Quantifying the resistance of mixed-forests against natural hazards in the Pyrenees.

Mountain protection forests can prevent natural hazards by reducing their onset and propagation probabilities. In fact, individual trees act as natural barriers against hydrogeomorphic events. However, assessing the structural strength of trees against these hazards is challenging, especially in a context of climate change due to the intensification of extreme events and changes in forest dynamics. Here, we focus on the mechanical analyses of two of the most common tree species across the Pyrenees (Abies alba Mill. and Fagus sylvatica L.) growing in two different areas (Spain and France), and affected by recurrent snow avalanche and rockfall events. We first performed 53 pulling test on mature trees, where the root-plate stiffness and the modulus of elasticity of the stems were evaluated. To further analyse the impact of forest management and climate on protective forests, we yielded information on tree growth using dendroecology techniques. Then, we assessed structure and neighbourhood characteristics for each target tree to account for the surrounding forest structure. Finally, using linear and structured equation models we tested if the mechanical capacity of the trees is determined either by functional traits (e.g. species, tree growth, diameter and height) or forest structural traits (e.g. tree density, tree structure and slenderness) or both. Our results suggest that the forest neighbourhood influences tree mechanical capacity through two pathways, including both functional and structural traits. The individual stiffness parameter of trees is influenced by their functional traits, while their structural traits are more closely related with changes in the modulus of elasticity. Both species exhibit varying levels of dominance in different locations, which is related to their resilience to the diverse natural hazards they confront. Our findings provide relevant insights to anticipating management strategies for forests that serve as a protective barrier against natural hazards in the context of a changing climate.

Global patterns of tree density are contingent upon local determinants in the world's natural forests.

Previous attempts to quantify tree abundance at global scale have largely neglected the role of local competition in modulating the influence of climate and soils on tree density. Here, we evaluated whether mean tree size in the world's natural forests alters the effect of global productivity on tree density. In doing so, we gathered a vast set of forest inventories including >3000 sampling plots from 23 well-conserved areas worldwide to encompass (as much as possible) the main forest biomes on Earth. We evidence that latitudinal productivity patterns of tree density become evident as large trees become dominant. Global estimates of tree abundance should, therefore, consider dependencies of latitudinal sources of variability on local biotic influences to avoid underestimating the number of trees on Earth and to properly evaluate the functional and social consequences.

Warming reverses directionality in the richness-abundance relationship in ephemeral Mediterranean plant communities.

Recent findings in forests worldwide have demonstrated how directionality in the richness-abundance causality shifts along global climate gradients: The so-called more-species hypothesis (richness determines abundance) prevails in Earth's most productive climates, whereas the opposite, the so-called more-individuals hypothesis (abundance determines richness), is more likely to prevail in climatically harsh conditions. Since temporal variability is the norm, a critical question is whether this directionality shift is also a function of temporal climatic fluctuations locally. Here, we analyze whether directionality in the richness-abundance relationship is contingent on temporal variability over 10 annual consecutive realizations in ephemeral plant assemblages. Our results support the idea that the more-species hypothesis prevailed in the most benign years, whereas the more-individuals hypothesis did so during less productive years, which were significantly linked to the warmest years. These results support the idea that rising temperatures can reverse directionality in the richness-abundance relationship in these annual plant communities, and therefore, climate warming can have a significant effect on the relationship between diversity and ecosystem functions, such as productivity, by altering the prevalence of primary mechanisms involved in species assembly.

Disentangling the Legacies of Climate and Management on Tree Growth.

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. SUPPLEMENTARY INFORMATION: The online version contains supplementary material available at 10.1007/s10021-021-00650-8.

Climate reverses directionality in the richness-abundance relationship across the World's main forest biomes.

