Martínez-Vilalta J., Lloret F. (2016) Drought-induced vegetation shifts in terrestrial ecosystems: The key role of regeneration dynamics. Global and Planetary Change. 144: 94-108.EnllaçDoi: 10.1016/j.gloplacha.2016.07.009
Ongoing climate change is modifying climatic conditions worldwide, with a trend towards drier conditions in most regions. Vegetation will respond to these changes, eventually adjusting to the new climate. It is unclear, however, how close different ecosystems are to climate-related tipping points and, thus, how dramatic these vegetation changes will be in the short- to mid-term, given the existence of strong stabilizing processes. Here, we review the published evidence for recent drought-induced vegetation shifts worldwide, addressing the following questions: (i) what are the necessary conditions for vegetation shifts to occur? (ii) How much evidence of drought-induced vegetation shifts do we have at present and where are they occurring? (iii) What are the main processes that favor/oppose the occurrence of shifts at different ecological scales? (iv) What are the complications in detecting and attributing drought-induced vegetation shifts? (v) What ecological factors can interact with drought to promote shifts or stability? We propose a demographic framework to classify the likely outcome of instances of drought-induced mortality, based upon the survival of adults of potential replacement species and the regeneration of both formerly dominant affected species and potential replacement species. Out of 35 selected case studies only eight were clearly consistent with the occurrence of a vegetation shift (species or biome shift), whereas three corresponded to self-replacements in which the affected, formerly dominant species was able to regenerate after suffering drought-induced mortality. The other 24 cases were classified as uncertain, either due to lack of information or, more commonly, because the initially affected and potential replacement species all showed similar levels of regeneration after the mortality event. Overall, potential vegetation transitions were consistent with more drought-resistant species replacing less resistant ones. However, almost half (44%) of the vegetation trajectories associated to the 35 case studies implied no change in the functional type of vegetation. Of those cases implying a functional type change, the most common one was a transition from tree- to shrub-dominated communities. Overall, evidence for drought-induced vegetation shifts is still limited. In this context, we stress the need for improved, long-term monitoring programs with sufficient temporal resolution. We also highlight the critical importance of regeneration in determining the outcome of drought-induced mortality events, and the crucial role of co-drivers, particularly management. Finally, we illustrate how placing vegetation shifts in a biogeographical and successional context may support progress in our understanding of the underlying processes and the ecosystem-level implications. © 2016 Elsevier B.V.
Martínez-Vilalta, J., Garcia-Forner, N. (2016) Water potential regulation, stomatal behaviour and hydraulic transport under drought: Deconstructing the iso/anisohydric concept. Plant Cell and Environment. : 0-0.EnllaçDoi: 10.1111/pce.12846
Martínez-Vilalta, J., Sala, A., Asensio, D., Galiano, L., Hoch, G., Palacio, S., Piper, F.I., Lloret, F. (2016) Dynamics of non-structural carbohydrates in terrestrial plants: A global synthesis. Ecological Monographs. 86: 495-516.EnllaçDoi: 10.1002/ecm.1231
Vayreda, J., Martinez-Vilalta, J., Gracia, M., Canadell, J.G., Retana, J. (2016) Anthropogenic-driven rapid shifts in tree distribution lead to increased dominance of broadleaf species. Global Change Biology. 22: 3984-3995.EnllaçDoi: 10.1111/gcb.13394
Vayreda, J., Martínez-Vilalta, J., Vilà-Cabrera, A. (2016) The Ecological Forest Inventory of Catalonia: A tool for functional ecology [El Inventario Ecológico y Forestal de Cataluña: una herramienta para la ecología funcional]. Ecosistemas. 25: 70-79.EnllaçDoi: 10.7818/ECOS.2016.25-3.08
Aguade D., Poyatos R., Gomez M., Oliva J., Martinez-Vilalta J. (2015) The role of defoliation and root rot pathogen infection in driving the mode of drought-related physiological decline in Scots pine (Pinus sylvestris L.). Tree Physiology. 35: 229-242.EnllaçDoi: 10.1093/treephys/tpv005
Drought-related tree die-off episodes have been observed in all vegetated continents. Despite much research effort, however, the multiple interactions between carbon starvation, hydraulic failure and biotic agents in driving tree mortality under field conditions are still not well understood. We analysed the seasonal variability of non-structural carbohydrates (NSCs) in four organs (leaves, branches, trunk and roots), the vulnerability to embolism in roots and branches, native embolism (percentage loss of hydraulic conductivity (PLC)) in branches and the presence of root rot pathogens in defoliated and non-defoliated individuals in a declining Scots pine (Pinus sylvestris L.) population in the NE Iberian Peninsula in 2012, which included a particularly dry and warm summer. No differences were observed between defoliated and non-defoliated pines in hydraulic parameters, except for a higher vulnerability to embolism at pressures below-2 MPa in roots of defoliated pines. No differences were found between defoliation classes in branch PLC. Total NSC (TNSC, soluble sugars plus starch) values decreased during drought, particularly in leaves. Defoliation reduced TNSC levels across