra-dominated areas of the ecotone are included. This is unfortunate and somewhat problematic since the treed part that serves as a source of seeds for new seedlings and saplings in the tundra-dominated part is excluded. This inconsistency hampers monitoring efficiency and biodiversity conservation efforts. During the International Polar Year, a large international research project on the forest–tundra ecotone established numerous sites around the circumpolar north where causes and consequences of vegetation change were analyzed. This network of sites and data forms an excellent basis for necessary monitoring of the spatial and temporal complexity of forest encroachment into tundra and its relation to arctic biodiversity

Is subarctic forest advance able to keep pace with climate change?

Published VersionRecent climate warming and scenarios for further warming have led to expectations of rapid movement of ecological boundaries. Here we focus on the circumarctic forest–tundra ecotone (FTE), which represents an important bioclimatic zone with feedbacks from forest advance and corresponding tundra disappearance (up to 50% loss predicted this century) driving widespread ecological and climatic changes. We address FTE advance and climate history relations over the 20th century, using FTE response data from 151 sites across the circumarctic area and site-specific climate data. Specifically, we investigate spatial uniformity of FTE advance, statistical asso ciations with 20th century climate trends, and whether advance rates match climate change velocities (CCVs). Study sites diverged into four regions (Eastern Canada; Central and Western Canada and Alaska; Siberia; and Western Eurasia) based on their climate history, although all were characterized by similar qualitative patterns of behaviour (with about half of the sites showing advancing behaviour). The main associations between climate trend variables and behaviour indicate the importance of precipitation rather than temperature for both qualitative and quantitative behav iours, and the importance of non-growing season as well as growing season months. Poleward latitudinal advance rates differed significantly among regions, being small est in Eastern Canada (~10 m/year) and largest in Western Eurasia (~100 m/year). These rates were 1–2 orders of magnitude smaller than expected if vegetation dis tribution remained in equilibrium with climate. The many biotic and abiotic factors influencing FTE behaviour make poleward advance rates matching predicted 21st century CCVs (~103–104 m/year) unlikely. The lack of empirical evidence for swift forest relocation and the discrepancy between CCV and FTE response contradict equilibrium model-based assumptions and warrant caution when assessing global change-related biotic and abiotic implications, including land–atmosphere feedbacks and carbon sequestration

Experiment, monitoring, and gradient methods used to infer climate change effects on plant communities yield consistent patterns

Inference about future climate change impacts typically relies on one of three approaches: manipulative experiments, historical comparisons (broadly defined to include monitoring the response to ambient climate fluctuations using repeat sampling of plots, dendroecology, and paleoecology techniques), and space-for-time substitutions derived from sampling along environmental gradients. Potential limitations of all three approaches are recognized. Here we address the congruence among these three main approaches by comparing the degree to which tundra plant community composition changes (i) in response to in situ experimental warming, (ii) with interannual variability in summer temperature within sites, and (iii) over spatial gradients in summer temperature. We analyzed changes in plant community composition from repeat sampling (85 plant communities in 28 regions) and experimental warming studies (28 experiments in 14 regions) throughout arctic and alpine North America and Europe. Increases in the relative abundance of species with a warmer thermal niche were observed in response to warmer summer temperatures using all three methods; however, effect sizes were greater over broad-scale spatial gradients relative to either temporal variability in summer temperature within a site or summer temperature increases induced by experimental warming. The effect sizes for change over time within a site and with experimental warming were nearly identical. These results support the view that inferences based on space-for-time substitution overestimate the magnitude of responses to contemporary climate warming, because spatial gradients reflect long-term processes. In contrast, in situ experimental warming and monitoring approaches yield consistent estimates of the magnitude of response of plant communities to climate warming

Reproduction as a bottleneck to treeline advance across the circumarctic forest tundra ecotone

