Environmental drivers of increased ecosystem respiration in a warming tundra.

Arctic and alpine tundra ecosystems are large reservoirs of organic carbon1,2. Climate warming may stimulate ecosystem respiration and release carbon into the atmosphere3,4. The magnitude and persistency of this stimulation and the environmental mechanisms that drive its variation remain uncertain5-7. This hampers the accuracy of global land carbon-climate feedback projections7,8. Here we synthesize 136 datasets from 56 open-top chamber in situ warming experiments located at 28 arctic and alpine tundra sites which have been running for less than 1 year up to 25 years. We show that a mean rise of 1.4 °C [confidence interval (CI) 0.9-2.0 °C] in air and 0.4 °C [CI 0.2-0.7 °C] in soil temperature results in an increase in growing season ecosystem respiration by 30% [CI 22-38%] (n = 136). Our findings indicate that the stimulation of ecosystem respiration was due to increases in both plant-related and microbial respiration (n = 9) and continued for at least 25 years (n = 136). The magnitude of the warming effects on respiration was driven by variation in warming-induced changes in local soil conditions, that is, changes in total nitrogen concentration and pH and by context-dependent spatial variation in these conditions, in particular total nitrogen concentration and the carbon:nitrogen ratio. Tundra sites with stronger nitrogen limitations and sites in which warming had stimulated plant and microbial nutrient turnover seemed particularly sensitive in their respiration response to warming. The results highlight the importance of local soil conditions and warming-induced changes therein for future climatic impacts on respiration.

Traditional plant functional groups explain variation in economic but not size-related traits across the tundra biome.

AIM: Plant functional groups are widely used in community ecology and earth system modelling to describe trait variation within and across plant communities. However, this approach rests on the assumption that functional groups explain a large proportion of trait variation among species. We test whether four commonly used plant functional groups represent variation in six ecologically important plant traits. LOCATION: Tundra biome. TIME PERIOD: Data collected between 1964 and 2016. MAJOR TAXA STUDIED: 295 tundra vascular plant species. METHODS: We compiled a database of six plant traits (plant height, leaf area, specific leaf area, leaf dry matter content, leaf nitrogen, seed mass) for tundra species. We examined the variation in species-level trait expression explained by four traditional functional groups (evergreen shrubs, deciduous shrubs, graminoids, forbs), and whether variation explained was dependent upon the traits included in analysis. We further compared the explanatory power and species composition of functional groups to alternative classifications generated using post hoc clustering of species-level traits. RESULTS: Traditional functional groups explained significant differences in trait expression, particularly amongst traits associated with resource economics, which were consistent across sites and at the biome scale. However, functional groups explained 19% of overall trait variation and poorly represented differences in traits associated with plant size. Post hoc classification of species did not correspond well with traditional functional groups, and explained twice as much variation in species-level trait expression. MAIN CONCLUSIONS: Traditional functional groups only coarsely represent variation in well-measured traits within tundra plant communities, and better explain resource economic traits than size-related traits. We recommend caution when using functional group approaches to predict tundra vegetation change, or ecosystem functions relating to plant size, such as albedo or carbon storage. We argue that alternative classifications or direct use of specific plant traits could provide new insights for ecological prediction and modelling.

Intra- and interspecific trait variations reveal functional relationships between specific leaf area and soil niche within a subtropical forest.

