In situ seasonal patterns of root auxin concentrations and meristem length in an arctic sedge.

Seasonal dynamics of root growth play an important role in large-scale ecosystem processes; they are largely governed by growth regulatory compounds and influenced by environmental conditions. Yet, our knowledge about physiological drivers of root growth is mostly limited to laboratory-based studies on model plant species. We sampled root tips of Eriophorum vaginatum and analyzed their auxin concentrations and meristem lengths biweekly over a growing season in situ in a subarctic peatland, both in surface soil and at the permafrost thawfront. Auxin concentrations were almost five times higher in surface than in thawfront soils and increased over the season, especially at the thawfront. Surprisingly, meristem length showed an opposite pattern and was almost double in thawfront compared with surface soils. Meristem length increased from peak to late season in the surface soils but decreased at the thawfront. Our study of in situ seasonal dynamics in root physiological parameters illustrates the potential for physiological methods to be applied in ecological studies and emphasizes the importance of in situ measurements. The strong effect of root location and the unexpected opposite patterns of meristem length and auxin concentrations likely show that auxin actively governs root growth to ensure a high potential for nutrient uptake at the thawfront.

Regional invasion history and land use shape the prevalence of non-native species in local assemblages.

The ecological impact of non-native species arises from their establishment in local assemblages. However, the rates of non-native spread in new regions and their determinants have not been comprehensively studied. Here, we combined global databases documenting the occurrence of non-native species and residence of non-native birds, mammals, and vascular plants at regional and local scales to describe how the likelihood of non-native occurrence and their proportion in local assemblages relate with their residence time and levels of human usage in different ecosystems. Our findings reveal that local non-native occurrence generally increases with residence time. Colonization is most rapid in croplands and urban areas, while it is slower and variable in natural or semi-natural ecosystems. Notably, non-native occurrence continues to rise even 200 years after introduction, especially for birds and vascular plants, and in other land-use types rather than croplands and urban areas. The impact of residence time on non-native proportions is significant only for mammals. We conclude that the continental exchange of biotas requires considerable time for effects to manifest at the local scale across taxa and land-use types. The unpredictability of future impacts, implied by the slow spread of non-native species, strengthens the call for stronger regulations on the exchange of non-native species to reduce the long-lasting invasion debt looming on ecosystems' future.

Human appropriation of net primary production as driver of change in landscape-scale vertebrate richness.

Aim: Land use is the most pervasive driver of biodiversity loss. Predicting its impact on species richness (SR) is often based on indicators of habitat loss. However, the degradation of habitats, especially through land-use intensification, also affects species. Here, we evaluate whether an integrative metric of land-use intensity, the human appropriation of net primary production, is correlated with the decline of SR in used landscapes across the globe. Location: Global. Time period: Present. Major taxa studied: Birds, mammals and amphibians. Methods: Based on species range maps (spatial resolution: 20 km × 20 km) and an area-of-habitat approach, we calibrated a "species-energy model" by correlating the SR of three groups of vertebrates with net primary production and biogeographical covariables in "wilderness" areas (i.e., those where available energy is assumed to be still at pristine levels). We used this model to project the difference between pristine SR and the SR corresponding to the energy remaining in used landscapes (i.e., SR loss expected owing to human energy extraction outside wilderness areas). We validated the projected species loss by comparison with the realized and impending loss reconstructed from habitat conversion and documented by national Red Lists. Results: Species-energy models largely explained landscape-scale variation of mapped SR in wilderness areas (adjusted R 2-values: 0.79-0.93). Model-based projections of SR loss were lower, on average, than reconstructed and documented ones, but the spatial patterns were correlated significantly, with stronger correlation in mammals (Pearson's r = 0.68) than in amphibians (r = 0.60) and birds (r = 0.57). Main conclusions: Our results suggest that the human appropriation of net primary production is a useful indicator of heterotrophic species loss in used landscapes, hence we recommend its inclusion in models based on species-area relationships to improve predictions of land-use-driven biodiversity loss.

