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

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

Differences in temperature sensitivity and drought recovery between natural stands and plantations of conifers are species-specific

Forests are being impacted by climate and land-use changes which have altered their productivity and growth. Understanding how tree growth responds to climate in natural and planted stands may provide valuable information to prepare management in sight of climate change. Plantations are expected to show higher sensitivity to climate and lower post-drought resilience than natural stands, due to their lower compositional and structural diversity. We reconstructed and compared the radial growth of six conifers with contrasting ecological and climatic niches (Abies pinsapo, Cedrus atlantica, Pinus sylvestris, Pinus nigra, Pinus pinea, Pinus pinaster) in natural and planted stands subjected to seasonal drought in 40 sites. We quantified the relationships between individual growth variability and climate variables (temperature, precipitation and the SPEI drought index), as well as post-drought resilience. Elevated precipitation during the previous autumn-winter and current spring to early summer enhanced growth in both natural and planted stands of all species. Temperature effects on growth were less consistent: only plantations of A. pinsapo, C. atlantica, P. nigra, P. pinea, P. sylvetris and a natural stand of P. nigra showed negative impacts of summer temperature on growth. Drought reduced growth of all species in both plantations and natural stands, with variations in the temporal scale of the response. Drought constrained growth more severely in natural stands than in plantations of C. atlantica, P. pinaster and P. nigra, whereas the inverse pattern was found for A. pinsapo. Resilience to drought varied between species: natural stands of A. pinsapo, C. atlantica and P. pinaster recovered faster than plantations, while P. pinea plantations recovered faster than natural stands. Overall, plantations did not consistently show a higher sensitivity to climate and a lower capacity to recover after drought. Therefore, plantations are potential tools for mitigating climate warming. © 2021 The AuthorsThis study was supported by project FORMAL ( RTI2018-096884-B-C31 ) from the Spanish Ministry of Science, Innovation and Universities . GS-B was supported by a Spanish Ministry of Economy, Industry and Competitiveness Postdoctoral grant ( IJC2019-040571-I ; FEDER funds)

Papel de los factores bióticos y las sequías en el decaimiento del bosque: aportaciones desde la dendroecología

[ES] El aumento de la mortalidad y la pérdida de vitalidad de los bosques son procesos que afectan a diversas especies de árboles y biomas. Estos fenómenos de decaimiento forestal se han asociado a la interacción de múltiples factores relacionados con el cambio global, como el aumento de las sequías o la mayor incidencia de factores bióticos que generan estrés. Sin embargo, el papel que juegan estos organismos en el decaimiento del bosque en relación a la sequía no está del todo comprendido. En este trabajo caracterizamos retrospectivamente los efectos que ejercen dichos organismos sobre el crecimiento radial de árboles estresados por sequía utilizando dendrocronología. El efecto combinado del muérdago y las sequías reducen el crecimiento del pino silvestre y aumentan la defoliación, siendo estos efectos más evidentes en el ápice del árbol. La procesionaria induce pérdidas de crecimiento en pino laricio, pero en general se observa una recuperación dos años después. No encontramos un papel determinante de los hongos patógenos de raíz en el decaimiento del abeto. Finalmente, pinos carrasco muertos que presentaban ataques de insectos perforadores mostraron patrones de crecimiento similares a pinos vivos. Este estudio nos permite caracterizar algunos de los complejos y múltiples efectos de diferentes factores bióticos de estrés sobre el crecimiento y la vulnerabilidad de los bosques susceptibles a los efectos negativos de las sequías.[EN] Increasing mortality and decline of forests are affecting several tree species and biomes. These events of forest decline have been related to the interaction of several global change elements, like the increase in drought stress or a higher incidence of biotic stress factors. However, the roles played by biotic stressors have been scarcely explored as related to drought stress. In this paper we retrospectively characterize the effects of these organisms on radial growth of trees already stressed by drought using dendrochronology. The combined effect of mistletoe and droughts reduce tree growth in Scots pine and increase defoliation, being these effects more noticeable in the apex. The pine processionary moth induces growth losses in the affected stands of black pine, but they usually recover two years after the outbreak. We did not find a determinate role of root pathogens on silver fir dieback. Finally, dead Aleppo pine trees infested by bark beetles showed similar growth patterns than living pine trees. This study allows characterizing some of the complex and multiple effects of different biotic stressors on growth and vulnerability of forests subjected to the negative effects exerted by droughts.Este estudio ha sido financiado por los proyectos CGL2007- 66066-C04-02/BOS, CGL2008-04847-C02-01, CGL2011-26654 (MINECO), 387/2011 (Organismo Autónomo de Parques Nacionales) y PROCLIM (La Caixa-Gobierno de Aragón). Agradecemos la revisión realizada por dos revisores anónimos de una versión previa del texto. También queremos agradecer la ayuda prestada por todos los miembros del Laboratorio de Sanidad Forestal de Mora de Rubielos (Gobierno de Aragón), a todo el personal del Parque Nacional de Ordesa y Monte Perdido, al servicio de parques y jardines del Ayuntamiento de Zaragoza (Montserrat Hernández y José Bellosta), y a todos los APNs que nos han ayudado en las numerosas jornadas de campo. Finalmente, agradecemos a nuestros compañeros del IPE (CSIC) J.D. Galván y A.Q. Alla por su ayuda en los muestreos y la discusión de las ideas presentadas.Peer Reviewe