Background and Aims: How functional traits vary with environmental conditions is of fundamental importance in trait-based community ecology. However, how intraspecific variability in functional traits is connected to species distribution is not well understood. This study investigated inter- and intraspecific variation of a key functional trait, i.e. specific leaf area (leaf area per unit dry mass; SLA), in relation to soil factors and tested if trait variation is more closely associated with specific environmental regimes for low-variability species than for high-variability species. Methods: In a subtropical evergreen forest plot (50 ha, southern China), 106 700 leaves from 5335 individuals of 207 woody species were intensively collected, with 30 individuals sampled for most species to ensure a sufficient sample size representative of intraspecific variability. Soil conditions for each plant were estimated by kriging from more than 1700 observational soil locations across the plot. Intra- and interspecific variation in SLA were separately related to environmental factors. Based on the species-specific variation of SLA, species were categorized into three groups: low-, intermediate- and high-intraspecific variability. Intraspecific habitat ranges and the strength of SLA-habitat relationships were compared among these three groups. Key Results: Interspecific variation in SLA overrides the intraspecific variation (77 % vs. 8 %). Total soil nitrogen (TN, positively) and total organic carbon (TOC, negatively) are the most important explanatory factors for SLA variation at both intra- and interspecific levels. SLA, both within and between species, decreases with decreasing soil nitrogen availability. As predicted, species with low intraspecific variability in SLA have narrower habitat ranges with respect to soil TOC and TN and show a stronger SLA-habitat association than high-variability species. Conclusions: For woody plants low SLA is a phenotypic and probably adaptive response to nitrogen stress, which drives the predominance of species with ever-decreasing SLA towards less fertile habitats. Intraspecific variability in SLA is positively connected to species' niche breadth, suggesting that low-variability species may play a more deterministic role in structuring plant assemblages than high-variability species. This study highlights the importance of quantifying intraspecific trait variation to improve our understanding of species distributions across a vegetated landscape.

Downed woody debris carbon emissions in a European temperate virgin forest as driven by species, decay classes, diameter and microclimate.

Downed woody debris (DWD) plays an important role as regulator of nutrient and carbon (C) cycling in forests, accounting for up to the 20 % of the total C stocks in primary forests. DWD persistence is highly influenced by microbial decomposition, which is determined by various environmental factors, including fluctuations in temperature and moisture, as well as in intrinsic DWD properties determined by species, diameter, or decay classes (DCs). The relative importance of these different drivers, as well as their interactions, remains largely unknown. Moreover, the importance of DWD for C cycling in virgin forests remains poorly understood, due to their scarcity and poor accessibility. To address this research gap, we conducted a study on DWD respiration (RDWD), in a temperate virgin forest dominated by European beech and silver fir. Our investigation analysed the correlation between RDWD of these two dominant tree species and the seasonal changes in climate (temperature and moisture), considering other intrinsic DWD traits such as DCs (1, 2 and 4) and diameters (1, 10 and 25 cm). As anticipated, RDWD (normalized per gram of dry DWD) increased with air temperature. Surprisingly, DWD diameter also had a strong positive correlation with RDWD. Nonetheless, the sensitivity to both variables and other intrinsic traits (DC and density) was greatly modulated by the species. On the contrary, water content, which exhibited a considerable spatial variation, had an overall negative effect on RDWD. Virgin forests are generally seen as ineffective C sinks due to their lack of net productivity and high respiration and nutrient turnover. However, the rates of RDWD in this virgin forest were significantly lower than those previously estimated for managed forests. This suggests that DWD in virgin forests may be buffering forest CO2 emissions to the atmosphere more than previously thought.

Litter stoichiometric traits of plant species of high-latitude ecosystems show high responsiveness to global change without causing strong variation in litter decomposition.

• High-latitude ecosystems are important carbon accumulators, mainly as a result of low decomposition rates of litter and soil organic matter. We investigated whether global change impacts on litter decomposition rates are constrained by litter stoichiometry. • Thereto, we investigated the interspecific natural variation in litter stoichiometric traits (LSTs) in high-latitude ecosystems, and compared it with climate change-induced LST variation measured in the Meeting of Litters (MOL) experiment. This experiment includes leaf litters originating from 33 circumpolar and high-altitude global change experiments. Two-year decomposition rates of litters from these experiments were measured earlier in two common litter beds in sub-Arctic Sweden. • Response ratios of LSTs in plants of high-latitude ecosystems in the global change treatments showed a three-fold variation, and this was in the same range as the natural variation among species. However, response ratios of decomposition were about an order of magnitude lower than those of litter carbon/nitrogen ratios. • This implies that litter stoichiometry does not constrain the response of plant litter decomposition to global change. We suggest that responsiveness is rather constrained by the less responsive traits of the Plant Economics Spectrum of litter decomposability, such as lignin and dry matter content and specific leaf area.

