Barba J., Curiel Yuste J., Martinez-Vilalta J., Lloret F. (2013) Drought-induced tree species replacement is reflected in the spatial variability of soil respiration in a mixed Mediterranean forest. Forest Ecology and Management. 306: 79-87.EnllaçDoi: 10.1016/j.foreco.2013.06.025
As episodes of drought-induced forest mortality are being increasingly reported worldwide and may become more frequent in the future as a result of climate change, it is essential to characterize their functional implications in terms of ecosystem carbon and water fluxes. We investigated the spatial variability of soil respiration in a mixed Mediterranean forest located on rugged terrain, where Scots pine (Pinus sylvestris) is affected by drought-induced dieback and appears to have been replaced by Holm oak (Quercus ilex) as the dominant tree species. Soil respiration was measured in spring 2010 on two plots (16.2×16.2m) using a static closed chamber method (soda lime technique) and a systematic sampling (1.8-m grid) including 100 points per plot. Biotic and abiotic variables, such as soil moisture, soil temperature, soil organic matter content, stoniness, pH, fine root C:N ratio and biomass, tree basal area and tree species and health condition of nearest neighbouring tree were also recorded. Our results showed that the spatial variability of soil respiration under optimal environmental conditions (spring) was high and showed no spatial autocorrelation on the scale studied (1-18m). A mixed-effects model applied to explain the spatial variability of soil respiration indicated that only the variables related to forest structure (i.e., health condition and basal area) explained any of the observed variability of soil respiration (R2=0.45). Our model revealed that soil respiration was highest in soils close to dead pines and under Holm oak trees, suggesting that tree mortality and species replacement of pine trees by Holm oak may lead to higher soil respiration fluxes. The direct effect of tree mortality on soil respiration may be a transitory response caused by fine root mortality. Furthermore, the fact that tree species replacement as a result of drought-induced die-off is accompanied by concomitant changes in soil respiration has important implications for soil and ecosystem carbon balance. © 2013 Elsevier B.V.
Doblas-Miranda E., Rovira P., Brotons L., Martinez-Vilalta J., Retana J., Pla M., Vayreda J. (2013) Soil carbon stocks and their variability across the forests, shrublands and grasslands of peninsular Spain. Biogeosciences. 10: 8353-8361.EnllaçDoi: 10.5194/bg-10-8353-2013
Accurate estimates of C stocks and fluxes of soil organic carbon (SOC) are needed to assess the impact of climate and land use change on soil C uptake and soil C emissions to the atmosphere. Here, we present an assessment of SOC stocks in forests, shrublands and grasslands of peninsular Spain based on field measurements in more than 900 soil profiles. SOC to a depth of 1 m was modelled as a function of vegetation cover, mean annual temperature, total annual precipitation, elevation and the interaction between temperature and elevation, while latitude and longitude were used to model the correlation structure of the errors. The resulting statistical model was used to estimate SOC in the ∼8 million pixels of the Spanish Forest Map (29.3 × 106 ha). We present what we believe is the most reliable estimation of current SOC in forests, shrublands and grasslands of peninsular Spain thus far, based on the use of spatial modelling, the high number of profiles and the validity and refinement of the data layers employed. Mean concentration of SOC was 8.7 kg m-2, ranging from 2.3 kg m-2 in dry Mediterranean areas to 20.4 kg m -2 in wetter northern locations. This value corresponds to a total stock of 2.544 Tg SOC, which is four times the amount of C estimated to be stored in the biomass of Spanish forests. Climate and vegetation cover were the main variables influencing SOC, with important ecological implications for peninsular Spanish ecosystems in the face of global change. The fact that SOC was positively related to annual precipitation and negatively related to mean annual temperature suggests that future climate change predictions of increased temperature and reduced precipitation may strongly reduce the potential of Spanish soils as C sinks. However, this may be mediated by changes in vegetation cover (e.g. by favouring the development of forests associated to higher SOC values) and exacerbated by perturbations such as fire. The estimations presented here provide a baseline to estimate future changes in soil C stocks and to assess their vulnerability to key global change drivers, and should inform future actions aimed at the conservation and management of C stocks. © 2013 Author(s).