Climate change leads to accelerated transformation of high-elevation vegetation in the central Alps.

High mountain ecosystems and their biota are governed by low-temperature conditions and thus can be used as indicators for climate warming impacts on natural ecosystems, provided that long-term data exist. We used data from the largest alpine to nival permanent plot site in the Alps, established in the frame of the Global Observation Research Initiative in Alpine Environments (GLORIA) on Schrankogel in the Tyrolean Alps, Austria, in 1994, and resurveyed in 2004 and 2014. Vascular plant species richness per plot increased over the entire period, albeit to a lesser extent in the second decade, because disappearance events increased markedly in the latter period. Although presence/absence data could only marginally explain range shift dynamics, changes in species cover and plant community composition indicate an accelerating transformation towards a more warmth-demanding and more drought-adapted vegetation, which is strongest at the lowest, least rugged subsite. Divergent responses of vertical distribution groups of species suggest that direct warming effects, rather than competitive displacement, are the primary causes of the observed patterns. The continued decrease in cryophilic species could imply that trailing edge dynamics proceed more rapidly than successful colonisation, which would favour a period of accelerated species declines.

The global biodiversity footprint of urban consumption: A spatially explicit assessment for the city of Vienna.

With ongoing global urbanization processes and consumption patterns increasingly recognized as key determinants of environmental change, a better understanding of the links between urban consumption and biodiversity loss is paramount. Here we quantify the global biodiversity footprint (BDF) of Vienna's (Austria) biomass consumption. We present a state-of-the-art product specific approach to (a) locate the production areas required for Vienna's consumption and map Vienna's BDF by (b) linking them with data taken from a previously published countryside Species-Area-Relationship (cSAR) model with a representation of land-use intensity. We found that food has the largest share in Vienna's BDF (58 %), followed by biomass for material applications (28 %) and bioenergy (13 %). The total BDF occurs predominantly within Austria and in its neighbouring countries, with ~20 % located outside Europe. Although the per capita biomass consumption in Vienna is above the global average, global and Viennese per capita BDFs are roughly equal, indicating that Vienna sources its products from high-yield regions with efficient production systems and comparatively low native species richness. We conclude that, among others, dietary changes offer a key leverage point for reducing the urban BDF, while expanding the use of biomass for material and energy use may increase the BDF and requires appropriate monitoring.

The impact of land use on non-native species incidence and number in local assemblages worldwide.

While the regional distribution of non-native species is increasingly well documented for some taxa, global analyses of non-native species in local assemblages are still missing. Here, we use a worldwide collection of assemblages from five taxa - ants, birds, mammals, spiders and vascular plants - to assess whether the incidence, frequency and proportions of naturalised non-native species depend on type and intensity of land use. In plants, assemblages of primary vegetation are least invaded. In the other taxa, primary vegetation is among the least invaded land-use types, but one or several other types have equally low levels of occurrence, frequency and proportions of non-native species. High land use intensity is associated with higher non-native incidence and frequency in primary vegetation, while intensity effects are inconsistent for other land-use types. These findings highlight the potential dual role of unused primary vegetation in preserving native biodiversity and in conferring resistance against biological invasions.

Effects of chronic crude oil exposure on the fitness of polar cod (Boreogadus saida) through changes in growth, energy reserves and survival.

Climate models predict extended periods with sea-ice free Arctic waters during the next decade, which will allow more shipping activity and easier access to petroleum resources. Increased industrial activities raise concerns about the biological effects of accidental petroleum release on key species of the Arctic marine ecosystem, such as the polar cod (Boreogadus saida). This study examines effects on physiological traits related to the fitness of adult polar cod, such as growth, survival, and lipid parameters. Fish were exposed to environmentally-relevant crude oil doses through their diet over an 8-month period, concurrent with reproductive development. In liver tissue, lipid class composition differed between treatments while in gonad tissue, lipid class composition varied between sexes, but not treatments. Crude oil did not affect growth and survival, which indicated that polar cod were relatively robust to dietary crude oil exposure at doses tested (0.11-1.14 µg crude oil/g fish/day) in this study.

