The relative importance of global versus local environmental factors for growth and thus carbon uptake of the bryophyte genusSphagnum-the main peat-former and ecosystem engineer in northern peatlands-remains unclear. We measured length growth and net primary production (NPP) of two abundantSphagnumspecies across 99 Holarctic peatlands. We tested the importance of previously proposed abiotic and biotic drivers for peatland carbon uptake (climate, N deposition, water table depth and vascular plant cover) on these two responses. Employing structural equation models (SEMs), we explored both indirect and direct effects of drivers onSphagnumgrowth. Variation in growth was large, but similar within and between peatlands. Length growth showed a stronger response to predictors than NPP. Moreover, the smaller and denserSphagnum fuscumgrowing on hummocks had weaker responses to climatic variation than the larger and looserSphagnum magellanicumgrowing in the wetter conditions. Growth decreased with increasing vascular plant cover within a site. Between sites, precipitation and temperature increased growth forS. magellanicum. The SEMs indicate that indirect effects are important. For example, vascular plant cover increased with a deeper water table, increased nitrogen deposition, precipitation and temperature. These factors also influencedSphagnumgrowth indirectly by affecting moss shoot density. Synthesis. Our results imply that in a warmer climate,S. magellanicumwill increase length growth as long as precipitation is not reduced, whileS. fuscumis more resistant to decreased precipitation, but also less able to take advantage of increased precipitation and temperature. Such species-specific sensitivity to climate may affect competitive outcomes in a changing environment, and potentially the future carbon sink function of peatlands.

Altitudinal gradients offer attractive opportunities for studies of population differentiation in response to environmental heterogeneity. In this thesis, I examined population differentiation along altitudinal gradients by combining common-garden experiments with field studies and experiments in alpine, subalpine and boreal populations of the perennial herb Solidago virgaurea. More specifically, I determined whether leaf physiology in terms of nitrogen concentration and resorption, flowering phenology, flower production and reproductive effort vary along altitudinal gradients.

Nitrogen concentration in green leaves were higher in alpine than in subalpine and boreal populations. These differences persisted when plants were grown from seeds in a common-garden experiment at two sites, suggesting that the differences have a genetic component. There was mixed support for a trade-off between maximized carbon gain through the maintenance of high nitrogen concentration, and minimized nitrogen loss through high resorption.

In their natural habitats alpine populations began flowering later than subalpine populations, but this difference was reversed when plants were grown in a common environment. This suggests that genetic differences among populations counteract environmental effects and reduce phenotypic variation in flowering time among populations. Flowering time thus shows countergradient genetic variation in S. virgaurea.

In a common-garden experiment, boreal populations produced more flowers and had a higher reproductive effort than subalpine and alpine populations indicating habitat-specific genetic differences in reproductive allocation. In a field study, which included three populations, seed set was close to zero in the alpine population, intermediate in the subalpine population, and high in the boreal population. Experimental flower removal showed that seed production was associated with a considerable cost in terms of reduced flowering propensity the following year, but did not support the hypothesis that a large floral display is important for pollination success.

Questions: Roots represent a considerable proportion of biomass, primary production and litter input in arctic tundra, and plant allocation of biomass to above- or below-ground tissue in response to climate change is a key factor in the future C balance of these ecosystems. According to optimality theory plants allocate C to the above- or below-ground structure that captures the most limiting resource. We used an elevational gradient to test this theory and as a space-for-time substitution to inform on tundra carbon allocation patterns under a shifting climate, by exploring if increasing elevation was positively related to the root:shoot ratio, as well as a larger plant allocation to adsorptive over storage roots.

Location: Arctic tundra heath dominated by Empetrum hermaphroditum close to Abisko, Sweden.

Methods: We measured root:shoot and fine:coarse root ratios of the plant communities along an elevational gradient by sampling above- and below-ground biomass, further separating root biomass into fine (<1 mm) and coarse roots.

Results: Plant biomass was higher at the lower elevations, but the root:shoot ratio did not vary with elevation. Resource allocation to fine relative to coarse roots increased with elevation, resulting in a fine:coarse root ratio that more than doubled with increasing elevation.

Conclusions: Contrary to previous works, the root:shoot ratio along this elevational gradient remained stable. However, communities along our study system were dominated by the same species at each elevation, which suggests that when changes in the root:shoot ratio occur with elevation these changes may be driven by differences in allocation patterns among species and thus turnover in plant community structure. Our results further reveal that the allocation of biomass to fine relative to coarse roots can differ between locations along an elevational gradient, even when overall above- vs below-ground biomass allocation does not. Given the functionally different roles of fine vs coarse roots this could have large implications for below-ground C cycling. Our results highlight the importance of direct effects vs indirect effects (such as changes in plant community composition and nutrient availability) of climate change for future C allocation above and below ground.

