T Matthew Robson

Abstract:

Abstract: Subalpine grasslands are highly seasonal environments and likely subject to strong variability in nitrogen (N) dynamics. Plants and microbes typically compete for N acquisition during the growing season and particularly at plant peak biomass. During snowmelt, plants could potentially benefit from a decrease in competition by microbes because of greater plant N uptake associated with active growth and freeze-thaw cycles restricting microbial growth. In managed subalpine grasslands, we expect these interactions to be influenced by recent changes in agricultural land-use, and associated modifications in plant and microbial communities. At a subalpine grassland site in the Central French Alps, a pulse of 15N was added to the soil at the end of snowmelt, allowing us to compare the dynamics of inorganic N uptake in plants and microbes during this period with that previously reported at the peak biomass in July. In all grasslands, specific plant (per g of biomass) dissolved inorganic N (DIN) uptake was two to five times greater at snow-melt than at peak biomass, whereas the specific microbial DIN uptakes were similar between the two sampling dates. On an area basis, plant communities took more DIN than microbial communities at the end of snowmelt, and the intensity of this DIN uptake by plants differed across land use types. Consequently, N partitioning after snowmelt switches in favor of plant communities allowing them to support their growing capacities at this period of the year. Seasonal differences in microbial and plant N-related dynamics were also affected by past (terraced vs. unterraced) rather than current (mown vs. unmown) land use. In terraced grasslands, microbial biomass N remained similar across seasons, whereas in unterraced grasslands, microbial biomass N was higher and microbial C : N lower at the end of snowmelt as compared to peak biomass. Further investigations on microbial community composition and their organic N uptake dynamics are required to better understand the decrease in microbial DIN uptake.

Abstract: European beech (Fagus sylvatica) reaches the south-western limit of its distribution in northern Spain, beyond which the Mediterranean climate is thought to restrict further expansion of the species range. Consequently, current and future climate change in the region is expected to push back the range margin and threaten the survival of local beech populations. In a provenance trial of pan-European beech populations growing under harsh conditions in La Rioja, we tested whether associations between the timing of spring phenology assessed over three years affected the performance of beeches at the site, and whether they exhibited a trade off between growth rate and survival. In particular, we considered whether the relationship between performance under conditions of summer drought and spring frost at the trial site was dependent on the climate at the site of provenance origin. We report that early-flushing provenances from continental climates in the south-east and parts of central Europe were among the tallest after ten years of growth in the trial: for instance from Val di Sella, northern Italy (mean bud burst day 114 and height 173 cm); Gotze Delchev, Bulgaria (day 115, height 135 cm); and Aarberg, Switzerland (day 118, height 151 cm). While late-flushing provenances from maritime climates in northern and western Europe were among the shortest in the trial: for instance from Soignes, Belgium (day 124, height 73 cm); Gullmarsberg, Sweden (day 122, height 69 cm); and Bathurst Estate, southern England (day 122, height 85 cm). There was no evidence that early flushing increased the mortality of trees at the trial site. The large-scale geographical patterns in flushing strategy reflected a trade off between pre-emptive growth before the summer drought and susceptibility to late frosts. Our findings emphasise the need to conserve populations from the range edge in the south of Europe, the Balkans and western Alps, whose combination of early flushing and drought resistance may become desirable traits for the improved future performance of beech in response to climate change.

Abstract: Patterns of intraspecific variation in functional traits have been widely studied across plant species to find out what general suites of traits provide functional advantage under specific environmental conditions. Much less is known about this variation within tree species and, in particular, about its relationship with performance variables such as photosynthetic rates under water deficit. Nevertheless, this knowledge is fundamental to understand the adaptive potential of drought sensitive tree species to increased aridity as predicted in the context of climate change. Intraspecific variation in photosynthetic performance and other leaf functional traits in response to water availability were examined in a glasshouse experiment using seedlings of six European beech populations. The physiological response of seedlings to a �water stress� treatment was compared to a �control� treatment along an experimental cycle of progressive soil water deficit and recovery. We found evidence of intraspecific variation in beech�s photosynthetic performance and other leaf functional traits in response to water availability. We also detected intraspecific variation in leaf-level tolerance of water deficit and phenotypic plasticity to water availability suggesting a pattern shaped by both regional and local scale effects. The Swedish population was particularly sensitive to water deficit, being the only population showing impaired photochemical efficiency under the experimental water deficit. Leaf-level tolerance of water deficit was related to PNUE, but not to other functional traits, such as WUE, SLA or leaf nitrogen content, that have been described to vary across species in adaptation to drought tolerance. Our results support the idea that general trends for variation in functional traits across species do not necessarily reflect a similar pattern when observed at the intraspecific level. The observed functional variation between beech populations reaffirms the importance of local adaptation to water deficit in the context of climate change.

