Plant performance in a warmer world: General responses of plants from cold, northern biomes and the importance of winter and spring events

During the past three decades the Earth has warmed with a rate unprecedented during the past 1000 years. There is already ample evidence that this fast climate warming has affected a broad range of organisms, including plants. Plants from high-latitude and high-altitude sites ('cold biomes') are especially sensitive to climate warming. In this paper we (1) review the response in the phenology of plants, changes in their range and distribution, soil nutrient availability, and the effects on the structure and dynamics of plant communities for cold, northern biomes; and (2) we show, by using data from an ongoing snow and temperature manipulation experiment in northern Sweden, that also winter and spring events have a profound influence on plant performance. Both long-term phenological data sets, experimental warming studies (performed in summer or year-round), natural gradient studies and satellite images show that key phenological events are responsive to temperature increases and that recent climate warming does indeed lead to changes in plant phenology. However, data from a warming and snow manipulation study that we are conducting in northern Sweden show that plants respond differently to the various climatic scenarios that we had imposed on these species and that especially winter and spring events have a profound impact. This indicates that it is necessary to include several scenarios of both summer and winter climate change in experimental climate change studies, and that we need detailed projections of future climate at a regional scale to be able to assess their impacts on natural ecosystems. There is also ample evidence that the range shift of herbs and shrubs to more northern regions is for the vast majority of species mainly caused by changes in the climate. This is in line with the observed 'up-greening' of northern tundra sites. These rapid northern shifts in distribution of plants as a result of climate warming may have substantial consequences for the structure and dynamics of high-latitude ecosystems. An analysis of warming studies at 9 tundra sites shows that heating during at least 3 years increased net N-mineralization from 0.32 +/- 0.31 (SE) g N m(-2) yr(-1) in the controls to 0.53 +/- 0.31 (SE) g N m(2) yr(-1) in the heated plots (p < 0.05), an increase of about 70%. Thus, warming leads to higher N availability in high-latitude northern tundra sites, but the variability is substantial. Higher nutrient availability affects in turn the species composition of high-latitude sites, which has important consequences for the carbon and water balance of these systems.