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About 10% of the anthropogenic CO2 emissions have been absorbed by northern terrestrial ecosystems during the past decades. It has been hypothesized that part of this increasing carbon (C) sink is caused by the alleviation of nitrogen (N) limitation by increasing anthropogenic N inputs. However, little is known about this N-dependent C sink. Here, we studied the effect of chronic seabird-derived N inputs (47–67 kg N ha−1 year−1) on the net soil organic C (SOC) storage rate of unmanaged Icelandic grasslands on the volcanic Vestmannaeyjar archipelago by using a stock change approach in combination with soil dating. We studied both early developmental (young) soils that had been receiving increased N inputs over a decadal timescale since an eruption in 1963, and well-developed soils, that had been receiving N inputs over a millennial timescale. For the latter, however, the effects on both decadal (topsoil; 40 years) and millennial (total soil profile; 1600 years) SOC storage could be studied, as the age of topsoil and the total soil profile could be determined from volcanic ash layers deposited in 1973 and 395 AD. We found that enhanced N availability—either from accumulation over time, or seabird derived—increased the net SOC storage rate. Under low N inputs, early developmental soils were weak decadal C sinks (0.018 ton SOC ha−1 year−1), but this increased quickly under ca. 30 years of elevated N inputs to 0.29 ton SOC ha−1 year−1, thereby equalling the decadal SOC storage rate of the unfertilized well-developed soils. Furthermore, for the well-developed soils, chronically elevated N inputs not only stimulated the decadal SOC storage rate in the topsoil, but also the total millennial SOC storage was consistently higher. Hence, our study suggests that Icelandic grasslands, if not disturbed, can remain C sinks for many centuries under current climatic conditions and that chronically elevated N inputs can induce a permanent strengthening of this sink. more...

Research in warm-climate biomes has shown that invasion by symbiotic dinitrogen (N2)-fixing plants can transform ecosystems in ways analogous to the transformations observed as a consequence of anthropogenic, atmospheric nitrogen (N) deposition: declines in biodiversity, soil acidification, and alterations to carbon and nutrient cycling, including increased N losses through nitrate leaching and emissions of the powerful greenhouse gas nitrous oxide (N2O). Here, we used literature review and case study approaches to assess the evidence for similar transformations in cold-climate ecosystems of the boreal, subarctic and upper montane-temperate life zones. Our assessment focuses on the plant genera Lupinus and Alnus, which have become invasive largely as a consequence of deliberate introductions and/or reduced land management. These cold biomes are commonly located in remote areas with low anthropogenic N inputs, and the environmental impacts of N2-fixer invasion appear to be as severe as those from anthropogenic N deposition in highly N polluted areas. Hence, inputs of N from N2 fixation can affect ecosystems as dramatically or even more strongly than N inputs from atmospheric deposition, and biomes in cold climates represent no exception with regard to the risk of being invaded by N2-fixing species. In particular, the cold biomes studied here show both a strong potential to be transformed by N2-fixing plants and a rapid subsequent saturation in the ecosystem's capacity to retain N. Therefore, analogous to increases in N deposition, N2-fixing plant invasions must be deemed significant threats to biodiversity and to environmental quality. more...

Temperature is increasing at unprecedented rates across most of the tundra biome. Remote-sensing data indicate that contemporary climate warming has already resulted in increased productivity over much of the Arctic, but plot-based evidence for vegetation transformation is not widespread. We analysed change in tundra vegetation surveyed between 1980 and 2010 in 158 plant communities spread across 46 locations. We found biome-wide trends of increased height of the plant canopy and maximum observed plant height for most vascular growth forms; increased abundance of litter; increased abundance of evergreen, low-growing and tall shrubs; and decreased abundance of bare ground. Intersite comparisons indicated an association between the degree of summer warming and change in vascular plant abundance, with shrubs, forbs and rushes increasing with warming. However, the association was dependent on the climate zone, the moisture regime and the presence of permafrost. Our data provide plot-scale evidence linking changes in vascular plant abundance to local summer warming in widely dispersed tundra locations across the globe. © 2012 Macmillan Publishers Limited. All rights reserved. more...

Facing an increased threat of rapid climate change in cold-climate regions, it is important to understand the sensitivity of plant communities both in terms of degree and direction of community change. We studied responses to 3–5 years of moderate experimental warming by open-top chambers in two widespread but contrasting tundra communities in Iceland. In a species-poor and nutrient-deficient moss heath, dominated by Racomitrium lanuginosum, mean daily air temperatures at surface were 1–2°C higher in the warmed plots than the controls whereas soil temperatures tended to be lower in the warmed plots throughout the season. In a species-rich dwarf shrub heath on relatively rich soils at a cooler site, dominated by Betula nana and R. lanuginosum, temperature changes were in the same direction although more moderate. In the moss heath, there were no detectable community changes while significant changes were detected in the dwarf shrub heath: the abundance of deciduous and evergreen dwarf shrubs significantly increased (>50%), bryophytes decreased (18%) and canopy height increased (100%). Contrary to some other studies of tundra communities, we detected no changes in species richness or other diversity measures in either community and the abundance of lichens did not change. It is concluded that the sensitivity of Icelandic tundra communities to climate warming varies greatly depending on initial conditions in terms of species diversity, dominant species, soil and climatic conditions as well as land-use history. more...

