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.

Rapid upwards spread of non-native plants in mountains across continents.

High-elevation ecosystems are among the few ecosystems worldwide that are not yet heavily invaded by non-native plants. This is expected to change as species expand their range limits upwards to fill their climatic niches and respond to ongoing anthropogenic disturbances. Yet, whether and how quickly these changes are happening has only been assessed in a few isolated cases. Starting in 2007, we conducted repeated surveys of non-native plant distributions along mountain roads in 11 regions from 5 continents. We show that over a 5- to 10-year period, the number of non-native species increased on average by approximately 16% per decade across regions. The direction and magnitude of upper range limit shifts depended on elevation across all regions. Supported by a null-model approach accounting for range changes expected by chance alone, we found greater than expected upward shifts at lower/mid elevations in at least seven regions. After accounting for elevation dependence, significant average upward shifts were detected in a further three regions (revealing evidence for upward shifts in 10 of 11 regions). Together, our results show that mountain environments are becoming increasingly exposed to biological invasions, emphasizing the need to monitor and prevent potential biosecurity issues emerging in high-elevation ecosystems.

Think globally, measure locally: The MIREN standardized protocol for monitoring plant species distributions along elevation gradients.

Climate change and other global change drivers threaten plant diversity in mountains worldwide. A widely documented response to such environmental modifications is for plant species to change their elevational ranges. Range shifts are often idiosyncratic and difficult to generalize, partly due to variation in sampling methods. There is thus a need for a standardized monitoring strategy that can be applied across mountain regions to assess distribution changes and community turnover of native and non-native plant species over space and time. Here, we present a conceptually intuitive and standardized protocol developed by the Mountain Invasion Research Network (MIREN) to systematically quantify global patterns of native and non-native species distributions along elevation gradients and shifts arising from interactive effects of climate change and human disturbance. Usually repeated every five years, surveys consist of 20 sample sites located at equal elevation increments along three replicate roads per sampling region. At each site, three plots extend from the side of a mountain road into surrounding natural vegetation. The protocol has been successfully used in 18 regions worldwide from 2007 to present. Analyses of one point in time already generated some salient results, and revealed region-specific elevational patterns of native plant species richness, but a globally consistent elevational decline in non-native species richness. Non-native plants were also more abundant directly adjacent to road edges, suggesting that disturbed roadsides serve as a vector for invasions into mountains. From the upcoming analyses of time series, even more exciting results can be expected, especially about range shifts. Implementing the protocol in more mountain regions globally would help to generate a more complete picture of how global change alters species distributions. This would inform conservation policy in mountain ecosystems, where some conservation policies remain poorly implemented.

How does a wetland plant respond to increasing temperature along a latitudinal gradient?

Global warming affects plant fitness through changes in functional traits and thereby ecosystem function. Wetlands are declining worldwide, and hence, ecosystem functions linked to wetlands are threatened. We use Caltha palustris "a common wetland plant" to study whether warming affects growth and reproduction differently depending on origin of source population, potentially affecting phenotypic response to local climate. We conducted a 2-year in situ temperature manipulation experiment using clone pairs of C. palustris in four regions, along a 1300-km latitudinal gradient of Sweden. Open-top chambers were used to passively increase temperature, paired with controls. Growth and reproductive traits were measured from 320 plants (four regions × five sites × two treatments × eight plants) over two consecutive seasons to assess the effect of warming over time. We found that warming increased plant height, leaf area, number of leaves, and roots. High-latitude populations responded more strongly to warming than low-latitude populations, especially by increasing leaf area. Warming increased number of flowers in general, but only in the second year, while number of fruits increased in low-latitude populations the first year. Prolonged warming leads to an increase in both number of leaves and flowers over time. While reproduction shows varying and regional responses to warming, impacts on plant growth, especially in high-latitude populations, have more profound effects. Such effects could lead to changes in plant community composition with increased abundance of fast-growing plants with larger leaves and more clones, affecting plant competition and ecological functions such as decomposition and nutrient retention. Effects of warming were highly context dependent; thus, we encourage further use of warming experiments to predict changes in growth, reproduction, and community composition across wetland types and climate gradients targeting different plant forms.

The role of arbuscular mycorrhizal fungi in nonnative plant invasion along mountain roads.

