Winter drought impairs xylem phenology, anatomy and growth in Mediterranean Scots pine forests

39 páginas, 5 tablas, 6 figuras, 2 figuras suplementarias. -- This is a pre-copyedited, author-produced version of an article accepted for publication in Tree Physiology following peer review. The version of record [J. J. Camarero, G. Guada, R. Sánchez-Salguero, E. Cervantes; Winter drought impairs xylem phenology, anatomy and growth in Mediterranean Scots pine forests. Tree Physiol 2016; 36 (12): 1536-1549] is available online at: http://dx.doi.org/10.1093/treephys/tpw077Continental Mediterranean forests face drought and cold stress. Drought and cold spells are climate extremes which can impair the recovery and resilience capacity of some Mediterranean forests. Climate warming could amplify the negative effects on forests of climate extremes by altering phenology phases (premature dehardening) and by magnifying drought stress. Here we capitalize on a winter-drought induced dieback triggered by a cold spell which occurred in December 2001 and affected Scots pine forests in E. Spain. We assessed post-dieback recovery by quantifying and comparing radial growth and xylem anatomy of non-declining (crown cover > 50%) and declining (crown cover ≤ 50%) trees in two sites (VP, Villarroya de los Pinares; TO, Torrijas). We also characterized aboveground productivity in site VP and xylogenesis in both sites during 2005. Dieback caused legacy effects since needle loss, a 60% reduction in aboveground biomass and radial-growth decline characterized declining trees three years after these dieback symptoms started in spring 2002. Declining trees formed collapsed tracheids in the 2002-ring earlywood, particularly in the most affected VP site where xylogenesis differences between non-declining and declining trees were most noticeable. The lower growth rates of declining trees were explained by a shorter duration of their major xylogenesis phases. In the site VP the radial-enlargement and wall-thickening of tracheids were significantly reduced in declining trees as compared to non-declining trees, and this was because these phases tended to start earlier and end later in non-declining than in declining trees. Gompertz models fitted to tracheid production predicted that maximum growth rates occurred 11-12 days earlier in non-declining than in declining trees. The formation of radially-enlarging tracheids was enhanced by longer days in both study sites and also by wetter conditions in the driest TO site, but this xylogenesis sensitivity to climate was reduced in declining trees. Winter-drought induced dieback impairs xylem anatomy and phenology, aboveground productivity, xylogenesis and growth in Mediterranean Scots pine populations. Affected stands show a costly post-dieback recovery challenging their resilience ability.JJC acknowledges the support of the Excellence Network “Red de Ecología Terrestre para afrontar los retos del Cambio Global—ECOMETAS” (CGL2014-53840-REDT) of the Spanish Ministry of Economy.Peer reviewe

Blue is the fashion in Mediterranean pines: New drought signals from tree-ring density in southern Europe

Identificador de proyecto: Marie Sklodowska-Curie grant agreement No 101029581Long-term records of tree-ring width (TRW), latewood maximum density (MXD) and blue intensity (BI) measurements on conifers have been largely used to develop high-resolution temperature reconstructions in cool temperate forests. However, the potential of latewood blue intensity (LWBI), less commonly used earlywood blue intensity (EWBI), and delta (difference between EWBI and LWBI, dBI) blue intensity in Mediterranean tree species is still unexplored. Here we developed BI chronologies in moist-elevation limits of the most southwestern European distribution of Pinus nigra subsp. salzmanii Arnold. We tested whether BI variables derived from tree rings of black pine are better proxies than ring-width variables to reconstruct long-term changes in climatic factors and water availability. For this we applied correlations and regression analyses with daily and monthly climate data, a spatial and temporal drought index (Standardized Precipitation-Evapotranspiration Index-SPEI) and Vapour Pressure Deficit (VPD), as well as atmospheric circulation patterns: North Atlantic Oscillation (NAO), Southern Oscillation Index (SOI) and Western Mediterranean Oscillation (WeMO). We found a positive relation between black pine growth (RW) and temperature during the winter preceding the growing season. Among all variables LWBI and dBI were found to be more sensitive than TRW to SPEI at low-elevation site, with EWBI series containing an opposite climatic signal. LWBI and dBI were significantly related to June and September precipitation at high-elevation site. Winter VPD was related with higher EWI and LWI series, whereas dBI and EWBI were related with January SOI and February NAO. We confirm the potential of long-term dBI series to reconstruct climate in drought-prone regions. This novel study in combination with other wood anatomical measurements has wide implications for further use of BI to understand and reconstruct environmental changes in Mediterranean conifer forests.Wageningen University & ResearchUniversidade de Santiago de CompostelaUniversity of Amsterda