Abstract: Lolium perenne (cv. AberDart) was grown at 14 locations along a latitudinal gradient across Europe (37-68°N) to study the impact of ultraviolet radiation (UV) and climate on aboveground growth and foliar UV-B absorbing compounds. At each location, plants were grown outdoors for 5 weeks in a replicated UV-B filtration experiment consisting of open, UV-B transparent (cellulose diacetate) and UV-B opaque (polyester) environments. Fourier transform-infrared spectroscopy was used to compare plant metabolite profiles in relation to treatment and location. UV radiation and climatic parameters were determined for each location from online sources and the data were assessed using a combination of anova and multiple regression analyses. Most of the variation in growth between the locations was attributable to the combination of climatic parameters, with minimum temperature identified as an important growth constraint. However, no single environmental parameter could consistently account for the variability in plant growth. Concentrations of foliar UV-B absorbing compounds showed a positive trend with solar UV across the latitudinal gradient; however, this relationship was not consistent in all treatments. The most striking experimental outcome from this study was the effect of presence or absence of filtration frames on UV-absorbing compounds. Overall, the study demonstrates the value of an European approach in studying the impacts of natural UV across a large latitudinal gradient. We have shown the feasibility of coordinated UV filtration at multiple sites but have also highlighted the need for open controls and careful interpretation of plant responses.

Abstract: Recent molecular and physiological studies have demonstrated that ultraviolet-B radiation (UV-B) can affect some of the processes involved in leaf growth, but the phases of leaf growth affected have not been clearly delimited. We used functional growth analysis to assess the effects of UV-B radiation on the time course of leaf growth in seedlings of two birch species (Betula pendula and Betula pubescens). Our aim was to identify the phase(s) of leaf development affected by UV-B radiation. In a greenhouse study, 1-year-old birch seedlings were subjected to three daily doses of supplemental UV-B radiation treatments (UV-B+) and no UV-B radiation controls (UV-B�). Leaf growth measurements every 2 days were complemented by assessment of other functional traits over a 4-week period at the start of the growing season. Using fitted curves, we were able to determine that the rate of leaf expansion was slowed by the UV-B+ treatment in leaves of B. pendula because of a slower maximum leaf growth rate compared with plants under the UV-B� controls, but that compensation toward the end of the period of expansion negated this difference when leaves reached their final size. UV-B+ had little effect on the rate of B. pubescens leaf growth despite a larger reduction in leaf final size due to UV-B+ than occurred in B. pendula leaves. In conclusion, effective regulation ameliorated the effects of UV-B radiation on leaf and seedling growth in B. pendula, whereas in B. pubescens, reductions in leaf final size under UV-B+ were consistent with a slightly reduced rate of height growth.

Abstract: Atmospheric nitrogen (N) deposition increasingly impacts remote ecosystems. At high altitudes, snow is a key carrier of water and nutrients from the atmosphere to the soil. Medium-sized subalpine grassland terraces are characteristic of agricultural landscapes in the French Alps and influence spatial and temporal snow pack variables. At the Lautaret Pass, we investigated snow and soil characteristics along mesotopographic gradients across the terraces before and during snowmelt. Total N concentrations in the snowpack did not vary spatially and were dominated by organic N forms either brought by dry deposition trapped by the snow, or due to snowmicrobial immobilization and turnover. As expected, snowpack depth, total N deposited with snow and snowmelt followed the terrace toposequence; more snow-N accumulated towards the bank over longer periods. However, direct effects of snow-N on soil-N cycling seem unlikely since the amount of nitrogen released into the soil from the snowpack was very small relative to soil-N pools and N mineralization rates. Nevertheless, some snow-N reached the soil at thaw where it underwent biotic and abiotic processes. In situ soil-N mineralization rates did not vary along the terrace toposequence but soil-N cycling was indirectly affected by the snowpack. Indeed, N mineralization responded to the snowmelt dynamic via induced temporal changes in soil characteristics (i.e. moisture and T) which cascaded down to affect N-related microbial activities and soil pH. Soil-NH4 and DON accumulated towards the bank during snowmelt while soil-NO3 followed a pulse-release pattern. At the end of the snowmelt season, organic substrate limitation might be accountable for the decrease in N mineralization in general, and in NH4 ? production in particular. Possibly, during snowmelt, other biotic or abiotic processes (nitrification, denitrification, plant uptake, leaching) were involved in the transformation and transfer of snow and soil-N pools. Finally, subalpine soils at the Lautaret Pass during snowmelt experienced strong biotic and abiotic changes and switched between a source and a sink of N.

Abstract: Seasonal variations and the differential effect of drought on photosynthetic parameters involved in carbon assimilation in forest species have been poorly studied under natural conditions. Limitations to diffusion and biochemical factors affecting leaf carbon uptake were analyzed in young beech seedlings (Fagus sylvatica L.) growing in natural gaps of a beech-wood at the southern limit of the species. Half of the seedlings received periodic watering in addition to natural rainfall to reduce the severity of the summer drought which is typical in the area. Plant water status was evaluated by measuring predawn water potential. Basic biochemical parameters were inferred from photosynthesis-CO2 curves (A-Cc) under saturating light and chlorophyll fluorescence. The curves were established from data collected on three dates during the summer months. The main variables studied included: stomatal and mesophyll conductance to CO2 (gs and gm respectively), maximum velocity of carboxylation (Vcmax) and maximum electron transport capacity (Jmax). The gm was estimated by two methodologies: the curve-fitting and J constant methods. Seedlings withstood moderate water stress as the leaf predawn water potential (Ψpd) measured during the study was within the range -0.2 to -0.5 MPa. Mild drought caused gs and gm to decrease only slightly in response to Ψpd. However both diffusional parameters explained most of the limitations to CO2 uptake. Furthermore, non-stomatal limitations were also important, since they limited net photosynthesis as water stress increased. In addition, it should be highlighted that biochemical limitations, prompted by Vcmax and Jmax, were related mainly to ontogenic factors, without any clear relationship with drought under the moderate water stress experienced by beech seedlings through the study. The results may help to further understanding of the functional mechanisms influencing the carbon fixation capacity of beech seedlings. This capacity may influence the total carbon balance of beech seedlings, and feed forward to impact on their growth and survival in drought-prone environments such as those in the south of Europe.

