Dr. Elisabet Martinez Sancho

Forests are undergoing increasing risks of drought-induced tree mortality. Species replacement patterns following mortality may have a significant impact on the global carbon cycle. Among major hardwoods, deciduous oaks (Quercus spp.) are increasingly reported as replacing dying conifers across the Northern Hemisphere. Yet, our knowledge on the growth responses of these oaks to drought is incomplete, especially regarding post-drought legacy effects. The objectives of this study were to determine the occurrence, duration, and magnitude of legacy effects of extreme droughts and how that vary across species, sites, and drought characteristics. The legacy effects were quantified by the deviation of observed from expected radial growth indices in the period 1940–2016. We used stand-level chronologies from 458 sites and 21 oak species primarily from Europe, north-eastern America, and eastern Asia. We found that legacy effects of droughts could last from 1 to 5 years after the drought and were more prolonged in dry sites. Negative legacy effects (i.e., lower growth than expected) were more prevalent after repetitive droughts in dry sites. The effect of repetitive drought was stronger in Mediterranean oaks especially in Quercus faginea. Species-specific analyses revealed that Q. petraea and Q. macrocarpa from dry sites were more negatively affected by the droughts while growth of several oak species from mesic sites increased during post-drought years. Sites showing positive correlations to winter temperature showed little to no growth depression after drought, whereas sites with a positive correlation to previous summer water balance showed decreased growth. This may indicate that although winter warming favors tree growth during droughts, previous-year summer precipitation may predispose oak trees to current-year extreme droughts. Our results revealed a massive role of repetitive droughts in determining legacy effects and highlighted how growth sensitivity to climate, drought seasonality and species-specific traits drive the legacy effects in deciduous oak species.

Cellulose of tree rings is often assumed to be predominantly formed by direct assimilation of CO₂ by photosynthesis and consequently can be used to reconstruct past atmospheric ¹⁴C concentrations at annual resolution. Yet little is known about the extent and the age of stored carbon from previous years used in addition to the direct assimilation in tree rings. Here, we studied ¹⁴C in earlywood and latewood cellulose of four different species (oak, pine, larch and spruce), which are commonly used for radiocarbon calibration and dating. These trees were still growing during the radiocarbon bomb peak period (1958-1972). We compared cellulose ¹⁴C measured in tree-ring subdivisions with the atmospheric ¹⁴C corresponding to the time of ring formation. We observed that cellulose ¹⁴C carried up to about 50% of the atmospheric ¹⁴C signal from the previous 1-2 years only in the earlywood of oak, whereas in conifers it was up to 20% in the earlywood and in the case of spruce also in the latewood. The bias in using the full ring of trees growing in a temperate oceanic climate to estimate atmospheric ¹⁴C concentration might be minimal considering that earlywood has a low mass contribution and that the variability in atmospheric ¹⁴C over a few years is usually less than 3‰.

The vapor pressure deficit reflects the difference between how much moisture the atmosphere could and actually does hold, a factor that fundamentally affects evapotranspiration, ecosystem functioning, and vegetation carbon uptake. Its spatial variability and long-term trends under natural versus human-influenced climate are poorly known despite being essential for predicting future effects on natural ecosystems and human societies such as crop yield, wildfires, and health. Here we combine regionally distinct reconstructions of pre-industrial summer vapor pressure deficit variability from Europe’s largest oxygen-isotope network of tree-ring cellulose with observational records and Earth system model simulations with and without human forcing included. We demonstrate that an intensification of atmospheric drying during the recent decades across different European target regions is unprecedented in a pre-industrial context and that it is attributed to human influence with more than 98% probability. The magnitude of this trend is largest in Western and Central Europe, the Alps and Pyrenees region, and the smallest in southern Fennoscandia. In view of the extreme drought and compound events of the recent years, further atmospheric drying poses an enhanced risk to vegetation, specifically in the densely populated areas of the European temperate lowlands.

New mutations provide the raw material for evolution and adaptation. The distribution of fitness effects (DFE) describes the spectrum of effects of new mutations that can occur along a genome, and is, therefore, of vital interest in evolutionary biology. Recent work has uncovered striking similarities in the DFE between closely related species, prompting us to ask whether there is variation in the DFE among populations of the same species, or among species with different degrees of divergence, that is whether there is variation in the DFE at different levels of evolution. Using exome capture data from six tree species sampled across Europe we characterized the DFE for multiple species, and for each species, multiple populations, and investigated the factors potentially influencing the DFE, such as demography, population divergence, and genetic background. We find statistical support for the presence of variation in the DFE at the species level, even among relatively closely related species. However, we find very little difference at the population level, suggesting that differences in the DFE are primarily driven by deep features of species biology, and those evolutionarily recent events, such as demographic changes and local adaptation, have little impact.

