Functional traits and climate drive interspecific differences in disturbance-induced tree mortality.

With climate change, natural disturbances such as storm or fire are reshuffled, inducing pervasive shifts in forest dynamics. To predict how it will impact forest structure and composition, it is crucial to understand how tree species differ in their sensitivity to disturbances. In this study, we investigated how functional traits and species mean climate affect their sensitivity to disturbances while controlling for tree size and stand structure. With data on 130,594 trees located on 7617 plots that were disturbed by storm, fire, snow, biotic or other disturbances from the French, Spanish, and Finnish National Forest Inventory, we modeled annual mortality probability for 40 European tree species as a function of tree size, dominance status, disturbance type, and intensity. We tested the correlation of our estimated species probability of disturbance mortality with their traits and their mean climate niches. We found that different trait combinations controlled species sensitivity to disturbances. Storm-sensitive species had a high height-dbh ratio, low wood density and high maximum growth, while fire-sensitive species had low bark thickness and high P50. Species from warmer and drier climates, where fires are more frequent, were more resistant to fire. The ranking in disturbance sensitivity between species was overall consistent across disturbance types. Productive conifer species were the most disturbance sensitive, while Mediterranean oaks were the least disturbance sensitive. Our study identified key relations between species functional traits and disturbance sensitivity, that allows more reliable predictions of how changing climate and disturbance regimes will impact future forest structure and species composition at large spatial scales.

Tree growth response to drought partially explains regional-scale growth and mortality patterns in Iberian forests.

Tree-ring data has been widely used to inform about tree growth responses to drought at the individual scale, but less is known about how tree growth sensitivity to drought scales up driving changes in forest dynamics. Here, we related tree-ring growth chronologies and stand-level forest changes in basal area from two independent data sets to test if tree-ring responses to drought match stand forest dynamics (stand basal area growth, ingrowth, and mortality). We assessed if tree growth and changes in forest basal area covary as a function of spatial scale and tree taxa (gymnosperm or angiosperm). To this end, we compared a tree-ring network with stand data from the Spanish National Forest Inventory. We focused on the cumulative impact of drought on tree growth and demography in the period 1981-2005. Drought years were identified by the Standardized Precipitation Evapotranspiration Index, and their impacts on tree growth by quantifying tree-ring width reductions. We hypothesized that forests with greater drought impacts on tree growth will also show reduced stand basal area growth and ingrowth and enhanced mortality. This is expected to occur in forests dominated by gymnosperms on drought-prone regions. Cumulative growth reductions during dry years were higher in forests dominated by gymnosperms and presented a greater magnitude and spatial autocorrelation than for angiosperms. Cumulative drought-induced tree growth reductions and changes in forest basal area were related, but initial stand density and basal area were the main factors driving changes in basal area. In drought-prone gymnosperm forests, we observed that sites with greater growth reductions had lower stand basal area growth and greater mortality. Consequently, stand basal area, forest growth, and ingrowth in regions with large drought impacts was significantly lower than in regions less impacted by drought. Tree growth sensitivity to drought can be used as a predictor of gymnosperm demographic rates in terms of stand basal area growth and ingrowth at regional scales, but further studies may try to disentangle how initial stand density modulates such relationships. Drought-induced growth reductions and their cumulative impacts have strong potential to be used as early-warning indicators of regional forest vulnerability.

Plant functional traits have globally consistent effects on competition.

Phenotypic traits and their associated trade-offs have been shown to have globally consistent effects on individual plant physiological functions, but how these effects scale up to influence competition, a key driver of community assembly in terrestrial vegetation, has remained unclear. Here we use growth data from more than 3 million trees in over 140,000 plots across the world to show how three key functional traits--wood density, specific leaf area and maximum height--consistently influence competitive interactions. Fast maximum growth of a species was correlated negatively with its wood density in all biomes, and positively with its specific leaf area in most biomes. Low wood density was also correlated with a low ability to tolerate competition and a low competitive effect on neighbours, while high specific leaf area was correlated with a low competitive effect. Thus, traits generate trade-offs between performance with competition versus performance without competition, a fundamental ingredient in the classical hypothesis that the coexistence of plant species is enabled via differentiation in their successional strategies. Competition within species was stronger than between species, but an increase in trait dissimilarity between species had little influence in weakening competition. No benefit of dissimilarity was detected for specific leaf area or wood density, and only a weak benefit for maximum height. Our trait-based approach to modelling competition makes generalization possible across the forest ecosystems of the world and their highly diverse species composition.