Seasonal climate manipulations have only minor effects on litter decomposition rates and N dynamics but strong effects on litter P dynamics of sub-arctic bog species.

Litter decomposition and nutrient mineralization in high-latitude peatlands are constrained by low temperatures. So far, little is known about the effects of seasonal components of climate change (higher spring and summer temperatures, more snow which leads to higher winter soil temperatures) on these processes. In a 4-year field experiment, we manipulated these seasonal components in a sub-arctic bog and studied the effects on the decomposition and N and P dynamics of leaf litter of Calamagrostis lapponica, Betula nana, and Rubus chamaemorus, incubated both in a common ambient environment and in the treatment plots. Mass loss in the controls increased in the order Calamagrostis < Betula < Rubus. After 4 years, overall mass loss in the climate-treatment plots was 10 % higher compared to the ambient incubation environment. Litter chemistry showed within each incubation environment only a few and species-specific responses. Compared to the interspecific differences, they resulted in only moderate climate treatment effects on mass loss and these differed among seasons and species. Neither N nor P mineralization in the litter were affected by the incubation environment. Remarkably, for all species, no net N mineralization had occurred in any of the treatments during 4 years. Species differed in P-release patterns, and summer warming strongly stimulated P release for all species. Thus, moderate changes in summer temperatures and/or winter snow addition have limited effects on litter decomposition rates and N dynamics, but summer warming does stimulate litter P release. As a result, N-limitation of plant growth in this sub-arctic bog may be sustained or even further promoted.

Climatic and soil factors explain the two-dimensional spectrum of global plant trait variation.

Plant functional traits can predict community assembly and ecosystem functioning and are thus widely used in global models of vegetation dynamics and land-climate feedbacks. Still, we lack a global understanding of how land and climate affect plant traits. A previous global analysis of six traits observed two main axes of variation: (1) size variation at the organ and plant level and (2) leaf economics balancing leaf persistence against plant growth potential. The orthogonality of these two axes suggests they are differently influenced by environmental drivers. We find that these axes persist in a global dataset of 17 traits across more than 20,000 species. We find a dominant joint effect of climate and soil on trait variation. Additional independent climate effects are also observed across most traits, whereas independent soil effects are almost exclusively observed for economics traits. Variation in size traits correlates well with a latitudinal gradient related to water or energy limitation. In contrast, variation in economics traits is better explained by interactions of climate with soil fertility. These findings have the potential to improve our understanding of biodiversity patterns and our predictions of climate change impacts on biogeochemical cycles.

Is leaf dry matter content a better predictor of soil fertility than specific leaf area?

BACKGROUND AND AIMS: Specific leaf area (SLA), a key element of the 'worldwide leaf economics spectrum', is the preferred 'soft' plant trait for assessing soil fertility. SLA is a function of leaf dry matter content (LDMC) and leaf thickness (LT). The first, LDMC, defines leaf construction costs and can be used instead of SLA. However, LT identifies shade at its lowest extreme and succulence at its highest, and is not related to soil fertility. Why then is SLA more frequently used as a predictor of soil fertility than LDMC? METHODS: SLA, LDMC and LT were measured and leaf density (LD) estimated for almost 2000 species, and the capacity of LD to predict LDMC was examined, as was the relative contribution of LDMC and LT to the expression of SLA. Subsequently, the relationships between SLA, LDMC and LT with respect to soil fertility and shade were described. KEY RESULTS: Although LD is strongly related to LDMC, and LDMC and LT each contribute equally to the expression of SLA, the exact relationships differ between ecological groupings. LDMC predicts leaf nitrogen content and soil fertility but, because LT primarily varies with light intensity, SLA increases in response to both increased shade and increased fertility. CONCLUSIONS: Gradients of soil fertility are frequently also gradients of biomass accumulation with reduced irradiance lower in the canopy. Therefore, SLA, which includes both fertility and shade components, may often discriminate better between communities or treatments than LDMC. However, LDMC should always be the preferred trait for assessing gradients of soil fertility uncoupled from shade. Nevertheless, because leaves multitask, individual leaf traits do not necessarily exhibit exact functional equivalence between species. In consequence, rather than using a single stand-alone predictor, multivariate analyses using several leaf traits is recommended.

