Growth models can be used to assess forest vulnerability to climate warming. If global warming amplifies water deficit in drought‐prone areas, tree populations located at the driest and southernmost ...distribution limits (rear‐edges) should be particularly threatened. Here, we address these statements by analyzing and projecting growth responses to climate of three major tree species (silver fir, Abies alba; Scots pine, Pinus sylvestris; and mountain pine, Pinus uncinata) in mountainous areas of NE Spain. This region is subjected to Mediterranean continental conditions, it encompasses wide climatic, topographic and environmental gradients, and, more importantly, it includes rear‐edges of the continuous distributions of these tree species. We used tree‐ring width data from a network of 110 forests in combination with the process‐based Vaganov–Shashkin‐Lite growth model and climate–growth analyses to forecast changes in tree growth during the 21st century. Climatic projections were based on four ensembles CO2 emission scenarios. Warm and dry conditions during the growing season constrain silver fir and Scots pine growth, particularly at the species rear‐edge. By contrast, growth of high‐elevation mountain pine forests is enhanced by climate warming. The emission scenario (RCP 8.5) corresponding to the most pronounced warming (+1.4 to 4.8 °C) forecasted mean growth reductions of −10.7% and −16.4% in silver fir and Scots pine, respectively, after 2050. This indicates that rising temperatures could amplify drought stress and thus constrain the growth of silver fir and Scots pine rear‐edge populations growing at xeric sites. Contrastingly, mountain pine growth is expected to increase by +12.5% due to a longer and warmer growing season. The projections of growth reduction in silver fir and Scots pine portend dieback and a contraction of their species distribution areas through potential local extinctions of the most vulnerable driest rear‐edge stands. Our modeling approach provides accessible tools to evaluate forest vulnerability to warmer conditions.
Forecasted increase drought frequency and severity may drive worldwide declines in forest productivity. Species‐level responses to a drier world are likely to be influenced by their functional ...traits. Here, we analyse forest resilience to drought using an extensive network of tree‐ring width data and satellite imagery. We compiled proxies of forest growth and productivity (TRWi, absolutely dated ring‐width indices; NDVI, Normalized Difference Vegetation Index) for 11 tree species and 502 forests in Spain corresponding to Mediterranean, temperate, and continental biomes. Four different components of forest resilience to drought were calculated based on TRWi and NDVI data before, during, and after four major droughts (1986, 1994–1995, 1999, and 2005), and pointed out that TRWi data were more sensitive metrics of forest resilience to drought than NDVI data. Resilience was related to both drought severity and forest composition. Evergreen gymnosperms dominating semi‐arid Mediterranean forests showed the lowest resistance to drought, but higher recovery than deciduous angiosperms dominating humid temperate forests. Moreover, semi‐arid gymnosperm forests presented a negative temporal trend in the resistance to drought, but this pattern was absent in continental and temperate forests. Although gymnosperms in dry Mediterranean forests showed a faster recovery after drought, their recovery potential could be constrained if droughts become more frequent. Conversely, angiosperms and gymnosperms inhabiting temperate and continental sites might have problems to recover after more intense droughts since they resist drought but are less able to recover afterwards.
In this study, we analysed the resistance and resilience to drought of forests dominated by 11 species across wide climatic and environmental gradients in the Mediterranean basin using proxies of forest productivity (NDVI) and carbon accumulation (ring‐width indices, TRWi) and considering four extreme drought events recorded between 1980 and 2005. Our results indicate that drought intensity is a major driver of forest resilience to drought but that species inhabiting different regions present different strategies to cope with drought and thus they can respond differently to more frequent and severe droughts.
Global climate change is expected to further raise the frequency and severity of extreme events, such as droughts. The effects of extreme droughts on trees are difficult to disentangle given the ...inherent complexity of drought events (frequency, severity, duration, and timing during the growing season). Besides, drought effects might be modulated by trees’ phenotypic variability, which is, in turn, affected by long‐term local selective pressures and management legacies. Here we investigated the magnitude and the temporal changes of tree‐level resilience (i.e., resistance, recovery, and resilience) to extreme droughts. Moreover, we assessed the tree‐, site‐, and drought‐related factors and their interactions driving the tree‐level resilience to extreme droughts. We used a tree‐ring network of the widely distributed Scots pine (Pinus sylvestris) along a 2,800 km latitudinal gradient from southern Spain to northern Germany. We found that the resilience to extreme drought decreased in mid‐elevation and low productivity sites from 1980–1999 to 2000–2011 likely due to more frequent and severe droughts in the later period. Our study showed that the impact of drought on tree‐level resilience was not dependent on its latitudinal location, but rather on the type of sites trees were growing at and on their growth performances (i.e., magnitude and variability of growth) during the predrought period. We found significant interactive effects between drought duration and tree growth prior to drought, suggesting that Scots pine trees with higher magnitude and variability of growth in the long term are more vulnerable to long and severe droughts. Moreover, our results indicate that Scots pine trees that experienced more frequent droughts over the long‐term were less resistant to extreme droughts. We, therefore, conclude that the physiological resilience to extreme droughts might be constrained by their growth prior to drought, and that more frequent and longer drought periods may overstrain their potential for acclimation.
