Drought-induced Mortality Research Articles

Synchrotron X-ray microtomography of xylem embolism in Sequoia sempervirens saplings during cycles of drought and recovery.

The formation of emboli in xylem conduits can dramatically reduce hydraulic capacity and represents one of the principal mechanisms of drought-induced mortality in woody plants. However, our understanding of embolism formation and repair is constrained by a lack of tools to directly and nondestructively measure these processes at high spatial resolution. Using synchrotron-based microcomputed tomography (microCT), we examined embolism in the xylem of coast redwood (Sequoia sempervirens) saplings that were subjected to cycles of drought and rewatering. Embolism formation was observed occurring by three different mechanisms: as tracheids embolizing in wide tangential bands; as isolated tracheids in seemingly random events; and as functional groups connected to photosynthetic organs. Upon rewatering, stem water potential recovered to predrought stress levels within 24h; however, no evidence of embolism repair was observed even after a further 2wk under well-watered conditions. The results indicate that intertracheid air seeding is the primary mechanism by which embolism spreads in the xylem of S.sempervirens, but also show that a small number of tracheids initially become gas-filled via another mechanism. The inability of S.sempervirens saplings to reverse drought-induced embolism is likely to have important ecological impacts on this species.

Genetic and environmental characterization of <em>Abies alba</em> Mill. populations at its western rear edge

The genetic structure of ten Abies alba populations at the western rear edge in the Spanish Pyrenees was characterized and compared with two German populations, looking for the role of climatic factors in the fir decline. Growth, defoliation, aridity and cumulative summer water deficit summer were also characterized. Spanish populations show a lower genetic diversity and a high genetic differentiation than German ones, mainly established by an East (higher diversity)-West (lower diversity) gradient. The three defoliated populations are the western ones, with higher summer aridity. Contrastingly, the southern population (Guara) shows climatic variables close to western sites but without defoliation and with a higher genetic diversity, indicating a possible adaptation to the sub-Mediterranean conditions. Silver fir in the Spanish Pyrenees constitutes a “stable” rear edge because of their isolation, small sized and small genetic diversity. Western Pyrenean sites subjected to dryer conditions and presenting lower genetic diversity are prone to drought-induced mortality in the context of global warming.

Carbohydrate dynamics and mortality in a piñon-juniper woodland under three future precipitation scenarios.

Drought-induced forest mortality is an increasing global problem with wide-ranging consequences, yet mortality mechanisms remain poorly understood. Depletion of non-structural carbohydrate (NSC) stores has been implicated as an important mechanism in drought-induced mortality, but experimental field tests are rare. We used an ecosystem-scale precipitation manipulation experiment to evaluate leaf and twig NSC dynamics of two co-occurring conifers that differ in patterns of stomatal regulation of water loss and recent mortality: the relatively desiccation-avoiding piñon pine (Pinus edulis) and the relatively desiccation-tolerant one-seed juniper (Juniperus monosperma). Piñon pine experienced 72% mortality after 13-25 months of experimental drought and juniper experienced 20% mortality after 32-47 months. Juniper maintained three times more NSC in the foliage than twigs, and converted NSC to glucose and fructose under drought, consistent with osmoregulation requirements to maintain higher stomatal conductance during drought than piñon. Despite these species differences, experimental drought caused decreased leaf starch content in dying trees of both species (P < 0.001). Average dry-season leaf starch content was also a good predictor of drought-survival time for both species (R(2) = 0.93). These results, along with observations of drought-induced reductions to photosynthesis and growth, support carbon limitation as an important process during mortality of these two conifer species.