Wood anatomy and tree growth covary in riparian ash forests along climatic and ecological gradients

Riparian ash forests subjected to seasonal drought are among the most endangered ecosystems in Europe. They are threatened by climate warming causing aridification and by land-use changes modifying river flow. To assess the impacts of these two stress factors on riparian forests, we studied radial growth and xylem anatomical traits in five narrow-leaved ash (Fraxinus angustifolia) stands across wide climatic and ecological gradients from northern Italy to southern Portugal. Radial growth rates and earlywood hydraulic diameter (Dh) were directly correlated, whilst earlywood vessel density and growth rates were inversely associated. Ash growth positively responded to precipitation. Higher and lower rates of growth increase in response to precipitation were found in dry (annual precipitation 357–750 mm, annual water balance -39 to -48 mm) and wet (annual precipitation 1030 mm, annual water balance 27 mm) sites, respectively. Wet conditions in autumn and winter of the year prior to tree-ring formation lead to larger Dh values, except in the wet site where warmer conditions from prior autumn to current spring were positively associated to wider vessels. Growth was also enhanced by a higher river flow, reflecting higher soil moisture due to elevated groundwater table levels. Peaks in river flow from late winter to early spring increased Dh in dry-continental sites. Growth and potential hydraulic conductivity in drought-prone riparian ash forests are differently impacted by climate variability and river flow depending on site and hydrological conditions. Nevertheless, covariation between radial growth and the earlywood vessel diameter was found, regardless of site specific differences. Wood production and hydraulic conductivity are coordinated through the production of large earlywood vessels which may allow reaching higher growth rates. © 2021 The Author

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

Scots 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

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)

Contrasting growth and water use efficiency after thinning in mixed Abies pinsapo–Pinus pinaster–Pinus sylvestris forests

Foresters frequently lack sufficient information about thinning intensity effects to optimize semi-natural forest management and their effects and interaction with climate are still poorly understood. In an Abies pinsapo–Pinus pinaster–Pinus sylvestris forest with three thinning intensities, a dendrochronologial approach was used to evaluate the short-term responses of basal area increment (BAI), carbon isotope (δ13C) and water use efficiency (iWUE) to thinning intensity and climate. Thinning generally increased BAI in all species, except for the heavy thinning in P. sylvestris. Across all the plots, thinning increased 13C-derived water-use efficiency on average by 14.49% for A. pinsapo, 9.78% for P. sylvestris and 6.68% for P. pinaster, but through different ecophysiological mechanisms. Our findings provide a robust mean of predicting water use efficiency responses from three coniferous species exposed to different thinning strategies which have been modulated by climatic conditions over time

When a tree dies in the forest : scaling climate-driven tree mortality to ecosystem water and carbon fluxes

Altres ajuts: COST FP1106 network STReESS.Drought- and heat-driven tree mortality, along with associated insect outbreaks, have been observed globally in recent decades and are expected to increase in future climates. Despite its potential to profoundly alter ecosystem carbon and water cycles, how tree mortality scales up to ecosystem functions and fluxes is uncertain. We describe a framework for this scaling where the effects of mortality are a function of the mortality attributes, such as spatial clustering and functional role of the trees killed, and ecosystem properties, such as productivity and diversity. We draw upon remote-sensing data and ecosystem flux data to illustrate this framework and place climate-driven tree mortality in the context of other major disturbances. We find that emerging evidence suggests that climate-driven tree mortality impacts may be relatively small and recovery times are remarkably fast (~4 years for net ecosystem production). We review the key processes in ecosystem models necessary to simulate the effects of mortality on ecosystem fluxes and highlight key research gaps in modeling. Overall, our results highlight the key axes of variation needed for better monitoring and modeling of the impacts of tree mortality and provide a foundation for including climate-driven tree mortality in a disturbance framework