Plant colonization and succession on the volcanic island of Surtsey, formed in 1963, have been closely followed. In 2013, a total of 69 vascular plant species had been discovered on the island; of these, 59 were present and 39 had established viable populations. Surtsey had more than twice the species of any of the comparable neighbouring islands, and all of their common species had established on Surtsey. The first colonizers were dispersed by sea, but, after 1985, bird dispersal became the principal pathway with the formation of a seagull colony on the island and consequent site amelioration. This allowed wind-dispersed species to establish after 1990. Since 2007, there has been a net loss of species on the island. A study of plant succession, soil formation and invertebrate communities in permanent plots on Surtsey and on two older neighbouring islands (plants and soil) has revealed that seabirds, through their transfer of nutrients from sea to land, are major drivers of development of these ecosystems. In the area impacted by seagulls, dense grassland swards have developed and plant cover, species richness, diversity, plant biomass and soil carbon become significantly higher than in low-impact areas, which remained relatively barren. A similar difference was found for the invertebrate fauna. After 2000, the vegetation of the oldest part of the seagull colony became increasingly dominated by long-lived, rhizomatous grasses (Festuca, Poa, Leymus) with a decline in species richness and diversity. Old grasslands of the neighbouring islands Elliđaey (puffin colony, high nutrient input) and Heimaey (no seabirds, low nutrient input) contrasted sharply. The puffin grassland of Elliđaey was very dense and species-poor. It was dominated by Festuca and Poa, and very similar to the seagull grassland developing on Surtsey. The Heimaey grassland was significantly higher in species richness and diversity, and had a more even cover of dominants (Festuca/Agrostis/Ranunculus). We forecast that, with continued erosion of Surtsey, loss of habitats and increasing impact from seabirds a lush, species-poor grassland will develop and persist, as on the old neighbouring islands. more...

What happens during primary succession after the first colonizers have occupied a pristine surface largely depends on how they ameliorate living conditions for other species. For vascular plants the onset of soil development and associated increase in nutrient (mainly nitrogen; N) and water availability is especially important. Here, we report the relationship between N accumulation and biomass and ecosystem carbon (C) stocks in a 50-year-old volcanic island, Surtsey, Iceland, where N stocks are still exceptionally low. However, a 28-year-old seagull colony on the island provided nutrient-enriched areas, which enabled us to assess the relationship between N stock and biomass and ecosystem C stocks across a much larger range in N stock. Further, we compared areas on shallow and deep tephra sands as we expected that deep-rooted systems would be more efficient in retaining N. The sparsely vegetated area outside the colony had accumulated 0.7 kg N ha−1 yr−1, which was ca. 50–60% of the estimated N input rate from wet deposition. This approximates values for systems under low N input and bare dune habitats. The seagulls have added, on average, 47 kg N ha−1 yr−1, which induced a shift from belowground to aboveground in ecosystem N and C stocks and doubled the ecosystem N-use efficiency, determined as the ratio of biomass and C storage per unit N input. Soil depth did not significantly affect total N stocks, which suggests a high N retention potential. Both total ecosystem biomass and C stocks were strongly correlated with N stock inside the colony, which indicated the important role of N during the first steps of primary succession. Inside the colony, the ecosystem biomass C stocks (17–27 ton C ha−1) had reached normal values for grasslands, while the soil organic carbon (SOC) stocks (4–10 ton C ha−1 were only a fraction of normal grassland values. Thus, it will take a long time until the SOC stock reaches equilibrium with the current primary production, during which conditions for new colonists may change. more...

Recent observations of changes in some tundra ecosystems appear to be responses to a warming climate. Several experimental studies have shown that tundra plants and ecosystems can respond strongly to environmental change, including warming; however, most studies were limited to a single location and were of short duration and based on a variety of experimental designs. In addition, comparisons among studies are difficult because a variety of techniques have been used to achieve experimental warming and different measurements have been used to assess responses. We used metaanalysis on plant community measurements from standardized warming experiments at 11 locations across the tundra biome involved in the International Tundra Experiment. The passive warming treatment increased plant-level air temperature by 1-3°C, which is in the range of predicted and observed warming for tundra regions. Responses were rapid and detected in whole plant communities after only two growing seasons. Overall, warming increased height and cover of deciduous shrubs and graminoids, decreased cover of mosses and lichens, and decreased species diversity and evenness. These results predict that warming will cause a decline in biodiversity across a wide variety of tundra, at least in the short term. They also provide rigorous experimental evidence that recently observed increases in shrub cover in many tundra regions are in response to climate warming. These changes have important implications for processes and interactions within tundra ecosystems and between tundra and the atmosphere. more...

Changes in understory biomass, forest floor carbon (C) stock and vegetation composition were studied in six age-classes of Siberian larch (Larix sibirica) and two age-classes of native birch (Betula pubescens) in Iceland. The ground vegetation was less in the larch during the thicket stage and in the old-growth birch compared to a treeless pasture. Understory biomass was strongly related to canopy gap fraction across forest stands (P, Biomasse et composition de la végétation de sous-bois, et stock de carbone du sol dans une chronoséquence de mélèze de Sibérie et de bouleau pubescent en Islande. Nous avons étudié en Islande les modifications de biomasse, de composition floristique, et de stock de carbone en sous bois dans des peuplements de mélèze de Sibérie correspondant à 6 classes d'âge, et dans des peuplements de bouleau pubescent correspondant à deux classes d'âge. En comparaison avec une pâture non boisée, la végétation au sol était moins développée dans le peuplement de mélèze au stade gaulis et dans celui de bouleau mature. La biomasse au sol était fortement corrélée à la fraction de trouées dans l'ensemble des peuplements (p more...