Integrating ecophysiology and forest landscape models to improve projections of drought effects under climate change

Fundamental drivers of ecosystem processes such as temperature and precipitation are rapidly changing and creating novel environmental conditions. Forest landscape models (FLM) are used by managers and policy-makers to make projections of future ecosystem dynamics under alternative management or policy options, but the links between the fundamental drivers and projected responses are weak and indirect, limiting their reliability for projecting the impacts of climate change. We developed and tested a relatively mechanistic method to simulate the effects of changing precipitation on species competition within the LANDIS-II FLM. Using data from a field precipitation manipulation experiment in a piñon pine (Pinus edulis) and juniper (Juniperus monosperma) ecosystem in New Mexico (USA), we calibrated our model to measurements from ambient control plots and tested predictions under the drought and irrigation treatments against empirical measurements. The model successfully predicted behavior of physiological variables under the treatments. Discrepancies between model output and empirical data occurred when the monthly time step of the model failed to capture the short-term dynamics of the ecosystem as recorded by instantaneous field measurements. We applied the model to heuristically assess the effect of alternative climate scenarios on the piñon-juniper ecosystem and found that warmer and drier climate reduced productivity and increased the risk of drought-induced mortality, especially for piñon. We concluded that the direct links between fundamental drivers and growth rates in our model hold great promise to improve our understanding of ecosystem processes under climate change and improve management decisions because of its greater reliance on first principles.

Declining hydraulic performances and low carbon investments in tree rings predate Scots pine drought-induced mortality

The retrospective analysis of wood anatomical features evidences how a long-term deterioration of hydraulic performance and carbon use portend drought-induced mortality in Scots pine. Widespread episodes of drought-induced tree mortality are predicted to become more frequent as climate becomes warmer and drier. Nevertheless, growth trends and their links to changes in wood anatomy before tree dies are still poorly understood. Wood anatomical features provide valuable information that can be extracted to infer the mechanisms leading to tree death. In this study, we characterize drought-induced mortality affecting two Scots pine (Pinus sylvestris) sites (Prades and Arcalís) located in the North Eastern Iberian Peninsula. Co-occurring now-dead and living Scots pine trees were sampled and their wood anatomical features were measured and compared. We aimed to detect differences in anatomical features between living and dead trees, and to infer past physiological performances that might have determined their subsequent death or survival. Now-dead trees showed lower tracheid and resin duct production, and smaller radial lumen diameters than co-occurring living trees. At the more xeric Prades site, these anatomical differences were larger and chronic, i.e. were observed over the three studied decades, whilst they were less pronounced at the other, more mesic Arcalís site, where tree mortality episodes were more recent. This indicates that dead trees’ hydraulic conductivity was severely affected and that carbon investment in xylem formation and resin duct production was constrained prior to tree death. Our findings show that both hydraulic deterioration and low carbon allocation to xylem formation were associated to drought-induced mortality in Scots pine. Nevertheless, the temporal dynamics of these processes differed between populations as a function of site climatic conditions.

Functional response of U.S. grasslands to the early 21st-century drought.

Grasslands across the United States play a key role in regional livelihood and national food security. Yet, it is still unclear how this important resource will respond to the prolonged warm droughts and more intense rainfall events predicted with climate change. The early 21st-century drought in the southwestern United States resulted in hydroclimatic conditions that are similar to those expected with future climate change. We investigated the impact of the early 21st-century drought on aboveground net primary production (ANPP) of six desert and plains grasslands dominated by C4 (warm season) grasses in terms of significant deviations between observed and expected ANPP. In desert grasslands, drought-induced grass mortality led to shifts in the functional response to annual total precipitation (P(T)), and in some cases, new species assemblages occurred that included invasive species. In contrast, the ANPP in plains grasslands exhibited a strong linear function of the current-year P(T) and the previous-year ANPP, despite prolonged warm drought. We used these results to disentangle the impacts of interannual total precipitation, intra-annual precipitation patterns, and grassland abundance on ANPP, and thus generalize the functional response of C4 grasslands to predicted climate change. This will allow managers to plan for predictable shifts in resources associated with climate change related to fire risk, loss of forage, and ecosystem services.

