Timing of Drought Triggers Distinct Growth Responses in Holm Oak: Implications to Predict Warming-Induced Forest Defoliation and Growth Decline

Droughts negatively impact forests by reducing growth and increasing defoliation leading to forest dieback as the climate becomes warmer and drier. However, the timing and severity of droughts determine how differently or intensively water shortage affects primary (shoot and leaf formation) and secondary growth (stem radial growth based on tree-ring widths). We compare the impact of two severe droughts (2005, 2012), showing different climatic characteristics on the growth responses of three Mediterranean holm oak stands in northeastern Spain. We also quantify climate trends and drought severity. Then, we use remote sensing data to infer how those droughts impacted forest productivity. Both droughts were characterized by warm and dry spring conditions leading to reduced budburst, low shoot production, asynchrony in primary growth and decreased productivity and scarce radial growth, particularly in 2005. However, defoliation peaked in 2012 when radial growth showed minimum values and early spring and late summer temperatures reached maximum values. We discuss how uncoupled and resilient are the responses of primary and secondary growth to drought. Finally, these findings are used to gain insight into the drought-related drivers of defoliation in Spanish holm oak forests.We acknowledge support by the CSIC Open Access Publication Initiative through its Unit of Information Resources for Research (URICI).Peer reviewe

Sex and tree rings: Females neither grow less nor are less water-use efficient than males in four dioecious tree species

Females of woody dioecious species usually expend more resources on reproduction than males. Therefore, it is expected that females incur greater costs of reproduction than males, and, as a result, trade-offs between reproduction and growth should emerge. The aim of this study is to test those hypotheses by analyzing the differences between genders regarding radial growth (basal area increment) and wood carbon isotope composition (d13C), a proxy of water-use efficiency. We compared these two variables in males and females of four dioecious tree species inhabiting drought-prone Mediterranean sites in Spain (Pistacia terebinthus, Ilex aquifolium, Juniperus thurifera and Ailanthus altissima). We analyzed the influence of sex on the radial growth patterns throughout the tree life considering the growth stage of individuals, the differences in the response of genders to climate variables (air temperature, precipitation and drought severity), and the d13C during a severe drought period. One site was studied for each species and 21–33 trees per species were sampled in each site. No differences in growth were found between genders for any of the four species throughout their life span. No significant interactions between gender and precipitation were found, although A. altissima males were more responsive to summer (June-July) temperature. No differences in d13C were found between genders excepting for P. terebinthus, indicating that the males of this species show a less efficient water use during drought events than the females. These results do not support the broad assumption that females of woody dioecious plants show lower growth and are less water-use efficient than males or that they respond differently to precipitation variability, except for P. terebinthus during drought events. Further analyses could be performed in other dioecious species inhabiting seasonally dry regions to confirm or reject our conclusions

Prior height, growth, and wood anatomy differently predispose to drought-induced dieback in two Mediterranean oak speciesk

International audienceAbstractKey messageCoexisting Mediterranean oaks are differently predisposed to heat- and drought-induced dieback as a function of height, prior growth, wood anatomy, and growth responsiveness to temperatures. To forecast post-dieback damage, the variability of species and individual traits must be considered.ContextForests are susceptible to drought-induced dieback. However, considerable variability in how drought translates into tree damage exists in coexisting species.AimsThis study aimed to assess if tree size, radial growth, and wood anatomy predisposed trees to drought damage, measured as defoliation and changes in non-structural carbohydrate (NSC) concentrations.MethodsWe measured radial growth, wood anatomy, and post-drought NSC concentrations in highly defoliated and less defoliated holm oak (Quercus ilex) and Portuguese oak (Quercus faginea) trees co-occurring in a coppice stand.ResultsHighly defoliated trees showed a lower height than less defoliated individuals. In holm oak, a reduced previous growth and the formation of vessels with smaller lumen areas predisposed to drought damage, which suggests hydraulic deterioration. In Portuguese oak, most defoliated trees grew less in response to elevated growing season temperatures. Sapwood starch and NSC concentrations decrease in defoliated holm oaks.ConclusionA height-dependent predisposition modulates the responses to drought-induced dieback in Mediterranean oak coppices. Coexisting oak species presented different predisposing factors to drought-induced dieback related to growth (holm oak) and its sensitivity to temperature (Portuguese oak). To forecast post-dieback damage, we should consider the variability of traits between and within species