Abstract: The interactive effects of photosynthetically active radiation (PAR) on plants� acclimation to ultraviolet (UV) radiation were examined under field conditions in two barley varieties (Barke, Bonus). Plants, pre-treated under UV exclusion and low PAR intensities, were subsequently exposed to four radiation treatments representing the combination of low [�] and high [+] UV and PAR intensities. Selective UV and PAR filters were used for UV exclusion and reduction of PAR to ca 25% of ambient irradiance. A system of modulated lamps was used to enhance UV to ca 200% of ambient. Changes in flavonol and chlorophyll content, chlorophyll fluorescence, gas-exchange and leaf hyperspectral reflectance were studied during seven days of acclimation to the new treatments. At the end of this period morphological analysis of aboveground biomass was carried out. The [UV+PAR�] treatment significantly reduced the photosynthetic activity of barley leaves; the reduction was more pronounced in old than young leaves and greater in the variety Barke than Bonus. Whereas, [PAR+] treatment triggered photoprotective mechanisms which partially ameliorated the UV effects on photochemistry and carbon assimilation. The [PAR+] treatment induced accumulation of flavonols, mainly in young leaves, whereas in old leaves UV-induced accumulation was more pronounced. An inverse proportion was found between flavonol content and specific leaf area irrespective of barley variety and UV/PAR treatment. Enhanced UV radiation reduced the final leaf length, particularly in [PAR�] plants, in young leaves and in variety Barke. However, [PAR+] mitigated the morphological effects induced by the [UV+] treatment, particularly changes in SLA.

Abstract: We assessed the response of 11-year-old saplings from six beech provenances growing in a common-garden trial at the southwestern range limit. Provenances from distinct climatic regions across the European beech distribution were selected. The local Spanish provenance appeared well suited to the site conditions, maintaining high rates of assimilation even in midsummer, but so did the provenance of southern continental origin, from Gotze-Delchev, Bulgaria. Those provenances from cooler sites in central Europe, a continental mountain climate in the Czech Republic and a continental range-edge site in eastern Poland, along with a German provenance of mild maritime origin, had good physiological functionality in early summer but reduced carbon assimilation (Aarea) and apparent soil�leaf hydraulic conductivity (KL) in midsummer. The northern maritime provenance from Sweden demonstrated severely-reduced photosynthetic capacity. These groupings of provenances according to their photosynthetic performance, stable carbon isotope composition (δ13C; a proxy for water-use efficiency) and leaf water potential under marginal conditions, during late summer in the trial, suggest that they have divergent strategies for water use. The research highlights large intraspecific differences among beech provenances of distinct origin and strategies which are expected to modify their response to drought, requiring future genetic studies to explicitly determine the basis of this ecophysiological differentiation.

Abstract: Aim To assess the effect of local adaptation and phenotypic plasticity on the potential distribution of species under future climate changes. Trees may be adapted to specific climatic conditions; however, species range predictions have classically been assessed by species distributionmodels (SDMs) that do not account for intra-specific genetic variability and phenotypic plasticity, because SDMs rely on the assumption that species respond homogeneously to climate change across their range, i.e. a species is equally adapted throughout its range, and all species are equally plastic. These assumptions could cause SDMs to exaggerate or underestimate species at risk under future climate change. Location The Iberian Peninsula. Methods Species distributions are predicted by integrating experimental data and modelling techniques. We incorporate plasticity and local adaptation into a SDM by calibrating models of tree survivorship with adaptive traits in provenance trials. Phenotypic plasticity was incorporated by calibrating our model with a climatic index that provides a measure of the differences between sites and provenances. Results We present a new modelling approach that is easy to implement and makes use of existing tree provenance trials to predict species distribution models under global warming. Our results indicate that the incorporation of intrapopulation genetic diversity and phenotypic plasticity in SDMs significantly altered their outcome. In comparing species range predictions, the decrease in area occupancy under global warming conditions is smaller when considering our survival� adaptationmodel than that predicted by a �classical SDM� calibrated with presence� absence data. These differences in survivorship are due to both local adaptation and plasticity. Differences due to the use of experimental data in the model calibration are also expressed in our results: we incorporate a nullmodel that uses survival data from all provenances together. This model always predicts less reduction in area occupancy for both species than the SDM calibrated with presence�absence. Main conclusions We reaffirm the importance of considering adaptive traits when predicting species distributions and avoiding the use of occurrence data as a predictive variable. In light of these recommendations, we advise that existing predictions of future species distributions and their component populations must be reconsidered.