We examined tree growth resilience of Scots pine along a 2,800 km latitudinal gradient from southern Spain to north‐eastern Germany using 615 adult trees from 30 different sites. We found that the resilience of Scots pine to extreme drought decreased in mid‐elevation and low productivity sites from 1980–1999 to 2000–2011 due to more frequent and severe droughts in the later period. We showed that the impact of drought on tree‐level resilience was not dependent on its latitudinal location, but rather on the type of sites trees were growing at and on their growth performances during the pre‐drought period.
Aim: We investigate the effects of the environmental and geographical processes driving growth resilience and recovery in response to drought in Mediterranean Pinus pinaster forests. We explicitly ...consider how intraspecific variability modulates growth resilience to drought. Location: Western Mediterranean basin. Methods: We analysed tree rings from a large network of 48 forests (836 trees) encompassing wide ecological and climatic gradients, including six provenances. To characterize the major constraints of P. pinaster growth under extremely dry conditions, we simulated growth responses to temperature and soil moisture using a process-based growth model coupled with the quantification of climate–growth relationships. Then, we related growth–resilience indices to provenance and site variables considering different drought events. Results: Pinus pinaster displayed strong variation in growth resilience across its distributional range, but common patterns were found within each provenance. Post-drought resilience increased with elevation and drier conditions but decreased with spring precipitation. Trees from dry sites were less resistant to drought but recovered faster than trees from wet sites. Main conclusions: Resilience strategies differed among tree provenances: wet forests showed higher growth resistance to drought, while dry forests presented faster growth recovery, suggesting different impacts of climate warming on forest productivity. We detected geographically structured resilience patterns corresponding to different provenances, confirming high intraspecific variability in response to drought. This information should be included in species distribution models to simulate forest responses to climate warming and forecasted aridification.
Rear‐edge populations at the xeric distribution limit of tree species are particularly vulnerable to forest dieback triggered by drought. This is the case of silver fir (Abies alba) forests located ...in Southwestern Europe. While silver fir drought‐induced dieback patterns have been previously explored, information on the role played by nutritional impairment is lacking despite its potential interactions with tree carbon‐water balances. We performed a comparative analysis of radial growth, intrinsic water‐use efficiency (iWUE), oxygen isotopes (δ18O) and nutrient concentrations in leaves of declining (DD) and non‐declining (ND) trees in silver fir in four forests in the Spanish Pyrenees. We also evaluated the relationships among dieback predisposition, intraspecific trait variation (wood density and leaf traits) and rhizosphere soil physical–chemical properties. The onset of growth decline in DD trees occurred more than two decades ago, and they subsequently showed low growth resilience against droughts. The DD trees presented consistently lower foliar concentrations of nutrients such as P, K, Cu and Ni than ND trees. The strong effects of foliar nutrient status on growth resilience indices support the key role played by mineral nutrition in tree functioning and growth before, during and after drought. In contrast, variability in wood density and leaf morphological traits, as well as soil properties, showed weak relationships with tree nutritional status and drought performance. At the low elevation, warmer sites, DD trees showed stronger climate–growth relationships and lower δ18O than ND trees. The uncoupling between iWUE and δ18O, together with the positive correlations between P and K leaf concentrations and δ18O, point to deeper soil/bedrock water sources and vertical decoupling between nutrient and water uptake in DD trees. This study provides novel insights into the mechanisms driving silver fir dieback and highlights the need to incorporate tree nutrition into forest dieback studies.
