Aim
Plant growth and phenology respond plastically to changing climatic conditions in both space and time. Species‐specific levels of growth plasticity determine biogeographical patterns and the ...adaptive capacity of species to climate change. However, a direct assessment of spatial and temporal variability in radial growth dynamics is complicated, because long records of cambial phenology do not exist.
Location
Sixteen sites across European distribution margins of Juniperus communis L. (the Mediterranean, the Arctic, the Alps and the Urals).
Time period
1940–2016.
Major taxa studied
Juniperus communis.
Methods
We applied the Vaganov–Shashkin process‐based model of wood formation to estimate trends in growing season duration and growth kinetics since 1940. We assumed that J. communis would exhibit spatially and temporally variable growth patterns reflecting local climatic conditions.
Results
Our simulations indicate regional differences in growth dynamics and plastic responses to climate warming. The mean growing season duration is the longest at Mediterranean sites and, recently, there has been a significant trend towards its extension of up to 0.44 days/year. However, this stimulating effect of a longer growing season is counteracted by declining summer growth rates caused by amplified drought stress. Consequently, overall trends in simulated ring widths are marginal in the Mediterranean. In contrast, durations of growing seasons in the Arctic show lower and mostly non‐significant trends. However, spring and summer growth rates follow increasing temperatures, leading to a growth increase of up to 0.32 %/year.
Main conclusions
This study highlights the plasticity in growth phenology of widely distributed shrubs to climate warming: an earlier onset of cambial activity that offsets the negative effects of summer droughts in the Mediterranean and, conversely, an intensification of growth rates during the short growing seasons in the Arctic. Such plastic growth responsiveness allows woody plants to adapt to the local pace of climate change.
Severe droughts have the potential to reduce forest productivity and trigger tree mortality. Most trees face several drought events during their life and therefore resilience to dry conditions may be ...crucial to long-term survival. We assessed how growth resilience to severe droughts, including its components resistance and recovery, is related to the ability to survive future droughts by using a tree-ring database of surviving and now-dead trees from 118 sites (22 species, >3,500 trees). We found that, across the variety of regions and species sampled, trees that died during water shortages were less resilient to previous non-lethal droughts, relative to coexisting surviving trees of the same species. In angiosperms, drought-related mortality risk is associated with lower resistance (low capacity to reduce impact of the initial drought), while it is related to reduced recovery (low capacity to attain pre-drought growth rates) in gymnosperms. The different resilience strategies in these two taxonomic groups open new avenues to improve our understanding and prediction of drought-induced mortality.
Hydraulic impairment due to xylem embolism and carbon starvation are the two proposed mechanisms explaining drought‐induced forest dieback and tree death. Here, we evaluate the relative role played ...by these two mechanisms in the long‐term by quantifying wood‐anatomical traits (tracheid size and area of parenchyma rays) and estimating the intrinsic water‐use efficiency (iWUE) from carbon isotopic discrimination. We selected silver fir and Scots pine stands in NE Spain with ongoing dieback processes and compared trees showing contrasting vigour (declining vs nondeclining trees). In both species earlywood tracheids in declining trees showed smaller lumen area with thicker cell wall, inducing a lower theoretical hydraulic conductivity. Parenchyma ray area was similar between the two vigour classes. Wet spring and summer conditions promoted the formation of larger lumen areas, particularly in the case of nondeclining trees. Declining silver firs presented a lower iWUE than conspecific nondeclining trees, but the reverse pattern was observed in Scots pine. The described patterns in wood anatomical traits and iWUE are coherent with a long‐lasting deterioration of the hydraulic system in declining trees prior to their dieback. Retrospective quantifications of lumen area permit to forecast dieback in declining trees 2–5 decades before growth decline started. Wood anatomical traits provide a robust tool to reconstruct the long‐term capacity of trees to withstand drought‐induced dieback.
Long‐term tree recruitment dynamics of subalpine forests mainly depend on temperature changes, but little is known about the feedbacks between historical land use and climate. Here, we analyze a ...southern European, millennium‐long dataset of tree recruitment from three high‐elevation pine forests located in Mediterranean mountains (Pyrenees, northeastern Spain; Pollino, southern Italy; and Mt. Smolikas, northern Greece). We identify synchronized recruitment peaks in the late 15th and early 16th centuries, following prolonged periods of societal and climate instability. Major European population crises in the 14th and 15th centuries associated with recurrent famines, the Black Death pandemic, and political turmoil are likely to have reduced the deforestation of subalpine environments and caused widespread rewilding. We suggest that a distinct cold phase in the Little Ice Age around 1450 ce could also have accelerated the cessation of grazing pressure, particularly in the Pyrenees, where the demographic crisis was less severe. Most pronounced in the Pyrenees, the enhanced pine recruitment from around 1500–1550 ce coincides with temporarily warmer temperatures associated with a positive phase of the North Atlantic Oscillation. We diagnose that a mixture of human and climate factors has influenced past forest recruitment dynamics in Mediterranean subalpine ecosystems. Our results highlight how complex human–climate interactions shaped forest dynamics during pre‐industrial times and provide historical analogies to recent rewilding.