tree organs, especially just before (June) and during (August) drought. Root rot infection by the fungal pathogen Onnia P. Karst spp. was detected but it did not appear to be associated to tree defoliation. However, Onnia infection was associated with reduced leaf-specific hydraulic conductivity and sapwood depth, and thus contributed to hydraulic impairment, especially in defoliated pines. Infection was also associated with virtually depleted root starch reserves during and after drought in defoliated pines. Moreover, defoliated and infected trees tended to show lower basal area increment. Overall, our results show the intertwined nature of physiological mechanisms leading to drought-induced mortality and the inherent difficulty of isolating their contribution under field conditions. © The Author 2015. Published by Oxford University Press. All rights reserved.
Aguade D., Poyatos R., Rosas T., Martinez-Vilalta J. (2015) Comparative drought responses of Quercus ilex L. and Pinus sylvestris L. In a montane forest undergoing a vegetation shift. Forests. 6: 2505-2529.EnllaçDoi: 10.3390/f6082505
Different functional and structural strategies to cope with water shortage exist both within and across plant communities. The current trend towards increasing drought in many regions could drive some species to their physiological limits of drought tolerance, potentially leading to mortality episodes and vegetation shifts. In this paper, we study the drought responses of Quercus ilex and Pinus sylvestris in a montane Mediterranean forest where the former species is replacing the latter in association with recent episodes of drought-induced mortality. Our aim was to compare the physiological responses to variations in soil water content (SWC) and vapor pressure deficit (VPD) of the two species when living together in a mixed stand or separately in pure stands, where the canopies of both species are completely exposed to high radiation and VPD. P. sylvestris showed typical isohydric behavior, with greater losses of stomatal conductance with declining SWC and greater reductions of stored non-structural carbohydrates during drought, consistent with carbon starvation being an important factor in the mortality of this species. On the other hand, Q. ilex trees showed a more anisohydric behavior, experiencing more negative water potentials and higher levels of xylem embolism under extreme drought, presumably putting them at higher risk of hydraulic failure. In addition, our results show relatively small changes in the physiological responses of Q. ilex in mixed vs. pure stands, suggesting that the current replacement of P. sylvestris by Q. ilex will continue. © 2015 by the authors.
Caceres M.D., Martinez-Vilalta J., Coll L., Llorens P., Casals P., Poyatos R., Pausas J.G., Brotons L. (2015) Coupling a water balance model with forest inventory data to predict drought stress: The role of forest structural changes vs. climate changes. Agricultural and Forest Meteorology. 213: 77-90.EnllaçDoi: 10.1016/j.agrformet.2015.06.012
Mechanistic water balance models can be used to predict soil moisture dynamics and drought stress in individual forest stands. Predicting current and future levels of plant drought stress is important not only at the local scale, but also at larger, landscape to regional, scales, because these are the management scales at which adaptation and mitigation strategies are implemented. To obtain reliable predictions of soil moisture and plant drought stress over large extents, water balance models need to be complemented with detailed information about the spatial variation of vegetation and soil attributes. We designed, calibrated and validated a water balance model that produces annual estimates of drought intensity and duration for all plant cohorts in a forest stand. Taking Catalonia (NE Spain) as a case study, we coupled this model with plot records from two Spanish forest inventories in which species identity, diameter and height of plant cohorts were available. Leaf area index of each plant cohort was estimated from basal area using species-specific relationships. Vertical root distribution for each species in each forest plot was estimated by determining the distribution that maximized transpiration in the model, given average climatic conditions, soil attributes and stand density. We determined recent trends (period 1980-2010) in drought stress for the main tree species in Catalonia; where forest growth and densification occurs in many areas as a result of rural abandonment and decrease of forest management. Regional increases in drought stress were detected for most tree species, although we found high variation in stress changes among individual forest plots. Moreover, predicted trends in tree drought stress were mainly due to changes in leaf area occurred between the two forest inventories rather than to climatic trends. We conclude that forest structure needs to be explicitly considered in assessments of plant drought stress patterns and trends over large geographic areas, and that forest inventories are useful sources of data provided that reasonably good estimates of soil attributes and root distribution are available. Our approach coupled with recent improvements in forest survey technologies may allow obtaining spatially continuous and precise assessments of drought stress. Further efforts are needed to calibrate drought-related demographic processes before water balance and drought stress estimates can be fully used for the accurate prediction of drought impacts. © 2015 Elsevier B.V.
