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.
Cork is one of the main non-timber forest products in the world. Most of its production is concentrated in the Iberian Peninsula, a climate change hotspot. Climate warming may lead to increased ...aridification and reduce cork production in that region. However, we still lack assessments of climate-cork relationships across ample geographical and climatic gradients explicitly considering site aridity. We quantified cork growth by measuring cork ring width and related it to climate variables and a drought index using dendrochronology. Four cork oak (Quercus suber) forests located from north eastern Spain to south western Morocco (31.5–41.5° N) and subjected to different aridity levels were sampled. Warm conditions in spring to early summer, when cork is formed, reduced cork width, whereas high precipitation in winter and spring enhanced it. The response of cork to increased water availability in summer peaked (r = 0.89, p = 0.00002) in the most arid and continental site considering 14-month long droughts. A severe drought caused a disproportionate loss of cork production in this site, where for every five-fold decrease in the drought index, the cork-width index declined by a factor of thirteen. Therefore, site aridity determines the responses of cork growth to the soil water availability resulting from accumulated precipitation during winter and spring previous to cork growth and until summer. In general, this cumulative water balance, which is very dependent on temperature and evapotranspiration rate, is critical for cork production, especially in continental, dry sites. The precipitation during the hydrological year can be used as a proxy of cork production in similar sites. Assessments of climate-cork relationships in the western Mediterranean basin could be used as analogues to forecast the impacts of aridification on future cork production.
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•High variability in space and time of tree-growth response to climate in Spain.•Negative impact of low summer rainfall on growth increases at low elevations and high latitudes.•Growth at low ...elevations is more dependent on spring climate than at high elevations.•Negative impact of summer heat and drought has increased during the past decades.•Local adaptation to climate change is a key factor for future Mediterranean forests.
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.
•We analyzed climate, drought and hydrology effects on Fraxinus angustifolia growth.•Diverse growth responses to climate and drought reflected the large bioclimatic range of the study.•Growth ...responses intensified in warmer sites and at longer time scales.•Growth was more coupled with hydrologic regime in sites with least human alteration.•Climate warming will negatively impact Mediterranean riparian ash forests.
Mediterranean riparian forests are among the most threatened ecosystems in Europe. These ecosystems are exposed to land-use changes threatening their reduced habitat and by global warming, which is already triggering aridification processes. To assess the impact of these major threats, we studied the radial-growth responses to climate and drought in the narrow-leaved ash (Fraxinus angustifolia). This riparian tree species presents a relatively large ecological spectrum in its habitat preference in the Mediterranean Basin. We studied five sites arranged across a wide geographical range from Iberia to Italy, subjected to contrasting climatic conditions and located in hydrographic basins with different sizes and water regimes. We found diverse growth responses to climate and drought across the Mediterranean distribution range of the narrow-leaved ash at the individual and site levels. The growth of this species increased in response to wet and cool conditions in the prior winter and spring. The response to summer conditions was only observed in the coldest and wettest site (Ticino). Growth responded negatively to 2–14 month droughts that occurred from previous winter up to summer, particularly in the warmest-driest sites. Growth responses to drought peaked in the warmest-driest sites in terms of climate water balance (Odelouca, Doñana), but not in the driest sites in terms of annual precipitation (Tudela, Zaragoza). Hydrological conditions also affected the narrow-leaved ash with high discharges in the prior winter and early spring enhancing wood production. Considering projected aridification and increased hydrological alteration, implying limited water supply in the Mediterranean region, climate warming will negatively impact productivity of narrow-leaved ash riparian forests. Further research should combine analyses of growth responses to climate and hydrology from tree to basin scales to disentangle their relative roles as drivers of productivity under different scenarios of climate and hydrological changes, in order to aid adaptive management of these key ecosystems.
•Forest structure modulates Scots pine growth responses to climate.•Tree-to-tree competition and radial growth were negatively related.•Tree growth is strongly modulated by competition.•Competition ...overrides climate as a driver of tree growth.•Climate–growth sensitivity increases as competition decreases.