• The oxygen isotope composition (δ¹⁸O) of tree-ring cellulose is used to evaluate tree physiological responses to climate, but their interpretation is still limited due to the complexity of the isotope fractionation pathways.
• We assessed the relative contribution of seasonal needle and xylem water δ¹⁸O variations to the intra-annual tree-ring cellulose δ¹⁸O signature of larch trees at two sites with contrasting soil water availability in the Swiss Alps. We combined biweekly δ¹⁸O measurements of soil water, needle water, and twig xylem water with intra-annual δ¹⁸O measurements of tree-ring cellulose, xylogenesis analysis, and mechanistic and structural equation modeling.
• Intra-annual cellulose δ¹⁸O values resembled source water δ¹⁸O mean levels better than needle water δ¹⁸O. Large parts of the rings were formed under high proportional exchange with unenriched xylem water (p_ex). Maximum p_ex values were achieved in August and imprinted on sections at 50–75% of the ring. High p_ex values were associated with periods of high atmospheric evaporative demand (VPD). While VPD governed needle water δ¹⁸O variability, we estimated a limited Péclet effect at both sites.
• Due to a variable p_ex, source water has a strong influence over large parts of the intra-annual tree-ring cellulose δ¹⁸O variations, potentially masking signals coming from needle-level processes.

The European Dendroecological Fieldweek (EDF) is a one-week course that takes place every year at varying locations in Europe according to the principle "Bring tree-ring research to the people". The EDF welcomes early-career to advanced researchers, but also forest service and other federal agency employees and private people interested in tree-ring sciences from all over the world. It encompasses a large spectrum of dendrochronological field, laboratory and data analysis methods and scientific fields including climatology, ecology, physiology, geomorphology and archaeology. Multiple scales of observations from the individual cell to the ecosystem level and from seasonal to multi-centennial periods are covered. Work on mini research projects in topic groups alternates with keynote lectures and individual participants’ presentations.
As one of the first in-person tree-ring meetings since the start of the COVID-19 pandemic the 31st EDF was held in summer 2021 in Val Müstair, Switzerland. Topics included i) Tree age and climate sensitivity of a relict, old-growth Scots pine stand, ii) Blue intensity-based climate sensitivity of Norway spruce growth, iii) Tree rings as indicators of grey larch budmoth outbreaks, iv) Growth of larch trees along an abandoned irrigation channel, v) Wood anatomical characteristics of two alpine creeping shrub species, and vi) Historical dating of a stable and a residential house. Alongside with their educational value these projects allowed novel insight into the age structure and growth dynamics of the sub-alpine forests and beyond in the valley and provided valuable outcome to the local stakeholders such as the Nature Park Biosfera Val Müstair, the local forest service and the public of Val Müstair.
Under hindered conditions due to the pandemic, the 31st EDF still demonstrated its strength as an international educational and interdisciplinary scientific field and lab course, combining teaching with the application of cutting-edge technologies.

The mechanistic pathways connecting ocean-atmosphere variability and terrestrial productivity are well-established theoretically, but remain challenging to quantify empirically. Such quantification will greatly improve the assessment and prediction of changes in terrestrial carbon sequestration in response to dynamically induced climatic extremes. The jet stream latitude (JSL) over the North Atlantic-European domain provides a synthetic and robust physical framework that integrates climate variability not accounted for by atmospheric circulation patterns alone. Surface climate impacts of north-south summer JSL displacements are not uniform across Europe, but rather create a northwestern-southeastern dipole in forest productivity and radial-growth anomalies. Summer JSL variability over the eastern North Atlantic-European domain (5-40E) exerts the strongest impact on European beech, inducing anomalies of up to 30% in modelled gross primary productivity and 50% in radial tree growth. The net effects of JSL movements on terrestrial carbon fluxes depend on forest density, carbon stocks, and productivity imbalances across biogeographic regions.

Klimawandel, Trockenheit, Tannensterben: Forscherinnen und Forscher debattieren über die Zukunftsaussichten der Weisstanne. Jüngste Forschungsergebnisse deuten an, dass diese Baumart grösseres Potenzial hat als oft angenommen.

Changement climatique, sécheresse et dépérissement du sapin suscitent le débat scientifique sur les perspectives d’avenir de cette essence. Des résultats récents suggèrent que son potentiel est meilleur qu’on ne le pense.