Evidence of non-stationary relationships between climate and forest responses: Increased sensitivity to climate change in Iberian forests.

Climate and forest structure are considered major drivers of forest demography and productivity. However, recent evidence suggests that the relationships between climate and tree growth are generally non-stationary (i.e. non-time stable), and it remains uncertain whether the relationships between climate, forest structure, demography and productivity are stationary or are being altered by recent climatic and structural changes. Here we analysed three surveys from the Spanish Forest Inventory covering c. 30 years of information and we applied mixed and structural equation models to assess temporal trends in forest structure (stand density, basal area, tree size and tree size inequality), forest demography (ingrowth, growth and mortality) and above-ground forest productivity. We also quantified whether the interactive effects of climate and forest structure on forest demography and above-ground forest productivity were stationary over two consecutive time periods. Since the 1980s, density, basal area and tree size increased in Iberian forests, and tree size inequality decreased. In addition, we observed reductions in ingrowth and growth, and increases in mortality. Initial forest structure and water availability mainly modulated the temporal trends in forest structure and demography. The magnitude and direction of the interactive effects of climate and forest structure on forest demography changed over the two time periods analysed indicating non-stationary relationships between climate, forest structure and demography. Above-ground forest productivity increased due to a positive balance between ingrowth, growth and mortality. Despite increasing productivity over time, we observed an aggravation of the negative effects of climate change and increased competition on forest demography, reducing ingrowth and growth, and increasing mortality. Interestingly, our results suggest that the negative effects of climate change on forest demography could be ameliorated through forest management, which has profound implications for forest adaptation to climate change.

Biotic homogenization can decrease landscape-scale forest multifunctionality.

Many experiments have shown that local biodiversity loss impairs the ability of ecosystems to maintain multiple ecosystem functions at high levels (multifunctionality). In contrast, the role of biodiversity in driving ecosystem multifunctionality at landscape scales remains unresolved. We used a comprehensive pan-European dataset, including 16 ecosystem functions measured in 209 forest plots across six European countries, and performed simulations to investigate how local plot-scale richness of tree species (α-diversity) and their turnover between plots (ß-diversity) are related to landscape-scale multifunctionality. After accounting for variation in environmental conditions, we found that relationships between α-diversity and landscape-scale multifunctionality varied from positive to negative depending on the multifunctionality metric used. In contrast, when significant, relationships between ß-diversity and landscape-scale multifunctionality were always positive, because a high spatial turnover in species composition was closely related to a high spatial turnover in functions that were supported at high levels. Our findings have major implications for forest management and indicate that biotic homogenization can have previously unrecognized and negative consequences for large-scale ecosystem multifunctionality.

Continental mapping of forest ecosystem functions reveals a high but unrealised potential for forest multifunctionality.

Humans require multiple services from ecosystems, but it is largely unknown whether trade-offs between ecosystem functions prevent the realisation of high ecosystem multifunctionality across spatial scales. Here, we combined a comprehensive dataset (28 ecosystem functions measured on 209 forest plots) with a forest inventory dataset (105,316 plots) to extrapolate and map relationships between various ecosystem multifunctionality measures across Europe. These multifunctionality measures reflected different management objectives, related to timber production, climate regulation and biodiversity conservation/recreation. We found that trade-offs among them were rare across Europe, at both local and continental scales. This suggests a high potential for 'win-win' forest management strategies, where overall multifunctionality is maximised. However, across sites, multifunctionality was on average 45.8-49.8% below maximum levels and not necessarily highest in protected areas. Therefore, using one of the most comprehensive assessments so far, our study suggests a high but largely unrealised potential for management to promote multifunctional forests.

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.

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.

Effects of climate extremes on the terrestrial carbon cycle: concepts, processes and potential future impacts.