Colomer-Ventura F., Martínez-Vilalta J., Zuccarini P., Escolà A., Armengot L., Castells E. (2015) Contemporary evolution of an invasive plant is associated with climate but not with herbivory. Functional Ecology. 29: 1475-1485.EnllaçDoi: 10.1111/1365-2435.12463
Divergence in plant traits and trait plasticity after invasion has been proposed as mechanisms favouring invasion success. Current hypotheses predict a rapid evolution in response to changes in the abiotic conditions in the area of introduction or to differences in the herbivore consumption pressure caused by a decrease in the enemies associated with the area of origin [e.g. evolution of increased competitive ability (EICA) hypothesis]. The importance of these factors in determining plant geographical divergence has not been yet simultaneously evaluated. Senecio pterophorus (Asteraceae) is a perennial shrub native to eastern South Africa and a recent invader in western South Africa (since ~100 years ago), Australia (>70-100 years) and Europe (>30 years). These areas differ in their summer drought stress [measured as the ratio of summer precipitation to potential evapotranspiration (P/PET)] and their interactions with herbivores. We performed a common garden experiment with S. pterophorus sampled throughout its entire known distributional area to determine (i) whether native and non-native populations diverge in their traits, as well as the plasticity of these traits in response to water availability and (ii) whether climate and herbivory play a role in the genetic differentiation across regions. Plants from the non-native regions were smaller and had a lower reproductive output than plants from the indigenous area. No geographical differences in phenotypic plasticity were found in response to water availability. Herbivory was not related to the plant geographical divergence. In contrast, our results are consistent with the role of climate as a driver for postinvasive evolution, as suggested by adaptation of plants to a drought cline in the native range, the analogous change in plant traits in independently invaded regions and the convergence of vegetative traits between non-native plants and native plants under similar drought conditions. Native and non-native populations of S. pterophorus differed in plant traits, but not in trait plasticity, in response to their local climatic conditions. Our results are contrary to the role of herbivory as a selective factor after invasion and highlight the importance of climate driving rapid evolution of exotic plants. © 2015 The Authors.
Garcia-Forner N., Adams H.D., Sevanto S., Collins A.D., Dickman L.T., Hudson P.J., Zeppel M.J., Jenkins M.W., Powers H., Martinez-Vilalta J., Mcdowell N.G. (2015) Responses of two semiarid conifer tree species to reduced precipitation and warming reveal new perspectives for stomatal regulation. Plant, Cell and Environment. : 0-0.EnllaçDoi: 10.1111/pce.12588
Relatively anisohydric species are predicted to be more predisposed to hydraulic failure than relatively isohydric species, as they operate with narrower hydraulic safety margins. We subjected co-occurring anisohydric Juniperus monosperma and isohydric Pinus edulis trees to warming, reduced precipitation, or both, and measured their gas exchange and hydraulic responses. We found that reductions in stomatal conductance and assimilation by heat and drought were more frequent during relatively moist periods, but these effects were not exacerbated in the combined heat and drought treatment. Counter to expectations, both species exhibited similar gs temporal dynamics in response to drought. Further, whereas P.edulis exhibited chronic embolism, J.monosperma showed very little embolism due to its conservative stomatal regulation and maintenance of xylem water potential above the embolism entry point. This tight stomatal control and low levels of embolism experienced by juniper refuted the notion that very low water potentials during drought are associated with loose stomatal control and with the hypothesis that anisohydric species are more prone to hydraulic failure than isohydric species. Because direct association of stomatal behaviour with embolism resistance can be misleading, we advocate consideration of stomatal behaviour relative to embolism resistance for classifying species drought response strategies. © 2015 John Wiley & Sons Ltd.
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