We analyse a southern European, millennium‐long dataset of tree recruitment from three high‐elevation pine forests located in Mediterranean mountains. We identify synchronized recruitment peaks in the late 15th and early 16th centuries, following prolonged periods of societal and climate instability. Major European population crises in the 14th and 15th centuries associated with recurrent famines, the Black Death pandemic, and political turmoil are likely to have reduced the deforestation of subalpine environments and caused widespread rewilding.
1. Mixed conifer-hardwood forests can be more productive than pure forests and they are increasingly considered as ecosystems that could provide adaptation strategies in the face of global change. ...However, the combined effects of tree-to-tree competition, rising atmospheric CO₂ concentrations and climate on such mixtures remain poorly characterized and understood. 2. To fill this research gap, we reconstructed 34-year series (1980-2013) of growth (basal area increment, BAI) and intrinsic water-use efficiency (iWUE) of Scots pine (Pinus sylvestris L.)-European beech (Fagus sylvatica L.) mixed stands at two climatically contrasting sites located in the southwestern Pyrenees. We also gathered data on tree-to-tree competition and climate variables in order to test the hypotheses that (1) radial growth will be greater when exposed to inter- than to intraspecific competition, that is, when species complementarity occurs and (2) enhanced iWUE could be linked to improved stem radial growth. 3. Growth of both species was reduced when intraspecific competition increased. Species complementarity was linked to improved growth of Scots pine at the continental site, while competition overrode any complementarity advantage at the drought-prone Mediterranean site. Beech growth did not show any significant response to pine admixture likely due to shade tolerance and the highly competitive nature of this species. Increasing interspecific competition drove recent iWUE changes, which increased in Scots pine but decreased in European beech. The iWUE enhancement did not involve any growth improvement in Scots pine. However, the positive BAI-iWUE relationship found for beech suggests an enhanced beech growth in drought-prone sites due to improved water use. 4. Synthesis. Complementarity may enhance growth in mixed forests. However, water scarcity can constrict light-related complementarity for shade intolerant species (Scots pine) in drought-prone sites. Basal area increment-intrinsic water-use efficiency relationships were negative for Scots pine and positive for European beech. These contrasting behaviours have got implications for coping with the expected increasing drought events in Scots pine-European beech mixtures located near the ecological limit of the two species. Complementarity effects between tree species should be considered to avoid overestimating the degree of future carbon uptake by mixed conifer-broadleaf forests.
Theory predicts that the postindustrial rise in the concentration of CO₂in 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 cₐor 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 cₐby 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 cₐin 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 cₐon 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 cₐdid not compensate for the reduction in water availability. Thus, warming‐triggered drought stress may become a more important direct driver of growth than rising cₐin similar subalpine forests. We argue that broad approaches in biogeographical and temporal terms are required to adequately evaluate any effect of rising cₐon forest growth.
Drought-triggered declines in forest productivity and associated die-off events have increased considerably due to climate warming in the last decades. There is an increasing interest in quantifying ...the resilience capacity of forests against climate warming and drought to uncover how different stands and tree species will resist and recover after more frequent and intense droughts. Trees form annual growth rings that represent an accurate record of how forest growth responded to past droughts. Here we use dendrochronology to quantify the radial growth of different forests subjected to contrasting climatic conditions in Spain during the last half century. Particularly, we considered four climatically contrasting areas where dominant forests showed clear signs of drought-induced dieback. Studied forests included wet sites dominated by silver fir (Abies alba) in the Pyrenees and beech (Fagus sylvatica) stands in northern Spain, and drought-prone sites dominated by Scots pine (Pinus sylvestris) in eastern Spain and black pine (Pinus nigra) in the semi-arid south-eastern Spain. We quantified the growth reduction caused by different droughts and assessed the short-and long-term resilience capacity of declining vs. non-declining trees in each forest. In all cases, drought induced a marked growth reduction regardless tree vigor. However, the capacity to recover after drought (resilience) at short- and long-term scales varied greatly between declining and non-declining individuals. In the case of beech and silver fir, non-declining individuals presented greater growth rates and capacity to recover after drought than declining individuals. For Scots pine, the resilience to drought was found to be lower in recent years regardless the tree vigor, but the growth reduction caused by successive droughts was more pronounced in declining than in non-declining individuals. In the black pine forest an extreme drought induced a marked growth reduction in declining individuals when accounting for age effects on growth rates. We demonstrate the potential of tree-ring data to record short- and long-term impacts of drought on forest growth and to quantify the resilience capacity of trees.