Plasticity in dendroclimatic response across the distribution range of Aleppo pine (Pinus halepensis)

We investigated the variability of the climate-growth relationship of Aleppo pine across its distribution range in the Mediterranean Basin. We constructed a network of tree-ring index chronologies from 63 sites across the region. Correlation function analysis identified the relationships of tree-ring index to climate factors for each site. We also estimated the dominant climatic gradients of the region using principal component analysis of monthly, seasonal, and annual mean temperature and total precipitation from 1,068 climatic gridpoints. Variation in ring width index was primarily related to precipitation and secondarily to temperature. However, we found that the dendroclimatic relationship depended on the position of the site along the climatic gradient. In the southern part of the distribution range, where temperature was generally higher and precipitation lower than the regional average, reduced growth was also associated with warm and dry conditions. In the northern part, where the average temperature was lower and the precipitation more abundant than the regional average, reduced growth was associated with cool conditions. Thus, our study highlights the substantial plasticity of Aleppo pine in response to different climatic conditions. These results do not resolve the source of response variability as being due to either genetic variation in provenance, to phenotypic plasticity, or a combination of factors. However, as current growth responses to inter-annual climate variability vary spatially across existing climate gradients, future climate-growth relationships will also likely be determined by differential adaptation and/or acclimation responses to spatial climatic variation. The contribution of local adaptation and/or phenotypic plasticity across populations to the persistence of species under global warming could be decisive for prediction of climate change impacts across populations. In this sense, a more complex forest dynamics modeling approach that includes the contribution of genetic variation and phenotypic plasticity can improve the reliability of the ecological inferences derived from the climate-growth relationships.This work was partially supported by Spanish Ministry of Education and Science co-funded by FEDER program (CGL2012-31668), the European Union and the National Ministry of Education and Religion of Greece (EPEAEK- Environment – Archimedes), the Slovenian Research Agency (program P4-0015), and the USDA Forest Service. The cooperation among international partners was supported by the COST Action FP1106, STREeSS

Long-term irrigation effects on Spanish holm oak growth and its black truffle symbiont

Ulf Büntgen...[et.al.]The Périgord black truffle is an exclusive culinary delicacy, but its Mediterranean harvests have declined, despite cultivation efforts since the 1970s. The role of long-term irrigation, symbiotic fungus-host interaction, and microbial belowground progression remain poorly understood, because generally too short experimental settings miss the necessary degree of real world complexity and reliable information from truffle orchards is limited. Here, we conduct the first dendrochronological and wood anatomical assessment of 295 holm oaks, which have been growing under different irrigation intensities in the world's largest truffle orchard in Spain. The relationships between different climatic variables (monthly temperature means and precipitation totals) and dendro-parameters (ring width, vessel count and vessel size) of the oak hosts are utilized to disentangle direct and indirect drivers of truffle fruit body production. Irrigation at medium - instead of high - intensity is most beneficial for oak growth. Non-irrigated trees reveal overall lower stem increments. Warmer temperatures from February to April and wetter conditions from May to July enhance host vitality and possibly also its interplay with fungi symbionts via increased fine root production and mycorrhizal colonization. Adequately irrigated Mediterranean orchards may counteract some of the drought-induced natural truffle decline, and help stabilizing rural tourism, regional agriculture and global markets. © 2014 Elsevier B.V.Supported by the WSL-internal DITREC project, the Ernst Göhner Foundation, the ClimFun project of the Norwegian RC (No. 225043), the project AGL2012-40035-C03 (Government of Spain), the project Micosylva+ (Interreg IVB SUDOE SOE3/P2/E533), the Government of Castilla y León, ARAID, the project Xilva (CGL2011-26654, Economy and Competitiveness Ministry), as well as the Operational Program of Education for Competitiveness of Ministry of Education, Youth and Sports of the Czech Republic (No. CZ.1.07/2.3.00/20.0248). We are particularly thankful to the Arotz Food Company for helping us to develop this research (via José Cuenca). José Miguel Altelarrea and to José Antonio Vega Borjabad (Cesefor Foundation) contributed to the fieldwork.Peer Reviewe

What drives growth of Scots pine in continental Mediterranean climates: Drought, low temperatures or both?