Relative effects of land conversion and land-use intensity on terrestrial vertebrate diversity.

Land-use has transformed ecosystems over three quarters of the terrestrial surface, with massive repercussions on biodiversity. Land-use intensity is known to contribute to the effects of land-use on biodiversity, but the magnitude of this contribution remains uncertain. Here, we use a modified countryside species-area model to compute a global account of the impending biodiversity loss caused by current land-use patterns, explicitly addressing the role of land-use intensity based on two sets of intensity indicators. We find that land-use entails the loss of ~15% of terrestrial vertebrate species from the average 5 × 5 arcmin-landscape outside remaining wilderness areas and ~14% of their average native area-of-habitat, with a risk of global extinction for 556 individual species. Given the large fraction of global land currently used under low land-use intensity, we find its contribution to biodiversity loss to be substantial (~25%). While both sets of intensity indicators yield similar global average results, we find regional differences between them and discuss data gaps. Our results support calls for improved sustainable intensification strategies and demand-side actions to reduce trade-offs between food security and biodiversity conservation.

Experimental warming differentially affects vegetative and reproductive phenology of tundra plants.

Rapid climate warming is altering Arctic and alpine tundra ecosystem structure and function, including shifts in plant phenology. While the advancement of green up and flowering are well-documented, it remains unclear whether all phenophases, particularly those later in the season, will shift in unison or respond divergently to warming. Here, we present the largest synthesis to our knowledge of experimental warming effects on tundra plant phenology from the International Tundra Experiment. We examine the effect of warming on a suite of season-wide plant phenophases. Results challenge the expectation that all phenophases will advance in unison to warming. Instead, we find that experimental warming caused: (1) larger phenological shifts in reproductive versus vegetative phenophases and (2) advanced reproductive phenophases and green up but delayed leaf senescence which translated to a lengthening of the growing season by approximately 3%. Patterns were consistent across sites, plant species and over time. The advancement of reproductive seasons and lengthening of growing seasons may have significant consequences for trophic interactions and ecosystem function across the tundra.

Soil organic carbon depletion and degradation in surface soil after long-term non-growing season warming in High Arctic Svalbard.

Arctic tundra active-layer soils are at risk of soil organic carbon (SOC) depletion and degradation upon global climate warming because they are in a stage of relatively early decomposition. Non-growing season (NGS) warming is particularly pronounced, and observed increases of CO2 emissions during experimentally warmed NGSs give concern for great SOC losses to the atmosphere. Here, we used snow fences in Arctic Spitsbergen dwarf shrub tundra to simulate 1.86 °C NGS warming for 9 consecutive years, while growing season temperatures remained unchanged. In the snow fence treatment, the 4-11 cm thick A-horizon had a 2% lower SOC concentration and a 0.48 kg C m-2 smaller pool size than the controls, indicating SOC pool depletion. The snow fence treatment's A-horizon's alkyl/O-alkyl ratio was also significantly increased, indicating an advance of SOC degradation. The underlying 5 cm of B/C-horizon did not show these effects. Our results support the hypothesis that SOC depletion and degradation are connected to the long-term transience of observed ecosystem respiration (ER) increases upon soil warming. We suggest that the bulk of warming induced ER increases may originate from surface and not deep active layer or permafrost horizons. The observed losses of SOC might be significant for the ecosystem in question, but are in magnitude comparatively small relative to anthropogenic greenhouse gas enrichment of the atmosphere. We conclude that a positive feedback of carbon losses from surface soils of Arctic dwarf shrub tundra to anthropogenic forcing will be minor, but not negligible.

Author Correction: Warming shortens flowering seasons of tundra plant communities.

In the version of this Article originally published, the following sentence was missing from the Acknowledgements: "This work was supported by the Norwegian Research Council SnoEco project, grant number 230970". This text has now been added.