Legacies of past climate conditions and historical management govern forest productivity and tree growth. Understanding how these processes interact and the timescales over which they influence tree ...growth is critical to assess forest vulnerability to climate change. Yet, few studies address this issue, likely because integrated long-term records of both growth and forest management are uncommon. We applied the stochastic antecedent modelling (SAM) framework to annual tree-ring widths from mixed forests to recover the ecological memory of tree growth. We quantified the effects of antecedent temperature and precipitation up to 4 years preceding the year of ring formation and integrated management effects with records of harvesting intensity from historical forest management archives. The SAM approach uncovered important time periods most influential to growth, typically the warmer and drier months or seasons, but variation among species and sites emerged. Silver fir responded primarily to past climate conditions (25–50 months prior to the year of ring formation), while European beech and Scots pine responded mostly to climate conditions during the year of ring formation and the previous year, although these responses varied among sites. Past management and climate interacted in such a way that harvesting promoted growth in young silver fir under wet and warm conditions and in old European beech under drier and cooler conditions. Our study shows that the ecological memory associated with climate legacies and historical forest management is species-specific and context-dependent, suggesting that both aspects are needed to properly evaluate forest functioning under climate change.
Climate warming is driving an advance of leaf unfolding date in temperate deciduous forests, promoting longer growing seasons and higher carbon gains. However, an earlier leaf phenology also ...increases the risk of late frost defoliation (LFD) events. Compiling the spatiotemporal patterns of defoliations caused by spring frost events is critical to unveil whether the balance between the current advance in leaf unfolding dates and the frequency of LFD occurrence is changing and represents a threaten for the future viability and persistence of deciduous forests. We combined satellite imagery with machine learning techniques to reconstruct the spatiotemporal patterns of LFD events for the 2003–2018 period in the Iberian range of European beech (Fagus sylvatica), at the drier distribution edge of the species. We used MODIS Vegetation Index Products to generate a Normalized Difference Vegetation Index (NDVI) time series for each 250 × 250 m pixel in a total area of 1,013 km² (16,218 pixels). A semi-supervised approach was used to train a machine learning model, in which a binary classifier called Support Vector Machine with Global Alignment Kernel was used to differentiate between late frost and non-late frost pixels. We verified the obtained estimates with photointerpretation and existing beech tree-ring chronologies to iteratively improve the model. Then, we used the model output to identify topographical and climatic factors that determined the spatial incidence of LFD. During the study period, LFD was a low recurrence phenomenon that occurred every 15.2 yr on average and showed high spatiotemporal heterogeneity. Most LFD events were condensed in 5 yr and clustered in western forests (86.5% in one-fifth of the pixels) located at high elevation with lower than average precipitation. Elevation and longitude were the major LFD risk factors, followed by annual precipitation. The synergistic effects of increasing drought intensity and rising temperature combined with more frequent late frost events may determine the future performance and distribution of beech forests. This interaction might be critical at the beech drier range edge, where the concentration of LFD at high elevations could constrain beech altitudinal shifts and/or favor species with higher resistance to late frosts.
1. Biotic stressors such as hemiparasites have a profound impact on forest functioning. However, predicting the future incidence of these stressors on forests remains challenging because ...climate-based distribution does not consider tree-hemiparasite interactions or the impacts of extreme climate events on stressors' performance. 2. We use species distribution models (SDMs) and ecophysiological and demographic field data to assess whether climatic suitability is a proxy for the performance of the hemiparasite mistletoe (Viscum album) in two forests with contrasting climatic conditions. Two host tree species representing wet-cold (Scots pine) and drywarm (Aleppo pine) conditions were selected. We fitted SDMs based on climate variables, and measured different ecophysiological variables capturing cold- (photoinhibition) and drought tolerance (intrinsic water-use efficiency, iWUE). We also assessed demographic variables related to seed germination and establishment rates of mistletoe through a translocation experiment. 3. Species distribution models showed a high climatic suitability of mistletoe in both forests. Mistletoes living in the Scots pines site presented a higher cold tolerance, while those inhabiting the Aleppo pine site showed a higher iWUE. Seedlings coming from local seeds showed a lower mortality than seedlings coming from translocated seeds. Germination and seedling establishment showed temporal mismatches when comparing local and translocated seeds. 4. Synthesis. Habitat suitability predicted by SDMs based on climate data and field performance were related in this mistletoe species. However, ecophysiological and demographic variables indicated a lower fitness of mistletoe in the drywarm site associated with drought stress. In conclusion, predicted climate suitability based on SDMs forecasts should be refined using field data on actual performance and considering plant-to-plant interactions and extreme climate events.
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