Plant associated mutualists can mediate invasion success by affecting the ecological niche of nonnative plant species. Anthropogenic disturbance is also key in facilitating invasion success through changes in biotic and abiotic conditions, but the combined effect of these two factors in natural environments is understudied. To better understand this interaction, we investigated how disturbance and its interaction with mycorrhizas could impact range dynamics of nonnative plant species in the mountains of Norway. Therefore, we studied the root colonisation and community composition of arbuscular mycorrhizal (AM) fungi in disturbed vs undisturbed plots along mountain roads. We found that roadside disturbance strongly increases fungal diversity and richness while also promoting AM fungal root colonisation in an otherwise ecto-mycorrhiza and ericoid-mycorrhiza dominated environment. Surprisingly, AM fungi associating with nonnative plant species were present across the whole elevation gradient, even above the highest elevational limit of nonnative plants, indicating that mycorrhizal fungi are not currently limiting the upward movement of nonnative plants. We conclude that roadside disturbance has a positive effect on AM fungal colonisation and richness, possibly supporting the spread of nonnative plants, but that there is no absolute limitation of belowground mutualists, even at high elevation.

Dwelling in the deep - strongly increased root growth and rooting depth enhance plant interactions with thawing permafrost soil.

Climate-warming-induced permafrost thaw exposes large amounts of carbon and nitrogen in soil at considerable depths, below the seasonally thawing active layer. The extent to which plant roots can reach and interact with these hitherto detached, deep carbon and nitrogen stores remains unknown. We aimed to quantify how permafrost thaw affects root dynamics across soil depths and plant functional types compared with above-ground abundance, and potential consequences for plant-soil interactions. A decade of experimental permafrost thaw strongly increased total root length and growth in the active layer, and deep roots invaded the newly thawed permafrost underneath. Root litter input to soil across all depths was 10 times greater with permafrost thaw. Root growth timing was unaffected by experimental permafrost thaw but peaked later in deeper soil, reflecting the seasonally receding thaw front. Deep-rooting species could sequester 15 N added at the base of the ambient active layer in October, which was after root growth had ceased. Deep soil organic matter that has long been locked up in permafrost is thus no longer detached from plant processes upon thaw. Whether via nutrient uptake, carbon storage, or rhizosphere priming, plant root interactions with thawing permafrost soils may feed back on our climate both positively and negatively.

Habitat properties and plant traits interact as drivers of non-native plant species' seed production at the local scale.

To answer the long-standing question if we can predict plant invader success based on characteristics of the environment (invasibility) or the invasive species (invasiveness), or the combination of both, there is a need for detailed observational studies in which habitat properties, non-native plant traits, and the resulting invader success are locally measured. In this study, we assess the interaction of gradients in the environmental and trait space on non-native species fitness, expressed as seed production, for a set of 10 invasive and noninvasive non-native species along a wide range of invaded sites in Flanders. In our multidimensional approach, most of the single environmental gradients (temperature, light availability, native plant species diversity, and soil fertility) and sets of non-native plant traits (plant size, photosynthesis, and foliar chemical attributes) related positively with invader seed production. Yet correlation with seed production was much stronger when several environmental gradients were assessed in interaction, and even more so when we combined plant traits and habitat properties. The latter increased explanatory power of the models on average by 25% for invasive and by 7% for noninvasive species. Additionally, we report a 70-fold higher seed production in invasive than in noninvasive species and fundamentally different correlations of seed production with plant traits and habitat properties in noninvasive versus invasive species. We conclude that locally measured traits and properties deserve much more attention than they currently get in invasion literature and thus encourage further studies combining this level of detail with the generality of a multiregion and multispecies approach across different stages of invasion.

Lags in the response of mountain plant communities to climate change.

Rapid climatic changes and increasing human influence at high elevations around the world will have profound impacts on mountain biodiversity. However, forecasts from statistical models (e.g. species distribution models) rarely consider that plant community changes could substantially lag behind climatic changes, hindering our ability to make temporally realistic projections for the coming century. Indeed, the magnitudes of lags, and the relative importance of the different factors giving rise to them, remain poorly understood. We review evidence for three types of lag: "dispersal lags" affecting plant species' spread along elevational gradients, "establishment lags" following their arrival in recipient communities, and "extinction lags" of resident species. Variation in lags is explained by variation among species in physiological and demographic responses, by effects of altered biotic interactions, and by aspects of the physical environment. Of these, altered biotic interactions could contribute substantially to establishment and extinction lags, yet impacts of biotic interactions on range dynamics are poorly understood. We develop a mechanistic community model to illustrate how species turnover in future communities might lag behind simple expectations based on species' range shifts with unlimited dispersal. The model shows a combined contribution of altered biotic interactions and dispersal lags to plant community turnover along an elevational gradient following climate warming. Our review and simulation support the view that accounting for disequilibrium range dynamics will be essential for realistic forecasts of patterns of biodiversity under climate change, with implications for the conservation of mountain species and the ecosystem functions they provide.

