Juan Carlos Linares

Climate change impacts and vulnerability of the southern populations of Pinus nigra subsp. salzmannii

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.

Plastic responses of Abies pinsapo xylogenesis to drought and competition

Radial growth and xylogenesis were studied to investigate the influence of climate variability and intraspecific competition on secondary growth in Abies pinsapo Boiss., a relic Mediterranean fir. We monitored the responses to three thinning treatments (unthinned control -C-, 30% -T30- and 60% -T60- of basal area removed) to test the hypothesis that they may improve the adaptation capacity of tree growth to climatic stress. We also assessed whether xylogenesis was differentially affected by tree-to-tree competition. Secondary growth was assessed using manual band dendrometers from 2005 to 2007. In 2006, xylogenesis (phases of tracheid formation) was also investigated by taking microcores and performing histological analyses. Seasonal dynamics of radial increment were modeled using Gompertz functions and correlations with microclimate and radiation were performed. Histological analyses revealed it as fundamental to calibrate the dendrometer estimates of radial increment and to establish the actual onset and end dates of tracheid production. The lower radial-increment rates and number of produced tracheids were observed in the trees subjected to high competition in the unthinned plots. The growing season differed among the plots, and its duration ranged from an average of 78 days in unthinned plots to 115 days in thinned ones (T60). Variations in the beginning of the growing season (13 April to 22 May) and earlywood-latewood transition (early August) were mainly determined by the temperature pattern, while the onset and the end of the growing season were related to both annual precipitation and tree-to-tree competition. The tracheid-formation phases of radial enlargement and cell-wall thickening showed similar patterns in the trees from thinned and unthinned plots subjected to low and high competition, respectively, but the mean number of tracheids in each phase was always higher in the trees from the thinned plots. The reduction of competition through thinning induced a longer growing season and enhanced the radial growth in A. pinsapo.

Forests synchronize their growth in contrasting Eurasian regions in response to climate warming

Significance Forests dominate carbon fluxes in terrestrial ecosystems. We demonstrate how an intensified climatic influence on tree growth during the last 120 y has increased spatial synchrony in annual ring-width patterns within contrasting (boreal and Mediterranean) Eurasian biomes and on broad spatial scales. Current trends in tree growth synchrony are related to regional changes in climate factors controlling productivity, overriding local and taxonomic imprints on forest carbon dynamics. Enhanced synchrony is becoming a widespread, although regionally dependent, phenomenon related to warmer springs and increased temperature variability in high latitudes and to warmer winters and drier growing seasons in mid-latitudes. Forests play a key role in the carbon balance of terrestrial ecosystems. One of the main uncertainties in global change predictions lies in how the spatiotemporal dynamics of forest productivity will be affected by climate warming. Here we show an increasing influence of climate on the spatial variability of tree growth during the last 120 y, ultimately leading to unprecedented temporal coherence in ring-width records over wide geographical scales (spatial synchrony). Synchrony in growth patterns across cold-constrained (central Siberia) and drought-constrained (Spain) Eurasian conifer forests have peaked in the early 21st century at subcontinental scales (∼1,000 km). Such enhanced synchrony is similar to that observed in trees co-occurring within a stand. In boreal forests, the combined effects of recent warming and increasing intensity of climate extremes are enhancing synchrony through an earlier start of wood formation and a stronger impact of year-to-year fluctuations of growing-season temperatures on growth. In Mediterranean forests, the impact of warming on synchrony is related mainly to an advanced onset of growth and the strengthening of drought-induced growth limitations. Spatial patterns of enhanced synchrony represent early warning signals of climate change impacts on forest ecosystems at subcontinental scales.

Mistletoe effects on Scots pine decline following drought events: insights from within-tree spatial patterns, growth and carbohydrates

Forest decline has been attributed to the interaction of several stressors including biotic factors such as mistletoes and climate-induced drought stress. However, few data exist on how mistletoes are spatially arranged within trees and how this spatial pattern is related to changes in radial growth, responses to drought stress and carbon use. We used dendrochronology to quantify how mistletoe (Viscum album L.) infestation and drought stress affected long-term growth patterns in Pinus sylvestris L. at different heights. Basal area increment (BAI) trends and comparisons between trees of three different infestation degrees (without mistletoe, ID1; moderately infested trees, ID2; and severely infested trees, ID3) were performed using linear mixed-effects models. To identify the main climatic drivers of tree growth tree-ring widths were converted into indexed chronologies and related to climate data using correlation functions. We performed spatial analyses of the 3D distribution of mistletoe individuals and their ages within the crowns of three severely infested pines to describe their patterns. Lastly, we quantified carbohydrate and nitrogen concentrations in needles and sapwood of branches from severely infested trees and from trees without mistletoe. Mistletoe individuals formed strongly clustered groups of similar age within tree crowns and their age increased towards the crown apex. Mistletoe infestation negatively impacted growth but this effect was stronger near the tree apex than in the rest of sampled heights, causing an average loss of 64% in BAI (loss of BAI was ∼51% at 1.3 m or near the tree base). We found that BAI of severely infested trees and moderately or non-infested trees diverged since 2001 and such divergence was magnified by drought. Infested trees had lower concentrations of soluble sugars in their needles than non-infested ones. We conclude that mistletoe infestation causes growth decline and increases the sensitivity of trees to drought stress.