Similar cation exchange capacities among bryophyte species refute a presumed mechanism of peatland acidification.

Fen-bog succession is accompanied by strong increases of carbon accumulation rates. We tested the prevailing hypothesis that living Sphagna have extraordinarily high cation exchange capacity (CEC) and therefore acidify their environment by exchanging tissue-bound protons for basic cations in soil water. As Sphagnum invasion in a peatland usually coincides with succession from a brown moss-dominated alkaline fen to an acidic bog, the CEC of Sphagna is widely believed to play an important role in this acidification process. However, Sphagnum CEC has never been compared explicitly to that of a wide range of other bryophyte taxa. Whether high CEC directly leads to the ability to acidify the environment in situ also remains to be tested. We screened 20 predominant subarctic bryophyte species, including fen brown mosses and bog Sphagna for CEC, in situ soil water acidification capacity (AC), and peat acid neutralizing capacity (ANC). All these bryophyte species possessed substantial CEC, which was remarkably similar for brown mosses and Sphagna. This refutes the commonly accepted idea of living Sphagnum CEC being responsible for peatland acidification, as Sphagnum's ecological predecessors, brown mosses, can do the same job. Sphagnum AC was several times higher than that of other bryophytes, suggesting that CE (cation exchange) sites of Sphagna in situ are not saturated with basic cations, probably due to the virtual absence of these cations in the bog water. Together, these results suggest that Sphagna can not realize their CEC in bogs, while fen mosses can do so in fens. The fen peat ANC was 65% higher than bog ANC, indicating that acidity released by brown mosses in the CE process was neutralized, maintaining an alkaline environment. We propose two successional pathways indicating boundaries for a fen-bog shift with respect to bryophyte CEC. In neither of them is Sphagnum CE an important factor. We conclude that living Sphagnum CEC does not play any considerable role in the fen-bog shift. Alternatively, we propose that exclusively indirect effects of Sphagnum expansion such as peat accumulation and subsequent blocking of upward alkaline soil water transport are keys to the fen-bog succession and therefore for bog-associated carbon accumulation.

Stomatal vs. genome size in angiosperms: the somatic tail wagging the genomic dog?

BACKGROUND AND AIMS: Genome size is a function, and the product, of cell volume. As such it is contingent on ecological circumstance. The nature of 'this ecological circumstance' is, however, hotly debated. Here, we investigate for angiosperms whether stomatal size may be this 'missing link': the primary determinant of genome size. Stomata are crucial for photosynthesis and their size affects functional efficiency. METHODS: Stomatal and leaf characteristics were measured for 1442 species from Argentina, Iran, Spain and the UK and, using PCA, some emergent ecological and taxonomic patterns identified. Subsequently, an assessment of the relationship between genome-size values obtained from the Plant DNA C-values database and measurements of stomatal size was carried out. KEY RESULTS: Stomatal size is an ecologically important attribute. It varies with life-history (woody species < herbaceous species < vernal geophytes) and contributes to ecologically and physiologically important axes of leaf specialization. Moreover, it is positively correlated with genome size across a wide range of major taxa. CONCLUSIONS: Stomatal size predicts genome size within angiosperms. Correlation is not, however, proof of causality and here our interpretation is hampered by unexpected deficiencies in the scientific literature. Firstly, there are discrepancies between our own observations and established ideas about the ecological significance of stomatal size; very large stomata, theoretically facilitating photosynthesis in deep shade, were, in this study (and in other studies), primarily associated with vernal geophytes of unshaded habitats. Secondly, the lower size limit at which stomata can function efficiently, and the ecological circumstances under which these minute stomata might occur, have not been satisfactorally resolved. Thus, our hypothesis, that the optimization of stomatal size for functional efficiency is a major ecological determinant of genome size, remains unproven.

Global plant trait relationships extend to the climatic extremes of the tundra biome.

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.

Functional leaf attributes predict litter decomposition rate in herbaceous plants.