Pinyon Pine Mortality Alters Communities of Ground-Dwelling Arthropods

We documented the effect of drought-induced mortality of pinyon pine (Pinus edulis Engelm.) on communities of ground-dwelling arthropods. Tree mortality alters microhabitats utilized by ground-dwelling arthropods by increasing solar radiation, dead woody debris, and understory vegetation. Our major objectives were to determine (1) whether there were changes in community composition, species richness, and abundance of ground-dwelling arthropods associated with pinyon mortality and (2) whether specific habitat characteristics and microhabitats accounted for these changes. We predicted shifts in community composition and increases in arthropod diversity and abundance due to the presumed increased complexity of microhabitats from both standing dead and fallen dead trees. We found significant differences in arthropod community composition between high and low pinyon mortality environments, despite no differences in arthropod abundance or richness. Overall, 22% (51 taxa) of the arthropod community were identified as being indicators of either high or low mortality. Our study corroborates other research indicating that arthropods are responsive to even moderate disturbance events leading to changes in the environment. These arthropod responses can be explained in part due to the increase in woody debris and reduced canopy cover created by tree mortality.

Plant diversity and drought: The role of deep roots

Extreme temperatures and droughts in the wake of climate change potentially threaten plant diversity. A strategy that plants use to improve survival during seasonal drought is to establish deep roots, aptly named tap roots for their ability to tap into groundwater. Little is known, however, about the role of deep roots in maintaining plant diversity. Here, we extend an established model of plants canopies by Iwasa et al. (1985), in which plants of different heights compete for light, to allow strategic investments in an optional tap root. We investigate how emerging diversity varies with water table depth, soil water gradient and drought-induced mortality rate. Having a tap root enables plants to reach deep water, thus reducing mortality, but also carries a construction cost, thus inducing a tradeoff. We find (1) that tap roots maintain plant diversity under increasing drought mortality, (2) that tap roots evolve when ground water is accessible at low to intermediate depths, (3) no viable strategies at high drought mortality and deep water table, and (4) Red Queen evolutionary dynamics in mixed communities with and without tap root.

Applicability of Predictive Models of Drought-Induced Tree Mortality between the Midwest and Northeast United States

Regression models developed in the upper Midwest (United States) to predict drought-induced tree mortality from measures of drought (Palmer Drought Severity Index) were tested in the northeastern United States and found inadequate. The most likely cause of this result is that long drought events were rare in the Northeast during the period when inventory data were available. Therefore, new predictive models of drought mortality for the Northeast were built using USDA Forest Service inventory data and national climate data from 1969 to 2007. The Standardized Precipitation Index was better correlated with tree mortality in the Northeast than the Palmer Drought Severity Index, and new models were estimated. The reliability of the northeast models varied considerably by drought-sensitivity class, with the model for drought-intolerant species being particularly suspect. I argue that the Midwest models may nevertheless have some value in the Northeast because my tests were unable to cover the range of drought conditions under which the models were developed, there is no obvious reason why the same species should respond differently in a very similar ecological province, and some northeast models are very weak.

Recent advances in tree hydraulics highlight the ecological significance of the hydraulic safety margin

Drought-induced cavitation resistance varies considerably between tree species and forest ecosystems (Maherali et al., 2004; Delzon et al., 2010) and is closely linked to survival under severe drought in both conifers (Brodribb & Cochard, 2009; Brodribb et al., 2010) and angiosperms (Kursar et al., 2009; Anderegg et al., 2012; Barigah et al., 2013; Urli et al., 2013). Choat et al. (2012) recently reported that most trees operate very close to their threshold of cavitation, leaving them potentially vulnerable to drought-induced mortality in a warmer/drier world (Engelbrecht, 2012). Indeed, species growing in dry environments are more resistant to droughtinduced cavitation (more negative water potential at 50% cavitation, P50) but experience a more negative minimumwater potential (Pmin) than those growing in wet environments. The so-called hydraulic safety margin, the difference between the level of water stress experienced by a species in the field (Pmin) and the level of water stress leading to hydraulic failure, is, therefore, remarkably narrow, whatever the forest species and biome considered (Choat et al., 2012). This pattern provides clues to the global droughtinduced mortality currently observed, even in very wet environments, such as tropical forests (Allen et al., 2010). Klein et al. (2014) play down the functional significance of the hydraulic safety margin in the vulnerability of forests to drought, pointing out the important role played by additional mechanisms, such as the ability of trees to repair embolism. While it is obvious that drought-induced forest dieback is a complex process involving a number of biotic and abiotic factors, we would like to draw the attention of scientists to the state of evidence for embolism repair, thereby guiding research on tree drought resistance into the most relevant and fruitful directions