Galiano L., Martinez-Vilalta J., Eugenio M., Granzow-de la Cerda I., Lloret F. (2013) Seedling emergence and growth of Quercus spp. following severe drought effects on a Pinus sylvestris canopy. Journal of Vegetation Science. 24: 580-588.EnllaçDoi: 10.1111/j.1654-1103.2012.01485.x
Questions: We addressed the following questions: (1) did defoliation and die-off of the dominant Pinus sylvestris, induced by an extreme drought episode, favour emergence of other tree species; (2) did the defoliated canopies of P. sylvestris resulting from drought promote radial growth among other pre-existing tree species seedlings under them? Location: P. sylvestris forest in Central Pyrenees (NE Spain) affected by a severe drought in 2004-2005. Methods: Despite increased focus on climate-related forest die-off, studies of the effects on regeneration processes following extreme drought remain scarce. We analysed whether an episode of drought-induced mortality on the dominant P. sylvestris L. may act as a driver of vegetation shift. Seedlings of Quercus humilis Mill. and Q. ilex L. from 27 plots were sampled under P. sylvestris canopies with
Lloret F., Martinez-Vilalta J., Serra-Diaz J.M., Ninyerola M. (2013) Relationship between projected changes in future climatic suitability and demographic and functional traits of forest tree species in Spain. Climatic Change. 120: 449-462.EnllaçDoi: 10.1007/s10584-013-0820-6
The response of plant species to future climate conditions is probably dependent on their ecological characteristics, including climatic niche, demographic rates and functional traits. Using forest inventory data from 27 dominant woody species in Spanish forests, we explore the relationships between species characteristics and projected changes in their average climatic suitability (occurrence of suitable climatic conditions for a species in a given territory) obtained by empirical niche-based models, under a business-as-usual climate change scenario (A1, HadCM3, 2001-2100). We hypothesize that most species will suffer a decline in climatic suitability, with a less severe for species (i) currently living in more arid climates or exhibiting a broader current climatic niche; (ii) with higher current growth rates; (iii) with functional traits related to resistance to water deficits. The analysis confirm our hypothesis since apart from a few Mediterranean species, most species decrease their climatic suitability in the region under future climate, characterized by increased aridity. Also, species living in warmer locations or under a wider range of climatic conditions tend to experience less decrease in climatic suitability. As hypothesized, a positive relationship was detected between current relative growth rates and increase in future climatic suitability. Nevertheless, current tree mortality did not correlate with changes in future climatic suitability. In contrast with our hypothesis, functional traits did not show a clear relationship with changes in climate suitability; instead species often presented idiosyncratic responses that, in some cases, could reflect past management. These results suggest that the extrapolation of species performance to future climatic scenarios based on current patterns of dominance is constrained by factors other than species autoecology, particularly human activity. © 2013 Springer Science+Business Media Dordrecht.
