Ratcliffe S., Liebergesell M., Ruiz-Benito P., Madrigal González J., Muñoz Castañeda J.M., Kändler G., Lehtonen A., Dahlgren J., Kattge J., Peñuelas J., Zavala M.A., Wirth C. (2015) Modes of functional biodiversity control on tree productivity across the European continent. Global Ecology and Biogeography. : 0-0.LinkDoi: 10.1111/geb.12406
Aim: The relative contribution of community functional diversity and composition to ecosystem functioning is a critical question in ecology in order to enable better predictions of how ecosystems may respond to a changing climate. However, there is little consensus about which modes of functional biodiversity are most important for tree growth at large spatial scales. Here we assessed the relative importance of climate, functional diversity and functional identity (i.e. the community mean values of four key functional traits) for tree growth across the European continent, spanning the northern boreal to the southern Mediterranean forests. Location: Finland, Germany, Sweden, Spain and Wallonia (Belgium). Methods: Using data from five European national forest inventories we applied a hierarchical linear model to estimate the sensitivity of tree growth to changes in climate, functional diversity and functional identity along a latitudinal gradient. Results: Functional diversity was weakly related to tree growth in the temperate and boreal regions and more strongly in the Mediterranean region. In the temperate region, where climate was the most important predictor, functional diversity and identity had a similar importance for tree growth. Functional identity was strongest at the latitudinal extremes of the continent, largely driven by strong changes in the importance of maximum height along the latitudinal gradient. Main conclusions: Functional diversity is an important driver of tree growth in the Mediterranean region, providing evidence that niche complementarity may be more important for tree growth in water-limited forests. The strong influence of functional identity at the latitudinal extremes indicates the importance of a particular trait composition for tree growth in harsh climates. Furthermore, we speculate that this functional identity signal may reflect a trait-based differentiation of successional stages rather than abiotic filtering due to water or energy limitation. © 2015 John Wiley & Sons Ltd.
Rivas-Ubach A., Sardans J., Hódar J.A., Garcia-Porta J., Guenther A., Oravec M., Urban O., Peñuelas J. (2015) Similar local, but different systemic, metabolomic responses of closely related pine subspecies to folivory by caterpillars of the processionary moth. Plant Biology. : 0-0.LinkDoi: 10.1111/plb.12422
Plants respond locally and systemically to herbivore attack. Most of the research conducted on plant-herbivore relationships at element and molecular levels have focused on the elemental composition or/and certain molecular compounds or specific families of defence metabolites showing that herbivores tend to select plant individuals or species with higher nutrient concentrations and avoid those with higher levels of defence compounds. We performed stoichiometric and metabolomics, both local and systemic, analyses in two subspecies of Pinus sylvestris under attack from caterpillars of the pine processionary moth, an important pest in the Mediterranean Basin. Both pine subspecies responded locally to folivory mainly by increasing relative concentrations of terpenes and some phenolics. Systemic responses differed between pine subspecies, and most of the metabolites presented intermediate concentrations between those of the affected parts and unattacked trees. Our results support the hypothesis that foliar nutrient concentrations are not a key factor for plant selection by adult female processionary moths for oviposition, since folivory was not associated with any of the elements analysed. Phenolic compounds generally did not increase in the attacked trees, questioning the suggestion of induction of phenolics following folivory attack and the anti-feeding properties of phenolics. Herbivory attack produced a general systemic shift in pines, in both primary and secondary metabolism, which was less intense and chemically different from the local responses. Local pine responses were similar between pine subspecies, while systemic responses were more distant. © 2015 German Botanical Society and The Royal Botanical Society of the Netherlands.
Sakschewski B., von Bloh W., Boit A., Rammig A., Kattge J., Poorter L., Penuelas J., Thonicke K. (2015) Leaf and stem economics spectra drive diversity of functional plant traits in a dynamic global vegetation model. Global Change Biology. : 0-0.LinkDoi: 10.1111/gcb.12870
Functional diversity is critical for ecosystem dynamics, stability and productivity. However, dynamic global vegetation models (DGVMs) which are increasingly used to simulate ecosystem functions under global change, condense functional diversity to plant functional types (PFTs) with constant parameters. Here, we develop an individual- and trait-based version of the DGVM LPJmL (Lund-Potsdam-Jena managed Land) called LPJmL- flexible individual traits (LPJmL-FIT) with flexible individual traits) which we apply to generate plant trait maps for the Amazon basin. LPJmL-FIT incorporates empirical ranges of five traits of tropical trees extracted from the TRY global plant trait database, namely specific leaf area (SLA), leaf longevity (LL), leaf nitrogen content (Narea), the maximum carboxylation rate of Rubisco per leaf area (vcmaxarea), and wood density (WD). To scale the individual growth performance of trees, the leaf traits are linked by trade-offs based on the leaf economics spectrum, whereas wood density is linked to tree mortality. No preselection of growth strategies is taking place, because individuals with unique trait combinations are uniformly distributed at tree establishment. We validate the modeled trait distributions by empirical trait data and the modeled biomass by a remote sensing product along a climatic gradient. Including trait variability and trade-offs successfully predicts natural trait distributions and achieves a more realistic representation of functional diversity at the local to regional scale. As sites of high climatic variability, the fringes of the Amazon promote trait divergence and the coexistence of multiple tree growth strategies, while lower plant trait diversity is found in the species-rich center of the region with relatively low climatic variability. LPJmL-FIT enables to test hypotheses on the effects of functional biodiversity on ecosystem functioning and to apply the DGVM to current challenges in ecosystem management from local to global scales, that is, deforestation and climate change effects. © 2015 John Wiley & Sons Ltd.