Both seed germination and seedling mortality increase with experimental warming and fertilization in a subarctic tundra.

Climate change is expected to force many species in arctic regions to migrate and track their climatic niche. This requires recruitment from seed, which currently shows very low rates in arctic regions, where long-lived and vegetatively reproducing plants dominate. Therefore, we pose the question whether recruitment (germination and seedling establishment) in arctic regions will significantly improve in a warmer world, and thus allow species to follow their climatic niche. We used a full factorial experiment to examine if realistic warmer temperatures (+3 °C; infrared radiation) and increased nitrogen availability (+1.4 g N m-2 year-1) affected germination, seedling survival and above- and below-ground seedling biomass in five species common in subarctic regions (Anthoxanthum odoratum, Betula nana, Pinus sylvestris, Solidago virgaurea, Vaccinium myrtillus). We found that warming increased seedling emergence in all species, but that subsequent mortality also increased, resulting in no net warming effect on seedling establishment. Warming slightly increased above-ground seedling biomass. Fertilization, on the other hand, did not influence seedling biomass, but it increased seedling establishment in B. nana while it reduced establishment in V. myrtillus. This may help B. nana dominate over V. myrtillus in warmer tundra. Surprisingly, no interactive effects between warming and fertilization were found. The lack of a general positive response of seedling establishment to warmer and more nutrient-rich conditions suggests that (sub)arctic species may experience difficulties in tracking their climatic niche. Predictions of future species distributions in arctic regions solely based on abiotic factors may therefore overestimate species' ranges due to their poor establishment. Also, the opposite response to fertilization of two key (sub)arctic dwarf shrubs, i.e. B. nana and V. myrtillus, could have important implications for the future development of arctic plant communities and argues for more research into the role of fertilization for plant establishment.

Potential contributions of root decomposition to the nitrogen cycle in arctic forest and tundra.

Plant contributions to the nitrogen (N) cycle from decomposition are likely to be altered by vegetation shifts associated with climate change. Roots account for the majority of soil organic matter input from vegetation, but little is known about differences between vegetation types in their root contributions to nutrient cycling. Here, we examine the potential contribution of fine roots to the N cycle in forest and tundra to gain insight into belowground consequences of the widely observed increase in woody vegetation that accompanies climate change in the Arctic. We combined measurements of root production from minirhizotron images with tissue analysis of roots from differing root diameter and color classes to obtain potential N input following decomposition. In addition, we tested for changes in N concentration of roots during early stages of decomposition, and investigated whether vegetation type (forest or tundra) affected changes in tissue N concentration during decomposition. For completeness, we also present respective measurements of leaves. The potential N input from roots was twofold greater in forest than in tundra, mainly due to greater root production in forest. Potential N input varied with root diameter and color, but this variation tended to be similar in forest and tundra. As for roots, the potential N input from leaves was significantly greater in forest than in tundra. Vegetation type had no effect on changes in root or leaf N concentration after 1 year of decomposition. Our results suggest that shifts in vegetation that accompany climate change in the Arctic will likely increase plant-associated potential N input both belowground and aboveground. In contrast, shifts in vegetation might not alter changes in tissue N concentration during early stages of decomposition. Overall, differences between forest and tundra in potential contribution of decomposing roots to the N cycle reinforce differences between habitats that occur for leaves.

Disturbance is the key to plant invasions in cold environments.