Sánchez-Salguero, R. et al.- 37 páginasScots pine forests subjected to continental Mediterranean climates undergo cold winter temperatures and drought stress. Recent climatic trends towards warmer and drier conditions across the Mediterranean Basin might render some of these pine populations more vulnerable to drought-induced growth decline at the Southernmost limit of the species distribution. We investigated how cold winters and dry growing seasons drive the radial growth of Scots pine subject to continental Mediterranean climates by relating growth to climate variables at local (elevational gradient) and regional (latitudinal gradient) scales. Local climate-growth relationships were quantified on different time scales (5-, 10- and 15-days) to evaluate the relative role of elevation and specific site characteristics. A negative water balance driven by high maximum temperatures in June (low-elevation sites) and July (high-elevation sites) was the major constraint on growth, particularly on a 5- to 10-day time scale. Warm nocturnal conditions in January were associated with wider rings at the high-elevation sites. At the regional scale, Scots pine growth mainly responded positively to July precipitation, with a stronger association at lower elevations and higher latitudes. January minimum temperatures showed similar patterns but played a secondary role as a driver of tree growth. The balance between positive and negative effects of summer precipitation and winter temperature on radial growth depends on elevation and latitude, with low-elevation populations being more prone to suffer drought and heat stress; whereas, high-elevation populations may be favoured by warmer winter conditions. This negative impact of summer heat and drought has increased during the past decades. This interaction between climate and site conditions and local adaptations is therefore decisive for the future performance and persistence of Scots pine populations in continental Mediterranean climates. Forecasting changes in the Scots pine range due to climate change should include this site-related information to obtain more realistic predictions, particularly in Mediterranean rear-edge areas. © 2015 Elsevier B.V.We were glad to be able to use the E-OBS dataset (EU- project ENSEMBLES) and the data providers in the ECA&D project (http://www.ecad.eu), and thank Dr. Geert Jan van Oldenborgh for his assistance with these data. We are grateful to the researchers who provided data on the International Tree-Ring Data Bank. This study was funded by the projects CoMo-ReAdapt (CGL2013-48843-C2-1-R) and FORRISK (Interreg IV B SUDOE 2007-2013). The authors thank CENEAM, Montes de Valsaín and OAPN and all participants involved in the International Tree-Ring Summer School carried out in 2012 in Valsaín (Segovia-Spain), especially to Kenza Garah, María Tabakova, Virginia Garófano-Gómez and Clara Rodriguez-Morata. R. Sánchez-Salguero is grateful for the financial support of University of Córdoba-Campus de Excelencia ceiA3 and “Fondo Europeo de Desarrollo Regional (FEDER)-Programa de Fortalecimiento de las capacidades en I + D + i de las Universidades 2014–2015, Junta de Andalucía”. We are grateful to Silvia Dingwall for the language review. This work has been carried out under the framework of the COST FP1106 network STReESS.Peer Reviewe

Mistletoe effects on Scots pine decline following drought events: Insights from within-tree spatial patterns, growth and carbohydrates

[EN] Forest decline has been attributed to the interaction of several stressors including biotic factors such as mistletoes and climate-induced drought stress. However, few data exist on how mistletoes are spatially arranged within trees and how this spatial pattern is related to changes in radial growth, responses to drought stress and carbon use. We used dendrochronology to quantify how mistletoe (Viscum album L.) infestation and drought stress affected long-term growth patterns in Pinus sylvestris L. at different heights. Basal area increment (BAI) trends and comparisons between trees of three different infestation degrees (without mistletoe, ID1; moderately infested trees, ID2; and severely infested trees, ID3) were performed using linear mixed-effects models. To identify the main climatic drivers of tree growth tree-ring widths were converted into indexed chronologies and related to climate data using correlation functions. We performed spatial analyses of the 3D distribution of mistletoe individuals and their ages within the crowns of three severely infested pines to describe their patterns. Lastly, we quantified carbohydrate and nitrogen concentrations in needles and sapwood of branches from severely infested trees and from trees without mistletoe. Mistletoe individuals formed strongly clustered groups of similar age within tree crowns and their age increased towards the crown apex. Mistletoe infestation negatively impacted growth but this effect was stronger near the tree apex than in the rest of sampled heights, causing an average loss of 64 in BAI (loss of BAI was ∼51 at 1.3m or near the tree base). We found that BAI of severely infested trees and moderately or non-infested trees diverged since 2001 and such divergence was magnified by drought. Infested trees had lower concentrations of soluble sugars in their needles than non-infested ones. We conclude that mistletoe infestation causes growth decline and increases the sensitivity of trees to drought stress. © The Author 2012. Published by Oxford University Press. All rights reserved.J.J.C. acknowledges collaborative efforts within the Globimed network (www.globimed.net). We thank the Spanish Meteorological Agency (AEMET) for providing climate data. We sincerely thank Melchor Maestro and Elena Lahoz for performing the chemical analyses. We thank the colleagues of the Forest Health Laboratory for their support (Mora de Rubielos, Gobierno de Aragón). We sincerely thank the three anonymous reviewers and the editor (M. Abrams) for their comments and improvements.Peer Reviewe