Aragon R., Sardans J., Penuelas J. (2014) Soil enzymes associated with carbon and nitrogen cycling in invaded and native secondary forests of northwestern Argentina. Plant and Soil. : 0-0.LinkDoi: 10.1007/s11104-014-2192-8
Background and aims Alien success has frequently been associated with changes in the concentrations of soil nutrients. We aim to investigate the effects of plant invasion on soil nutrients, potential enzyme activity and litter elemental composition and stoichiometry. Methods We compared stands of secondary forest invaded by Ligustrum lucidum and those dominated by natives, and performed litter chemical analyses on 3 native and 2 exotic tree species. Results Soils of invaded sites had 20 and 30 % increase in β-glucosidase and alkaline phosphatase activity, higher Olsen-phosphorus (P) and potassium (K) concentrations and lower nitrogen (N) concentration and N:P, N:K and ammonium:Olsen-P ratios. Invaded and non-invaded sites differed in their overall nutrient composition and enzyme activity. Natives and exotics differed in nine of the 16 litter elemental composition and stoichiometry variables analyzed. Conclusions The low N:P ratio in litter, the decrease in soil N in invaded stands and the low N concentration of exotics suggest that N is the limiting nutrient and that exotic success is related to higher N uptake and use efficiency. The higher investment in the acquisition of soil resources, higher nutrient uptake and use efficiency of limiting nutrients contribute to the success of exotics in this subtropical forest. © 2014 Springer International Publishing Switzerland.
Barbeta A., Mejia-Chang M., Ogaya R., Voltas J., Dawson T.E., Penuelas J. (2014) The combined effects of a long-term experimental drought and an extreme drought on the use of plant-water sources in a Mediterranean forest. Global Change Biology. : 0-0.LinkDoi: 10.1111/gcb.12785
Vegetation in water-limited ecosystems relies strongly on access to deep water reserves to withstand dry periods. Most of these ecosystems have shallow soils over deep groundwater reserves. Understanding the functioning and functional plasticity of species-specific root systems and the patterns of or differences in the use of water sources under more frequent or intense droughts is therefore necessary to properly predict the responses of seasonally dry ecosystems to future climate. We used stable isotopes to investigate the seasonal patterns of water uptake by a sclerophyll forest on sloped terrain with shallow soils. We assessed the effect of a long-term experimental drought (12 years) and the added impact of an extreme natural drought that produced widespread tree mortality and crown defoliation. The dominant species, Quercus ilex, Arbutus unedo and Phillyrea latifolia, all have dimorphic root systems enabling them to access different water sources in space and time. The plants extracted water mainly from the soil in the cold and wet seasons but increased their use of groundwater during the summer drought. Interestingly, the plants subjected to the long-term experimental drought shifted water uptake toward deeper (10-35 cm) soil layers during the wet season and reduced groundwater uptake in summer, indicating plasticity in the functional distribution of fine roots that dampened the effect of our experimental drought over the long term. An extreme drought in 2011, however, further reduced the contribution of deep soil layers and groundwater to transpiration, which resulted in greater crown defoliation in the drought-affected plants. This study suggests that extreme droughts aggravate moderate but persistent drier conditions (simulated by our manipulation) and may lead to the depletion of water from groundwater reservoirs and weathered bedrock, threatening the preservation of these Mediterranean ecosystems in their current structures and compositions.