Extreme droughts, heat waves, frosts, precipitation, wind storms and other climate extremes may impact the structure, composition and functioning of terrestrial ecosystems, and thus carbon cycling and its feedbacks to the climate system. Yet, the interconnected avenues through which climate extremes drive ecological and physiological processes and alter the carbon balance are poorly understood. Here, we review the literature on carbon cycle relevant responses of ecosystems to extreme climatic events. Given that impacts of climate extremes are considered disturbances, we assume the respective general disturbance-induced mechanisms and processes to also operate in an extreme context. The paucity of well-defined studies currently renders a quantitative meta-analysis impossible, but permits us to develop a deductive framework for identifying the main mechanisms (and coupling thereof) through which climate extremes may act on the carbon cycle. We find that ecosystem responses can exceed the duration of the climate impacts via lagged effects on the carbon cycle. The expected regional impacts of future climate extremes will depend on changes in the probability and severity of their occurrence, on the compound effects and timing of different climate extremes, and on the vulnerability of each land-cover type modulated by management. Although processes and sensitivities differ among biomes, based on expert opinion, we expect forests to exhibit the largest net effect of extremes due to their large carbon pools and fluxes, potentially large indirect and lagged impacts, and long recovery time to regain previous stocks. At the global scale, we presume that droughts have the strongest and most widespread effects on terrestrial carbon cycling. Comparing impacts of climate extremes identified via remote sensing vs. ground-based observational case studies reveals that many regions in the (sub-)tropics are understudied. Hence, regional investigations are needed to allow a global upscaling of the impacts of climate extremes on global carbon-climate feedbacks.

The effects of phenotypic plasticity and local adaptation on forecasts of species range shifts under climate change.

Species are the unit of analysis in many global change and conservation biology studies; however, species are not uniform entities but are composed of different, sometimes locally adapted, populations differing in plasticity. We examined how intraspecific variation in thermal niches and phenotypic plasticity will affect species distributions in a warming climate. We first developed a conceptual model linking plasticity and niche breadth, providing five alternative intraspecific scenarios that are consistent with existing literature. Secondly, we used ecological niche-modeling techniques to quantify the impact of each intraspecific scenario on the distribution of a virtual species across a geographically realistic setting. Finally, we performed an analogous modeling exercise using real data on the climatic niches of different tree provenances. We show that when population differentiation is accounted for and dispersal is restricted, forecasts of species range shifts under climate change are even more pessimistic than those using the conventional assumption of homogeneously high plasticity across a species' range. Suitable population-level data are not available for most species so identifying general patterns of population differentiation could fill this gap. However, the literature review revealed contrasting patterns among species, urging greater levels of integration among empirical, modeling and theoretical research on intraspecific phenotypic variation.

Stand dynamics and tree coexistence in an analytical structured model: the role of recruitment.

Understanding the mechanisms of coexistence and niche partitioning in plant communities is a central question in ecology. Current theories of forest dynamics range between the so-called neutral theories which assume functional equivalence among coexisting species to forest simulators that explain species assemblages as the result of tradeoffs in species individual strategies at several ontogenetic stages. Progress in these questions has been hindered by the inherent difficulties of developing analytical size-structured models of stand dynamics. This precludes examination of the relative importance of each mechanism on tree coexistence. In previous simulation and analytical studies emphasis has been given to interspecific differences at the sapling stage, and less so to interspecific variation in seedling recruitment. In this study we develop a partial differential equation model of stand dynamics in which competition takes place at the recruitment stage. Species differ in their size-dependent growth rates and constant mortality rates. Recruitment is described as proportional to the basal area of conspecifics, to account for fecundity and seed supply per unit of basal area, and is corrected with a decreasing function of species specific basal area to account for competition. We first analyze conditions for population persistence in monospecific stands and second we investigate conditions of coexistence for two species. In the monospecific case we found a stationary stand structure based on an inequality between mortality rate and seed supply. In turn, intra-specific competition does not play any role on the asymptotic extinction or population persistence. In the two-species case we found that coexistence can be attained when the reciprocal negative effect on recruitment follows a given relation with respect to intraspecific competition. Specifically a tradeoff between recruitment potential (i.e. shade tolerance or predation avoidance) and fecundity or growth rate. This is to our knowledge the first study that describes coexistence mechanisms in an analytical size-structured model in terms of competitive differences at the regeneration state.