Warming shortens flowering seasons of tundra plant communities.

Advancing phenology is one of the most visible effects of climate change on plant communities, and has been especially pronounced in temperature-limited tundra ecosystems. However, phenological responses have been shown to differ greatly between species, with some species shifting phenology more than others. We analysed a database of 42,689 tundra plant phenological observations to show that warmer temperatures are leading to a contraction of community-level flowering seasons in tundra ecosystems due to a greater advancement in the flowering times of late-flowering species than early-flowering species. Shorter flowering seasons with a changing climate have the potential to alter trophic interactions in tundra ecosystems. Interestingly, these findings differ from those of warmer ecosystems, where early-flowering species have been found to be more sensitive to temperature change, suggesting that community-level phenological responses to warming can vary greatly between biomes.

Daphnia magna negatively affected by chronic exposure to purified Cry-toxins.

Cry-toxin genes originating from Bacillus thuringiensis are inserted into genetically modified (GM) plants, often called Bt-plants, to provide insect resistance to pests. Significant amounts of Bt-plant residues, and thus Cry-toxins, will be shed to soil and aquatic environments. We exposed Daphnia magna to purified Cry1Ab and Cry2Aa toxins for the full life-span of the animals. We used single toxins in different doses and combinations of toxins and Roundup(®), another potential stressor on the rise in agricultural ecosystems. Animals exposed to 4.5 mg/L (ppm) of Cry1Ab, Cry2Aa and the combination of both showed markedly higher mortality, smaller body size and very low juvenile production compared to controls. Animals exposed to 0.75 mg/L also showed a tendency towards increased mortality but with increased early fecundity compared to the controls. Roundup(®) stimulated animals to strong early reproductive output at the cost of later rapid mortality. We conclude that i) purified Cry-toxins in high concentrations are toxic to D. magna, indicating alternative modes-of-action for these Cry-toxins; ii) Cry-toxins act in combination, indicating that 'stacked events' may have stronger effects on non-target organisms; iii) further studies need to be done on combinatorial effects of multiple Cry-toxins and herbicides that co-occur in the environment.

Distinct summer and winter bacterial communities in the active layer of Svalbard permafrost revealed by DNA- and RNA-based analyses.

The active layer of soil overlaying permafrost in the Arctic is subjected to dramatic annual changes in temperature and soil chemistry, which likely affect bacterial activity and community structure. We studied seasonal variations in the bacterial community of active layer soil from Svalbard (78°N) by co-extracting DNA and RNA from 12 soil cores collected monthly over a year. PCR amplicons of 16S rRNA genes (DNA) and reverse transcribed transcripts (cDNA) were quantified and sequenced to test for the effect of low winter temperature and seasonal variation in concentration of easily degradable organic matter on the bacterial communities. The copy number of 16S rRNA genes and transcripts revealed no distinct seasonal changes indicating potential bacterial activity during winter despite soil temperatures well below -10°C. Multivariate statistical analysis of the bacterial diversity data (DNA and cDNA libraries) revealed a season-based clustering of the samples, and, e.g., the relative abundance of potentially active Cyanobacteria peaked in June and Alphaproteobacteria increased over the summer and then declined from October to November. The structure of the bulk (DNA-based) community was significantly correlated with pH and dissolved organic carbon, while the potentially active (RNA-based) community structure was not significantly correlated with any of the measured soil parameters. A large fraction of the 16S rRNA transcripts was assigned to nitrogen-fixing bacteria (up to 24% in June) and phototrophic organisms (up to 48% in June) illustrating the potential importance of nitrogen fixation in otherwise nitrogen poor Arctic ecosystems and of phototrophic bacterial activity on the soil surface.

Idiosyncratic responses of high Arctic plants to changing snow regimes.