Abstract: Little is known about how environmental factors shape the short- and long-term responses of leaf respiration to temperature under field conditions despite the importance of respiration for plant and stand carbon balances. Impacts of water availability and canopy cover on leaf dark respiration (R) and temperature sensitivity were assessed in beech (Fagus sylvatica L.) seedlings in a sub-Mediterranean population. We studied seedlings established within canopy gaps (39% global site factor; GSF) that were subject to either no watering (unwatered plants; UW) or regular watering (2� 10% higher volumetric topsoil water content as summer progressed; W plants) and seedlings established beneath the adjacent understorey (12% GSF). Leaf R rose exponentially with diurnal increases in temperature; the same temperature sensitivity (Q10: 2.2) was found for understorey and gap plants, irrespective of watering treatment. Respiration estimated at 25 °C (R25) was lower in the understorey than the gaps and was significantly lower in the unwatered than in the watered gap plants by the end of summer (0.65 versus 0.80 μmol m�2 s�1). R25 declined with increasing summer temperature in all plants; however, respiration estimated at the prevailing ambient temperature did not change through the summer. There were parallel declines in R25 and concentrations of starch and soluble sugars with increasing summer temperature for gap plants. We conclude that seasonal shifts in temperature�response curves of beech leaf R occur in both low- and high-light environments; since leaf R decreased with increasing plant water deficit, such shifts are likely to be greater whenever plants experience summer drought compared to scenarios where plants experience high rainfall in summer.

Abstract: 1. Nitrogen (N) cycling is a key process determining ecosystem functioning in subalpine grasslands where traditional mowing and manuring are being abandoned. However, the roles of the plant and microbial communities in mediating changes inNavailability are still poorly understood. 2. We inoculated 15 subalpine grassland fields with dual-labelled ammonium nitrate (15NH4 + , 15NO3 )) during July 2005 and used pool dilutions over 1 month to calculate inorganic N fluxes into the microbial pool and uptake in plant communities by grasses, forbs and legumes. The effects of current land abandonment were assessed by comparing manured and mown terraces (ancient croplands) with other terraces where these practices have ceased, and mown versus unmown unterraced meadows. 3. Rapid cycling of inorganicNand high soilNavailability in forb-dominated manured and mown terraces resulted from fast plant N uptake and low microbial C:N ratio. In grass-dominated unmown terraces, N cycling was slower and N retention was greater; microbial N uptake remained similar to that in the other terraces, although a higher C:N ratio suggested a shift towards fungal dominance. 4. In unterraced meadows, pH was low due to reduced mixing of soil with the underlying calcareous rock. Soil [NH4 + ] was high and [NO3 ) ] low, but current management had no effect on N pool size, although plant N uptake was greater in the mown than unmown fields. This may be partially explained by high N retention by dominant Festuca paniculata tussocks. The microbial N pool and N uptake were both low and the microbial C:N ratio was high, suggesting that fungi slowed N cycling and reduced the influence of mowing onNturnover. 5. Synthesis. In these marginal long-term grasslands, with low productivity and high biodiversity value, changes in ecosystem function associated with reduced management intensity were mediated through slowerN cycling. This response was expressed as more gradual nutrient uptake but greater retention by unmown plant communities, slower microbial uptake and smaller soilNpools. In contrast to more productive ecosystems, such as north-western European grasslands, reduced management is detrimental to both biodiversity and the maintenance of soil-related ecosystem services. These costs will need to be balanced against potential benefits, such as carbon storage.

Abstract: Land use and climate changes induce shifts in plant functional diversity and community structure, thereby modifying ecosystem processes. This is particularly true for litter decomposition, an essential process in the biogeochemical cycles of carbon and nutrients. In this study, we asked whether changes in functional traits of living leaves in response to changes in land use and climate were related to rates of litter potential decomposition, hereafter denoted litter decomposability, across a range of 10 contrasting sites. To disentangle the different control factors on litter decomposition, we conducted a microcosm experiment to determine the decomposability under standard conditions of litters collected in herbaceous communities from Europe and Israel. We tested how environmental factors (disturbance and climate) affected functional traits of living leaves and how these traits then modified litter quality and subsequent litter decomposability. Litter decomposability appeared proximately linked to initial litter quality, with particularly clear negative correlations with lignin-dependent indices (litter lignin concentration, lignin : nitrogen ratio, and fiber component). Litter quality was directly related to community-weighted mean traits. Lignin-dependent indices of litter quality were positively correlated with community-weighted mean leaf dry matter content (LDMC), and negatively correlated with community-weighted mean leaf nitrogen concentration (LNC). Consequently, litter decomposability was correlated negatively with community-weighted mean LDMC, and positively with community-weighted mean LNC. Environmental factors (disturbance and climate) influenced community-weighted mean traits. Plant communities experiencing less frequent or less intense disturbance exhibited higher community-weighted mean LDMC, and therefore higher litter lignin content and slower litter decomposability. LDMC therefore appears as a powerful marker of both changes in land use and of the pace of nutrient cycling across 10 contrasting sites.