Mcdowell N.G., Fisher R.A., Xu C., Domec J.C., Holtta T., Mackay D.S., Sperry J.S., Boutz A., Dickman L., Gehres N., Limousin J.M., Macalady A., Martinez-Vilalta J., Mencuccini M., Plaut J.A., Ogee J., Pangle R.E., Rasse D.P., Ryan M.G., Sevanto S., Waring R.H., Williams A.P., Yepez E.A., Pockman W.T. (2013) Evaluating theories of drought-induced vegetation mortality using a multimodel-experiment framework. New Phytologist. 200: 304-321.EnllaçDoi: 10.1111/nph.12465
Summary: Model-data comparisons of plant physiological processes provide an understanding of mechanisms underlying vegetation responses to climate. We simulated the physiology of a piñon pine-juniper woodland (Pinus edulis-Juniperus monosperma) that experienced mortality during a 5 yr precipitation-reduction experiment, allowing a framework with which to examine our knowledge of drought-induced tree mortality. We used six models designed for scales ranging from individual plants to a global level, all containing state-of-the-art representations of the internal hydraulic and carbohydrate dynamics of woody plants. Despite the large range of model structures, tuning, and parameterization employed, all simulations predicted hydraulic failure and carbon starvation processes co-occurring in dying trees of both species, with the time spent with severe hydraulic failure and carbon starvation, rather than absolute thresholds per se, being a better predictor of impending mortality. Model and empirical data suggest that limited carbon and water exchanges at stomatal, phloem, and below-ground interfaces were associated with mortality of both species. The model-data comparison suggests that the introduction of a mechanistic process into physiology-based models provides equal or improved predictive power over traditional process-model or empirical thresholds. Both biophysical and empirical modeling approaches are useful in understanding processes, particularly when the models fail, because they reveal mechanisms that are likely to underlie mortality. We suggest that for some ecosystems, integration of mechanistic pathogen models into current vegetation models, and evaluation against observations, could result in a breakthrough capability to simulate vegetation dynamics. © 2013 New Phytologist Trust.
Poyatos R., Aguade D., Galiano L., Mencuccini M., Martinez-Vilalta J. (2013) Drought-induced defoliation and long periods of near-zero gas exchange play a key role in accentuating metabolic decline of Scots pine. New Phytologist. 200: 388-401.EnllaçDoi: 10.1111/nph.12278
Summary: Drought-induced defoliation has recently been associated with the depletion of carbon reserves and increased mortality risk in Scots pine (Pinus sylvestris). We hypothesize that defoliated individuals are more sensitive to drought, implying that potentially higher gas exchange (per unit of leaf area) during wet periods may not compensate for their reduced photosynthetic area. We measured sap flow, needle water potentials and whole-tree hydraulic conductance to analyse the drought responses of co-occurring defoliated and nondefoliated Scots pines in northeast Spain during typical (2010) and extreme (2011) drought conditions. Defoliated Scots pines showed higher sap flow per unit leaf area during spring, but were more sensitive to summer drought, relative to nondefoliated pines. This pattern was associated with a steeper decline in soil-to-leaf hydraulic conductance with drought and an enhanced sensitivity of canopy conductance to soil water availability. Near-homeostasis in midday water potentials was observed across years and defoliation classes, with minimum values of -2.5 MPa. Enhanced sensitivity to drought and prolonged periods of near-zero gas exchange were consistent with low levels of carbohydrate reserves in defoliated trees. Our results support the critical links between defoliation, water and carbon availability, and their key roles in determining tree survival and recovery under drought. © 2013 New Phytologist Trust.
Rosas T., Galiano L., Ogaya R., Penuelas J., Martinez-Vilalta J. (2013) Dynamics of non-structural carbohydrates in three mediterranean woody species following long-term experimental drought. Frontiers in Plant Science. 4: 0-0.EnllaçDoi: 10.3389/fpls.2013.00400
Stored non-structural carbohydrates (NSC) have been proposed as a key determinant of drought resistance in plants. However, the evidence for this role is controversial, as it comes mostly from observational, short-term studies. Here, we take advantage of a long-term experimental throughfall reduction to elucidate the response of NSC to increased drought 14 years after the beginning of the treatment in three Mediterranean resprouter trees (Quercus ilex L., Arbutus unedo L. and Phillyrea latifclia L.). In addition, we selected 20. Q. ilex individuals outside the experimental plots to directly assess the relationship between defoliation and NSC at the individual level. We measured the seasonal course of NSC concentrations in leaves, branches and lignotuber in late winter, late spring, summer, and autumn 2012. Total concentrations of NSC were highest in the lignotuber for all species. In the long-term drought experiment we found significant depletion in concentrations of total NSC in treatment plots only in the lignotuber of A. unedo. At the same time, A. unedo was the only species showing a significant reduction in BAI under the drought treatment during the 14 years of the experiment. By contrast, Q. ilex just reduced stem growth only during the first 4 years of treatment and P latifclia remained unaffected over the whole study period. However, we found a clear association between the concentrations of NSC and defoliation in Q. ilex individuals sampled outside the experimental plots, with lower total concentrations of NSC and lower proportion of starch in defoliated individuals. Taken together, our results suggest that stabilizing processes, probably at the stand level, may have been operating in the long-term to mitigate any impact of drought on NSC levels, and highlight the necessity to incorporate long-term experimental studies of plant responses to drought. © 2013 Rosas, Galiano, Ogaya, Peñuelas and Martínez-Vilalta.