• The impact of climate extremes on forest ecosystems is poorly understood but important for predicting carbon and water cycle feedbacks to climate. Some knowledge gaps still remain regarding how drought-related adjustments in intra-annual tree-ring characteristics directly impact tree carbon and water use.
In this, study we quantified the impact of an extreme summer drought on the water-use efficiency and carbon sequestration of four mature Norway spruce trees. We used detailed observations of wood formation (xylogenesis) and intra-annual tree-ring properties (quantitative wood anatomy and stable carbon isotopes) combined with physiological water-stress monitoring.
• During 41 days of tree water deficit, we observed an increase in ¹³C but a reduction in cell enlargement and wall thickening processes, which impacted the anatomical characteristics. These adjustments diminished carbon sequestration by 67% despite an 11% increase in water-use efficiency during drought. However, with the resumption of a positive hydric state in the stem, we observed a fast recovery of cell formation rates based on the accumulated assimilates produced during drought.
• Our findings enhance our understanding of carbon and water fluxes between the atmosphere and forest ecosystems, providing observational evidence on the tree intra-annual carbon sequestration and water-use efficiency dynamics to improve future generations of vegetation models.

This is the first Europe-wide comprehensive assessment of the climatological and physiological information recorded by hydrogen isotope ratios in tree-ring cellulose (δ²H_c) based on a unique collection of annually resolved 100-year tree-ring records of two genera (Pinus and Quercus) from 17 sites (36°N to 68°N). We observed that the high-frequency climate signals in the δ²H_c chronologies were weaker than those recorded in carbon (δ¹³C_c) and oxygen isotope signals (δ¹⁸O_c) but similar to the tree-ring width ones (TRW). The δ²H_c climate signal strength varied across the continent and was stronger and more consistent for Pinus than for Quercus. For both genera, years with extremely dry summer conditions caused a significant ²H-enrichment in tree-ring cellulose.
The δ²H_c inter-annual variability was strongly site-specific, as a result of the imprinting of climate and hydrology, but also physiological mechanisms and tree growth. To differentiate between environmental and physiological signals in δ²H_c, we investigated its relationships with δ¹⁸O_c and TRW. We found significant negative relationships between δ²H_c and TRW (7 sites), and positive ones between δ²H_c and δ¹⁸O_c (10 sites). The strength of these relationships was nonlinearly related to temperature and precipitation. Mechanistic δ²H_c models performed well for both genera at continental scale simulating average values, but they failed on capturing year-to-year δ²H_c variations. Our results suggest that the information recorded by δ²H_c is significantly different from that of δ¹⁸O_c, and has a stronger physiological component independent from climate, possibly related to the use of carbohydrate reserves for growth. Advancements in the understanding of ²H-fractionations and their relationships with climate, physiology, and species-specific traits are needed to improve the modelling and interpretation accuracy of δ²H_c. Such advancements could lead to new insights into trees' carbon allocation mechanisms, and responses to abiotic and biotic stress conditions.

1. Plant species that grow across environmental gradients show a range of trait expression, but traits do not vary independently. In fact, phenotypes are integrated expressions of multiple traits that covary due to trade‐offs among functions and processes. Understanding trait covariation structures will ultimately help with predicting species' responses to change and developing management actions.
2. We measured variation and covariation (a proxy of phenotypic integration) among functional traits of Pinus sylvestris from paired populations across its European distribution. Populations within a pair were close enough to be in gene flow contact but located in contrasting environmental conditions across a local gradient. Measured traits represented three axes of variation (groups of traits) related to a tree's competitive ability and the trade‐off between resource acquisition and conservation, namely plant size measures and stem and foliar traits.
3. Results revealed important intra‐ and inter‐population trait variability. In particular, at the population level, trait means shifted across the climatic gradient mainly described by mean annual temperature. Moreover, we found a higher degree of trait covariation in populations under harsher environments (i.e. lower environmental suitability for the species). This pattern was consistent within population pairs, suggesting that higher trait covariation may be adaptive, being more coordinated in sites with harsher conditions. At larger spatial scales, we found a less conclusive pattern with a trend of increasing covariation at the northern edge of the species distribution. This result suggests that at larger scales different processes may be involved in the trade‐off between the adaptive value of phenotypic covariation versus its constraints on trait combinations that may limit plant's response capability.
4. Synthesis. Trait covariation varies at different spatial scales, increasing under harsher conditions, and the robustness and repeatability of this pattern suggests its adaptive role for the species' responses to different environments.