Plant chemical and structural defence compounds are well known to impact upon herbivory of fresh leaves and influence decomposition rates after leaf senescence. A number of theories predict that alleviating nutrient limitation and reducing other environmental stressors will result in decreased production of plant chemical defences. In this study, we measured plant defence properties [total polyphenols (TP), condensed tannins (CT) and lignin concentrations, and protein complexation capacity (PCC)] in both fresh and senesced plant leaves in a fully factorial N and P fertilization experiment set-up at each of three elevations along an elevational gradient in Swedish subarctic tundra heath vegetation. Further, we performed a decomposition of variance analysis on community-weighted averages (CWAs) of plant defence properties to determine the relative contributions of interspecific and intraspecific variation to the total variation observed in response to elevation and nutrient addition. We hypothesized that N fertilization would reduce plant defence properties and that this reduction would be greater at higher elevations, while the effects of P fertilization would have no effect at any elevation. At the community level, N addition reduced CT and PCC in both fresh and senesced leaves and TP in senesced leaves, while P addition had few effects, broadly in line with our hypothesis. The effects of N addition frequently varied with elevation, but in contrast to our hypothesis, the said effects were strongest at the lowest elevations. The effects of N addition and the interactive effect of N with elevation were primarily driven by intraspecific, rather than interspecific, variation. Our findings suggest that as temperatures warm and N availability increases due to global climate change, secondary metabolites in subarctic heath vegetation will decline particularly within species. Our results highlight the need to consider the effects of both nutrient availability and temperature, and their interaction, in driving subarctic plant defence.

Introduction. Within Racomitrium Sect. Racomitrium, several species are distinguished in the Southern Hemisphere, but only the variable R. lanuginosum (Hedw.) Brid. exists in the north. Plants deviating from the common R. lanuginosum phenotype in leaf orientation and in almost entire or weakly dentate hair-point margins suggest that more than one species may exist in this region.

Methods. Sixty-five specimens of R. lanuginosum, including eleven deviating ones (four growing intermixed with ‘normal’ plants), and two of each of the Southern Hemisphere R. geronticum Müll.Hal. and R. pruinosum (Wilson) Müll.Hal. were studied, based on the nuclear ITS and the plastid rpl16 and trnG. Relationships among specimens were explored with NeighborNet split networks, maximum parsimony, and Jacknife analyses.

Key results. The molecular analyses revealed one grade and four distinct lineages, two of which correspond with the Southern Hemisphere species. Two of the three R. lanuginosum entities are widespread in Scandinavia, whereas the third occurs only in the mountains. Morphologically deviating plants do not form their own lineage but appeared in all three R. lanuginosum entities.

Conclusions. The three entities within R. lanuginosum do not form a monophyletic group, good support exists for their recognition, and they are molecularly as distinct as the two morphologically recognisable Southern Hemisphere species. They should therefore be recognised as cryptic species. The morphologically deviating plants represent genotypic differentiation, which was further confirmed by molecular evidence in three out of four occurrences where they grew intermixed with ‘normal’ plants. Since deviating plants occur in all three cryptic species they should not be taxonomically recognised.

Question

Dark diversity refers to the set of species that are not observed in an area but could potentially occur based on suitable local environmental conditions. In this paper, we applied both niche-based and co-occurrence-based methods to estimate the dark diversity of vascular plant species in the subarctic mountains. We then aimed to unravel the drivers explaining (a) why some locations were missing relatively more suitable species than others, and (b) why certain plant species were more often absent from suitable locations than others.

Location

The Scandinavian mountains around Abisko, northern Sweden.

Methods

We calculated the dark diversity in 107 plots spread out across four mountain trails using four different methods: two co-occurrence-based (Beals? index and the hypergeometric method) and two niche-based (the climatic niche model and climatic niche model followed by species-specific threshold). We then applied multiple Generalized Linear Mixed-Effects Models and General Linear Models to determine which habitat characteristics and species traits contributed the most to dark diversity.

Results

The study showed a notable divergence in the predicted drivers of dark diversity depending on the method used. Nevertheless, we can conclude that plot-level dark diversity was generally 17% higher in areas at low elevations and 31% higher in areas with a low species richness.

Conclusion

Our findings call for caution when interpreting statistical findings of dark-diversity estimates. Even so, all analyses point toward an important role for natural processes such as competitive dominance as the main driver of the spatial patterns found in dark diversity in the northern Scandes.

Data i denna studie är baserad på en tidigare studie som undersökt om snödjup påverkar medelvärdet för tillväxt och reproduktion hos Empetrum hermaphroditum över en latitud- och klimatgradient (Bienau et al. 2014). Jag testade effekterna för region och snödjup på intraspecifik variation av tillväxtrelaterade variabler istället, för att klargöra om arten har möjlighet att klara av förändringar i snödjup och ökad växtsäsong i framtiden. Tidigare forskning menar att inomartsvariation beror på resurser i habitatet och väcker frågeställningen om det är större variation hos Empetrum i gynnsamma habitat som björkskog och habitat med tjockt snötäcke än i ogynnsamma habitat som de med tunt snötäcke. Vid analys av tidigare samlad data fanns signifikanta skillnader i variation mellan habitat i några morfologiska karaktärer. Däremot är det inte alla dessa karaktärer som följer det förväntade mönstret om att de gynnsamma habitaten skulle ha större variation. Sammantaget visade analysen signifikant skillnad rörande huvudskotten och sidoskottens längd, levande blad på huvudskotten och torrvikten på stammen. Dessa resultat medför att hypotesen stämmer för några tillväxtrelaterade variabler. 