Carnicer J., Coll M., Pons X., Ninyerola M., Vayreda J., Penuelas J. (2014) Large-scale recruitment limitation in Mediterranean pines: The role of Quercus ilex and forest successional advance as key regional drivers. Global Ecology and Biogeography. 23: 371-384.LinkDoi: 10.1111/geb.12111
Aim: Large-scale patterns of limitations in tree recruitment remain poorly described in the Mediterranean Basin, and this information is required to assess the impacts of global warming on forests. Here, we unveil the existence of opposite trends of recruitment limitation between the dominant genera Quercus and Pinus on a large scale and identify the key ecological drivers of these diverging trends. Location: Spain Methods: We gathered data from the Spanish National Forest inventory to assess recruitment trends for the dominant species (Pinus halepensis, Pinus pinea, Pinus pinaster, Pinus nigra, Pinus sylvestris, Pinus uncinata, Quercus suber, Quercus ilex, Quercus petraea, Quercus robur, Quercus faginea and Quercus pyrenaica). We assessed the direct and indirect drivers of recruitment by applying Bayesian structural equation modelling techniques. Results: Severe limitations in recruitment were observed across extensive areas for all Pinus species studied, with recruitment failure affecting 54-71% of the surveyed plots. In striking contrast, Quercus species expanded into 41% of the plots surveyed compared to only 10% for Pinus and had a lower local recruitment failure (29% of Quercus localities compared to 63% for Pinus species). Bayesian structural equation models highlighted the key role of the presence of Q.ilex saplings and the increase in the basal area of Q.ilex in limiting recruitment in five Pinus species. The recruitment of P.sylvestris and P.nigra showed the most negative trends and was negatively associated with the impacts of fire. Main conclusions: This study identified Q.ilex, the most widespread species in this area, as a key driver of recruitment shifts on a large scale, negatively affecting most pine species with the advance of forest succession. These results highlight that the future expansion/contraction of Q.ilex stands with ongoing climate change will be a key process indirectly controlling the demographic responses of Pinus species in the Mediterranean Basin. © 2013 John Wiley & Sons Ltd.
Carnicer J., Sardans J., Stefanescu C., Ubach A., Bartons M., Asensio D., Penuelas J. (2014) Global biodiversity, stoichiometry and ecosystem function responses to human-induced C-N-P imbalances. Journal of Plant Physiology. : 0-0.LinkDoi: 10.1016/j.jplph.2014.07.022
Global change analyses usually consider biodiversity as a global asset that needs to be preserved. Biodiversity is frequently analysed mainly as a response variable affected by diverse environmental drivers. However, recent studies highlight that gradients of biodiversity are associated with gradual changes in the distribution of key dominant functional groups characterized by distinctive traits and stoichiometry, which in turn often define the rates of ecosystem processes and nutrient cycling. Moreover, pervasive links have been reported between biodiversity, food web structure, ecosystem function and species stoichiometry. Here we review current global stoichiometric gradients and how future distributional shifts in key functional groups may in turn influence basic ecosystem functions (production, nutrient cycling, decomposition) and therefore could exert a feedback effect on stoichiometric gradients. The C-N-P stoichiometry of most primary producers (phytoplankton, algae, plants) has been linked to functional trait continua (i.e. to major axes of phenotypic variation observed in inter-specific analyses of multiple traits). In contrast, the C-N-P stoichiometry of higher-level consumers remains less precisely quantified in many taxonomic groups. We show that significant links are observed between trait continua across trophic levels. In spite of recent advances, the future reciprocal feedbacks between key functional groups, biodiversity and ecosystem functions remain largely uncertain. The reported evidence, however, highlights the key role of stoichiometric traits and suggests the need of a progressive shift towards an ecosystemic and stoichiometric perspective in global biodiversity analyses.
Curiel Yuste J., Fernandez-Gonzalez A.J., Fernandez-Lopez M., Ogaya R., Penuelas J., Lloret F. (2014) Functional diversification within bacterial lineages promotes wide functional overlapping between taxonomic groups in a Mediterranean forest soil. FEMS Microbiology Ecology. : 0-0.LinkDoi: 10.1111/1574-6941.12373
We investigated the relationship between taxonomy and functioning of soil bacterial communities in soils from a Mediterranean holm oak forest using a high-throughput DNA pyrosequencing technique. We used nonparametric tests (Mann-Whitney U-test) to evaluate the sensitivity of each single bacterial genus within the community to the fluctuations of plant physiological and environmental abiotic variables, as well as to fluctuations in soil microbial respiration. Within-lineage (phylum/class) functional similarities were evaluated by the distribution of the Mann-Whitney U-test standardized coefficients (z) obtained for all genera within a given lineage. We further defined different ecological niches and within-lineage degree of functional diversification based on multivariate analyses (principal component analyses, PCA). Our results indicate that strong within-lineage functional diversification causes extensive functional overlapping between lineages, which hinders the translation of taxonomic diversity into a meaningful functional classification of bacteria. Our results further suggest a widespread colonization of possible ecological niches as taxonomic diversity increases. While no strong functional differentiation could be drawn from the analyses at the phylum/class level, our results suggest a strong ecological niche differentiation of bacteria based mainly on the distinct response of Gram-positive and Gram-negative bacteria to fluctuations in soil moisture. We investigated the relation between taxonomy and functioning of soil bacterial communities in soils from a Mediterranean Holm-oak forest using a high throughput DNA pyrosequencing technique. © 2014 Federation of European Microbiological Societies.