Vayreda J., Gracia M., Martinez-Vilalta J., Retana J. (2013) Patterns and drivers of regeneration of tree species in forests of peninsular Spain. Journal of Biogeography. 40: 1252-1265.EnllaçDoi: 10.1111/jbi.12105
Aim: Our study aimed to identify and explore the main factors that influence tree recruitment of multiple species at a regional scale across peninsular Spain, an understanding of which is essential for predicting future forest species composition in the face of ongoing environmental change. The study focused on the dynamics of the key transition phase from saplings to adult trees. Location: The forests of peninsular Spain. Methods: We used the extensive network of plots sampled in two consecutive Spanish national forest inventories (> 30,000 plots) to identify the factors that determine regeneration patterns of the 10 most abundant forest species of Spain at relatively large temporal (c. 10 years) and spatial scales (across Spain): five coniferous species of Pinus (pines) and five broadleaved species of the genera Fagus and Quercus. We fitted separate generalized linear models for the pine species and the broadleaved species to assess the response of sapling abundance and ingrowth rate to the spatial variability of climate (temperature, water availability and recent warming), forest structure (tree density, understorey and overstorey canopy cover, and basal area change) and disturbances (previous forest logging, wildfires and grazing). Results: Mean sapling abundance was four times higher for broadleaved species than for pines, while mean annual ingrowth was twice as high. Sapling abundance and ingrowth rate were mainly determined by stand structure, both in pines and broadleaved trees. The direct effects of disturbances and climate were comparatively smaller, and there was no detectable effect of recent warming. Main conclusions: The higher values of ingrowth rate of broadleaved species can be explained by their ability to maintain a higher sapling bank due to their greater shade tolerance. This differential response of pines and broadleaved species to canopy closure suggests a probable increase in broadleaved species at the expense of pines. This transition could occur earlier in stands with faster canopy closure dynamics. Spatially explicit, mixed-species demographic models incorporating both the ingrowth and the tree mortality components are needed for predicting the composition of future forests. © 2013 Blackwell Publishing Ltd.
Vilà-Cabrera A., Martínez-Vilalta J., Galiano L., Retana J. (2013) Patterns of Forest Decline and Regeneration Across Scots Pine Populations. Ecosystems. 16: 323-335.EnllaçDoi: 10.1007/s10021-012-9615-2
To predict future changes in forest ecosystems, it is crucial to understand the complex processes involved in decline of tree species populations and to evaluate the implications for potential vegetation shifts. Here, we study patterns of decline (canopy defoliation and mortality of adults) of four Scots pine populations at the southern edge of its distribution and characterized by different combinations of climate dryness and intensity of past management. General linear and structural equation modeling were used to assess how biotic, abiotic, and management components interacted to explain the spatial variability of Scots pine decline across and within populations. Regeneration patterns of Scots pine and co-occurring oak species were analyzed to assess potential vegetation shifts. Decline trends were related to climatic dryness at the regional scale, but, ultimately, within-population forest structure, local site conditions, and past human legacies could be the main underlying drivers of Scots pine decline. Overall, Scots pine regeneration was negatively related to decline both within and between populations, whereas oak species responded to decline idiosyncratically across populations. Taken together, our results suggest that (1) patterns of decline are the result of processes acting at the plot level that modulate forest responses to local environmental stress and (2) decline of adult Scots pine trees seems not to be compensated by self-recruitment so that the future dynamics of these forest ecosystems are uncertain. © 2012 Springer Science+Business Media New York.
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