We tested the hypothesis that functional attributes of living leaves provide a basis for predicting the decomposition rate of leaf litter. The data were obtained from standardized screening tests on 38 British herbaceous species. Graminoid monocots had physically tougher leaves with higher silicon contents than did herbaceous dicots, and this corresponded with the lesser decomposibility of the former. Total base content of living leaves was a good predictor of litter decomposition rate, but the evolutionary and ecological basis for this relationship appeared to differ between graminoid monocots and herbaceous dicots. In the monocots, litter decomposition rate was strongly predicted by leaf potassium content, which appeared to reflect other growth- related plant attributes such as seedling maximum relative growth rate, foliar nitrogen and phosphorus content, specific leaf area and short leaf lifespan. In the dicots the relationship between total leaf base content and litter decomposition rate was not unambiguously explained by growth-related leaf attributes, possibly because of the considerable calcium uptake by dicots, which varies according to calcium availability in the soil.

A triangular relationship between leaf size and seed size among woody species: allometry, ontogeny, ecology and taxonomy.

A hypothesized relationship between seed weight and leaf size was investigated for 58 diverse British (semi-)woody species. Interspecific variation in leaf size of adult plants corresponded allometrically with interspecific variation in the weight of an infructescence (seed-bearing inflorescence). The relationship between seed size and leaf size of adult plants was triangular. The corners of the triangle were interpreted in terms of ecological strategy. Medium-sized infructescences, small seeds and large leaves were seen among medium-sized, fast-growing, earlier-successional, mostly deciduous shrubs and trees; small infructescences, small seeds and small leaves mostly among low, slow-growing evergreens from stress-prone, proclimax habitats; and large infructescences, large seeds and large leaves among slow-growing, later-successional trees of potential competitive vigour. The hypothesis that the combination of large seeds and small leaves is allometrically unlikely was supported by the data. The roles of ontogeny and taxonomic relatedness in the seed size-leaf size relationship were examined by correlative and taxonomic analyses of seed, plant and leaf size during the unfolding of the life history from seed through two seedling phases to adulthood. Deciduous versus evergreen leaf habit was a source of deviation from the otherwise linear allometric relationships during ontogenetic development, none of which were, individually, confounded significantly with taxonomy.

Leaf structure and anatomy as related to leaf mass per area variation in seedlings of a wide range of woody plant species and types.

The structural causes of variation in leaf mass per area, and of variations in leaf structure accounted for by leaf habit and life form, were explored in a set of laboratory-grown seedlings of 52 European woody species. The leaf traits analysed included density, thickness, saturated mass/dry mass, and leaf nitrogen per mass and per area. Other traits described the anatomy of leaves, most of them relating to the lamina (proportions of palisade and spongy parenchymata, epidermis, air space and sclerified tissues, expressed as volume per leaf area, and per-cell transversal areas of epidermis and parenchymata), and another referring to the mid rib (transversal section of sclerified tissues). Across the whole set of species leaf mass per area was correlated with leaf density but not with thickness, and this was confirmed by taxonomic relatedness tests. Denser leaves corresponded with greater proportion of sclerified tissues in the lamina, smaller cells and lower water and N contents, but no relation was found with the proportion of air space in the lamina. Taxonomic relatedness analysis statistically supported the negative association of leaf density with saturated to dry leaf mass ratio. Thicker leaves also exhibited greater volume per leaf area and greater individual cell area in each of the tissues, particularly parenchyma. Mean leaf mass per area and leaf thickness were lower in deciduous than in evergreen species, but no significant differences in leaf density, proportion of sclerified tissues in the lamina or cell area were found between the two groups. Leaf mass per area was higher in trees and subshrubs than in shrubs and climbers-plus-scramblers, this rank being equal for leaf density and proportion of sclerified tissues in the lamina, and reversed for cell area. Given the standardised environment and ontogenetic phase of the seedlings, we conclude that variation in leaf structure and anatomy among species and species groups has a strong genetic basis, and is already expressed early in the development of woody plants. From an ecological viewpoint, we can interpret greater leaf mass per area across this species set as greater allocation to support and defence functions, as shown predominantly by species from resource-poor environments.