Until now, nonnative plant species were rarely found at high elevations and latitudes. However, partly because of climate warming, biological invasions are now on the rise in these extremely cold environments. These plant invasions make it timely to undertake a thorough experimental assessment of what has previously been holding them back. This knowledge is key to developing efficient management of the increasing risks of cold-climate invasions. Here, we integrate human interventions (i.e., disturbance, nutrient addition, and propagule input) and climatic factors (i.e., temperature) into one seed-addition experiment across two continents: the subantarctic Andes and subarctic Scandinavian mountains (Scandes), to disentangle their roles in limiting or favoring plant invasions. Disturbance was found as the main determinant of plant invader success (i.e., establishment, growth, and flowering) along the entire cold-climate gradient, explaining 40-60% of the total variance in our models, with no indication of any facilitative effect from the native vegetation. Higher nutrient levels additionally stimulated biomass production and flowering. Establishment and flowering displayed a hump-shaped response with increasing elevation, suggesting that competition is the main limit on invader success at low elevations, as opposed to low-growing-season temperatures at high elevations. Our experiment showed, however, that nonnative plants can establish, grow, and flower well above their current elevational limits in high-latitude mountains. We thus argue that cold-climate ecosystems are likely to see rapid increases in plant invasions in the near future as a result of a synergistic interaction between increasing human-mediated disturbances and climate warming.

The hidden season: growing season is 50% longer below than above ground along an arctic elevation gradient.

There is compelling evidence from experiments and observations that climate warming prolongs the growing season in arctic regions. Until now, the start, peak, and end of the growing season, which are used to model influences of vegetation on biogeochemical cycles, were commonly quantified using above-ground phenological data. Yet, over 80% of the plant biomass in arctic regions can be below ground, and the timing of root growth affects biogeochemical processes by influencing plant water and nutrient uptake, soil carbon input and microbial activity. We measured timing of above- and below-ground production in three plant communities along an arctic elevation gradient over two growing seasons. Below-ground production peaked later in the season and was more temporally uniform than above-ground production. Most importantly, the growing season continued c. 50% longer below than above ground. Our results strongly suggest that traditional above-ground estimates of phenology in arctic regions, including remotely sensed information, are not as complete a representation of whole-plant production intensity or duration, as studies that include root phenology. We therefore argue for explicit consideration of root phenology in studies of carbon and nutrient cycling, in terrestrial biosphere models, and scenarios of how arctic ecosystems will respond to climate warming.

Trade-off between competition and facilitation defines gap colonization in mountains.

Recent experimental observations show that gap colonization in small-stature (e.g. grassland and dwarf shrubs) vegetation strongly depends on the abiotic conditions within them. At the same time, within-gap variation in biotic interactions such as competition and facilitation, caused by distance to the gap edge, would affect colonizer performance, but a theoretical framework to explore such patterns is missing. Here, we model how competition, facilitation and environmental conditions together determine the small-scale patterns of gap colonization along a cold gradient in mountains, by simulating colonizer survival in gaps of various sizes. Our model adds another dimension to the known effects of biotic interactions along a stress gradient by focussing on the trade-off between competition and facilitation in the within-gap environment. We show that this trade-off defines a peak in colonizer survival at a specific distance from the gap edge, which progressively shifts closer to the edge as the environment gets colder, ultimately leaving a large fraction of gaps unsuitable for colonization in facilitation-dominated systems. This is reinforced when vegetation size and temperature amelioration are manipulated simultaneously with temperature in order to simulate an elevational gradient more realistically. Interestingly, all other conditions being equal, the magnitude of the realized survival peak was always lower in large than in small gaps, making large gaps harder to colonize. The model is relevant to predict effects of non-native plant invasions and climate warming on colonization processes in mountains.

Alien roadside species more easily invade alpine than lowland plant communities in a subarctic mountain ecosystem.

Effects of roads on plant communities are not well known in cold-climate mountain ecosystems, where road building and development are expected to increase in future decades. Knowledge of the sensitivity of mountain plant communities to disturbance by roads is however important for future conservation purposes. We investigate the effects of roads on species richness and composition, including the plant strategies that are most affected, along three elevational gradients in a subarctic mountain ecosystem. We also examine whether mountain roads promote the introduction and invasion of alien plant species from the lowlands to the alpine zone. Observations of plant community composition were made together with abiotic, biotic and anthropogenic factors in 60 T-shaped transects. Alpine plant communities reacted differently to road disturbances than their lowland counterparts. On high elevations, the roadside species composition was more similar to that of the local natural communities. Less competitive and ruderal species were present at high compared with lower elevation roadsides. While the effects of roads thus seem to be mitigated in the alpine environment for plant species in general, mountain plant communities are more invasible than lowland communities. More precisely, relatively more alien species present in the roadside were found to invade into the surrounding natural community at high compared to low elevations. We conclude that effects of roads and introduction of alien species in lowlands cannot simply be extrapolated to the alpine and subarctic environment.