Anthropogenic land-use legacies underpin climate change-related risks to forest ecosystems.

Forest ecosystems with long-lasting human imprints can emerge worldwide as outcomes of land-use cessation. However, the interaction of these anthropogenic legacies with climate change impacts on forests is not well understood. Here, we set out how anthropogenic land-use legacies that persist in forest properties, following alterations in forest distribution, structure, and composition, can interact with climate change stressors. We propose a risk-based framework to identify anthropogenic legacies of land uses in forest ecosystems and quantify the impact of their interaction with climate-related stress on forest responses. Considering anthropogenic land-use legacies alongside environmental drivers of forest ecosystem dynamics will improve our predictive capacity of climate-related risks to forests and our ability to promote ecosystem resilience to climate change.

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.

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.

Using spectral indices as early warning signals of forest dieback: The case of drought-prone Pinus pinaster forests.

Forest dieback processes linked to drought are expected to increase due to climate warming. Remotely sensed data offer several advantages over common field monitoring methods such as the ability to observe large areas on a systematic basis and monitoring their changes, making them increasingly used to assess changes in forest health. Here we aim to use a combined approximation of fieldwork and remote sensing to explore possible links between forest dieback and land surface phenological and trend variables derived from long Landsat time series. Forest dieback was evaluated in the field over 31 plots in a Mediterranean, xeric Pinus pinaster forest. Landsat 31-year time series of three greenness (EVI, NDVI, SAVI) and two wetness spectral indices (NMDI and TCW) were derived covering the period 1990-2020. Spectral indices from time series were decomposed into trend and seasonality using a Bayesian estimator while the relationships of the phenological and trend variables among levels of damage were assessed using linear and additive mixed models. We have not found any statistical pieces of evidence of extension or shortening patterns for the length of the phenological season over the examined 31-year period. Our results indicate that the dieback process was mainly related to the trend component of the spectral indices series whereas the phenological metrics were not related to forest dieback. We also found that plots with more dying or damaged trees displayed lower spectral indices trends after a severe drought event in the middle of the 1990s, which confirms the Landsat-derived spectral indices as indicators of early-warning signals. Drops in trends occurred earlier for wetness indices rather than for greenness indices which suggests that the former could be more appropriate for dieback detection, i.e. they could be used as early warning signals of impending loss of tree vigor.

Competition and species coexistence in a metapopulation model: can fast asymmetric migration reverse the outcome of competition in a homogeneous environment?

We investigate whether asymmetric fast migration can modify the predictions of classical competition theory and, in particular revert species dominance. We consider a model of two species competing for an implicit resource on a habitat divided into two patches. Both patches are connected through constant migration rates and in each patch local dynamics are driven by a Lotka-Volterra competition system. Local competition is asymmetric with the same superior competitor in both patches. Migration is asymmetric, species dependent and fast in comparison to local competitive interactions. The species and patches are taken to be otherwise similar: in both patches we assume the same carrying capacities for both species, and the same growth rates and pair-wise competition coefficients for each species. We show that global dynamics can be described by a classical Lotka-Volterra competition model. We found that by modifying the ratio of intraspecific migration rates for both species all possible combinations of global species relative dominance can be achieved. We find specific conditions for which the local superior competitor is globally excluded. This is to our knowledge the first study showing that fast asymmetric migration can lead to inferior competitor dominance in a homogeneous environment. We conclude that disparity of temporal scales between migration and local dynamics may have important consequences for the maintenance of biodiversity in spatially structured populations.

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.

Are pine plantations valid tools for restoring Mediterranean forests? An assessment along abiotic and biotic gradients.