Buffered climate change effects in a Mediterranean pine species: range limit implications from a tree-ring study.

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.

Drought-induced increase in water-use efficiency reduces secondary tree growth and tracheid wall thickness in a Mediterranean conifer

In order to understand the impact of drought and intrinsic water-use efficiency (iWUE) on tree growth, we evaluated the relative importance of direct and indirect effects of water availability on secondary growth and xylem anatomy of Juniperus thurifera, a Mediterranean anisohydric conifer. Dendrochronological techniques, quantitative xylem anatomy, and 13C/12C isotopic ratio were combined to develop standardized chronologies for iWUE, BAI (basal area increment), and anatomical variables on a 40-year-long annually resolved series for 20 trees. We tested the relationship between iWUE and secondary growth at short-term (annual) and long-term (decadal) temporal scales to evaluate whether gains in iWUE may lead to increases in secondary growth. We obtained a positive long-term correlation between iWUE and BAI, simultaneously with a negative short-term correlation between them. Furthermore, BAI and iWUE were correlated with anatomical traits related to carbon sink or storage (tracheid wall thickness and ray parenchyma amount), but no significant correlation with conductive traits (tracheid lumen) was found. Water availability during the growing season significantly modulated tree growth at the xylem level, where growth rates and wood anatomical traits were affected by June precipitation. Our results are consistent with a drought-induced limitation of tree growth response to rising CO2, despite the trend of rising iWUE being maintained. We also remark the usefulness of exploring this relationship at different temporal scales to fully understand the actual links between iWUE and secondary growth dynamics.

Human impacts drive forest structure and diversity. Insights from Mediterranean mountain forest dominated by Abies pinsapo (Boiss.)

Environmental factors and land-use are likely interacting to drive forest structure and species diversity, making it difficult to disentangle their separate impacts. Both components influence the resilience and stability of mountain forests in the Mediterranean Basin, since secular land-use has shaped stands structure and forests dynamics are constrained to the environment. Approaches covering environmental factors and contrasting land-uses are still needed to understand their implications on forest dynamics. We investigated patterns of environmental variables, stand structure and biodiversity in forest dominated by Abies pinsapo for 61 stands over their whole range. In addition, since land-use by local inhabitant could be a key factor, logging intensity was quantified by stumps basal area and recent land-use history was investigated. Partial redundancy analysis (pRDA) was conducted to assess to which degree the structural attributes are related to environmental characteristics and/or the land-use in the main A. pinsapo forests. Environmental characteristics accounted for 13% of the total variance, while the contrasting land-use (Spain vs. Morocco) and the stumps basal area, as indicator of human logging, accounted for 23% of the total variance. Trees species diversity was near 5 times higher in Moroccan stands than in stands from south Spain. According to recent human logging, Spanish stands have been strictly protected, but Moroccan stands showed widespread axe-logging by local inhabitant over the smaller tree sizes. Our results suggest that the preservation of a minor perturbation regimen by local populations enhance and help to preserve the whole biodiversity of this relict habitat.

Stand-structural effects on Heterobasidion abietinum-related mortality following drought events in Abies pinsapo

Climate change may affect tree–pathogen interactions. This possibility has important implications for drought-prone forests, where stand dynamics and disease pathogenicity are especially sensitive to climatic stress. In addition, stand structural attributes including density-dependent tree-to-tree competition may modulate the stands’ resistance to drought events and pathogen outbreaks. To assess the effects of stand structure on root-rot-related mortality after severe droughts, we focused on Heterobasidion abietinum mortality in relict Spanish stands of Abies pinsapo, a drought-sensitive fir. We compared stand attributes and tree spatial patterns in three plots with H. abietinum root-rot disease and three plots without root-rot. Point-pattern analyses were used to investigate the scale and extent of mortality patterns and to test hypotheses related to the spread of the disease. Dendrochronology was used to date the year of death and to assess the association between droughts and growth decline. We applied a structural equation modelling approach to test if tree mortality occurs more rapidly than predicted by a simple distance model when trees are subjected to high tree-to-tree competition and following drought events. Contrary to expectations of drought mortality, the effect of precipitation on the year of death was strong and negative, indicating that a period of high precipitation induced an earlier tree death. Competition intensity, related to the size and density of neighbour trees, also induced an earlier tree death. The effect of distance to the disease focus was negligible except in combination with intensive competition. Our results indicate that infected trees have decreased ability to withstand drought stress, and demonstrate that tree-to-tree competition and fungal infection act as predisposing factors of forest decline and mortality.