Local temperatures inferred from plant communities suggest strong spatial buffering of climate warming across Northern Europe.

Recent studies from mountainous areas of small spatial extent (<2500 km(2) ) suggest that fine-grained thermal variability over tens or hundreds of metres exceeds much of the climate warming expected for the coming decades. Such variability in temperature provides buffering to mitigate climate-change impacts. Is this local spatial buffering restricted to topographically complex terrains? To answer this, we here study fine-grained thermal variability across a 2500-km wide latitudinal gradient in Northern Europe encompassing a large array of topographic complexities. We first combined plant community data, Ellenberg temperature indicator values, locally measured temperatures (LmT) and globally interpolated temperatures (GiT) in a modelling framework to infer biologically relevant temperature conditions from plant assemblages within <1000-m(2) units (community-inferred temperatures: CiT). We then assessed: (1) CiT range (thermal variability) within 1-km(2) units; (2) the relationship between CiT range and topographically and geographically derived predictors at 1-km resolution; and (3) whether spatial turnover in CiT is greater than spatial turnover in GiT within 100-km(2) units. Ellenberg temperature indicator values in combination with plant assemblages explained 46-72% of variation in LmT and 92-96% of variation in GiT during the growing season (June, July, August). Growing-season CiT range within 1-km(2) units peaked at 60-65°N and increased with terrain roughness, averaging 1.97 °C (SD = 0.84 °C) and 2.68 °C (SD = 1.26 °C) within the flattest and roughest units respectively. Complex interactions between topography-related variables and latitude explained 35% of variation in growing-season CiT range when accounting for sampling effort and residual spatial autocorrelation. Spatial turnover in growing-season CiT within 100-km(2) units was, on average, 1.8 times greater (0.32 °C km(-1) ) than spatial turnover in growing-season GiT (0.18 °C km(-1) ). We conclude that thermal variability within 1-km(2) units strongly increases local spatial buffering of future climate warming across Northern Europe, even in the flattest terrains.

Plant community type and small-scale disturbances, but not altitude, influence the invasibility in subarctic ecosystems.

Little of our knowledge about invasibility comes from arctic and alpine ecosystems, despite increasing plant migration and invasion in those regions. Here, we examine how community type, altitude, and small-scale disturbances affect invasibility in a subarctic ecosystem. Over a period of 4 yr, we studied seedling emergence and establishment in 17 species sown in gaps or undisturbed vegetation in four subarctic community types (Salix scrub, meadow, rich heath, poor heath) along an elevation gradient. Invasibility was lowest in rich heath and highest in Salix scrub. Small disturbances significantly increased the invasibility in most communities, thereby showing the importance of biotic resistance to invasion in subarctic regions. Unexpectedly, invasibility did not decrease with increasing elevation, and it was also not related to summer temperature. Our data suggest that biotic resistance might be more important than abiotic stress for invasibility in subarctic tundra and that low temperatures do not necessarily limit seedling establishment at high altitudes. High elevations are therefore potentially more vulnerable to invasion than was originally thought. Changes in community composition as a result of species migration or invasion are most likely to occur in Salix scrub and meadow, whereas Empetrum-dominated rich heath will largely remain unchanged.

Environmental factors influencing soil testate amoebae in herbaceous and shrubby vegetation along an altitudinal gradient in subarctic tundra (Abisko, Sweden).

Shifts in community composition of soil protozoa in response to climate change may substantially influence microbial activity and thereby decomposition processes. However, effects of climate and vegetation on soil protozoa remain poorly understood. We studied the distribution of soil testate amoebae in herbaceous and shrubby vegetation along an altitudinal gradient (from below the treeline at 500 m to the mid-alpine region at 900 m a.s.l.) in subarctic tundra. To explain patterns in abundance, species diversity and assemblage composition of testate amoebae, a data set of microclimate and soil chemical characteristics was collected. Both elevation and vegetation influenced the assemblage composition of testate amoebae. The variation was regulated by interactive effects of summer soil moisture, winter soil temperature, soil pH and nitrate ion concentrations. Besides, soil moisture regulated non-linear patterns in species richness across the gradient. This is the first study showing the effects of winter soil temperatures on species composition of soil protozoa. The effects could be explained by specific adaptations of testate amoebae such as frost-resistant cysts allowing them to survive low winter temperatures. We conclude that the microclimate and soil chemical characteristics are the main drivers of changes in protozoan assemblage composition in response to elevation and vegetation.