Effects of simulated environmental change on bryophyte and lichen species richness and diversity in alpine tundra were investigated in a 5-year experiment at Latnjajaure, northern Sweden. The experiment had a factorial design including fertilisation and temperature enhancement in one meadow and one heath plant community. Responses in species richness, biodiversity, and species composition of bryophytes and lichens to experimental treatments were compared to the observed variation in six naturally occurring plant communities. The combination of fertilisation and enhanced temperature resulted in a species impoverishment, for bryophytes in the bryophyte-dominated community, and for lichens in the lichen-dominated communities, but the species composition stayed within the observed natural variation. During the course of the study, no species new to the investigated mid-alpine landscape were recorded, but that scenario is realistic within a decade when comparing with the processes seen in vascular plants.

In arctic ecosystems, climate change has increased plant productivity. As arctic carbon (C) stocks predominantly are located belowground, the effects of greater plant productivity on soil C storage will significantly determine the net sink/source potential of these ecosystems, but vegetation controls on soil CO2 efflux remain poorly resolved.

In order to identify the role of canopy‐forming species in belowground C dynamics, we conducted a girdling experiment with plots distributed across 1 km2 of treeline birch (Betula pubescens ) forest and willow (Salix lapponum ) patches in northern Sweden and quantified the contribution of canopy vegetation to soil CO2 fluxes and belowground productivity.

Girdling birches reduced total soil CO2 efflux in the peak growing season by 53%, which is double the expected amount, given that trees contribute only half of the total leaf area in the forest. Root and mycorrhizal mycelial production also decreased substantially. At peak season, willow shrubs contributed 38% to soil CO2 efflux in their patches.

Our findings indicate that C, recently fixed by trees and tall shrubs, makes a substantial contribution to soil respiration. It is critically important that these processes are taken into consideration in the context of a greening arctic because productivity and ecosystem C sequestration are not synonymous.

Alpine plants are assumed to be in particular danger as the climate changes rapidly worldwide. Specialist alpine species in Norrbotten County, northern Sweden have been surveyed over the last 20 years, providing insight to population dynamics and how the plants might respond to the changing climate. The main current threat to the species is habitat destruction as the climate changes. Variation in the number of plants among populations and years, and correlations with environmental variables were examined. Some species had increased while others had decreased over the years. No uniform relationship for all species and populations were discovered, but some of the species exhibited relationships between population size changes and temperature and precipitation. However, if the future climate in Norrbotten County changes according to the predictions, the habitats of the specialist alpine plants may be severely altered, leaving the species with no alternative places to establish and grow.

Bryophytes are abundant in tundra ecosystems, where they affect carbon and nitrogen cycling through primary production and associations with N2-fixing bacteria. Bryophyte responses to climate warming are inconclusive, likely because species-specific responses are poorly understood. Here we investigated how warming affects the growth and nitrogenase activity of 10 tundra bryophyte species in two tundra landscapes. Collected bryophyte samples were grown in temperature-controlled growth chambers for 12 weeks at five temperatures from 3 to 18 °C. We measured growth, N concentration, δ15N, and δ13C after 3 months and nitrogenase activity after 5 and 12 weeks. Bryophyte growth and associated nitrogenase activity generally increased with temperature, but species differed in their optima. Bryophyte N concentration and δ15N indicated that, for some species, increased N2-fixation could not compensate for growth-induced N limitation. High landscape coverage and large positive warming effects on feather mosses and Sphagnum species highlight their competitive advantages, confirm earlier field observations, and contribute to the mechanistic understanding of differential bryophyte growth in response to warming. We suggest that indirect effects of climate change, such as surface drying and shrub expansion, are likely main threats to slow-growing bryophytes across the Arctic, with consequences for biodiversity and C balance.

The majority of variation in six traits critical to the growth, survival and reproduction of plant species is thought to be organised along just two dimensions, corresponding to strategies of plant size and resource acquisition. However, it is unknown whether global plant trait relationships extend to climatic extremes, and if these interspecific relationships are confounded by trait variation within species. We test whether trait relationships extend to the cold extremes of life on Earth using the largest database of tundra plant traits yet compiled. We show that tundra plants demonstrate remarkably similar resource economic traits, but not size traits, compared to global distributions, and exhibit the same two dimensions of trait variation. Three quarters of trait variation occurs among species, mirroring global estimates of interspecific trait variation. Plant trait relationships are thus generalizable to the edge of global trait-space, informing prediction of plant community change in a warming world.