More tree species can increase the carbon storage capacity of forests (here referred to as the more species hypothesis) through increased tree productivity and tree abundance resulting from complementarity, but they can also be the consequence of increased tree abundance through increased available energy (more individuals hypothesis). To test these two contrasting hypotheses, we analyse the most plausible pathways in the richness-abundance relationship and its stability along global climatic gradients. We show that positive effect of species richness on tree abundance only prevails in eight of the twenty-three forest regions considered in this study. In the other forest regions, any benefit from having more species is just as likely (9 regions) or even less likely (6 regions) than the effects of having more individuals. We demonstrate that diversity effects prevail in the most productive environments, and abundance effects become dominant towards the most limiting conditions. These findings can contribute to refining cost-effective mitigation strategies based on fostering carbon storage through increased tree diversity. Specifically, in less productive environments, mitigation measures should promote abundance of locally adapted and stress tolerant tree species instead of increasing species richness.

Positive associations among rare species and their persistence in ecological assemblages.

According to the competitive exclusion principle, species with low competitive abilities should be excluded by more efficient competitors; yet, they generally remain as rare species. Here, we describe the positive and negative spatial association networks of 326 disparate assemblages, showing a general organization pattern that simultaneously supports the primacy of competition and the persistence of rare species. Abundant species monopolize negative associations in about 90% of the assemblages. On the other hand, rare species are mostly involved in positive associations, forming small network modules. Simulations suggest that positive interactions among rare species and microhabitat preferences are the most probable mechanisms underpinning this pattern and rare species persistence. The consistent results across taxa and geography suggest a general explanation for the maintenance of biodiversity in competitive environments.

Unveiling annual growth chronologies from inter-nodal branch elongations in a fruticose lichen in southern Europe.

Techniques for retrospective analysis of size dynamics at annual resolution remain poorly developed in lichens in general, and fruticose lichens in particular. Only a few attempts in very high latitudes suggested that growth might be studied as a chronosequence of inter-nodal branch elongations. Here we evaluated, for the first time, this hypothesis in a dry Mediterranean environment using the lichen Cladonia rangiformis as a case study. Mixed models supported a strong positive relationship between humidity measured as precipitation/PET and inter-nodal branch elongations. Importantly, model selection suggested that (i) the number of intermodal elongations were a major determinant of stem elongation, and (ii) a second-order temporal autocorrelation denoted legacies of environmental influences at least over the next 2 y. The strong growth-humidity relationship, along with the potential legacies observed, support the idea that inter-nodal branch elongations could be used to reconstruct growth chronologies at annual resolution in drylands. This finding highlights the high vulnerability of these organisms to rising aridity, and opens a new venue for climate reconstruction and other potential applications in Ecology and Earth Science disciplines.

Disentangling the relative role of climate change on tree growth in an extreme Mediterranean environment.

Climate change can impair ecosystem functions and services in extensive dry forests worldwide. However, attribution of climate change impacts on tree growth and forest productivity is challenging due to multiple inter-annual patterns of climatic variability associated with atmospheric and oceanic circulations. Moreover, growth responses to rising atmospheric CO2, namely carbon fertilization, as well as size ontogenetic changes can obscure the climate change signature as well. Here we apply Structural Equation Models (SEM) to investigate the relative role of climate change on tree growth in an extreme Mediterranean environment (i.e., extreme in terms of the combination of sandy-unconsolidated soils and climatic aridity). Specifically, we analyzed potential direct and indirect pathways by which different sources of climatic variability (i.e. warming and precipitation trends, the North Atlantic Oscillation, [NAO]; the Mediterranean Oscillation, [MOI]; the Atlantic Mediterranean Oscillation, [AMO]) affect aridity through their control on local climate (in terms of mean annual temperature and total annual precipitation), and subsequently tree productivity, in terms of basal area increments (BAI). Our results support the predominant role of Diameter at Breast Height (DHB) as the main growth driver. In terms of climate, NAO and AMO are the most important drivers of tree growth through their control of aridity (via effects of precipitation and temperature, respectively). Furthermore and contrary to current expectations, our findings also support a net positive role of climate warming on growth over the last 50 years and suggest that impacts of climate warming should be evaluated considering multi-annual and multi-decadal periods of local climate defined by atmospheric and oceanic circulation in the North Atlantic.

Erratum: Complementarity effects on tree growth are contingent on tree size and climatic conditions across Europe.