Elevated [CO2] does not ameliorate the negative effects of elevated temperature on drought‐induced mortality in Eucalyptus radiata seedlings

It has been reported that elevated temperature accelerates the time-to-mortality in plants exposed to prolonged drought, while elevated [CO(2)] acts as a mitigating factor because it can reduce stomatal conductance and thereby reduce water loss. We examined the interactive effects of elevated [CO(2)] and temperature on the inter-dependent carbon and hydraulic characteristics associated with drought-induced mortality in Eucalyptus radiata seedlings grown in two [CO(2)] (400 and 640 μL L(-1)) and two temperature (ambient and ambient +4 °C) treatments. Seedlings were exposed to two controlled drying and rewatering cycles, and then water was withheld until plants died. The extent of xylem cavitation was assessed as loss of stem hydraulic conductivity. Elevated temperature triggered more rapid mortality than ambient temperature through hydraulic failure, and was associated with larger water use, increased drought sensitivities of gas exchange traits and earlier occurrence of xylem cavitation. Elevated [CO(2)] had a negligible effect on seedling response to drought, and did not ameliorate the negative effects of elevated temperature on drought. Our findings suggest that elevated temperature and consequent higher vapour pressure deficit, but not elevated [CO(2)], may be the primary contributors to drought-induced seedling mortality under future climates.

Changing precipitation regimes and the water and carbon economies of trees

Climate change is predicted to affect the water balance of several ecosystems mostly through changes in the energy budget and hydrological input (rainfall frequency, intensity, and timing). Changes in rainfall patterns and cloudiness directly affect incoming radiation, atmospheric water vapor saturation deficit and soil water availability, the main variables controlling the rates of water uptake and transport along the soil–plant–atmosphere continuum (SPAC). Developing a predictive framework about vegetation responses to a changing climate is challenging because it involves complex non-linear interactions between these environmental variables and species-specific responses. By examining the hydraulic traits of functional groups within plant communities we can better predict the impacts of changes in rainfall regimes within functional groups and therefore, generate more realistic predictions of ecosystem carbon and water balance changes due to local and regional changes in precipitation regimes. In this review, we discuss several aspects of plant hydraulic functioning and then explore how predicted changes in precipitation regimes may affect tree water and carbon balance. We examine the impacts of changes in rainfall patterns on the SPAC and also explore the multiple ways that plants can absorb and transport water. Finally, we explore the diversity of hydraulic traits and potential mechanisms causing large-scale drought-induced mortality recently observed in several ecosystems.

Evaluating spatial patterns of drought-induced tree mortality in a coastal California pine forest

In a coastal, fog-influenced forest on Santa Cruz Island in southern California, we observed mortality of Bishop pine (Pinus muricata D.Don) trees following a brief (2year), yet intense, drought. While anecdotal evidence indicates that drought-induced Bishop pine mortality has occurred in the past in the stand we studied, this is the first attempt to capture the spatial distribution of mortality, and begin to understand the environmental drivers underlying these events. We used high spatial resolution remote sensing data to quantify the spatial extent of tree mortality using a 1m true color aerial photograph and a 1m LiDAR digital elevation model. We found the highest density of dead trees in the drier, more inland margins of the forest stand. We used the Random Forest decision tree algorithm to test which environmental variables (e.g., summertime cloud frequency, solar insolation, and geomorphic attributes) would best separate live and dead tree populations. We also included tree height as a variable in our analysis, which we used as a proxy for overall tree size and potential rooting distribution. Based on the Random Forest analysis, we generated a map of the probability of survival. We found tree survivorship after drought was best explained by the frequency of summertime clouds, elevation, and tree height. Specifically, survivorship was greatest for larger trees (∼8–10m tall) in more foggy parts of the stand located at moderate elevation. We found that probability of survival was lowest at the inland extent of the stand where trees occur at the upper limit of their elevation range (∼400m). The coexistence of these main factors with other landscape variables help identify areas of suitable habitat for Bishop pines across the stand, and extend our understanding of this species’ distribution.