Understanding the relative contributions of competition and climate on individual tree growth is critical to project realistic forest dynamics under projected climate scenarios. Furthermore, present competition levels may reflect legacies of past use. Here, we analyze the effects of climate, site conditions and competition on radial growth in three Scots pine stands located along an altitudinal gradient in central Spain. Current stand structure and retrospective analyses of radial growth (basal area increment, BAI) were used to model changes in tree growth as a function of a spatially-explicit competition index (CI) and climate. Linear mixed-effects models were employed to model BAI and to quantify the growth responses to climate of trees under low and high competition levels. Competition effects on growth were steady over time regardless of tree age. High competition levels negatively affected growth since negative exponential functions characterized the CI–BAI relationships. Tree growth sensitivity to climate increased with decreasing competition intensity. Growth at high elevations was mainly limited by low winter temperatures, whereas warm spring enhanced growth at middle elevations and warm late summer temperatures constrained growth at low elevation. Growth responsiveness to climate is enhanced under low competition levels. Overall, current competition is a more relevant driver of recent growth than climate. Proactive forest management should be adopted to reduce the vulnerability of Scots pine forests currently subjected to higher competition levels and warmer and drier conditions.
Intense debate surrounds the effects of post‐fire salvage logging (SL) versus nonintervention policies on forest regeneration, but scant support is available from experimental studies. We analyze the ...effect of three post‐fire management treatments on the recruitment of a serotinous pine (Pinus pinaster) at a Mediterranean mountain. Treatments were applied 7 months after the fire and differ in the degree of intervention, ranging from “no intervention” (NI, all trees left standing) to “partial cut plus lopping” (PCL, felling most of the trees, cutting the main branches, and leaving all the biomass in situ without mastication), and “SL” (felling and piling the logs, and masticating the woody debris). Seedling survival after 3 years was the highest in PCL (47.3% versus 38.7% in SL). This was associated with the amelioration of microclimatic conditions under the scattered branches, which reduced radiation and soil temperature while increasing soil moisture. Seedling density after 2 years was approximately 5.5 times higher in PCL than in SL, as in SL a large fraction of seedlings was lost as a consequence of mechanized mastication. The NI treatment showed the lowest seedling survival (17.3%). Nevertheless, seedling density was similar to SL. Seedling growth scarcely differed among treatments. Our results show that branches left onsite acted as nurse objects that improved key microclimatic conditions for seedling recruitment. This creates a facilitative interaction ideal for seedling establishment in moisture‐deficient ecosystems, as it provides the benefit of a shading overstory but without underground competition.
Disease emergence in northern and boreal forests has been mostly due to tree-pathogen encounters lacking a co-evolutionary past. However, outbreaks involving novel interactions of the host or the ...pathogen with the environment have been less well documented. Following an increase of records in Northern Europe, the first large outbreak of
on
was discovered in Sweden in 2016. By reconstructing the development of the epidemic, we found that the attacks started approx. 10 years back from several isolated trees in the stand and ended up affecting almost 90% of the trees in 2016. Limited damage was observed in other plantations in the surroundings of the affected stand, pointing to a new introduced pathogen as the cause of the outbreak. Nevertheless, no genetic differences based on SSR markers were found between isolates of the outbreak area and other Swedish isolates predating the outbreak or from other populations in Europe and Asia Minor. On a temporal scale, we saw that warm May and June temperatures were associated with higher damage and low tree growth, while cold and rainy conditions seemed to favor growth and deter disease. At a spatial scale, we saw that spread occurred predominantly in the SW aspect-area of the stand. Within that area and based on tree-ring and isotope (δ
C) analyses, we saw that disease occurred on trees that over the years had shown a lower water-use efficiency (WUE). Spore traps showed that highly infected trees were those producing the largest amount of inoculum.
impaired latewood growth and reduced C reserves in needles and branches.
attacks can cause serious economic damage by killing new shoots, disrupting the crown, and affecting the quality of stems. Our results show that
has no limitations in becoming a serious pathogen in Northern Europe. Management should focus on reducing inoculum, especially since climate change may bring more favorable conditions for this pathogen. Seedlings for planting should be carefully inspected as
may be present in a latent stage in asymptomatic tissues.
