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The on-going climate warming is promoting shrub abundance in high latitudes, but the effect of this phenomenon on ecosystem functioning is expected to depend on whether deciduous or evergreen species increase in response to warming. To explore effects of long-term warming on shrubs and further on ecosystem functioning, we analysed vegetation and ecosystem CO2 exchange after 20 years of warming in the forest–tundra ecotone in subarctic Sweden. A previous study conducted 9 years earlier had found increased evergreen Empetrum nigrum ssp. hermaphroditum in the forest and increased deciduous Betula nana in the tundra. Following current understanding, we expected continued increase in shrub abundance that would be stronger in tundra than in forest. We expected warming to increase ecosystem respiration (Re) and gross primary productivity (GPP), with a greater increase in Re in tundra due to increased deciduous shrub abundance, leading to a less negative net ecosystem exchange and reduced ecosystem C sink strength. As predicted, vascular plant abundances were higher in the warmed plots with a stronger response in tundra than in forest. However, whereas B. nana had increased in abundance since the last survey, E. hermaphroditum abundance had declined due to several moth and rodent outbreaks during the past decade. In contrast to predictions, Re was significantly lower in the warmed plots irrespective of habitat, and GPP increased marginally only in the forest. The lower Re and a higher GPP under warming in the forest together led to increased net C sink. Re was negatively associated with the total vascular plant abundance. Our results highlight the importance of disturbance regimes for vegetation responses to warming. Climate warming may promote species with both a high capacity to grow under warmer conditions and a resilience towards herbivore outbreaks. Negative correlation between Re and total vascular plant abundance further indicate that the indirect impacts of increase

Protected areas are considered fundamental to counter biodiversity loss. However, evidence for their effectiveness in averting local extinctions remains scarce and taxonomically biased. We employ a robust counterfactual multi-taxon approach to compare occupancy patterns of 638 species, including birds (150), mammals (23), plants (39) and phytoplankton (426) between protected and unprotected sites across four decades in Finland. We find mixed impacts of protected areas, with only a small proportion of species explicitly benefiting from protection-mainly through slower rates of decline inside protected areas. The benefits of protection are enhanced for larger protected areas and are traceable to when the sites were protected, but are mostly unrelated to species conservation status or traits (size, climatic niche and threat status). Our results suggest that the current protected area network can partly contribute to slow down declines in occupancy rates, but alone will not suffice to halt the biodiversity crisis. Efforts aimed at improving coverage, connectivity and management will be key to enhance the effectiveness of protected areas towards bending the curve of biodiversity loss.

This zipped folder includes all data and the scripts to replicate the analyses from the study by Santangeli et al. Mixed effects of protected areas on terrestrial and freshwater biodiversity. Specifically, we include in the main folder the script (R file, suitable for R software - "SiteMatchingBirdExample") and site level data ("BirdData.csv") used to run the matching analyses of protected and unportected sites (data and script provided for the birds data only due to data ownership agreements). This example is easily applicable to the other three taxonomic groups. We also include the data and scripts to run the Joint Species Distribution Modeling on the occurrence data separately for each of the four taxonomic groups studied in the folder "HMSC_models". Here the user will find four separate sub-folders, each including in turn a folder named data which hosts the occurrence data from sites selected after matching for that taxon (CSV file named SXY, includes also site level information), as well as a script file (named e.g. birds_HMSC in the case of birds folder) that can be used to replicate the joint species distribution model on that specific taxonomic group.

Climate change is leading to species redistributions. In the tundra biome, shrubs are generally expanding, but not all tundra shrub species will benefit from warming. Winner and loser species, and the characteristics that may determine success or failure, have not yet been fully identified. Here, we investigate whether past abundance changes, current range sizes and projected range shifts derived from species distribution models are related to plant trait values and intraspecific trait variation. We combined 17,921 trait records with observed past and modelled future distributions from 62 tundra shrub species across three continents. We found that species with greater variation in seed mass and specific leaf area had larger projected range shifts, and projected winner species had greater seed mass values. However, trait values and variation were not consistently related to current and projected ranges, nor to past abundance change. Overall, our findings indicate that abundance change and range shifts will not lead to directional modifications in shrub trait composition, since winner and loser species share relatively similar trait spaces.

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 it varies among and within taxa and functional groups. Using 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, 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 it varies among and within taxa and functional groups. Using 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

Identifying the ecological processes underlying community assembly remains an elusive goal in community ecology. We formalize assembly hypotheses as alternative models and apply each to predict 1,918 out-of-sample boreal forest understory communities to identify and separate the processes driving community assembly. Models are specified within a Bayesian joint species distribution framework that allows for the inclusion and separation of stochastic processes, environmental filtering, and two different species dependence structures. We found clear evidence that study communities are structured by both environmental filtering and compositional dependence highlighting the importance of selection in community assembly. The relative importance of environmental filtering was greater than compositional dependence in predicting both understory communities and the abundance of constituent species across broad suc-cessional and bioclimatic gradients. Contrary to ecological expectations, the inclusion of a flexible residual species dependence structure (accounting for more than compositional dependence) did not improve model predictions after accounting for the strong role of environmental filtering. Our results provide novel inference on the processes underlying community assembly facilitated by applying empirical approximations of alternative assembly processes to predict communities across a range of environmental conditions.

