46Scots pine forests subjected to continental Mediterranean climates undergo cold winter 47 temperatures and drought stress. Recent climatic trends towards warmer and drier 48 conditions across the Mediterranean Basin might render some of these pine populations 49 more vulnerable to drought-induced growth decline at the southernmost limit of the 50 species distribution. We investigated how cold winters and dry growing seasons drive 51 the radial growth of Scots pine subject to continental Mediterranean climates by relating 52 growth to climate variables at local (elevational gradient) and regional (latitudinal 53 gradient) scales. Local climate-growth relationships were quantified on different time 54 scales (5-, 10-and 15-days) to evaluate the relative role of elevation and specific site 55 characteristics. A negative water balance driven by high maximum temperatures in June 56 (low-elevation sites) and July (high-elevation sites) was the major constraint on growth, 57 particularly on a 5-to 10-day time scale. Warm nocturnal conditions in January were 58 associated with wider rings at the high-elevation sites. At the regional scale, Scots pine 59 growth mainly responded positively to July precipitation, with a stronger association at 60 lower elevations and higher latitudes. January minimum temperatures showed similar 61 patterns but played a secondary role as a driver of tree growth. The balance between 62 positive and negative effects of summer precipitation and winter temperature on radial 63 growth depends on elevation and latitude, with low-elevation populations being more 64 prone to suffer drought and heat stress, whereas high-elevation populations may be 65 favoured by warmer winter conditions. This negative impact of summer heat and 66 drought has increased during the past decades. This interaction between climate and site 67 conditions and local adaptations is therefore decisive for the future performance and 68 persistence of Scots pine populations in continental Mediterranean climates.
Climate, stand structure and local site conditions are potentially important determinants of forest dynamics. Understanding the relative contributions of competition and climate to tree growth is critical to project likely stand development under different climate change scenarios. Further, current competition levels and stand structure may reflect legacies of past forest management. Here, we analyze the effects of climate, site conditions and competition on radial growth in three Scots pine plots located along an altitudinal gradient. These stands are subjected to Mediterranean climate with continental influence, i.e., growth is limited by low winter temperatures and dry summer conditions. 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 competition (CI) and climate at an annual resolution. Negative exponential functions characterized the CI-BAI associations, whereas linear mixed-effects models were used to model BAI and to quantify the growth response to climate of trees under low and high competition. Competition effects on growth were steady over time regardless of the elevation and tree age. High competition levels negatively affected growth, with a proportionally stronger influence in suppressed trees than in dominant trees. Sensitivity of tree growth to climate increased with decreasing competition. Altitudinal gradientrelated growth responses to climate were found only for temperature variables. Specifically, growth at high elevations was mainly limited by low winter temperatures, whilst warm spring enhanced growth at middle elevations and late summer temperatures did it at low elevations. Since growth and its sensitivity to climate is more pronounced in low competition trees, we argue that the past management of the forest overrides site conditions and climate effects through the legacies on stand structure and competition. Pro-active forest management practices should be adopted to reduce the vulnerability of previously managed Scots pine forests currently threatened by the predicted warmer and drier conditions.
Europe is undergoing significant forest expansion due to the abandonment of rural areas driven by economic and demographic changes. Recently established forests provide key ecosystem services such as habitat provision and increased carbon stocks. However, we lack understanding of whether past land use might alter their resilience to climate change compared with long-established forests. Forests established in former agricultural areas may benefit from land use legacies resulting in higher fertility, yet such a benefit might turn into a disadvantage if it involves changes in functional attributes that lower their ability to cope with negative climatic events (e.g. droughts). Here we examined whether recently (post 1956) and long-established (pre 1956) beech forests in Catalonia (NE Spain) differ in their growth patterns, wood density, sensitivity to climate and response to extreme climatic events. Our results indicate higher growth (32%) and lower wood density (3%) in trees from recently established forests, even when controlling for tree age and competition. In addition, recently established forests showed a higher sensitivity to Standardised Precipitation-Evapotranspiration Index (SPEI), precipitation and temperature and to extreme negative and positive climatic events. In particularly wet years, recently established forests show twice the number of positive pointer years than long-established forests. Compensatory growth during positive years in recently established forests, may be driving the similar or even higher recovery and resilience detected after drought episodes. Nevertheless, the higher climatic sensitivity of the recently established forests, together with their greater growth and lower wood density indicates that they may be particularly vulnerable to future droughts. Such enhanced vulnerability might question their ability to contribute to carbon sequestration in the long term and emphasises the need to account for land use legacies to better predict future forest function as climate changes.
Tree rings provide an invaluable long‐term record for understanding how climate and other drivers shape tree growth and forest productivity. However, conventional tree‐ring analysis methods were not designed to simultaneously test effects of climate, tree size, and other drivers on individual growth. This has limited the potential to test ecologically relevant hypotheses on tree growth sensitivity to environmental drivers and their interactions with tree size. Here, we develop and apply a new method to simultaneously model nonlinear effects of primary climate drivers, reconstructed tree diameter at breast height (DBH), and calendar year in generalized least squares models that account for the temporal autocorrelation inherent to each individual tree's growth. We analyze data from 3811 trees representing 40 species at 10 globally distributed sites, showing that precipitation, temperature, DBH, and calendar year have additively, and often interactively, influenced annual growth over the past 120 years. Growth responses were predominantly positive to precipitation (usually over ≥3‐month seasonal windows) and negative to temperature (usually maximum temperature, over ≤3‐month seasonal windows), with concave‐down responses in 63% of relationships. Climate sensitivity commonly varied with DBH (45% of cases tested), with larger trees usually more sensitive. Trends in ring width at small DBH were linked to the light environment under which trees established, but basal area or biomass increments consistently reached maxima at intermediate DBH. Accounting for climate and DBH, growth rate declined over time for 92% of species in secondary or disturbed stands, whereas growth trends were mixed in older forests. These trends were largely attributable to stand dynamics as cohorts and stands age, which remain challenging to disentangle from global change drivers. By providing a parsimonious approach for characterizing multiple interacting drivers of tree growth, our method reveals a more complete picture of the factors influencing growth than has previously been possible.
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