Ongoing changes in global climate are altering ecological conditions for many species. The consequences of such changes are typically most evident at the edge of a species’ geographical distribution, ...where differences in growth or population dynamics may result in range expansions or contractions. Understanding population responses to different climatic drivers along wide latitudinal and altitudinal gradients is necessary in order to gain a better understanding of plant responses to ongoing increases in global temperature and drought severity. We selected Scots pine (Pinus sylvestris L.) as a model species to explore growth responses to climatic variability (seasonal temperature and precipitation) over the last century through dendrochronological methods. We developed linear models based on age, climate and previous growth to forecast growth trends up to year 2100 using climatic predictions. Populations were located at the treeline across a latitudinal gradient covering the northern, central and southernmost populations and across an altitudinal gradient at the southern edge of the distribution (treeline, medium and lower elevations). Radial growth was maximal at medium altitude and treeline of the southernmost populations. Temperature was the main factor controlling growth variability along the gradients, although the timing and strength of climatic variables affecting growth shifted with latitude and altitude. Predictive models forecast a general increase in Scots pine growth at treeline across the latitudinal distribution, with southern populations increasing growth up to year 2050, when it stabilizes. The highest responsiveness appeared at central latitude, and moderate growth increase is projected at the northern limit. Contrastingly, the model forecasted growth declines at lowland‐southern populations, suggesting an upslope range displacement over the coming decades. Our results give insight into the geographical responses of tree species to climate change and demonstrate the importance of incorporating biogeographical variability into predictive models for an accurate prediction of species dynamics as climate changes.
We used Scots pine populations located at the treeline across a latitudinal gradient covering the northern, central and southernmost populations and across an altitudinal gradient at the rear edge of the distribution to explore growth responses to climatic variability over the last century through dendrochronological methods and developed models based on age, climate and previous growth to forecast growth trends up to year 2100. Temperature was the main factor controlling growth variability, although the timing and strength of climatic variables affecting growth shifted with latitude and altitude. Predictive models forecast a general increase in Scots pine growth at treeline across the latitudinal distribution, with southern populations increasing growth up to year 2050, when it stabilizes. The highest responsiveness appeared at central latitudes, and moderate growth increase is projected at the northern limit. Contrastingly, the model forecasted growth decline at lowland‐southern populations, suggesting an upslope range displacement over the coming decades.
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.
•Mediterranean riparian forests are threatened but understudied ecosystems.•We quantified radial growth and earlywood anatomy in Fraxinus angustifolia.•Growth rate and the earlywood hydraulic diameter (Dh) covaried.•Growth increased in response to higher precipitation.•Earlywood Dh increased in response to wet prior autumn-winter conditions.
Tree species inhabiting riparian forests under Mediterranean climate have evolved to face summer water shortage but may fail to cope with future increases in drought severity. Thus, understanding ...tree growth phenological variations in response to environmental conditions is necessary to assess the impact of seasonal drought in riparian forests. In this study, we investigated the response of stem radial growth to climate in the narrow-leaved ash (Fraxinus angustifolia) over its distribution in southern Europe. We simulated intra- and inter-annual growth patterns using the Vaganov-Shashkin (VS) model considering five sites subjected to summer drought but showing different climate conditions. The growth pattern in this species varied from unimodal in cool-wet sites to facultative bimodal in warm-dry sites. Bimodal patterns were characterized by two growth peaks coinciding with favorable climate conditions in spring and autumn. The spring growth peak occurs earlier (May) in warm-dry sites than in wet-cool sites (June–July). The variation in the season growth length and growth timing suggests different strategies adopted by this species to cope with summer drought. The VS model revealed different growth patterns across which would be relevant in predicting the response of this and other riparian tree species to climate warming and aridification. Differences in the length of the growing season, timings of growth peaks and the shift from unimodal to bimodal growth patterns should be considered when assessing growth adjustments to future climate scenarios.
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•Ring-width and daily climatic data were used to adjust the VS model in narrow-leaved ash.•The ash growing season varied in length and pattern, but always showed a spring growth peak.•Ash growth shifted from unimodal in wet-cool sites to facultative bimodal in dry-warm sites.•The use of process-based models allows assessing intra-annual growth responses to climate.
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