Abstract: Refugia of mixed beech forest persist in the central mountains of the Iberian Peninsula at the southwestern limit of European beech (Fagus sylvatica L.) distribution. The lack of beech regeneration is a concern in this region that has experienced reduced rainfall and higher temperatures over the past 30 years. Beech is considered especially susceptible to climate change because of its conservative shade-tolerant growth strategy; hence seedling responses to drought stress in gaps and in the understory are of particular interest. During the summer of 2007, a watering treatment raised the soil water content by up to5%in gap and understory plots of beech seedlings in a mixed beech forest. Root-collar diameter was increased by our watering treatment in understory seedlings. Neither drought-avoidance through stomatal closure nor physiological drought-tolerance mechanisms were able to mitigate the effects of water stress in the understory seedlings, whereas osmotic adjustment enhanced the ability of the gap seedlings to tolerate water stress. Overall, high photosynthetic rates in the gaps, despite the photoinhibitory effects of high radiation, allowed gap seedlings to survive and grow better than the understory seedlings irrespective of water availability. Our results indicate that further intensification of summer drought, predicted for the Iberian Peninsula, will hinder the establishment of a beech seedling bank in the understory because of the conflicting seedling trait responses to simultaneously withstand water stress and to tolerate shade.

Abstract: This study was designed to test the hypothesis that the spectral composition of incident radiation, as defined by the relative proportions of blue (B; λmax = 455 nm) and red (R; λmax = 625 nm) photons, can affect photosynthetic induction, since B photons stimulate stomatal opening and are more effectively absorbed by leaves than R photons. Different stages of photosynthetic induction, primarily determined by the photo-modulation of Rubisco activity and stomata opening, were investigated in dark-adapted leaves of Fagus sylvatica transferred to saturating irradiance [800μmol(photon) m�2 s�1] at B/R ratios of 1/3, 1/1, or 3/1. In agreement with our hypothesis, photosynthesis was induced faster by irradiance with a high B/R ratio (3/1); as demontrated by a higher IS60 (induction state 60 s after leaf illumination) and lower T90 (the time period required to reach 90 % of maximum steady-state photosynthesis). However, there were no differences in induction between leaves receiving equal (1/1) and low (1/3) B/R ratios. Electron transport was highly sensitive to radiation quality, exhibiting faster induction kinetics with increasing B/R ratio. Such stimulation of carbon-assimilatory processes corresponds with faster activation of Rubisco and lower non-photochemical quenching (NPQ) as the proportion of B photons is increased. In contrast, the kinetics of stomatal opening was independent of the spectral composition of incoming radiation. Since slightly higher absorption efficiency of high B/R radiation does not fully explain the changes in induction kinetics, the other possible mechanisms contributing to the stimulation of electron transport and Rubisco activity are discussed.

Abstract: � In subalpine grasslands, changes in abiotic conditions with decreased management intensity alter the functional composition of plant communities, leading to modifications of ecosystem properties. Here, it is hypothesized that the nature of plant feedbacks on soil moisture is determined by the values of key traits at the community level. � As community functional parameters of grasslands change along a gradient of land uses, those traits that respond most to differences in abiotic conditions produced by land use changes were identified. A vegetation removal experiment was then conducted to determine how each plant community affected soil moisture. � Soil moisture was negatively correlated with community root length and positively correlated with canopy height, whereas average leaf area was associated with productivity. These traits were successfully used to predict the effects on soil moisture of each plant community in the removal experiment. This result was validated using data from an additional set of fields. � These findings demonstrate that the modification of soil moisture following land use change in subalpine grasslands can be mediated through those plant functional traits that respond to water availability.

Abstract: Climate change is expected to involve morefrequent and intense summer droughts in the Mediterranean region. This represents a threat for long-term persistence of woody species, such as European holly (Ilex aquifolium), that originated under humid climates during the Tertiary period. The capacity of this species to persist under increased water stress, both in gaps and in the understory of an oak-dominated woodland, was assessed by quantifying phenotypic plasticity in response to drought and shade. Physiological responses in plant-water relations and gas exchange were used as performance indicators under the different environments. Phenotypic plasticity of drought-stressed holly trees in response to changes in the light environment was low relative to the known response of co-occurring forest trees. Differences between morphological traits (e.g. specific leaf area and leaf: sapwood ratio in twigs) of sun- and shade-grown trees were small but significant while physiological traits were largely unresponsive to light availability. This supports the hypothesis that late-successional shade-tolerant species exhibit greater morphological than physiological plasticity. Sapling acclimation capacity through physiological mechanisms such as osmotic adjustment was insufficient to protect from summer drought. Holly mainly inhabits oceanic climates where extreme temperatures and droughts are unusual. Our results suggest that the species occupies a narrowing niche in continental Mediterranean habitats, and may lack the capacity to persist under more-severe future climate scenarios because of its low phenotypic plasticity in response to light and drought stresses.