Stem anatomy and relative growth rate in seedlings of a wide range of woody plant species and types.

Stem traits were analysed in laboratory-grown seedlings of 80 European woody and semiwoody species of known potential relative growth rate (RGR) and of similar ontogenetic phase. The objectives were, firstly, to assess the relation between stem structure and plant growth potential and, secondly, to explore how stem structure varies among species differing in life form and leaf habit. Hydraulic conductance was represented by the mean diameter of the widest xylem conduits (Dmax), and structural strength by the percentage of xylem tissue occupied by cell wall material (CWx) or stem tissue density (SD). Across all species RGR showed a weak positive correlation with Dmax and weak negative ones with CWx and SD, with slow-growers showing great dispersion of stem trait values. In the RGR-Dmax relationship this dispersion disappeared when trees were removed from the analysis. None of the relationships were significant among tree species alone. It was suggested that fast-growers require a xylem with wide conduits (high Dmax) to achieve high hydraulic conductivity, and "cheaply" constructed stems (low CWx and SD) to maximise allocation to leaves. However, the possession of such traits does not guarantee fast growth, as other factors may constrain RGR elsewhere in the plant. Deciduous seedlings showed higher Dmax and lower CWx than evergreens. Higher Dmax could reflect an innate higher tolerance of conductivity loss by freeze-induced embolism in deciduous plants, which are not burdened by the maintenance of foliage in winter. In contrast, life forms were differentiated most clearly by SD. For instance, shrub seedlings had less dense stem tissues than tree seedlings, possibly because they need less investment in long-term strength and stature.

Foliar nutrients in relation to growth, allocation and leaf traits in seedlings of a wide range of woody plant species and types.

This study aimed to identify functional correlates of seedling leaf nutrient content among woody species and to characterise functional species groups with respect to leaf nutrient attributes. Seedlings of 81 woody species from the temperate zone of western Europe were grown in a standard laboratory environment with standard, near-optimal nutrient availability. Weight-based leaf N content (Nwght) was positively correlated with mean relative growth rate (RGR), but the correlation with mean RGR was tighter when leaf N was expressed on a whole-plant weight basis: leaf nitrogen weight ratio (LNWR). Area-based leaf N content (Narea) was not associated with mean RGR, but was closely correlated with the quotient of saturated leaf weight and leaf area. Weight-based leaf K content (Kwght) was a close correlate of the saturated/dry weight ratio of the foliage. Within the lower range, Kwght corresponded with growth-related nutrient attributes, but higher values appeared to indicate succulence or remobilisable stored water. Functional groups of species and genera could be distinguished with respect to seedling leaf nutrient attributes. Deciduous woody climbers and scramblers had consistently higher leaf Nwght, LNWR and (apparently) leaf Kwght than other deciduous species or genera, and shrubs had higher values than trees. These differences seemed due partly to variation in specific leaf area. Evergreens had consistently higher leaf Narea than deciduous plants, but there were no significant differences in weight-based leaf nutrient attributes between these two groups, possibly because of `luxury nutrient consumption' by the slow-growing evergreens. Another functional group was that of the nitrogen-fixing species, which had consistently high innate leaf Nwght compared to non-N-fixers. The ecological significance of the leaf nutrient attributes in this study is discussed by comparing the seedling data with those from field-collected material, and by brief reference to the natural habitats of the species.

Meta-analysis reveals profound responses of plant traits to glacial CO2 levels.