Aim The diversity and composition of natural communities are rapidly changing due to anthropogenic disturbances. Magnitude of this compositional reorganization varies across the globe, but reasons behind the variation remain largely unknown. Disturbances induce temporal turnover by stimulating species colonizations, causing local extinctions, altering dominance structure, or all of these. We test which of these processes drive temporal community changes, and whether they are constrained by natural environmental gradients. Moreover, we assess to what degree identity shifts translate to changes in dominance structure. Location Finland. Time period Observations 1985–2006, disturbance history > 140 years. Major taxa studied Vascular plants. Methods We investigated temporal turnover of boreal forest understorey in response to disturbance, here forest management, along a soil fertility gradient. We disentangle the roles of species gains, losses and abundance changes in driving temporal turnover in response to and after disturbance by comparing turnover rates in different forest age categories along a fertility gradient. We quantify temporal turnover using richness‐based complement of Jaccard’s similarity index and proportional‐abundance based dissimilarity index. We also test whether disturbance history or fertility influence the relationship between identity shifts and dominance structure. Results We found that the impact of disturbance on temporal turnover depends on soil fertility. The greatest turnover occurred in the most fertile forests immediately after disturbance. There, species gains and losses strongly altered dominance structure leading to high turnover, whereas undisturbed old forests and nutrient‐poor habitats were characterized by stable dominant species even when the majority of species shifted their identity. Main conclusions Our results suggest that human impacts on temporal biodiversity change vary along environmental gradients. In boreal forests, the f

The circumpolar Arctic is currently facing multiple global changes that have the potential to alter the capacity of tundra soils to store carbon. Yet, predicting changes in soil carbon is hindered by the fact that multiple factors simultaneously control processes sustaining carbon storage and we do not understand how they act in concert. Here, we investigated the effects of warmer temperatures, enhanced soil nitrogen availability, and the combination of these on tundra carbon stocks at three different grazing regimes: on areas with over 50-yr history of either light or heavy reindeer grazing and in 5-yr-old exlosures in the heavily grazed area. In line with earlier reports, warming generally decreased soil carbon stocks. However, our results suggest that the mechanisms by which warming decreases carbon storage depend on grazing intensity: under long-term light grazing soil carbon losses were linked to higher shrub abundance and higher enzymatic activities, whereas under long-term heavy grazing, carbon losses were linked to drier soils and higher enzymatic activities. Importantly, under enhanced soil nitrogen availability, warming did not induce soil carbon losses under either of the long-term grazing regimes, whereas inside exclosures in the heavily grazed area, also the combination of warming and enhanced nutrient availability induced soil carbon loss. Grazing on its own did not influence the soil carbon stocks. These results reveal that accounting for the effect of warming or grazing alone is not sufficient to reliably predict future soil carbon storage in the tundra. Instead, the joint effects of multiple global changes need to be accounted for, with a special focus given to abrupt changes in grazing currently taking place in several parts of the Arctic.

Aim: Plant functional groups are widely used in community ecology and earth system modelling to describe trait variation within and across plant communities. However, this approach rests on the assumption that functional groups explain a large proportion of trait variation among species. We test whether four commonly used plant functional groups represent variation in six ecologically important plant traits. Location: Tundra biome. Time period: Data collected between 1964 and 2016. Major taxa studied: 295 tundra vascular plant species. Methods: We compiled a database of six plant traits (plant height, leaf area, specific leaf area, leaf dry matter content, leaf nitrogen, seed mass) for tundra species. We examined the variation in species-level trait expression explained by four traditional functional groups (evergreen shrubs, deciduous shrubs, graminoids, forbs), and whether variation explained was dependent upon the traits included in analysis. We further compared the explanatory power and species composition of functional groups to alternative classifications generated using post hoc clustering of species-level traits. Results: Traditional functional groups explained significant differences in trait expression, particularly amongst traits associated with resource economics, which were consistent across sites and at the biome scale. However, functional groups explained 19% of overall trait variation and poorly represented differences in traits associated with plant size. Post hoc classification of species did not correspond well with traditional functional groups, and explained twice as much variation in species-level trait expression. Main conclusions: Traditional functional groups only coarsely represent variation in well-measured traits within tundra plant communities, and better explain resource economic traits than size-related traits. We recommend caution when using functional group approaches to predict tundra vegetation change, or ecosystem functions relati