Abstract: � Background and Aims A standardized methodology to assess the impacts of land-use changes on vegetation and ecosystem functioning is presented. It assumes that species traits are central to these impacts, and is designed to be applicable in different historical, climatic contexts and local settings. Preliminary results are presented to show its applicability. � Methods Eleven sites, representative of various types of land-use changes occurring in marginal agro-ecosystems across Europe and Israel, were selected. Climatic data were obtained at the site level; soil data, disturbance and nutrition indices were described at the plot level within sites. Sixteen traits describing plant stature, leaf characteristics and reproductive phase were recorded on the most abundant species of each treatment. These data were combined with species abundance to calculate trait values weighed by the abundance of species in the communities. The ecosystem properties selected were components of above-ground net primary productivity and decomposition of litter. � Key Results The wide variety of land-use systems that characterize marginal landscapes across Europe was reflected by the different disturbance indices, and were also reflected in soil and/or nutrient availability gradients. The trait toolkit allowed us to describe adequately the functional response of vegetation to land-use changes, but we suggest that some traits (vegetative plant height, stem dry matter content) should be omitted in studies involving mainly herbaceous species. Using the example of the relationship between leaf dry matter content and aboveground dead material, we demonstrate how the data collected may be used to analyse direct effects of climate and land use on ecosystem properties vs. indirect effects via changes in plant traits. �Conclusions This work shows the applicability of a set of protocols that can be widely applied to assess the impacts of global change drivers on species, communities and ecosystems.

Abstract: Global environmental change affects the sustained provision of a wide set of ecosystem services. Although the delivery of ecosystem services is strongly affected by abiotic drivers and direct land use effects, it is also modulated by the functional diversity of biological communities (the value, range, and relative abundance of functional traits in a given ecosystem). The focus of this article is on integrating the different possible mechanisms by which functional diversity affects ecosystem properties that are directly relevant to ecosystem services. We propose a systematic way for progressing in understanding how land cover change affects these ecosystem properties through functional diversity modifications. Models on links between ecosystem properties and the local mean, range, and distribution of plant trait values are numerous, but they have been scattered in the literature, with varying degrees of empirical support and varying functional diversity components analyzed. Here we articulate these different components in a single conceptual and methodological framework that allows testing them in combination. We illustrate our approach with examples from the literature and apply the proposed framework to a grassland system in the central French Alps in which functional diversity, by responding to land use change, alters the provision of ecosystem services important to local stakeholders. We claim that our framework contributes to opening a new area of research at the interface of land change science and fundamental ecology.

Abstract: Nitrogen (N) availability in grasslands varies with agricultural land use. Traditional management regimes of mowing for hay and manuring in subalpine meadows maintain plant communities with exploitative functional strategies suited to fertile soils with fast turnover of nutrients. We investigated whether the neglect of traditional practices has led to a reduction in N availability in two parallel ecosystems (terraced and unterraced fields). Soil nitrate and ammonium contents were assessed using soil cores and ion exchange resins over a 1-year period, and assays of microbial nitrifying and denitrifying enzyme activities, made early in the growing season. A large difference in pH between the two ecosystems, caused by historical ploughing, facilitated greater N availability in terraced than unterraced fields. Abandonment of manuring and mowing caused a reduction in N availability and N transformation processes, which correlated with a shift in the plant community towards more-conservative functional strategies and greater dominance by grasses. We propose that positive feedback between the grassland management regime and dominant plant functional strategy maintained high N availability in these managed subalpine grasslands. When traditional practices of mowing and manuring are neglected, direct management effects combined with the spread of grass species with conservative strategies force down N availability in the soil, reduce microbial activity, change the pH, and lead to a long-term loss of characteristic herbaceous subalpine-meadow species.

Abstract: As a result of stratospheric ozone depletion, more solar ultraviolet-B radiation (UV-B, 280�315 nm) is reaching the Earth�s surface. Enhanced levels of UV-B may, in turn, alter ecosystem processes such as decomposition. Solar UV-B radiation could affect decomposition both indirectly, by changes in the chemical composition of leaves during growth, or directly by photochemical breakdown of litter and through changes in decomposer communities exposed to sunlight. In this experiment, we studied indirect and direct effects of solar UV-B radiation on decomposition of barley (Hordeum vulgare). We used barley straw and leaf litter grown under reduced UV-B (20% of ambient UV-B) or under near-ambient UV-B (90% of ambient UV-B) in Buenos Aires, Argentina, and decomposed the litter under reduced or near-ambient solar UV-B for 29 months in Tierra del Fuego, Argentina. We found that the UV-B treatment applied during growth decreased the decay rate. On the other hand, there was a marginally significant direct effect of elevated UV-B during the early stages of decomposition, suggesting increased mass loss. The effect of UV-B during growth on decomposition was likely the result of changes in plant litter chemical composition. Near-ambient UV-B received during plant growth decreased the concentrations of nitrogen, soluble carbohydrates, and N/P ratio, and increased the concentrations of phosphorus, cellulose, UV-B-absorbing compounds, and lignin/N ratio. Thus, solar UV-B radiation affects the decomposition of barley litter directly and indirectly, and indirect effects are persistent for the whole decomposition period.