Las poblaciones del límite xérico de distribución de las especies de árboles son particularmente vulnerables al decaimiento forestal inducido por sequía. Este es el caso de los bosques de abeto (Abies alba) situados en el suroeste de Europa. Si bien los patrones de decaimiento provocado por sequía del abeto se han explorado previamente, falta información sobre el papel que desempeña el deterioro nutricional a pesar de sus interacciones potenciales con los balances de agua y carbono de los árboles. En este estudio, hemos realizado un análisis comparativo del crecimiento radial, la eficiencia intrínseca del uso del agua (iWUE), los isótopos de oxígeno (δ18O) y las concentraciones de nutrientes en hojas de árboles decaídos (DD) y no decaídos (ND) en cuatro abetares de los Pirineos españoles. También evaluamos las relaciones entre la predisposición al decaimiento, la variación de rasgos intraespecíficos (densidad de la madera y rasgos de las hojas) y las propiedades físico‐químicas de la rizosfera. El inicio de la disminución del crecimiento en los árboles DD ocurrió hace más de dos décadas y posteriormente mostraron una baja resiliencia de crecimiento frente a las sequías. Los árboles DD presentaron concentraciones foliares consistentemente más bajas de nutrientes como P, K, Cu y Ni que los árboles ND. Los fuertes efectos del estado de los nutrientes foliares en los índices de resiliencia del crecimiento respaldan el papel clave que desempeña la nutrición mineral en el funcionamiento y el crecimiento de los árboles antes, durante y después de la sequía. En contraste, la variabilidad en la densidad de la madera y los rasgos morfológicos de las hojas, así como las propiedades del suelo, mostraron una relación débil con el estado nutricional de los árboles y la respuesta del crecimiento a la sequía. En los sitios más cálidos y de baja elevación, los árboles DD mostraron relaciones clima‐crecimiento más fuertes y un δ18O más bajo que los árboles ND. El desacoplamiento entre iWUE y δ18O, junto con las correlaciones positivas entre las concentraciones foliares de P y K y δ18O, apuntan a fuentes de agua más profundas del suelo/lecho rocoso y un desacoplamiento vertical entre la absorción de nutrientes y agua en los árboles DD. Este estudio proporciona información novedosa sobre los mecanismos que impulsan el decaimiento del abeto y destaca la necesidad de incorporar la nutrición de los árboles en los estudios de muerte regresiva del bosque.
Declining silver fir trees show lower foliar concentrations of essential nutrients such as P, K, Cu and Ni than non‐declining trees. This nutritional impairment exacerbates and amplifies the negative effects of drought on tree functioning and growth, considering the relationship of tree nutritional status with water‐use efficiency and growth resilience against drought. Incorporating nutritional aspects into forest die‐off studies can provide novel insights into the mechanisms driving drought‐related tree dieback.
Climate warming is expected to positively alter upward and poleward treelines which are controlled by low temperature and a short growing season. Despite the importance of treelines as a bioassay of ...climate change, a global field assessment and posterior forecasting of tree growth at annual scales is lacking. Using annually resolved tree‐ring data located across Eurasia and the Americas, we quantified and modeled the relationship between temperature and radial growth at treeline during the 20th century. We then tested whether this temperature–growth association will remain stable during the 21st century using a forward model under two climate scenarios (RCP 4.5 and 8.5). During the 20th century, growth enhancements were common in most sites, and temperature and growth showed positive trends. Interestingly, the relationship between temperature and growth trends was contingent on tree age suggesting biogeographic patterns in treeline growth are contingent on local factors besides climate warming. Simulations forecast temperature–growth decoupling during the 21st century. The growing season at treeline is projected to lengthen and growth rates would increase and become less dependent on temperature rise. These forecasts illustrate how growth may decouple from climate warming in cold regions and near the margins of tree existence. Such projected temperature–growth decoupling could impact ecosystem processes in mountain and polar biomes, with feedbacks on climate warming.
Treelines draw the low temperature limit for tree growth. These ecotones are monitors of the effects of climate warming on terrestrial ecosystems. Treelines constitute natural laboratories to shifts in temperature–growth coupling in response to climate change, but how will be tree growth at treeline during the 21st century? By using a new and tailored tree‐ring network from treelines across four continents, and by applying a forward, process‐based growth model, we found rising growth trends and projected a decoupling between temperature and growth with the warming climate during the 21st century. This will induce major changes in alpine and polar treelines.
Climate extremes, such as abnormally dry and wet conditions, generate abrupt shifts in tree growth, a situation which is expected to increase under predicted climate conditions. Thus, it is crucial ...to understand factors determining short‐ and long‐term tree performance in response to higher frequency and intensity of climate extremes.