This corrects the article DOI: 10.1038/srep32233.

Last-century forest productivity in a managed dry-edge Scots pine population: the two sides of climate warming.

Climate change in the Mediterranean, associated with warmer temperatures and more frequent droughts, is expected to impact forest productivity and the functioning of forests ecosystems as carbon reservoirs in the region. Climate warming can positively affect forest growth by extending the growing season, whereas increasing summer drought generally reduces forest productivity and may cause growth decline, trigger dieback, hamper regeneration, and increase mortality. Forest management could potentially counteract such negative effects by reducing stand density and thereby competition for water. The effectiveness of such interventions, however, has so far mostly been evaluated for short time periods at the tree and stand levels, which limits our confidence regarding the efficacy of thinning interventions over longer time scales under the complex interplay between climate, stand structure, and forest management. In this study, we use a century-long historical data set to assess the effects of climate and management on forest productivity. We consider rear-edge Scots pine (Pinus sylvestris) populations covering continental and Mediterranean conditions along an altitudinal gradient in Central Spain. We use linear mixed-effects models to disentangle the effects of altitude, climate, and stand volume on forest growth and ingrowth (recruitment and young trees' growth). We find that warming tends to benefit these tree populations, warmer winter temperature has a significant positive effect on both forest growth and ingrowth, and the effect is more pronounced at low elevations. However, drought conditions severely reduce growth and ingrowth, in particular when competition (stand volume) is high. We conclude that summer droughts are the main threat to Scots pine populations in the region, and that a reduction of stand volume can partially mitigate the negative impacts of more arid conditions. Mitigation and adaptation measures could therefore manage stand structure to adopt for the anticipated impacts of climate change in Mediterranean forest ecosystems.

Forest productivity in southwestern Europe is controlled by coupled North Atlantic and Atlantic Multidecadal Oscillations.

The North Atlantic Oscillation (NAO) depicts annual and decadal oscillatory modes of variability responsible for dry spells over the European continent. The NAO therefore holds a great potential to evaluate the role, as carbon sinks, of water-limited forests under climate change. However, uncertainties related to inconsistent responses of long-term forest productivity to NAO have so far hampered firm conclusions on its impacts. We hypothesize that, in part, such inconsistencies might have their origin in periodical sea surface temperature anomalies in the Atlantic Ocean (i.e., Atlantic Multidecadal Oscillation, AMO). Here we show strong empirical evidence in support of this hypothesis using 120 years of periodical inventory data from Iberian pine forests. Our results point to AMO+ NAO+ and AMO-NAO- phases as being critical for forest productivity, likely due to decreased winter water balance and abnormally low winter temperatures, respectively. Our findings could be essential for the evaluation of ecosystem functioning vulnerabilities associated with increased climatic anomalies under unprecedented warming conditions in the Mediterranean.

Climate- and successional-related changes in functional composition of European forests are strongly driven by tree mortality.

Intense droughts combined with increased temperatures are one of the major threats to forest persistence in the 21st century. Despite the direct impact of climate change on forest growth and shifts in species abundance, the effect of altered demography on changes in the composition of functional traits is not well known. We sought to (1) quantify the recent changes in functional composition of European forests; (2) identify the relative importance of climate change, mean climate and forest development for changes in functional composition; and (3) analyse the roles of tree mortality and growth underlying any functional changes in different forest types. We quantified changes in functional composition from the 1980s to the 2000s across Europe by two dimensions of functional trait variation: the first dimension was mainly related to changes in leaf mass per area and wood density (partially related to the trait differences between angiosperms and gymnosperms), and the second dimension was related to changes in maximum tree height. Our results indicate that climate change and mean climatic effects strongly interacted with forest development and it was not possible to completely disentangle their effects. Where recent climate change was not too extreme, the patterns of functional change generally followed the expected patterns under secondary succession (e.g. towards late-successional short-statured hardwoods in Mediterranean forests and taller gymnosperms in boreal forests) and latitudinal gradients (e.g. larger proportion of gymnosperm-like strategies at low water availability in forests formerly dominated by broad-leaved deciduous species). Recent climate change generally favoured the dominance of angiosperm-like related traits under increased temperature and intense droughts. Our results show functional composition changes over relatively short time scales in European forests. These changes are largely determined by tree mortality, which should be further investigated and modelled to adequately predict the impacts of climate change on forest function.