Temperate-like stand dynamics in relict Mediterranean-fir (Abies pinsapo, Boiss.) forests from southern Spain

Introduction and statement of the research questionsGap dynamics have been widely studied in forests of Abies spp. from temperate and boreal regions. The local microclimate and competition for light have been identified as the main factors controlling changes in species composition and canopy structure, however little is known on dynamics of such forests in Mediterranean.Experimental design and aimsWe studied forest structure and dynamics of Abies pinsapo stands in southern Spain, in contrasting habitats and successional status. In addition past regeneration patterns and their relationship to canopy structure, disturbances and forest-use history were investigated.ResultsStands structure attributes were within the range described for temperate conifer biomes. The age structure revealed two main cohorts comprised of a few > 100 year-old trees and abundant younger trees established in a single recruitment event after the stands were protected in the 1950s. Initial growth-rate analyses indicated that A. pinsapo regenerated mainly in small canopy gaps, while only 15% recruited from the forest understorey. For the last ten years, basal area increment was lower than 10 cm2 y−1 in 91% of studied trees and growth rate differences between trees narrowed.ConclusionStand dynamics in A. pinsapo forests maintain general features of temperate fir forests. Tree establishment over time and current stand structure fit to known changes in forest use. Widespread growth decline trends might be linked to stand stagnation and global warming.RésuméIntroduction et état des questions de rechercheLes dynamiques des trouées ont été largement étudiées dans les forêts d’Abies spp. à partir des régions tempérées et boréales. Le microclimat local et la compétition pour la lumière ont été identifiés comme les principaux facteurs de contrôle des changements dans la composition des espèces et la structure du couvert, mais on sait peu de choses sur la dynamique de ces forêts dans les écosystèmes méditerranéens.Plan expérimental et objectifsNous avons étudié la structure de la forêt et la dynamique de peuplements d’Abies pinsapo dans le sud de l’Espagne, dans des habitats contrastés et dans des états d’évolution successifs. En outre, les modèles passés de régénération et leurs relations avec la structure du couvert, les perturbations et l’histoire de l’utilisation des forêts ont été étudiés.RésultatsLes attributs de structure des peuplements se situent dans la gamme décrite pour les biomes de conifères tempérés. La structure des âges a révélé deux cohortes principales composées de quelques vieux arbres d’un âge supérieur à 100 ans, et de nombreux jeunes arbres établis dans un seul évènement de régénération après que les peuplements aient été protégés dans les années 1950. Les premières analyses des taux de croissance ont indiqué que A. pinsapo s’est régénéré principalement dans les petites trouées, tandis que seulement 15 % des arbres ont été recrutés dans le sous-bois. Au cours des dix dernières années, l’augmentation de la surface terrière a été inférieure à 10 cm2 an−1 chez 91 % des arbres étudiés et les différences entre les taux de croissance des arbres étaient réduites.ConclusionLes dynamique des peuplements d’A. pinsapo conservent les caractéristiques générales des forêts de sapins tempérés. L’installation des arbres et la structure actuelle des peuplements permettent de connaître les changements dans l’utilisation de la forêt. La baisse généralisée des tendances de croissance pourrait être liée à la stagnation du peuplement et au réchauffement de la planète.

Phenological and water-use patterns underlying maximum growing season length at the highest elevations: implications under climate change

Consequences of climate change on tree phenology are readily observable, but little is known about the variations in phenological sensitivity to drought between populations within a species. In this study, we compare the phenological sensitivity to temperature and water availability in Abies pinsapo Boiss., a drought-sensitive Mediterranean fir, across its altitudinal distribution gradient. Twig growth and needle fall were related to temperature, precipitation and plant water status on a daily scale. Stands located at the top edge of the distributional range showed the most favourable water balance, maximum growth rates and little summer defoliation. Towards higher elevations, the observed delay in budburst date due to lower spring temperatures was overcome by a stronger delay in growth cessation date due to the later onset of strong water-deficit conditions in the summer. This explains an extended growing season and the greatest mean growth at the highest elevation. Conversely, lower predawn xylem water potentials and early partial stomatal closure and growth cessation were found in low-elevation A. pinsapo trees. An earlier and higher summer peak of A. pinsapo litterfall was also observed at these water-limited sites. Our results illustrate the ecophysiological background of the ongoing altitudinal shifts reported for this relict tree species under current climatic conditions.