A general understanding of the links between atmospheric CO2 concentration and the functioning of the terrestrial biosphere requires not only an understanding of plant trait responses to the ongoing transition to higher CO2 but also the legacy effects of past low CO2. An interesting question is whether the transition from current to higher CO2 can be thought of as a continuation of the past trajectory of low to current CO2 levels. Determining this trajectory requires quantifying the effect sizes of plant response to low CO2. We performed a meta-analysis of low CO2 growth experiments on 34 studies with 54 species. We quantified how plant traits vary at reduced CO2 levels and whether C3 versus C4 and woody versus herbaceous plant species respond differently. At low CO2, plant functioning changed drastically: on average across all species, a 50% reduction in current atmospheric CO2 reduced net photosynthesis by 38%; increased stomatal conductance by 60% and decreased intrinsic water use efficiency by 48%. Total plant dry biomass decreased by 47%, while specific leaf area increased by 17%. Plant types responded similarly: the only significant differences being no increase in SLA for C4 species and a 16% smaller decrease in biomass for woody C3 species at glacial CO2. Quantitative comparison of low CO2 effect sizes to those from high CO2 studies showed that the magnitude of response of stomatal conductance, water use efficiency and SLA to increased CO2 can be thought of as continued shifts along the same line. However, net photosynthesis and dry weight responses to low CO2 were greater in magnitude than to high CO2. Understanding the causes for this discrepancy can lead to a general understanding of the links between atmospheric CO2 and plant responses with relevance for both the past and the future.

Climate change has only a minor impact on nutrient resorption parameters in a high-latitude peatland.

Nutrient resorption from senescing plant tissues is an important determinant of the fitness of plant populations in nutrient-poor ecosystems, because it makes plants less dependent on current nutrient uptake. Moreover, it can have significant "afterlife" effects through its impact on litter chemistry and litter decomposability. Little is known about the effects of climate change on nutrient resorption. We studied the effects of climate change treatments (including winter snow addition, and spring and/or summer warming) on nutrient resorption of four dominant species in a nutrient-poor subarctic peatland. These species were Betula nana (woody deciduous), Vaccinium uliginosum (woody deciduous), Calamagrostis lapponica (graminoid) and Rubus chamaemorus (forb). After five years of treatments both mature and senesced leaf N concentrations showed a small but significant overall reduction in response to the climate treatments. However, the effects were species-specific. For example, in the controls the N concentration in senesced leaves of Calamagrostis (3.0+/-0.2 mg N g(-1)) was about four times lower than for Rubus (11.2+/-0.2 mg N g(-1)). There were no significant treatment effects on N resorption efficiency (% of the N pool in mature leaves that is resorbed during senescence). The nitrogen resorption efficiency of Calamagrostis (about 80%) was higher than in the other three species (about 60%). Thus, climate change has only a minor impact on nutrient resorption parameters. However, given the substantial interspecific differences in these parameters, substantial changes in plant-soil feedbacks may be expected as a result of the observed changes in the species composition of high-latitude vegetation. These changes are species-specific and thus difficult to predict.

Foliar pH as a new plant trait: can it explain variation in foliar chemistry and carbon cycling processes among subarctic plant species and types?

Plant traits have become popular as predictors of interspecific variation in important ecosystem properties and processes. Here we introduce foliar pH as a possible new plant trait, and tested whether (1) green leaf pH or leaf litter pH correlates with biochemical and structural foliar traits that are linked to biogeochemical cycling; (2) there is consistent variation in green leaf pH or leaf litter pH among plant types as defined by nutrient uptake mode and higher taxonomy; (3) green leaf pH can predict a significant proportion of variation in leaf digestibility among plant species and types; (4) leaf litter pH can predict a significant proportion of variation in leaf litter decomposability among plant species and types. We found some evidence in support of all four hypotheses for a wide range of species in a subarctic flora, although cryptogams (fern allies and a moss) tended to weaken the patterns by showing relatively poor leaf digestibility or litter decomposability at a given pH. Among seed plant species, green leaf pH itself explained only up to a third of the interspecific variation in leaf digestibility and leaf litter up to a quarter of the interspecific variation in leaf litter decomposability. However, foliar pH substantially improved the power of foliar lignin and/or cellulose concentrations as predictors of these processes when added to regression models as a second variable. When species were aggregated into plant types as defined by higher taxonomy and nutrient uptake mode, green-specific leaf area was a more powerful predictor of digestibility or decomposability than any of the biochemical traits including pH. The usefulness of foliar pH as a new predictive trait, whether or not in combination with other traits, remains to be tested across more plant species, types and biomes, and also in relation to other plant or ecosystem traits and processes.