Drops in needle production are early-warning signals of drought-triggered dieback in scots pine

Producción CientíficaUnderstanding the mechanisms of drought-induced forest dieback and tree mortality is a priority for predicting forest responses to climate change. However, long-term information on drought impacts on crown dynamics and how it relates to tree water and carbon economy is still lacking. Comparing declining and non-declining Scots pines at the same site, we quantified primary and secondary growth and intrinsic water-use efficiency (WUEi). Further, we reconstructed the needle production and the annual total number of needles using the Needle Trace Method. Here, we reconstructed the apical needles through the assessment of 2460 longitudinal sections. We fitted partial least squares regression models to assess climate influence on primary and secondary growth and needle dynamics, and then linear mixed models using climate covariates and discerning between health status in the 1975–2016 period. Finally, we analyzed drought legacy effects applying a Bayesian hierarchical framework. We detected the highest differences between health tree classes in the annual total number of needles during the warm-dry 1980s. Declining trees responded more negatively to climate than non-declining conspecifics and showed a higher variability of drought legacy effects. We found significant drought legacy effects in the annual total number of needles up to 3 years after a drought. The warm, dry 1980s, that preceded the dieback, showed the highest δ13C values since the 1970s. Declining trees showed higher WUEi than non-declining trees from the 2000s onwards. Our study sheds light on the major role of needle dynamics in dieback episodes and illustrates how past drops in needle production may be interpreted as early warnings of drought-induced dieback in Scots pine.Ministerio de Ciencia e Innovación y Agencia Estatal de Investigación - MCIN/AEI/10.13039/501100011033 (grant IJC2019- 040571-I), and the Instituto de Estudios Turolenses (IET) project PROWARM (PID2020-118444GA-100)Publicación en abierto financiada por el Consorcio de Bibliotecas Universitarias de Castilla y León (BUCLE), con cargo al Programa Operativo 2014ES16RFOP009 FEDER 2014-2020 DE CASTILLA Y LEÓN, Actuación:20007-CL - Apoyo Consorcio BUCL

Post-drought Resilience After Forest Die-Off: Shifts in Regeneration, Composition, Growth and Productivity

A better understanding on the consequences of drought on forests can be reached by paying special attention to their resilience capacity, i.e., the ability to return to a state similar to pre-drought conditions. Nevertheless, extreme droughts may surpass the threshold for the resilience capacity triggering die-off causing multiple changes at varying spatial and temporal scales and affecting diverse processes (tree growth and regeneration, ecosystem productivity). Combining several methodological tools allows reaching a comprehensive characterization of post-drought forest resilience. We evaluated the changes in the abundance, regeneration capacity (seedling abundance), and radial growth (annual tree rings) of the main tree species. We also assessed if drought-induced reductions in growth and regeneration of the dominant tree species scale-up to drops in vegetation productivity by using the Normalized Difference Vegetation Index (NDVI). We studied two conifer forests located in north-eastern Spain which displayed drought-induced die-off during the last decades: a Scots pine (Pinus sylvestris) forest under continental Mediterranean conditions and a Silver fir (Abies alba) forest under more temperate conditions. We found a strong negative impact of a recent severe drought (2012) on Scots pine growth, whereas the coexisting Juniperus thurifera showed positive trends in basal area increment (0.02 ± 0.003 cm2 yr-1). No Scots pine recruitment was observed in sites with intense die-off, but J. thurifera and Quercus ilex recruited. The 2012 drought event translated into a strong NDVI reduction (32% lower than the 1982–2014 average). In Silver fir we found a negative impact of the 2012 drought on short-term radial growth, whilst long-term growth of Silver fir and the coexisting Fagus sylvatica showed positive trends. Growth rates were higher in F. sylvatica (0.04 ± 0.003 cm2 yr-1) than in A. alba (0.02 ± 0.004 cm2 yr-1). These two species recruited beneath declining and non-declining Silver fir trees. The 2012 drought translated into a strong NDVI reduction which lasted until 2013. The results presented here suggest two different post-drought vegetation pathways. In the Scots pine forest, the higher growth and recruitment rates of J. thurifera correspond to a vegetation shift where Scots pine is being replaced by the drought-tolerant juniper. Conversely, in the Silver fir forest there is an increase of F. sylvatica growth and abundance but no local extinction of the Silver fir. Further research is required to monitor the evolution of these forests in the forthcoming years to illustrate the cumulative impacts of drought on successional dynamics