Sardans J., Alonso R., Janssens I.A., Carnicer J., Vereseglou S., Rillig M.C., Fernández-Martínez M., Sanders T.G.M., Peñuelas J. (2015) Foliar and soil concentrations and stoichiometry of nitrogen and phosphorous across European Pinus sylvestris forests: Relationships with climate, N deposition and tree growth. Functional Ecology. : 0-0.LinkDoi: 10.1111/1365-2435.12541
This study investigated the factors underlying the variability of needle and soil elemental composition and stoichiometry and their relationships with growth in Pinus sylvestris forests throughout the species' distribution in Europe by analysing data from 2245 forest stands. Needle N concentrations and N:P ratios were positively correlated with total atmospheric N deposition, whereas needle P concentrations were negatively correlated. These relationships were especially pronounced at sites where high levels of N deposition coincided with both higher mean annual temperature and higher mean annual precipitation. Trends towards foliar P deficiency were thus more marked when high N deposition coincided with climatic conditions favourable to plant production. Atmospheric N deposition was positively correlated with soil solution NO3- , SO42- , K+, P and Ca2+ concentrations, the soil solution NO3-:P ratio, total soil N and the total soil N:Olsen P ratio and was negatively correlated with soil Olsen P concentration. Despite these nutrient imbalances, during the period studied (1990-2006), N deposition was positively related with Pinus sylvestris absolute basal diameter (BD) growth, although only accounting for the 10% of the total variance. However, neither N deposition nor needle N concentration was related with relative annual BD growth. In contrast, needle P concentration was positively related with both absolute and relative annual BD growth. These results thus indicate a tendency of European P. sylvestris forests to store N in trees and soil in response to N deposition and unveil a trend towards increased nutrient losses in run-off as a consequence of higher soil solution N concentrations. Overall, the data show increasing ecosystem nutrient imbalances with increasingly limiting roles of P and other nutrients such as K in European P. sylvestris forests, especially in the centre of their distribution where higher levels of N deposition are observed. Thus, although the data show that N deposition has had an overall positive effect on P. sylvestris growth, the effect of continuous N deposition, associated with decreasing P and K and increasing N:P in leaves and in soil, may in the future become detrimental for the growth and competitive ability of P. sylvestris trees. © 2015 British Ecological Society.
Sardans J., Janssens I.A., Alonso R., Veresoglou S.D., Rillig M.C., Sanders T.G.M., Carnicer J., Filella I., Farre-Armengol G., Penuelas J. (2015) Foliar elemental composition of European forest tree species associated with evolutionary traits and present environmental and competitive conditions. Global Ecology and Biogeography. 24: 240-255.LinkDoi: 10.1111/geb.12253
Aim: Plant elemental composition and stoichiometry are crucial for plant structure and function. We studied to what extent elemental stoichiometry in plants might be strongly related to environmental drivers and competition from coexisting species. Location: Europe. Methods: We analysed foliar N, P, K, Ca and Mg concentrations and their ratios among 50 species of European forest trees sampled in 5284 plots across Europe and their relationships with phylogeny, forest type, current climate and N deposition. Results: Phylogeny is strongly related to overall foliar elemental composition in European tree species. Species identity explained 56.7% of the overall foliar elemental composition and stoichiometry. Forest type and current climatic conditions also partially explained the differences in foliar elemental composition among species. In the same genus co-occurring species had overall higher differences in foliar elemental composition and stoichiometry than the non-co-occurring species. Main conclusions: The different foliar elemental compositions among species are related to phylogenetic distances, but they are also related to current climatic conditions, forest types, drivers of global change such as atmospheric N deposition, and to differences among co-occurring species as a probable consequence of niche specialization to reduce direct competition for the same resources. Different species have their own 'fixed' foliar elemental compositions but retain some degree of plasticity to the current climatic and competitive conditions. A wider set of elements beyond N and P better represent the biogeochemical niche and are highly sensitive to plant function. Foliar elemental composition can thus be useful for representing important aspects of plant species niches. © 2014 John Wiley & Sons Ltd.
