Forecasted increase drought frequency and severity may drive worldwide declines in forest productivity. Species-level responses to a drier world are likely to be influenced by their functional traits. Here, we analyse forest resilience to drought using an extensive network of tree-ring width data and satellite imagery. We compiled proxies of forest growth and productivity (TRWi, absolutely dated ring-width indices; NDVI, Normalized Difference Vegetation Index) for 11 tree species and 502 forests in Spain corresponding to Mediterranean, temperate, and continental biomes. Four different components of forest resilience to drought were calculated based on TRWi and NDVI data before, during, and after four major droughts (1986, 1994-1995, 1999, and 2005), and pointed out that TRWi data were more sensitive metrics of forest resilience to drought than NDVI data. Resilience was related to both drought severity and forest composition. Evergreen gymnosperms dominating semi-arid Mediterranean forests showed the lowest resistance to drought, but higher recovery than deciduous angiosperms dominating humid temperate forests. Moreover, semi-arid gymnosperm forests presented a negative temporal trend in the resistance to drought, but this pattern was absent in continental and temperate forests. Although gymnosperms in dry Mediterranean forests showed a faster recovery after drought, their recovery potential could be constrained if droughts become more frequent. Conversely, angiosperms and gymnosperms inhabiting temperate and continental sites might have problems to recover after more intense droughts since they resist drought but are less able to recover afterwards.

The understanding of regional vulnerability to climate change in Mediterranean mountain forests is not well developed. Climate change impacts on tree growth should be strongly related to the steep environmental gradients of mountainous areas, where a temperature-induced upward shift of the lower elevation limit is expected, particularly amongst drought-sensitive species. Trees will adapt not only to changes in mean climate variables but also to increased extreme events such as prolonged drought. In this paper, we investigate the sub-regional temperature and precipitation trends and measure the basal area increment (BAI) in Pinus nigra subsp. salzmannii (Dunal) Franco. Significant differences related to altitudinal and latitudinal gradients and stand-age structure were found in response to long-term trends in climate dryness. Old trees growing at higher elevations showed similar extreme drought sensitivity but maintained almost steady BAI. Declining BAI found in trees at lower elevations and drier sites may imply a higher vulnerability to temperature-induced drought stress, suggesting an impending growth decline and an enhanced die-off risk. Our results illustrate how the effects of long-term warming and short-term drought on tree BAI are influenced by both site conditions and mean stand age in a drought-sensitive Mediterranean pine.

1. The negative impacts of drought on forest growth and productivity last for several years generating legacies, although the factors that determine why such legacies vary across sites and tree species remain unclear. 2. We used an extensive network of tree-ring width (RWI, ring-width index) records of 16 tree species from 567 forests, and high-resolution climate and normalized difference vegetation index (NDVI) datasets across Spain during the common period 1982-2008 to test the hypothesis that climate conditions and growth features modulate legacy effects of drought on forests. Legacy effects of drought were calculated as the differences between detrended-only RWI and NDVI series (i.e. after removing long-term growth trends) and pre-whitened RWI and NDVI series predicted by a model including drought intensity. Superposed Epoch Analysis (SEA) was used to estimate whether legacy effects differed from random. Finally, legacy effects were related to water balance, growth persistence and variability, and tree species identity. 3. We found a widespread occurrence of drought legacy effects on both RWI and NDVI, but they were seldom significant. According to SEA, first-year drought legacies were negative and different from random in 9% and 5% of the RWI and

Eight Pinus nigra Arn. populations from Southern Spain and Northern Morocco were examined using inter-simple sequence repeat markers to characterize the genetic variability amongst populations. Pair-wise population genetic distance ranged from 0.031 to 0.283, with a mean of 0.150 between populations. The highest inter-population average distance was between PaCU from Cuenca and YeCA from Cazorla, while the lowest distance was between TaMO from Morocco and MA Sierra Mágina populations. Analysis of molecular variance (AMOVA) and Nei’s genetic diversity analyses revealed higher genetic variation within the same population than among different populations. Genetic differentiation (Gst) was 0.233. Cuenca showed the highest Nei’s genetic diversity followed by the Moroccan region, Sierra Mágina, and Cazorla region. However, clustering of populations was not in accordance with their geographical locations. Principal component analysis showed the presence of two major groups—Group 1 contained all populations from Cuenca while Group 2 contained populations from Cazorla, Sierra Mágina and Morocco—while Bayesian analysis revealed the presence of three clusters. The low genetic diversity observed in PaCU and YeCA is probably a consequence of inappropriate management since no estimation of genetic variability was performed before the silvicultural treatments. Data indicates that the inter-simple sequence repeat (ISSR) method is sufficiently informative and powerful to assess genetic variability among populations of P. nigra.

Within-range effects of climatic change on tree growth at the sub-regional scale remain poorly understood. The aim of this research was to use climate and radial-growth data to explain how long-term climatic trends affect tree growth patterns along the southern limit of the range of Pinus nigra ssp. salzmannii (Eastern Baetic Range, southern Spain). We used regional temperature and precipitation data and measured sub-regional radial growth variation in P. nigra forests over the past two centuries. A dynamic factor analysis was applied to test the hypothesis that trees subjected to different climates have experienced contrasting long-term growth variability. We defined four representative stand types based on average temperature and precipitation to evaluate climate-growth relationships using linear mixed-effect models and multi-model selection criteria. All four stand types experienced warming and declining precipitation throughout the twentieth century. From the onset of the twentieth century, synchronised basal-area increment decline was accounted for by dynamic factor analysis and was related to drought by climate-growth models; declining basal-area increment trends proved stronger at lower elevations, whereas temperature was positively related to growth in areas with high rainfall inputs. Given the contrasting sub-regional tree-growth responses to climate change, the role of drought becomes even more complex in shaping communities and affecting selection pressure in the Mediterranean mountain forests. Potential vegetation shifts will likely occur over the dry edge of species distributions, with major impacts on ecosystem structure and function.