Motivation: Trait variation within species can reveal plastic and/or genetic responses to environmental gradients, and may indicate where local adaptation has occurred. Here, we present a dataset of rangewide variation in leaf traits from seven of the most ecologically and economically important tree species in Europe. Sample collection and trait assessment are embedded in the GenTree project (EU‐Horizon 2020), which aims at characterizing the genetic and phenotypic variability of forest tree species to optimize the management and sustainable use of forest genetic resources. Our dataset captures substantial intra‐ and interspecific leaf phenotypic variability, and provides valuable information for studying the relationship between ecosystem functioning and trait variability of individuals, and the response and resilience of species to environmental changes.
Main types of variable contained: We chose morphological and chemical characters linked to trade‐offs between acquisition and conservation of resources and water use, namely specific leaf area, leaf size, carbon and nitrogen content and their ratio, and the isotopic signature of stable isotope ¹³C and ¹⁵N in leaves.
Spatial location and grain: We surveyed between 18 and 22 populations per species, 141 in total, across Europe.
Time period: Leaf sampling took place between 2016 and 2017.
Major taxa and level of measurement: We sampled at least 25 individuals in each population, 3,569 trees in total, and measured traits in 35,755 leaves from seven European tree species, i.e. the conifers Picea abies, Pinus pinaster and Pinus sylvestris, and the broadleaves Betula pendula, Fagus sylvatica, Populus nigra and Quercus petraea.
Software format: The data files are in ASCII text, tab delimited, not compressed.

Recent studies have identified strong relationships between delayed recovery of tree growth after drought and tree mortality caused by subsequent droughts. These observations raise concerns about forest ecosystem services and post-drought growth recovery given the projected increase in drought frequency and extremes. For quantifying the impact of extreme droughts on tree radial growth, we used a network of tree-ring width data of 1689 trees from 100 sites representing most of the distribution of two drought tolerant, deciduous oak species (Quercus petraea and Quercus robur). We first examined which climatic factors and seasons control growth of the two species and if there is any latitudinal, longitudinal or elevational trend. We then quantified the relative departure from pre-drought growth during droughts, and how fast trees were able to recover the pre-drought growth level. Our results showed that growth was more related to precipitation and climatic water balance (precipitation minus potential evapotranspiration) than to temperature. However, we did not detect any clear latitudinal, longitudinal or elevational trends except a decreasing influence of summer water balance on growth of Q. petraea with latitude. Neither species was able to maintain the pre-drought growth level during droughts. However, both species showed rapid recovery or even growth compensation after summer droughts but displayed slow recovery in response to spring droughts where none of the two species was able to fully recover the pre-drought growth-level over the three post-drought years. Collectively, our results indicate that oaks which are considered resilient to extreme droughts have also shown vulnerability when droughts occurred in spring especially at sites where long-term growth is not significantly correlated with climatic factors. This improved understanding of the role of drought seasonality and climate sensitivity of sites is key to better predict trajectories of post-drought growth recovery in response to the drier climate projected for Europe.

In this chapter we introduce the climate signal in stable isotope tree-ring records, with the emphasis on temperate forests. The development of the subdiscipline is recapped followed by an exploration of isotope dendroclimatic records by geography and, broadly, by isotopic species. Whilst there are still questions to be answered around signal strength and age-related effects in different environments and in different species, the proxy is now contributing to palaeoclimatology in a far greater way than in the days of the first hints of ‘isotope tree thermometers’. We include two summary tables. Table 19.1 exemplifies the range of climate information available from stable carbon isotope time series and Table 19.2 explores oxygen isotope proxy signals. Due to the greater complexity seen in stable carbon isotope interpretations we explore response groupings with example references given for each category of proxy response. Finally, we summarize the state of the art in isotope dendroclimatology and discuss possible future directions.

Temperature and precipitation variability throughout the year control the intra-annual dynamics of tree-ring formation. Physiological adaptation of trees to climate change is among the key issues to better understand and predict future forest performance and composition. In this study, we investigated the species' coexistence and performance of Scots pine and pubescent oak growing in a mixed sub-Mediterranean forest in the northeast of the Iberian Peninsula. We assessed intra-annual cumulative growth patterns derived from band dendrometers during four consecutive growing seasons and long-term changes in basal area increment for the period 1950-2014. Our results revealed that Scots pine followed an intra-annual bimodal growth pattern. Scots pine growth was mainly limited by water availability at intra-annual, interannual and decadal time scales, which resulted in a negative long-term growth trend. Conversely, oak displayed a unimodal growth pattern, which was less climatically constrained. A significant increase in basal area of oak denotes an overall better potential acclimation to prevailing climatic conditions at the expenses of a higher risk of physiological failure during extreme climate events.