Sardans J., Penuelas J. (2015) Potassium: A neglected nutrient in global change. Global Ecology and Biogeography. 24: 261-275.LinkDoi: 10.1111/geb.12259
Aim: Potassium (K) is the second most abundant nutrient in plant photosynthetic tissues after nitrogen (N). Thousands of physiological and metabolic studies in recent decades have established the fundamental role of K in plant function, especially in water-use efficiency and economy, and yet macroecological studies have mostly overlooked this nutrient. Methods: We have reviewed available studies on the content, stoichiometry and roles of K in the soil-plant system and in terrestrial ecosystems. We have also reviewed the impacts of global change drivers on K content, stoichiometry and roles. Conclusions: The current literature indicates that K, at a global level, is as limiting as N and phosphorus (P) for plant productivity in terrestrial ecosystems. Some degree of K limitation has been seen in up to 70% of all studied terrestrial ecosystems. However, in some areas atmospheric K deposition from human activities is greater than that from natural sources. We are far from understanding the K fluxes between the atmosphere and land, and the role of anthropogenic activities in these fluxes. The increasing aridity expected in wide areas of the world makes K more critical through its role in water-use efficiency. N deposition exerts a strong impact on the ecosystem K cycle, decreasing K availability and increasing K limitation. Plant invasive success is enhanced by higher soil K availability, especially in environments without strong abiotic stresses. The impacts of other drivers of global change, such as increasing atmospheric CO2 or changes in land use, remain to be elucidated. Current models of the responses of ecosystems and carbon storage to projected global climatic and atmospheric changes are now starting to consider N and P, but they should also consider K, mostly in arid and semi-arid ecosystems. © 2015 John Wiley & Sons Ltd.
Sardans J., Penuelas J. (2015) Trees increase their P: N ratio with size. Global Ecology and Biogeography. 24: 147-156.LinkDoi: 10.1111/geb.12231
Aim: Phosphorus (P) tends to become limiting in ageing terrestrial ecosystems, and its resorption efficiency is higher than for other elements such as nitrogen (N). We thus hypothesized that trees should store more P than those other elements such as N when tree size increases and that this process should be enhanced in slow-growing late-successional trees. Location: Catalan forests. Methods: We used data from the Catalan Forest Inventory that contains field data on the P and N contents of total aboveground, foliar and woody biomasses of the diverse mediterranean, temperate and alpine forests of Catalonia (1018 sites). We used correlation and general linear models to analyse the allometric relationships between the nutrient contents of different aboveground biomass fractions (foliar, branches and stems) and total aboveground biomass. Results: Aboveground forest P content increases proportionally more than aboveground forest N content with increasing aboveground biomass. Two mechanisms underlie this. First, woody biomass increases proportionally more than foliar biomass, with woody biomass having a higher P:N ratio than foliar biomass. Second, the P:N ratio of wood increases with tree size. These results are consistent with the generally higher foliar resorption of P than of N. Slow-growing species accumulate more total P aboveground with size than fast-growing species, mainly as a result of their large capacity to store P in wood. Main conclusions: Trees may have developed long-term adaptive mechanisms to store P in biomass, mainly in wood, thereby slowing the loss of P from ecosystems, reducing its availability for competitors and implying an increase in the P:N ratio in forest biomass with ageing. This trend to accumulate more P than N with size is more accentuated in slow-growing, large, long-lived species of late successional stages. In this way they partly counterbalance the gradual decrease of P in the soil. © 2014 John Wiley & Sons Ltd.