Published versionThe fundamental niche of many species is shifting with climate change, especially in sub-arctic ecosystems with pronounced recent warming. Ongoing warming in sub-arctic regions should lessen environmental constraints on tree growth and reproduction, leading to increased success of trees colonizing tundra. Nevertheless, variable responses of treeline ecotones have been documented in association with warming temperatures. One explanation for time lags between increasingly favourable environmental conditions and treeline ecotone movement is reproductive limitations caused by low seed availability. Our objective was to assess the reproductive constraints of the dominant tree species at the treeline ecotone in the circumpolar north. We sampled reproductive structures of trees (cones and catkins) and stand attributes across circumarctic treeline ecotones. We used generalized linear mixed models to estimate the sensitivity of seed production and the availability of viable seed to regional climate, stand structure, and species-specific characteristics. Both seed production and viability of available seed were strongly driven by specific, sequential seasonal climatic conditions, but in different ways. Seed production was greatest when growing seasons with more growing degree days coincided with years with high precipitation. Two consecutive years with more growing degree days and low precipitation resulted in low seed production. Seasonal climate effects on the viability of available seed depended on the physical characteristics of the reproductive structures. Large-coned and -seeded species take more time to develop mature embryos and were therefore more sensitive to increases in growing degree days in the year of flowering and embryo development. Our findings suggest that both moisture stress and abbreviated growing seasons can have a notable negative influence on the production and viability of available seed at treeline. Our synthesis revealed that constraints on pre-dispersal reproduction within the treeline ecotone might create a considerable time lag for range expansion of tree populations into tundra ecosystems

BioTIME: A database of biodiversity time series for the Anthropocene.

MotivationThe BioTIME database contains raw data on species identities and abundances in ecological assemblages through time. These data enable users to calculate temporal trends in biodiversity within and amongst assemblages using a broad range of metrics. BioTIME is being developed as a community-led open-source database of biodiversity time series. Our goal is to accelerate and facilitate quantitative analysis of temporal patterns of biodiversity in the Anthropocene.Main types of variables includedThe database contains 8,777,413 species abundance records, from assemblages consistently sampled for a minimum of 2 years, which need not necessarily be consecutive. In addition, the database contains metadata relating to sampling methodology and contextual information about each record.Spatial location and grainBioTIME is a global database of 547,161 unique sampling locations spanning the marine, freshwater and terrestrial realms. Grain size varies across datasets from 0.0000000158 km2 (158 cm2) to 100 km2 (1,000,000,000,000 cm2).Time period and grainBioTIME records span from 1874 to 2016. The minimal temporal grain across all datasets in BioTIME is a year.Major taxa and level of measurementBioTIME includes data from 44,440 species across the plant and animal kingdoms, ranging from plants, plankton and terrestrial invertebrates to small and large vertebrates.Software format.csv and .SQL

Circum-Arctic distribution of chemical anti-herbivore compounds suggests biome-wide trade-off in defence strategies in Arctic shrubs

Spatial variation in plant chemical defence towards herbivores can help us understand variation in herbivore top-down control of shrubs in the Arctic and possibly also shrub responses to global warming. Less defended, non-resinous shrubs could be more influenced by herbivores than more defended, resinous shrubs. However, sparse field measurements limit our current understanding of how much of the circum-Arctic variation in defence compounds is explained by taxa or defence functional groups (resinous/non-resinous). We measured circum-Arctic chemical defence and leaf digestibility in resinous (Betula glandulosa, B. nana ssp. exilis) and non-resinous (B. nana ssp. nana, B. pumila) shrub birches to see how they vary among and within taxa and functional groups. Using liquid chromatography-mass spectrometry (LC-MS) metabolomic analyses and in vitro leaf digestibility via incubation in cattle rumen fluid, we analysed defence composition and leaf digestibility in 128 samples from 44 tundra locations. We found biogeographical patterns in anti-herbivore defence where mean leaf triterpene concentrations and twig resin gland density were greater in resinous taxa and mean concentrations of condensing tannins were greater in non-resinous taxa. This indicates a biome-wide trade-off between triterpene- or tannin-dominated defences. However, we also found variations in chemical defence composition and resin gland density both within and among functional groups (resinous/non-resinous) and taxa, suggesting these categorisations only partly predict chemical herbivore defence. Complex tannins were the only defence compounds negatively related to in vitro digestibility, identifying this previously neglected tannin group as having a potential key role in birch anti-herbivore defence. We conclude that circum-Arctic variation in birch anti-herbivore defence can be partly derived from biogeographical distributions of birch taxa, although our detailed mapping of plant defence provides more information on this variation and can be used for better predictions of herbivore effects on Arctic vegetation