Abstract: The peatlands of Tierra del Fuego are subject to increased solar ultraviolet-B radiation (UV-B) due to the influence of the Antarctic �ozone hole�. Research into the effects of climate change and ozone depletion on peatlands has predominantly focused on the higher plant community and neglected other organisms. In the second 3-year portion of a 6-year experiment, we intensified our investigations of the response of the peatland surface microfaunal community to current and attenuated solar UV-B, and assessed possible links to changes in the microenvironment. Near-ambient UV-B and reduced UV-B treatments were realised by stretching plastic film filters that differentially attenuate UV-B over peatland sample plots. We extracted the microfauna and analysed the dissolved nutrients held within Sphagnum capitula removed from the top 1-cm of the peatland. In line with previous findings in this system, testate amoebae were more abundant under nearambient UV-B than under reduced UV-B. Populations of the most common genus, Assulina, and other less prominent amoebae species of Heleopera and Euglypha, were consistently increased under near-ambient UV-B. Overall diversity of testate amoebae was also higher under near-ambient UV-B than under reduced UV-B, whereas rotifers, nematodes and mites were less abundant under near-ambient UV-B. Concentrations of DOC and P were generally higher under near-ambient UV-B than under reduced UV-B. These changes, combined with the changes previously reported in the plant and fungal communities, have the potential to influence peatland C storage, and surface nutrient availability. The peatland microfaunal community under near-ambient solar UV-B may be regulated by the plant community through the leaching of nutrients from leaf cells, and changes in Sphagnum morphology that affect the capitulum microenvironment.

Abstract: The influence of near-ambient and reduced solar UV-B radiation on a peatland microfungal community was assessed by exposing experimental plots to UVselective filtration. Replicate plots were covered with special plastic films to effect treatments of near-ambient and attenuated solar UV-B. The microfungal community from the top 1 cm of Sphagnum capitulum in a Tierra del Fuego peatland was censused throughout three growing seasons, between 1999 and 2002. Sphagnum capitula under near-ambient UV-B were more compressed and held more water than capitula under reduced UV-B. This water had a greater conductivity and was more acidic under nearambient UV-B, as would be expected with increased leaching from the Sphagnum leaves. Nine regularly occurring hyphal fungi from the peatland were identified, at least to genus. Over three field seasons, no treatment effect on total fungal colony abundance was recorded, but individual species abundance was increased (Mortierella alpina), decreased (Penicillium frequentans), or was unaffected (P. thomii, Aureobasidium) by near-ambient UV-B. Species richness was also slightly lower under near-ambient UV-B. These treatment differences were smaller than seasonal or inter-annual fluctuations in abundance and species richness. In a growth chamber experiment, lamp UV-B treatments indicated that realistic fluxes of UV-B can inhibit fungal growth in some species. In addition to this direct UV-B effect, we suggest that changes in the peatland fungal community under nearambient solar UV-B may also result from increased nutrient and moisture availability in the Sphagnum capitulum. The subtle nature of the responses of peatland fungi to solar UV-B suggests that most fungal species we encountered are well adapted to current solar UV-B fluxes in Tierra del Fuego.

Abstract: � Tierra del Fuego is subject to increases in solar UV-B radiation in the austral spring and summer due to ozone depletion. � Plastic films were used to filter solar UV-B radiation over peatland plots through six field seasons, resulting in near-ambient (c. 90%) and reduced (c. 17%) solar UV-B treatments. � As in the first three field seasons of treatments, near-ambient UV-B caused reduced height growth but had no effect on biomass production of the moss Sphagnum magellanicum. It reduced leaf and rhizome growth of Tetroncium magellanicum. Height growth and morphology of Empetrum rubrum and Nothofagus antarctica were only affected by solar UV-B during the fourth to sixth field seasons. There was also a decrease in Tetroncium leaf nitrogen under near-ambient UV-B. � Growth of Sphagnum was less affected than that of most emergent vascular plants. This enabled the Sphagnum mat to engulf more Nothofagus, and limit the escape of Empetrum under near-ambient UV-B. Yet, differences in the response of species to solar UV-B were not expressed as changes in plant community composition.

Abstract: Tierra del Fuego, Argentina (551S), receives increased solar ultraviolet-B radiation (UV-B) as a result of Antarctic stratospheric ozone depletion. We conducted a field study to examine direct and indirect effects of solar UV-B radiation on decomposition of Gunnera magellanica, a native perennial herb, and on the native community of decomposer organisms. In general, indirect effects of UV-B mostly occur due to changes in the chemical composition of litter, whereas direct effects during decomposition result from changes in decomposer organisms and/or differences in the photochemical breakdown of litter.We designed a full-factorial experiment using senescent leaves that had received either near-ambient or attenuated UV-B during growth. The leaves were distributed in litterbags and allowed to decompose under near-ambient or reduced solar UV-B during the growing season. We evaluated initial litter quality, mass loss, and nutrient release of decomposing litter, and microbial colonization of both initial litter and decomposed litter. We found that litter that decomposed under near-ambient UV-B had significantly less mass loss than litter that decomposed under reduced UV-B. The UV-B conditions received by plants during growth, which did not affect mass loss and nutrient composition of litter, affected fungal species composition but in different ways throughout the decomposition period. Before the decomposition trial, Beauveria bassiana and Penicillium frequentans were higher under reduced UV-B, whereas Cladosporium herbarum and pigmented bacteria were more common under the nearambient compared to the reduced UV-B treatment. After the decomposition period, leaves that had grown under reduced UV-B showed higher frequency of Penicillium thomii and lower frequency of Trichoderma polysporum than leaves that had grown under near-ambient conditions. The UV-B condition received during decomposition also affected fungal colonization, with Penicillium chrysogenum being more frequent in leaves that had decomposed under reduced UV-B, while the other species were not affected. Our results demonstrate that, in this ecosystem, the effects of UV-B radiation on decomposition apparently occurred mostly through changes in the fungal community, while changes in photochemical breakdown appeared to be less important.