Detecting mortality induced structural and functional changes in a piñon-juniper woodland using Landsat and RapidEye time series

Piñon–juniper (PJ) woodlands have recently undergone dramatic drought-induced mortality, triggering broad scale structural changes in this extensive Southwestern US biome. Given that climate projections for the region suggest widespread conifer mortality is likely to continue into the next century, it is critical to better understand how this climate-induced change in vegetation structure alters ecosystem function (e.g., productivity, biogeochemical cycling, energy partitioning) in PJ woodlands. Data from satellite remote sensing could potentially help to gain some of this understanding. However, relatively little is known about the suitability of satellite remote sensing for monitoring mortality induced structural and functional changes in PJ woodlands, a semi-arid biome characterized by sparse vegetation that complicates remote sensing of vegetation properties. Here we examined the potential role of satellite remote sensing to better understand structural and functional changes that take place in PJ woodlands as they respond to and recover from climate induced mortality events. We used time series medium (30×30m) and high (≤5×5m) spatial resolution satellite data in concert with eddy covariance-based gross primary productivity (GPP) data collected in a PJ woodland mortality manipulation near Mountainair, NM. Our results showed that both high and medium resolution satellite remote sensing data can provide ecosystem level assessments of overall canopy mortality and subsequent regrowth in these semi-arid woodlands. However, high spatial resolution data allowed detecting the disturbance almost a full year earlier and enabled us to describe the spatially heterogeneous patterns of recovery typical for such complex. The ability of satellite remote sensing to be used to provide information about changes in ecosystem function was conditional on the presence of sufficient soil moisture; when soil moisture was present the strength of the linear relationship between GPP and spectral vegetation indices improved at the manipulated site (e.g. for high resolution NDVI data R2=0.13 during drought conditions, R2=0.46 when wet, P<.05) and less so at a nearby control site (R2=0.09 during drought conditions, compared to R2=0.22 during growing conditions, P<.05). Thus, although satellite time series can be used to monitor and improve our understanding of canopy function in both disturbed and undisturbed semi-arid coniferous biomes, combining in situ eddy covariance data with satellite time series demonstrated that the utility of these datasets is highest during periods of peak growing conditions. This study therefore provides a benchmark for understanding how spatial and temporal variation in vegetation structure and function can be best studied as semi-arid coniferous biomes respond to future climate variability.

Species assemblage patterns around a dominant emergent tree are associated with drought resistance

Water availability has long been recognized as an important driver of species distribution patterns in forests. The conifer Agathis australis (D. Don) Lindl. (kauri; Araucariaceae) grows in the species-rich forests of northern New Zealand. It is accompanied by distinctive species assemblages, and during summer the soil beneath A. australis is often significantly drier than soils beneath surrounding broadleaved angiosperm canopy species. We used a shade house dry-down experiment to determine whether species that grow close to A. australis differed in drought tolerance physiology compared with species that rarely grow close to A. australis. Stomatal conductance (g(s)) was plotted against leaf water potential (ψ) to identify drought tolerance strategies. Seedlings of species that occur in close spatial association with A. australis (including A. australis seedlings) were most resistant to drought stress, and all displayed a drought avoidance strategy of either declining gs to maintain ψ or simultaneous declines in g(s) and ψ. The species not commonly occurring beneath A. australis, but abundant in the surrounding forest, were the most drought-sensitive species and succumbed relatively quickly to drought-induced mortality with rapidly declining gs and ψ values. These results were confirmed with diurnal measurements of g(s) and assimilation rates throughout the day, and leaf wilting analysis. We conclude that the varied abilities of the species to survive periods of drought stress as seedlings shapes the composition of the plant communities beneath A. australis trees. Furthermore, forest diversity may be impacted by climate change as the predicted intensification of droughts in northern New Zealand is likely to select for drought-tolerant species over drought-intolerant species.