Circum-Arctic distribution of chemical anti-herbivore compounds suggests biome-wide trade-off in defence strategies in Arctic shrubs

Spatial variation in plant chemical defence towards herbivores can help us understand variation in herbivore top-down control of shrubs in the Arctic and possibly also shrub responses to global warming. Less defended, non-resinous shrubs could be more influenced by herbivores than more defended, resinous shrubs. However, sparse field measurements limit our current understanding of how much of the circum-Arctic variation in defence compounds is explained by taxa or defence functional groups (resinous/non-resinous). We measured circum-Arctic chemical defence and leaf digestibility in resinous (Betula glandulosa, B. nana ssp. exilis) and non-resinous (B. nana ssp. nana, B. pumila) shrub birches to see how they vary among and within taxa and functional groups. Using liquid chromatography-mass spectrometry (LC-MS) metabolomic analyses and in vitro leaf digestibility via incubation in cattle rumen fluid, we analysed defence composition and leaf digestibility in 128 samples from 44 tundra locations. We found biogeographical patterns in anti-herbivore defence where mean leaf triterpene concentrations and twig resin gland density were greater in resinous taxa and mean concentrations of condensing tannins were greater in non-resinous taxa. This indicates a biome-wide trade-off between triterpene- or tannin-dominated defences. However, we also found variations in chemical defence composition and resin gland density both within and among functional groups (resinous/non-resinous) and taxa, suggesting these categorisations only partly predict chemical herbivore defence. Complex tannins were the only defence compounds negatively related to in vitro digestibility, identifying this previously neglected tannin group as having a potential key role in birch anti-herbivore defence. We conclude that circum-Arctic variation in birch anti-herbivore defence can be partly derived from biogeographical distributions of birch taxa, although our detailed mapping of plant defence provides more information on this variation and can be used for better predictions of herbivore effects on Arctic vegetation.Peer reviewe

Psycho-social factors associated with mental resilience in the Corona lockdown.

The SARS-CoV-2 pandemic is not only a threat to physical health but is also having severe impacts on mental health. Although increases in stress-related symptomatology and other adverse psycho-social outcomes, as well as their most important risk factors have been described, hardly anything is known about potential protective factors. Resilience refers to the maintenance of mental health despite adversity. To gain mechanistic insights about the relationship between described psycho-social resilience factors and resilience specifically in the current crisis, we assessed resilience factors, exposure to Corona crisis-specific and general stressors, as well as internalizing symptoms in a cross-sectional online survey conducted in 24 languages during the most intense phase of the lockdown in Europe (22 March to 19 April) in a convenience sample of N = 15,970 adults. Resilience, as an outcome, was conceptualized as good mental health despite stressor exposure and measured as the inverse residual between actual and predicted symptom total score. Preregistered hypotheses (osf.io/r6btn) were tested with multiple regression models and mediation analyses. Results confirmed our primary hypothesis that positive appraisal style (PAS) is positively associated with resilience (p < 0.0001). The resilience factor PAS also partly mediated the positive association between perceived social support and resilience, and its association with resilience was in turn partly mediated by the ability to easily recover from stress (both p < 0.0001). In comparison with other resilience factors, good stress response recovery and positive appraisal specifically of the consequences of the Corona crisis were the strongest factors. Preregistered exploratory subgroup analyses (osf.io/thka9) showed that all tested resilience factors generalize across major socio-demographic categories. This research identifies modifiable protective factors that can be targeted by public mental health efforts in this and in future pandemics