The ecological impacts of forest plantations are a focus of intense debate, from studies that consider plantations as "biological deserts" to studies showing positive effects on plant diversity and dynamics. This lack of consensus might be influenced by the scarcity of studies that examine how the ecological characteristics of plantations vary along abiotic and biotic gradients. Here we conducted a large-scale assessment of plant regeneration and diversity in plantations of southern Spain. Tree seedling and sapling density, plant species richness, and Shannon's (H') diversity index were analyzed in 442 pine plantation plots covering a wide gradient of climatic conditions, stand density, and distance to natural forests that act as seed sources. Pronounced variation in regeneration and diversity was found in plantation understories along the gradients explored. Low- to mid-altitude plantations showed a diverse and abundant seedling bank dominated by Quercus ilex, whereas high-altitude plantations showed a virtually monospecific seeding bank of Pinus sylvestris. Regeneration was null in plantations with stand densities exceeding 1500 pines/ha. Moderate plantation densities (500-1000 pines/ha) promoted recruitment in comparison to low or null canopy cover, suggesting the existence of facilitative interactions. Quercus ilex recruitment diminished exponentially with distance to the nearest Q. ilex forest. Richness and H' index values showed a hump-shaped distribution along the altitudinal and radiation gradients and decreased monotonically along the stand density gradient. From a management perspective, different strategies will be necessary depending on where a plantation lies along the gradients explored. Active management will be required in high-density plantations with arrested succession and low diversity. Thinning could redirect plantations toward more natural densities where facilitation predominates. Passive management might be recommended for low- to moderate-density plantations with active successional dynamics (e.g., toward oak or pine-oak forests at low to mid altitudes). Enrichment planting will be required to overcome seed limitation, especially in plantations far from natural forests. We conclude that plantations should be perceived as dynamic systems where successional trajectories and diversity levels are determined by abiotic constraints, complex balances of competitive and facilitative interactions, the spatial configuration of native seed sources, and species life-history traits.

How do trees respond to species mixing in experimental compared to observational studies?

For decades, ecologists have investigated the effects of tree species diversity on tree productivity at different scales and with different approaches ranging from observational to experimental study designs. Using data from five European national forest inventories (16,773 plots), six tree species diversity experiments (584 plots), and six networks of comparative plots (169 plots), we tested whether tree species growth responses to species mixing are consistent and therefore transferrable between those different research approaches. Our results confirm the general positive effect of tree species mixing on species growth (16% on average) but we found no consistency in species-specific responses to mixing between any of the three approaches, even after restricting comparisons to only those plots that shared similar mixtures compositions and forest types. These findings highlight the necessity to consider results from different research approaches when selecting species mixtures that should maximize positive forest biodiversity and functioning relationships.

Distribution of pines in the Iberian Peninsula agrees with species differences in foliage frost tolerance, not with vulnerability to freezing-induced xylem embolism.

Drought and frosts are major determinants of plant functioning and distribution. Both stresses can cause xylem embolism and foliage damage. The objective of this study was to analyse if the distribution of six common pine species along latitudinal and altitudinal gradients in Europe is related to their interspecific differences in frost tolerance and to the physiological mechanisms underlying species-specific frost tolerance. We also evaluate if frost tolerance depends on plant water status. We studied survival to a range of freezing temperatures in 2-year-old plants and assessed the percentage loss of hydraulic conductivity (PLC) due xylem embolism formation and foliage damage determined by needle electrolyte leakage (EL) after a single frost cycle to -15 °C and over a range of predawn water potential (ψpd) values. Species experiencing cold winters in their range (Pinus nigra J.F. Arnold, Pinus sylvestris L. and Pinus uncinata Raymond ex A. DC.) had the highest frost survival rates and lowest needle EL and soluble sugar (SS) concentration. In contrast, the pines inhabiting mild or cool winter locations (especially Pinus halepensis Mill. and Pinus pinea L. and, to a lesser extent, Pinus pinaster Ait.) had the lowest frost survival and highest needle EL and SS values. Freezing-induced PLC was very low and differences among species were not related to frost damage. Reduction in ψpd decreased leaf frost damage in P. pinea and P. sylvestris, increased it in P. uncinata and had a neutral effect on the rest of the species. This study demonstrates that freezing temperatures are a major environmental driver for pine distribution and suggests that interspecific differences in leaf frost sensitivity rather than vulnerability to freezing-induced embolism or SS explain pine juvenile frost survival.