The Arctic is one of the ecosystems most affected by climate change; in particular, winter temperatures and precipitation are predicted to increase with consequent changes to snow cover depth and duration. Whether the snow-free period will be shortened or prolonged depends on the extent and temporal patterns of the temperature and precipitation rise; resulting changes will likely affect plant growth with cascading effects throughout the ecosystem. We experimentally manipulated snow regimes using snow fences and shoveling and assessed aboveground size of eight common high arctic plant species weekly throughout the summer. We demonstrated that plant growth responded to snow regime, and that air temperature sum during the snow free period was the best predictor for plant size. The majority of our studied species showed periodic growth; increases in plant size stopped after certain cumulative temperatures were obtained. Plants in early snow-free treatments without additional spring warming were smaller than controls. Response to deeper snow with later melt-out varied between species and categorizing responses by growth forms or habitat associations did not reveal generic trends. We therefore stress the importance of examining responses at the species level, since generalized predictions of aboveground growth responses to changing snow regimes cannot be made.

Snow cover and extreme winter warming events control flower abundance of some, but not all species in high arctic Svalbard.

The High Arctic winter is expected to be altered through ongoing and future climate change. Winter precipitation and snow depth are projected to increase and melt out dates change accordingly. Also, snow cover and depth will play an important role in protecting plant canopy from increasingly more frequent extreme winter warming events. Flower production of many Arctic plants is dependent on melt out timing, since season length determines resource availability for flower preformation. We erected snow fences to increase snow depth and shorten growing season, and counted flowers of six species over 5 years, during which we experienced two extreme winter warming events. Most species were resistant to snow cover increase, but two species reduced flower abundance due to shortened growing seasons. Cassiope tetragona responded strongly with fewer flowers in deep snow regimes during years without extreme events, while Stellaria crassipes responded partly. Snow pack thickness determined whether winter warming events had an effect on flower abundance of some species. Warming events clearly reduced flower abundance in shallow but not in deep snow regimes of Cassiope tetragona, but only marginally for Dryas octopetala. However, the affected species were resilient and individuals did not experience any long term effects. In the case of short or cold summers, a subset of species suffered reduced reproductive success, which may affect future plant composition through possible cascading competition effects. Extreme winter warming events were shown to expose the canopy to cold winter air. The following summer most of the overwintering flower buds could not produce flowers. Thus reproductive success is reduced if this occurs in subsequent years. We conclude that snow depth influences flower abundance by altering season length and by protecting or exposing flower buds to cold winter air, but most species studied are resistant to changes. Winter warming events, often occurring together with rain, can substantially remove snow cover and thereby expose plants to cold winter air. Depending on morphology, different parts of the plant can be directly exposed. On this picture, we see Dryas octopetala seed heads from the previous growing season protrude through the remaining ice layer after a warming event in early 2010. The rest of the plant, including meristems and flower primordia, are still somewhat protected by the ice. In the background we can see a patch of Cassiope tetragona protruding through the ice; in this case, the whole plant including flower primordia is exposed, which might be one reason why this species experienced a loss of flowers the following season. Photograph by Philipp Semenchuk.

Late snowmelt delays plant development and results in lower reproductive success in the High Arctic.

In tundra areas where the growing season is short, any delay in the start of summer may have a considerable effect on plant development, growth and reproductive success. Climate models suggest long-term changes in winter precipitation in the Arctic, which may lead to deeper snow cover and a resultant delay in date of snow melt. In this paper, we investigated the role of snow depth and melt out date on the phenological development and reproductive success of vascular plants in Adventdalen, Svalbard (78° 10'N, 16° 06'E). Effects of natural variations in snow accumulation were demonstrated using two vegetation types (snow depth: meadow 21 cm, heath 32 cm), and fences were used to experimentally increase snow depth by over 1m. Phenological delay was greatest directly after snowmelt in the earlier phenological phases, and had the largest effect on the early development of those species which normally green-up early (i.e. Dryas, Papaver, Salix, Saxifraga). Compressed growing seasons and length of the reproductive period led to a reduced reproductive success in some of the study species. There were fewer flowers, fewer plots with dispersing seeds, and lower germination rates. This can have consequences for plant establishment and community composition in the long-term.