Abstract: The southern part of Tierra del Fuego, in the southernmost tip of South America, is covered by dense Nothofagus spp. forests and Sphagnum-dominated peat bogs, which are subjected to the influence of ozone depletion and to increased levels of solar ultraviolet-B radiation (UV-B). Over the last 5 years we have studied some of the biological impacts of solar UV-B on natural ecosystems of this region.We have addressed two general problems: (i) do the fluctuations in UV-B levels under the influence of the Antarctic ozone �hole� have any measurable biological impact, and (ii) what are the long-term effects of solar (ambient) UV-B on the Tierra del Fuego ecosystems? In this paper, we provide an overview of the progress made during the first 4 years of the project.We highlight and discuss the following results: (1) ambient UV-B has subtle but significant inhibitory effects on the growth of herbaceous and graminoid species of this region (growth reduction #12%), whereas no consistent inhibitory effects could be detected in woody perennials; (2) in the species investigated in greatest detail, Gunnera magellanica, the inhibitory effect of solar UV-B is accompanied by increased levels of DNA damage in leaf tissue, and the DNA damage density in the early spring is clearly correlated with the dose of weighted UV-B measured at ground level; (3) the herbaceous species investigated thus far show little or no acclimation responses to ambient UV-B such as increased sunscreen levels and DNA repair capacity; and (4) ambient UV-B has significant effects on heterotrophic organisms, included marked inhibitory effects on insect herbivory. The results from the experiments summarized in this review clearly indicate that UV-B influences several potentially important processes and ecological interactions in the terrestrial ecosystems of Tierra del Fuego.

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Abstract: Seasonal variation and the differential effect of drought on photosynthetic parameters involved in carbon assimilation in forest species have been poorly studied under natural conditions. Limitations to diffusion and biochemical factors affecting leaf carbon uptake were analyzed in young beech seedlings (Fagus sylvtica L.) growing in natural gaps of a beech-wood at the southern limit of the species. Half of the seedling received periodic watering in addition to natural rainfall to reduce the severity of the summer drought which is typical in the area. Plant water status was evaluated by measuring predawn water potential. Basic biochemical parameters were inferred from photosynthesis-CO2 curves (A-Cc) under saturating light and chlorophyll fluorescence. The curves were established on three dates during the summer months Main variables studied included: stomatal and mesophyll conductance to CO2 (gs and gm respectively), maximum velocity of carboxylation (Vcmax) and maximum electron transport capacity (Jmax). The gm was estimated by two methodologies: the curve-fitting and J constant methods. Seedlings withstood a moderate water stress as the leaf predawn water potential (Ψpd) measured during the study was within the range �0.2 to �0.5 MPa. A mild drought caused gs and gm to decrease only slightly in response to Ψpd. However both diffusional parameters explained most of the limitations to CO2 uptake. Furthermore, non-stomatal limitations were also important limiting net photosynthesis as water stress increased. In addition, it should be highlighted that the biochemical limitations, prompted by Vcmax and Jmax, were related mainly to ontogenic factors, without any clear relationship with drought under the moderate water stress experienced by beech seedlings through the study. The results may help to further understand of the functional mechanisms influencing carbon fixation capacity of beech seedlings. This capacity may influence the total carbon balance of beech seedlings, and feed forward to their growth and survival under drought-prone environments such as those in the south of Europe.

Abstract: Spring phenology is considered one of the most important determinants of growth and survival in young stands. It is relatively easy to monitor and is expected to respond to climate changes that will affect the favourable period for growth in temperate regions. The response of trees to the environmental cues that govern spring phenology is largely under genetic control and inter-populational differences exist within species. This suggests that the trait undergoes site-specific selection. Data obtained through monitoring of bud burst at multiple beech provenance-trials were compared with specific site and weather data to reveal geographical clines in beech phenology.We fitted the Weibull function to harmonise phenology data collected using various flushing scales and at different intensities of monitoring. By comparing data from 20 annual census of phenology performed across 13 sites throughout Europe, we showed that accumulated temperature sum > 5°C modelled the timing and duration of flushing more consistently than other temperature sum models > 0°C or > 8°C, or simply Julian Day. Inconsistency in the number of degree hours required for flushing among sites, reinforced the need for testing of more complex mechanistic models that include photoperiod, chilling period, and summer drought in addition to temperature sum. South-North, East-West, and low-high elevational clines were confirmed from the analysis. These findings; reinforce the need for caution in planting provenances from the south-east of Europe, suited to warmer-drier summers, in more north-westerly sites; and highlight the location of some potentially valuable late-flushing populations that also tolerate warm dry temperatures.