Predicted changes in vegetation structure affect the susceptibility to invasion of bryophyte-dominated subarctic heath.

BACKGROUND AND AIMS: A meta-analysis of global change experiments in arctic tundra sites suggests that plant productivity and the cover of shrubs, grasses and dead plant material (i.e. litter) will increase and the cover of bryophytes will decrease in response to higher air temperatures. However, little is known about which effects these changes in vegetation structure will have on seedling recruitment of species and invasibility of arctic ecosystems. METHODS: A field experiment was done in a bryophyte-dominated, species-rich subarctic heath by manipulating the cover of bryophytes and litter in a factorial design. Three phases of seedling recruitment (seedling emergence, summer seedling survival, first-year recruitment) of the grass Anthoxanthum alpinum and the shrub Betula nana were analysed after they were sown into the experimental plots. KEY RESULTS: Bryophyte and litter removal significantly increased seedling emergence of both species but the effects of manipulations of vegetation structure varied strongly for the later phases of recruitment. Summer survival and first-year recruitment were significantly higher in Anthoxanthum. Although bryophyte removal generally increased summer survival and recruitment, seedlings of Betula showed high mortality in early August on plots where bryophytes had been removed. CONCLUSIONS: Large species-specific variation and significant effects of experimental manipulations on seedling recruitment suggest that changes in vegetation structure as a consequence of global warming will affect the abundance of grasses and shrubs, the species composition and the susceptibility to invasion of subarctic heath vegetation.

How do bryophytes govern generative recruitment of vascular plants?

Interactions between vascular plants and bryophytes determine plant community composition in many ecosystems. Yet, little is known about the importance of interspecific differences between bryophytes with respect to their effects on vascular plants. We compared the extent to which species-specific bryophyte effects on vascular plant generative recruitment depend on the following underlying mechanisms: allelopathy, mechanical obstruction, soil moisture and temperature control. We sowed 10 vascular plant species into monospecific mats of six chemically and structurally diverse bryophytes, and examined 1-yr seedling recruitment. Allelopathic effects were also assessed in a laboratory phyto-assay. Although all bryophytes suppressed vascular plant regeneration, there were significant differences between the bryophyte species. The lack of interactions indicated the absence of species-specific adaptations of vascular plants for recruitment in bryophyte mats. Differences between bryophyte species were best explained by alterations in temperature regime under bryophyte mats, mostly by reduced temperature amplitudes during germination. The temperature regime under bryophyte mats was well predicted by species-specific bryophyte cushion thickness. The fitness of established seedlings was not affected by the presence of bryophytes. Our results suggest that climatically or anthropogenically driven changes in the species' composition of bryophyte communities have knock-on effects on vascular plant populations via generative reproduction.

Effects of a warmer climate on seed germination in the subarctic.

BACKGROUND AND AIMS: In a future warmer subarctic climate, the soil temperatures experienced by dispersed seeds are likely to increase during summer but may decrease during winter due to expected changes in snow depth, duration and quality. Because little is known about the dormancy-breaking and germination requirements of subarctic species, how warming may influence the timing and level of germination in these species was examined. METHODS: Under controlled conditions, how colder winter and warmer summer soil temperatures influenced germination was tested in 23 subarctic species. The cold stratification and warm incubation temperatures were derived from real soil temperature measurements in subarctic tundra and the temperatures were gradually changed over time to simulate different months of the year. KEY RESULTS: Moderate summer warming (+2.5 degrees C) substantially accelerated germination in all but four species but did not affect germination percentages. Optimum germination temperatures (20/10 degrees C) further decreased germination time and increased germination percentages in three species. Colder winter soil temperatures delayed the germination in ten species and decreased the germination percentage in four species, whereas the opposite was found in Silene acaulis. In most species, the combined effect of a reduced snow cover and summer warming resulted in earlier germination and thus a longer first growing season, which improves the chance of seedling survival. In particular the recruitment of (dwarf) shrubs (Vaccinium myrtillus, V. vitis-idaea, Betula nana), trees (Alnus incana, Betula pubescens) and grasses (Calamagrostis lapponica, C. purpurea) is likely to benefit from a warmer subarctic climate. CONCLUSIONS: Seedling establishment is expected to improve in a future warmer subarctic climate, mainly by considerably earlier germination. The magnitudes of the responses are species-specific, which should be taken into account when modelling population growth and migration of subarctic species.