Detecting Nothofagus pumilio Growth Reductions Induced by Past Spring Frosts at the Northern Patagonian Andes

Extreme climatic events, such as late frosts in spring during leaf flush, have considerable impacts on the radial growth of temperate broadleaf trees. Albeit, all broadleaved species are potentially vulnerable, damage depends on the particularities of the local climate, the species, and its phenology. The impact of late spring frosts has been widely investigated in the Northern Hemisphere, but the potential incidence in Southern Hemisphere tree species is still poorly known. Here, we reconstruct spring frost occurrence at 30 stands of the deciduous tree Nothofagus pumilio in its northern range of distribution in the Patagonian Andes. We identified tree ring-width reductions at stand level not associated with regional or local drought events, matching unusual minimum spring temperatures during leaf unfolding. Several spring frosts were identified along the northern distribution of N. pumilio, being more frequent in the more continental Argentinean forests. Spring frost in 1980 had the largest spatial extent. The spring frosts in 1980 and 1992 also induced damages in regional orchards. Spring frost damage was associated with (i) a period of unusually warm temperatures at the beginning of leaf unfolding, followed by (ii) freezing temperatures. This study helps expand our understanding of the climatic constraints that could determine the future growth and dynamics of Andean deciduous forests and the potential use of tree-rings as archives of extreme events of spring frosts in northern Patagonia.Fil: Sangüesa Barreda, Gabriel. Universidad de Valladolid; EspañaFil: Villalba, Ricardo. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mendoza. Instituto Argentino de Nivología, Glaciología y Ciencias Ambientales. Provincia de Mendoza. Instituto Argentino de Nivología, Glaciología y Ciencias Ambientales. Universidad Nacional de Cuyo. Instituto Argentino de Nivología, Glaciología y Ciencias Ambientales; ArgentinaFil: Rozas, Vicente. Universidad de Valladolid; España. Universidad Austral de Chile; ChileFil: Christie, Duncan A.. Centro de Investigaciones sobre Clima y Resiliencia; Chile. Universidad Austral de Chile; ChileFil: Olano, José Miguel. Universidad de Valladolid; Españ

Disparate effects of global-change drivers on mountain conifer forests: warming-induced growth enhancement in young trees vs. CO2 fertilization in old trees from wet sites

Theory predicts that the postindustrial rise in the concentration of CO2 in the atmosphere (ca ) should enhance tree growth either through a direct fertilization effect or indirectly by improving water use efficiency in dry areas. However, this hypothesis has received little support in cold-limited and subalpine forests where positive growth responses to either rising ca or warmer temperatures are still under debate. In this study, we address this issue by analyzing an extensive dendrochronological network of high-elevation Pinus uncinata forests in Spain (28 sites, 544 trees) encompassing the whole biogeographical extent of the species. We determine if the basal area increment (BAI) trends are linked to climate warming and increased ca by focusing on region- and age-dependent responses. The largest improvement in BAI over the past six centuries occurred during the last 150 years affecting young trees and being driven by recent warming. Indeed, most studied regions and age classes presented BAI patterns mainly controlled by temperature trends, while growing-season precipitation was only relevant in the driest sites. Growth enhancement was linked to rising ca in mature (151-300 year-old trees) and old-mature trees (301-450 year-old trees) from the wettest sites only. This finding implies that any potential fertilization effect of elevated ca on forest growth is contingent on tree features that vary with ontogeny and it depends on site conditions (for instance water availability). Furthermore, we found widespread growth decline in drought-prone sites probably indicating that the rise in ca did not compensate for the reduction in water availability. Thus, warming-triggered drought stress may become a more important direct driver of growth than rising ca in similar subalpine forests. We argue that broad approaches in biogeographical and temporal terms are required to adequately evaluate any effect of rising ca on forest growth