Curiel Yuste J., Fernandez-Gonzalez A.J., Fernandez-Lopez M., Ogaya R., Penuelas J., Sardans J., Lloret F. (2014) Strong functional stability of soil microbial communities under semiarid Mediterranean conditions and subjected to long-term shifts in baseline precipitation. Soil Biology and Biochemistry. 69: 223-233.LinkDoi: 10.1016/j.soilbio.2013.10.045
We investigated the effect of soil microclimate on the structure and functioning of soil microbial communities in a Mediterranean Holm-oak forest subjected to 10 years of partial rain exclusion manipulations, simulating average drought conditions expected in Mediterranean areas for the following decades. We applied a high throughput DNA pyrosequencing technique coupled to parallel measurements of microbial respiration (RH) and temperature sensitivity of microbial respiration (Q10). Some consistent changes in the structure of bacterial communities suggest a slow process of community shifts parallel to the trend towards oligotrophy in response to long-term droughts. However, the structure of bacterial communities was mainly determined by short-term environmental fluctuations associated with sampling date (winter, spring and summer) rather than long-term (10 years) shifts in baseline precipitation. Moreover, long-term drought did not exert any chronic effect on the functioning of soil microbial communities (RH and Q10), emphasizing the functional stability of these communities to this long-term but mild shifts in water availability. We hypothesize that the particular conditions of the Mediterranean climate with strong seasonal shifts in both temperature and soil water availability but also characterized by very extreme environmental conditions during summer, was acting as a strong force in community assembling, selecting phenotypes adapted to the semiarid conditions characterizing Mediterranean ecosystems. Relations of climate with the phylogenetic structure and overall diversity of the communities as well as the distribution of the individual responses of different lineages (genera) to climate confirmed our hypotheses, evidencing communities dominated by thermotolerant and drought-tolerant phenotypes. © 2013 Elsevier Ltd.
Domenech M., Komac B., Penuelas J., Conesa J.A. (2014) Site-specific factors influence the richness and phenology of snowbed plants in the Pyrenees. Plant Biosystems. : 0-0.LinkDoi: 10.1080/11263504.2014.990941
Although the timing of snowmelt and growth temperatures appear to be the main factors that influence the species richness and phenology of snowbed plants, site-specific characteristics may also play a role in modifying the effects of the timing of snowmelt and temperature. In this study, the effects of site-specific factors (microtopography and snow origin) on species richness and plant phenology were evaluated in 72 plots in two snowbeds in the Andorran Pyrenees. Snowmelt patterns influenced the spatial distribution of species richness and abundance. Site-specific factors had significant effects on the responses of species (shortening or lengthening the duration of the phenophase) and on the extent to which the timing of snowmelt influenced leaf expansion and flowering. Notably, the highest rates of leaf expansion occurred on late snowmelt isoclines, where, nevertheless, the time taken to reach peak flowering was significantly longer than on the early snowmelt isoclines. The results of this study highlight the fact that, in addition to the effects of interannual variability in climate, site-specific factors have a significant effect on the phenology and reproductive success of the commonest plants in the snowbed communities of the Pyrenees.
Farre-Armengol G., Filella I., Llusia J., Niinemets U., Penuelas J. (2014) Changes in floral bouquets from compound-specific responses to increasing temperatures. Global Change Biology. : 0-0.LinkDoi: 10.1111/gcb.12628
We addressed the potential effects of changes in ambient temperature on the profiles of volatile emissions from flowers and tested whether warming could induce significant quantitative and qualitative changes in floral emissions, which would potentially interfere with plant-pollinator chemical communication. We measured the temperature responses of floral emissions of various common species of Mediterranean plants using dynamic headspace sampling and used GC-MS to identify and quantify the emitted terpenes. Floral emissions increased with temperature to an optimum and thereafter decreased. The responses to temperature modeled here predicted increases in the rates of floral terpene emission of 0.03-1.4-fold, depending on the species, in response to an increase of 1 °C in the mean global ambient temperature. Under the warmest projections that predict a maximum increase of 5 °C in the mean temperature of Mediterranean climates in the Northern Hemisphere by the end of the century, our models predicted increases in the rates of floral terpene emissions of 0.34-9.1-fold, depending on the species. The species with the lowest emission rates had the highest relative increases in floral terpene emissions with temperature increases of 1-5 °C. The response of floral emissions to temperature differed among species and among different compounds within the species. Warming not only increased the rates of total emissions, but also changed the ratios among compounds that constituted the floral scents, i.e. increased the signal for pollinators, but also importantly altered the signal fidelity and probability of identification by pollinators, especially for specialists with a strong reliance on species-specific floral blends. © 2014 John Wiley & Sons Ltd.