We evaluated how three successive droughts and wet years influenced short‐ and long‐term growth of six dominant Iberian tree species. Within species variation in growth response to repeated dry and wet years was evaluated as a function of individual traits related to resource and water use (diameter at breast height DBH, wood density WD and specific leaf area SLA) and tree‐to‐tree competition across climatically contrasted populations. Furthermore, we assessed how short‐term accumulated impacts of the repeated dry and wet years influenced long‐term growth performance.
All species showed strong short‐term growth decreases and enhancements due to repeated dry and wet years. However, patterns of accumulated growth decreases (AcGD) and enhancements (AcGE) across climatically contrasting populations were species‐specific. Furthermore, individual trait data were weakly associated with either AcGD or AcGE and the few relevant associations were found for conifers. Intraspecific variations in tree growth responses to repeated climates extremes were large, and not explained by intraspecific variability in SLA and WD. Accumulated impacts of repeated dry and wet years were related to long‐term growth trends, showing how the recurrence of climate extremes can determine growth trajectories. The relationships of AcGD and AcGE with long‐term growth trends were more common in conifers species.
Synthesis. Repeated climate extremes do not only cause short‐term growth reductions and enhancements but also determine long‐term tree growth trajectories. This result shows how repeated droughts can lead to growth decline. Conifers were more susceptible to the accumulated effects of extreme weather events indicating that in the future, more intense and frequent climate extremes will alter growth performance in forests dominated by these species.
Repeated climate extremes do not only cause short‐term growth reductions and enhancements but also determine long‐term tree growth trajectories. This result shows how repeated droughts can lead to growth decline. Conifers were more susceptible to the accumulated effects of extreme weather events indicating that in the future, more intense and frequent climate extremes will alter growth performance in forests dominated by these species.
Climate warming is expected to enhance tree growth at alpine treelines. A higher growth rate is forecasted as temperatures rise and growth becomes less dependent on the temperature rise. Since radial ...growth is just one component of treeline dynamics those forecasts do not necessarily apply to treeline elevation or latitude; treelines can shift upward or poleward or remain stable.
The negative impacts of severe drought on the growth and vigor of tree species and their relationship with forest decline have not been properly evaluated taking into account the differential ...responses to such stress of trees, sites and species. We evaluated these responses by quantifying the changes in radial growth of plantations of four pine species (
Pinus sylvestris
,
Pinus nigra
,
Pinus pinaster
,
Pinus halepensis
) which showed distinct decline and defoliation levels in southeastern Spain. We used dendrochronological methods, defoliation records, linear mixed models of basal area increment and dynamic factor analysis to quantify the responses of trees at the species and individual scales to site conditions and drought stress. In the region a temperature rise and a decrease in spring precipitation have led to drier conditions during the late twentieth century characterized by severe droughts in the 1990s and 2000s. As expected, the defoliation levels and the reductions in basal area increment were higher in those species more vulnerable to drought-induced xylem embolism (
P. sylvestris
) than in those more resistant (
P. halepensis
). Species adapted to xeric conditions but with high growth rates, such as
P. pinaster
, were also vulnerable to drought-induced decline. The reduction in basal area increment and the defoliation events occurred after consecutive severe droughts. A decrease in spring precipitation, which is the main driver of radial growth, is the most plausible cause of recent forest decline. The sharp growth reduction and widespread defoliation of the most affected pine plantations of Scots pine make their future persistence in drought-prone sites unlikely under the forecasted warmer and drier conditions.
Forests play a key role in the carbon balance of terrestrial ecosystems. One of the main uncertainties in global change predictions lies in how the spatiotemporal dynamics of forest productivity will ...be affected by climate warming. Here we show an increasing influence of climate on the spatial variability of tree growth during the last 120 y, ultimately leading to unprecedented temporal coherence in ring-width records over wide geographical scales (spatial synchrony). Synchrony in growth patterns across cold-constrained (central Siberia) and drought-constrained (Spain) Eurasian conifer forests have peaked in the early 21st century at subcontinental scales (∼1,000 km). Such enhanced synchrony is similar to that observed in trees co-occurring within a stand. In boreal forests, the combined effects of recent warming and increasing intensity of climate extremes are enhancing synchrony through an earlier start of wood formation and a stronger impact of year-to-year fluctuations of growing-season temperatures on growth. In Mediterranean forests, the impact of warming on synchrony is related mainly to an advanced onset of growth and the strengthening of drought-induced growth limitations. Spatial patterns of enhanced synchrony represent early warning signals of climate change impacts on forest ecosystems at subcontinental scales.
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