Evapotranspiration deficit controls net primary production and growth of silver fir: Implications for Circum-Mediterranean forests under forecasted warmer and drier conditions

Warming-induced drought stress has been hypothesized as a major driver of forest net primary production (NPP) reduction, but we lack reliable field data to assess if higher temperatures lead to forest NPP reduction, particularly in humid sites and at basin to landscape spatial scales. The use of a landscape approach would allow considering the feedbacks operating between climate, topography, soil vegetation and water resources. Here we follow that approach by simulating NPP using the regional hydro-ecologic simulation system (RHESSys) model and by comparing the results with radial growth data (tree-ring widths and intrinsic water-use efficiency - iWUE). We evaluate the relationships between climate, growth, NPP, atmospheric CO2 concentrations (ca) and iWUE in xeric and mesic silver fir forests subjected to contrasting water balances. The growth data successfully validated the 11-month NPP cumulated until spring. The main negative climatic driver of growth and NPP was the summer evapotranspiration deficit, which shows a negative association with tree-ring width indices. Sensitivity analyses indicate that rising ca do not compensate the severe NPP reduction associated to warmer and drier conditions. The positive effect of rising ca on NPP is mediated by climatic site conditions being detected only in mesic sites, whereas the negative effects of drought on NPP override any ca-related enhancement of NPP in xeric sites. Future warmer and drier conditions causing a higher evaporative demand by the atmosphere could lead to a NPP decline in temperate conifer forests subjected to episodic droughts. © 2015 Elsevier B.V.We would like to thank the Spanish Meteorological State Agency (AEMET) and the Confederación Hidrográfica del Ebro for providing the climatic and streamflow databases used in this study. This work has been supported by research projects CGL2011-27574-CO2-02, CGL2011-27536, CGL2014-52135-CO3-01 and Red de variabilidad y cambio climático RECLIM (CGL2014-517221-REDT) financed by the Spanish Commission of Science and Technology and FEDER, “LIFE12 ENV/ES/000536-Demonstration and validation of innovative methodology for regional climate change adaptation in the Mediterranean area (LIFE MEDACC)” financed by the LIFE programme of the European Commission and CTTP1/12 financed by the Comunidad de Trabajo de los Pirineos. JJC also acknowledges the support of ARAID and projects 012/2008, 387/2011 and 1012S (Organismo Autónomo Parques Nacionales, Spain).Peer Reviewe

Tree-ring density and carbon isotope composition are early-warning signals of drought-induced mortality in the drought tolerant Canary Island pine

Tree death is not always preceded by a visible decline in vigor (canopy dieback) or a progressive loss in crown volume. Identifying early-warning signals of incipient decline can help to implement the necessary measures to prevent tree death. The aim of this work is to understand what functional alterations preceded the massive drought-induced death of adult Pinus canariensis trees in an arid stand, located in the Canary Islands. To this aim, we analyzed interannual variations in earlywood and latewood width, wood density and anatomy, and carbon isotope composition, and the relationships among these variables, in dead and living trees from 1980 to 2013. Dead trees grew less since the 1990's, produced fewer parenchyma rays and resin canals, and exhibited a trend of decreasing latewood density and a marked shift in carbon isotope discrimination over the last 34 years that were reversed in living trees. Higher wood density in living trees resulted from thicker tracheid cell walls rather than narrower lumens. The intrinsic water use efficiency shifted from higher to lower values in dead trees after the 2000s. These results suggest a carbon limitation to maintain hydraulic safety under xylem tension, as well as to maintain storage and defense capacity, which can render trees more vulnerable to severe drought episodes. A long-term trend of decreasing tree-ring density and a reduced intrinsic water use efficiency in the short-term can be early-warning signals of carbon limitation and tree decline in drought-stressed P. canariensis. The analysis of these variables can be used to assess tree decline risks in similarly vulnerable conifer populations inhabiting drought-prone regions