Warmer climate and more frequent extreme droughts will pose major threats to forest ecosystems. Past demography processes due to post-glacial recolonization and adaptation to local environmental conditions are among the main contributors to genetic differentiation processes among provenances. Assessing the intra-specific variability of tree growth responses to such changes is crucial to explore a species' potential to cope with climate warming. We combined growth-related traits derived from tree-ring width series with neutral genetic information of 18 European provenances of silver fir (Abies alba Mill.) growing in two common garden experiments in Switzerland.
Analyses based on neutral single nucleotide polymorphisms revealed that the studied provenances grouped into three longitudinal clusters. These three genetic clusters showed differences in growth traits (height and DBH), with the provenances from the eastern cluster exhibiting the highest growth. The Pyrenees cluster showed significantly lower recovery and resilience to the extreme drought of 2003 as well as lower values of growth autocorrelation. Q_ST-F_ST and correlation analyses with climate of provenance origin suggest that the differences among provenances found in some traits result from natural selection.
Our study suggests that the last post-glacial re-colonization and natural selection are the major drivers explaining the intra-specific variability in growth of silver fir across Europe. These findings highlight the importance of combining dendroecology and genetic analyses on fitness-related traits to assess the potential of a species to cope with global environmental change and provide insights to support assisted gene flow to ensure the persistence of the species in European forests.

Background: Progress in the field of evolutionary forest ecology has been hampered by the huge challenge of phenotyping trees across their ranges in their natural environments, and the limitation in high-resolution environmental information. Findings: The GenTree Platform contains phenotypic and environmental data from 4,959 trees from 12 ecologically and economically important European forest tree species: Abies alba Mill. (silver fir), Betula pendula Roth. (silver birch), Fagus sylvatica L. (European beech), Picea abies (L.) H. Karst (Norway spruce), Pinus cembra L. (Swiss stone pine), Pinus halepensis Mill. (Aleppo pine), Pinus nigra Arnold (European black pine), Pinus pinaster Aiton (maritime pine), Pinus sylvestris L. (Scots pine), Populus nigra L. (European black poplar), Taxus baccata L. (English yew), and Quercus petraea (Matt.) Liebl. (sessile oak). Phenotypic (height, diameter at breast height, crown size, bark thickness, biomass, straightness, forking, branch angle, fructification), regeneration, environmental in situ measurements (soil depth, vegetation cover, competition indices), and environmental modeling data extracted by using bilinear interpolation accounting for surrounding conditions of each tree (precipitation, temperature, insolation, drought indices) were obtained from trees in 194 sites covering the species’ geographic ranges and reflecting local environmental gradients. Conclusion: The GenTree Platform is a new resource for investigating ecological and evolutionary processes in forest trees. The coherent phenotyping and environmental characterization across 12 species in their European ranges allow for a wide range of analyses from forest ecologists, conservationists, and macro-ecologists. Also, the data here presented can be linked to the GenTree Dendroecological collection, the GenTree Leaf Trait collection, and the GenTree Genomic collection presented elsewhere, which together build the largest evolutionary forest ecology data collection available.

During the last 25 years, the dendrochronology community has benefited from the application of stable carbon isotopes in tree rings to answer a number of research questions, which has resulted in an exponential burst in the number of publications related to this discipline. Here, we perform a literature review of the most influential topics (scope, methodology, targeted species, and climatic signals) that have shaped the research agenda and its impact on the scholarly output over the recent decades. Based on a total of 550 publications, we observed that: (1) conifers are the most investigated tree functional type, being included in over two-thirds of studies; (2) although being a subject of debate, the material of choice for carbon isotope analyses is still cellulose, for which the extraction procedure has been refined with methodological improvements over the years; (3) a shift in the application of carbon isotopes has occurred from proxies of climatic information for paleoclimatic reconstructions to tools for the understanding of short- and long-term tree functioning; and (4) such shift in research scope is apparently linked to an increasing number of publications showing drought-related signals governing isotope time series in the context of current climate change. Besides, we also assessed the number and outcome of publications analyzing ¹³C-derived intrinsic water-use efficiency (WUE_i) trends within the framework formulated by Saurer et al. (2004). We found that leaf intercellular CO₂ concentration responding proportionally to the increase in atmospheric CO₂ is the most common scenario reported across continents, but with changing importance relative to other scenarios depending on the biome examined. We also detected that climate plays a dominant role over any potential CO₂ fertilization linked to increasing WUE_i. Looking for new and thrilling research avenues, future studies should further examine δ¹³C fluctuations at intra-annual resolution, explore its links to wood characteristics, and disentangle δ¹³C signals of the distinct wood constituents for a better understanding of tree functioning at varying spatiotemporal scales, also benchmarking vegetation models of different complexities.