Drought-induced mortality selectively affects Scots pine trees that show limited intrinsic water-use efficiency responsiveness to raising atmospheric CO2.

Widespread drought-induced tree mortality has been documented around the world, and could increase in frequency and intensity under warmer and drier conditions. Ecophysiological differences between dying and surviving trees might underlie predispositions to mortality, but are poorly documented. Here we report a study of Scots pines (Pinus sylvestris L.) from two sites located in north-eastern Iberian Peninsula where drought-associated mortality episodes were registered during the last few decades. Time trends of discrimination against 13C (Δ13C) and intrinsic water-use efficiency (WUEi) in tree rings at an annual resolution and for a 34 year period were used to compare co-occurring now-dead and surviving pines. Results indicate that both surviving and now-dead pines significantly increased their WUEi over time, although this increase was significantly lower for now-dead individuals. These differential WUEi trends corresponded to different scenarios describing how plant gas exchange responds to increasing atmospheric CO2 (Ca): the estimated intercellular CO2 concentration was nearly constant in surviving pines but tended to increase proportionally to Ca in now-dead trees. Concurrently, the WUEi increase was not paralleled by a growth enhancement, regardless of tree state, suggesting that in water-limited areas like the Mediterranean, it cannot overcome the impact of an increasingly warmer and drier climate on tree growth.

Feature: Improving our knowledge of drought‐induced forest mortality through experiments, observations, and modeling

Feature: Improving our knowledge of drought‐induced forest mortality through experiments, observations, and modeling

Siberian pine decline and mortality in southern siberian mountains

The causes and resulting spatial patterns of Siberian pine mortality in eastern Kuznetzky Alatau Mountains, Siberia were analyzed based on satellite (Landsat, MODIS) and dendrochronology data. Climate variables studied included temperature, precipitation and Standardized Precipitation-Evapotranspiration Index (SPEI) drought index. Landsat data analysis showed that stand mortality was first detected in the year 2006 at an elevation of 650m, and extended up to 900m by the year 2012. Mortality was accompanied by a decrease in MODIS-derived vegetation index (EVI). The area of dead stands and the upper mortality line were correlated with increased drought. The uphill margin of mortality was limited by elevational precipitation gradients. Dead stands (i.e., >75% tree mortality) were located mainly on southern slopes. With respect to slope, mortality was observed within a 7–20° range with greatest mortality occurring on convex terrain. Tree radial increment measurements correlate and were synchronous with SPEI (r2=0.37, rs=80). The results also showed the primary role of drought stress on Siberian pine mortality. A secondary role may be played by bark beetles and root fungi attacks. The observed Siberian pine mortality is part of a broader phenomenon of “dark needle conifers” (DNC, i.e., Siberian pine, fir and spruce) decline and mortality in European Russia, Siberia, and the Russian Far East. All locations of DNC decline coincided with areas of observed drought increase. The results obtained are one of the first observations of drought-induced decline and mortality of DNC at the southern border of boreal forests. Meanwhile if model projections of increased aridity are correct DNC within the southern part of its areal may be replaced by drought-resistant Pinus silvestris and Larix sibirica.

Evaluating theories of drought‐induced vegetation mortality using a multimodel–experiment framework

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 5yr 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.

Drought-induced tree species replacement is reflected in the spatial variability of soil respiration in a mixed Mediterranean forest

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.