Sperlich D., Chang C.T., Penuelas J., Gracia C., Sabate S. (2015) Seasonal variability of foliar photosynthetic and morphological traits and drought impacts in a Mediterranean mixed forest. Tree Physiology. 35: 501-520.LinkDoi: 10.1093/treephys/tpv017
The Mediterranean region is a hot spot of climate change vulnerable to increased droughts and heat waves. Scaling carbon fluxes from leaf to landscape levels is particularly challenging under drought conditions. We aimed to improve the mechanistic understanding of the seasonal acclimation of photosynthesis and morphology in sunlit and shaded leaves of four Mediterranean trees (Quercus ilex L., Pinus halepensis Mill., Arbutus unedo L. and Quercus pubescens Willd.) under natural conditions. Vc,max and Jmax were not constant, and mesophyll conductance was not infinite, as assumed in most terrestrial biosphere models, but varied significantly between seasons, tree species and leaf position. Favourable conditions in winter led to photosynthetic recovery and growth in the evergreens. Under moderate drought, adjustments in the photo/biochemistry and stomatal/mesophyllic diffusion behaviour effectively protected the photosynthetic machineries. Severe drought, however, induced early leaf senescence mostly in A. unedo and Q. pubescens, and significantly increased leaf mass per area in Q. ilex and P. halepensis. Shaded leaves had lower photosynthetic potentials but cushioned negative effects during stress periods. Species-specificity, seasonal variations and leaf position are key factors to explain vegetation responses to abiotic stress and hold great potential to reduce uncertainties in terrestrial biosphere models especially under drought conditions. © The Author 2015. Published by Oxford University Press.
Urbina I., Sardans J., Beierkuhnlein C., Jentsch A., Backhaus S., Grant K., Kreyling J., Penuelas J. (2015) Shifts in the elemental composition of plants during a very severe drought. Environmental and Experimental Botany. 111: 63-73.LinkDoi: 10.1016/j.envexpbot.2014.10.005
Diverse plant functions (e.g., growth, storage, defense and anti-stress mechanisms) use elements disproportionally. We hypothesized that plants growing under different abiotic and biotic conditions would shift their elemental compositions in response to a very severe drought. We tested this hypothesis by investigating the changes in foliar stoichiometry and species composition from a very severe drought. We also tested the effects of previous droughts (acclimation) on this response. Different species growing in the same community responded more similarly to a very severe drought than did individual species growing in different communities. The stoichiometric shifts were thus more community-dependent than species-dependent. The results also suggested that plants grown in monoculture were less stoichiometrically plastic during the drought than plants grown in a more diverse community. Previous exposure to long-term drought treatments in the same communities did not significantly affect the stoichiometric shifts during the new drought. Differential use of resources may have been responsible for these responses. Monocultured plants, which used the same resources in similar proportions, had more difficulty avoiding direct competition when the resources became scarcer. Moreover, each species tested had a particular elemental composition in all communities and climatic treatments. The differences in foliar elemental compositions were largest between plant functional groups (shrubs and grasses) and smallest among species within the same functional group. Global principal components analyses (PCAs) identified a general tendency for all species, independently of the community in which they grew, toward lower concentrations of K, N, P, Mg and S, and to higher concentrations of C and Fe as the drought advanced. This study has demonstrated the utility of analyses of differences and shifts in plant elemental composition for understanding the processes underlying the responses of plants to changes in biotic and abiotic environmental conditions. © 2014.
Verger A., Baret F., Weiss M., Filella I., Penuelas J. (2015) GEOCLIM: A global climatology of LAI, FAPAR, and FCOVER from VEGETATION observations for 1999-2010. Remote Sensing of Environment. 166: 126-137.LinkDoi: 10.1016/j.rse.2015.05.027
Land-surface modelling would benefit significantly from improved characterisation of the seasonal variability of vegetation at a global scale. GEOCLIM, a global climatology of leaf area index (LAI), fraction of absorbed photosynthetically active radiation (FAPAR)-both essential climate variables-and fraction of vegetation cover (FCOVER), is here derived from observations from the SPOT VEGETATION programme. Interannual average values from the GEOV1 Copernicus Global Land time series of biophysical products at 1-km resolution and 10-day frequency are computed for 1999 to 2010. GEOCLIM provides the baseline characteristics of the seasonal cycle of the annual vegetation phenology for each 1-km. pixel on the globe. The associated standard deviation characterises the interannual variability. Temporal consistency and continuity is achieved by the accumulation of multi-year observations and the application of techniques for temporal smoothing and gap filling. Specific corrections are applied over cloudy tropical regions and high latitudes in the Northern Hemisphere where the low number of available observations compromises the reliability of estimates. Artefacts over evergreen broadleaf forests and areas of bare soil are corrected based on the expected limited seasonality. The GEOCLIM data set is demonstrated to be consistent, both spatially and temporally. GEOCLIM shows absolute differences lower than 0.5 compared with MODIS (GIMMS3g) climatology of LAI for more than 80% (90%) of land pixels, with higher discrepancies in tropical and boreal latitudes. ECOCLIMAP systematically produces higher LAI values. The phenological metric for the date of maximum foliar development derived from GEOCLIM is spatially consistent (correlation higher than 0.9) with those of MODIS, GIMMS3g, ECOCLIMAP and MCD12Q2 with average differences within 14. days at the global scale. © 2015 Elsevier Inc.
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