Geography and major host evolutionary transitions shape the resource use of plant parasites.

The evolution of resource use in herbivores has been conceptualized as an analog of the theory of island biogeography, assuming that plant species are islands separated by phylogenetic distances. Despite its usefulness, this analogy has paradoxically led to neglecting real biogeographical processes in the study of macroevolutionary patterns of herbivore-plant interactions. Here we show that host use is mostly determined by the geographical cooccurrence of hosts and parasites in spider mites (Tetranychidae), a globally distributed group of plant parasites. Strikingly, geography accounts for most of the phylogenetic signal in host use by these parasites. Beyond geography, only evolutionary transitions among major plant lineages (i.e., gymnosperms, commelinids, and eudicots) shape resource use patterns in these herbivores. Still, even these barriers have been repeatedly overcome in evolutionary time, resulting in phylogenetically diverse parasite communities feeding on similar hosts. Therefore, our results imply that patterns of apparent evolutionary conservatism may largely be a byproduct of the geographic cooccurrence of hosts and parasites.

Complementarity effects on tree growth are contingent on tree size and climatic conditions across Europe.

Neglecting tree size and stand structure dynamics might bias the interpretation of the diversity-productivity relationship in forests. Here we show evidence that complementarity is contingent on tree size across large-scale climatic gradients in Europe. We compiled growth data of the 14 most dominant tree species in 32,628 permanent plots covering boreal, temperate and Mediterranean forest biomes. Niche complementarity is expected to result in significant growth increments of trees surrounded by a larger proportion of functionally dissimilar neighbours. Functional dissimilarity at the tree level was assessed using four functional types: i.e. broad-leaved deciduous, broad-leaved evergreen, needle-leaved deciduous and needle-leaved evergreen. Using Linear Mixed Models we show that, complementarity effects depend on tree size along an energy availability gradient across Europe. Specifically: (i) complementarity effects at low and intermediate positions of the gradient (coldest-temperate areas) were stronger for small than for large trees; (ii) in contrast, at the upper end of the gradient (warmer regions), complementarity is more widespread in larger than smaller trees, which in turn showed negative growth responses to increased functional dissimilarity. Our findings suggest that the outcome of species mixing on stand productivity might critically depend on individual size distribution structure along gradients of environmental variation.

Climatic Stress during Stand Development Alters the Sign and Magnitude of Age-Related Growth Responses in a Subtropical Mountain Pine.

The modification of typical age-related growth by environmental changes is poorly understood, In part because there is a lack of consensus at individual tree level regarding age-dependent growth responses to climate warming as stands develop. To increase our current understanding about how multiple drivers of environmental change can modify growth responses as trees age we used tree ring data of a mountain subtropical pine species along an altitudinal gradient covering more than 2,200 m of altitude. We applied mixed-linear models to determine how absolute and relative age-dependent growth varies depending on stand development; and to quantify the relative importance of tree age and climate on individual tree growth responses. Tree age was the most important factor for tree growth in models parameterised using data from all forest developmental stages. Contrastingly, the relationship found between tree age and growth became non-significant in models parameterised using data corresponding to mature stages. These results suggest that although absolute tree growth can continuously increase along tree size when trees reach maturity age had no effect on growth. Tree growth was strongly reduced under increased annual temperature, leading to more constant age-related growth responses. Furthermore, young trees were the most sensitive to reductions in relative growth rates, but absolute growth was strongly reduced under increased temperature in old trees. Our results help to reconcile previous contrasting findings of age-related growth responses at the individual tree level, suggesting that the sign and magnitude of age-related growth responses vary with stand development. The different responses found to climate for absolute and relative growth rates suggest that young trees are particularly vulnerable under warming climate, but reduced absolute growth in old trees could alter the species' potential as a carbon sink in the future.