Fernandez-Martinez M., Vicca S., Janssens I.A., Luyssaert S., Campioli M., Sardans J., Estiarte M., Penuelas J. (2014) Spatial variability and controls over biomass stocks, carbon fluxes, and resource-use efficiencies across forest ecosystems. Trees - Structure and Function. 28: 597-611.LinkDoi: 10.1007/s00468-013-0975-9
Key message: Stand age, water availability, and the length of the warm period are the most influencing controls of forest structure, functioning, and efficiency. We aimed to discern the distribution and controls of plant biomass, carbon fluxes, and resource-use efficiencies of forest ecosystems ranging from boreal to tropical forests. We analysed a global forest database containing estimates of stand biomass and carbon fluxes (400 and 111 sites, respectively) from which we calculated resource-use efficiencies (biomass production, carbon sequestration, light, and water-use efficiencies). We used the WorldClim climatic database and remote-sensing data derived from the Moderate Resolution Imaging Spectroradiometer to analyse climatic controls of ecosystem functioning. The influences of forest type, stand age, management, and nitrogen deposition were also explored. Tropical forests exhibited the largest gross carbon fluxes (photosynthesis and ecosystem respiration), but rather low net ecosystem production, which peaks in temperate forests. Stand age, water availability, and length of the warm period were the main factors controlling forest structure (biomass) and functionality (carbon fluxes and efficiencies). The interaction between temperature and precipitation was the main climatic driver of gross primary production and ecosystem respiration. The mean resource-use efficiency varied little among biomes. The spatial variability of biomass stocks and their distribution among ecosystem compartments were strongly correlated with the variability in carbon fluxes, and both were strongly controlled by climate (water availability, temperature) and stand characteristics (age, type of leaf). Gross primary production and ecosystem respiration were strongly correlated with mean annual temperature and precipitation only when precipitation and temperature were not limiting factors. Finally, our results suggest a global convergence in mean resource-use efficiencies. © 2013 Springer-Verlag Berlin Heidelberg.
Fernandez-Martinez M., Vicca S., Janssens I.A., Sardans J., Luyssaert S., Campioli M., Chapin Iii F.S., Ciais P., Malhi Y., Obersteiner M., Papale D., Piao S.L., Reichstein M., Roda F., Penuelas J. (2014) Nutrient availability as the key regulator of global forest carbon balance. Nature Climate Change. 4: 471-476.LinkDoi: 10.1038/nclimate2177
Forests strongly affect climate through the exchange of large amounts of atmospheric CO 2 (ref.). The main drivers of spatial variability in net ecosystem production (NEP) on a global scale are, however, poorly known. As increasing nutrient availability increases the production of biomass per unit of photosynthesis and reduces heterotrophic respiration in forests, we expected nutrients to determine carbon sequestration in forests. Our synthesis study of 92 forests in different climate zones revealed that nutrient availability indeed plays a crucial role in determining NEP and ecosystem carbon-use efficiency (CUEe; that is, the ratio of NEP to gross primary production (GPP)). Forests with high GPP exhibited high NEP only in nutrient-rich forests (CUEe = 33 ± 4%; mean ± s.e.m.). In nutrient-poor forests, a much larger proportion of GPP was released through ecosystem respiration, resulting in lower CUEe (6 ± 4%). Our finding that nutrient availability exerts a stronger control on NEP than on carbon input (GPP) conflicts with assumptions of nearly all global coupled carbon cycle-climate models, which assume that carbon inputs through photosynthesis drive biomass production and carbon sequestration. An improved global understanding of nutrient availability would therefore greatly improve carbon cycle modelling and should become a critical focus for future research. © 2014 Macmillan Publishers Limited.
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