Relative decline in density of Northern Hemisphere tree species in warm and arid regions of their climate niches.

Although climate change is expected to drive tree species toward colder and wetter regions of their distribution, broadscale empirical evidence is lacking. One possibility is that past and present human activities in forests obscure or alter the effects of climate. Here, using data from more than two million monitored trees from 73 widely distributed species, we quantify changes in tree species density within their climatic niches across Northern Hemisphere forests. We observe a reduction in mean density across species, coupled with a tendency toward increasing tree size. However, the direction and magnitude of changes in density exhibit considerable variability between species, influenced by stand development that results from previous stand-level disturbances. Remarkably, when accounting for stand development, our findings show a significant change in density toward cold and wet climatic conditions for 43% of the species, compared to only 14% of species significantly changing their density toward warm and arid conditions in both early- and late-development stands. The observed changes in climate-driven density showed no clear association with species traits related to drought tolerance, recruitment and dispersal capacity, or resource use, nor with the temperature or aridity affiliation of the species, leaving the underlying mechanism uncertain. Forest conservation policies and associated management strategies might want to consider anticipated long-term species range shifts alongside the integration of contemporary within-distribution density changes.

The unexpected influence of legacy conspecific density dependence.

When plants die, neighbours escape competition. Living conspecifics could disproportionately benefit because they are freed from negative intraspecific processes; however, if the negative effects of past conspecific neighbours persist, other species might be advantaged, and diversity might be maintained through legacy effects. We examined legacy effects in a mapped forest by modelling the survival of 37,212 trees of 23 species using four neighbourhood properties: living conspecific, living heterospecific, legacy conspecific (dead conspecifics) and legacy heterospecific densities. Legacy conspecific effects proved nearly four times stronger than living conspecific effects; changes in annual survival associated with legacy conspecific density were 1.5% greater than living conspecific effects. Over 90% of species were negatively impacted by legacy conspecific density, compared to 47% by living conspecific density. Our results emphasize that legacies of trees alter community dynamics, revealing that prior research may have underestimated the strength of density dependent interactions by not considering legacy effects.

The interacting effect of climate change and herbivory can trigger large-scale transformations of European temperate forests.

In many regions of Europe, large wild herbivores alter forest community composition through their foraging preferences, hinder the forest's natural adaptive responses to climate change, and reduce ecosystem resilience. We investigated a widespread European forest type, a mixed forest dominated by Picea abies, which has recently experienced an unprecedented level of disturbance across the continent. Using the forest landscape model iLand, we investigated the combined effect of climate change and herbivory on forest structure, composition, and carbon and identified conditions leading to ecosystem transitions on a 300-year timescale. Eight climate change scenarios, driven by Representative Concentration Pathways 4.5 and 8.5, combined with three levels of regeneration browsing, were tested. We found that the persistence of the current level of browsing pressure impedes adaptive changes in community composition and sustains the presence of the vulnerable yet less palatable P. abies. These development trajectories were tortuous, characterized by a high disturbance intensity. On the contrary, reduced herbivory initiated a transformation towards the naturally dominant broadleaved species that was associated with an increased forest carbon and a considerably reduced disturbance. The conditions of RCP4.5 combined with high and moderate browsing levels preserved the forest within its reference range of variability, defining the actual boundaries of resilience. The remaining combinations of browsing and climate change led to ecosystem transitions. Under RCP4.5 with browsing effects excluded, the new equilibrium conditions were achieved within 120 years, whereas the stabilization was delayed by 50-100 years under RCP8.5 with higher browsing intensities. We conclude that forests dominated by P. abies are prone to transitions driven by climate change. However, reducing herbivory can set the forest on a stable and predictable trajectory, whereas sustaining the current browsing levels can lead to heightened disturbance activity, extended transition times, and high variability in the target conditions.

The anthropogenic imprint on temperate and boreal forest demography and carbon turnover.

Aim: The sweeping transformation of the biosphere by humans over the last millennia leaves only limited windows into its natural state. Much of the forests that dominated temperate and southern boreal regions have been lost and those that remain typically bear a strong imprint of forestry activities and past land-use change, which have changed forest age structure and composition. Here, we ask how would the dynamics, structure and function of temperate and boreal forests differ in the absence of forestry and the legacies of land-use change? Location: Global. Time Period: 2001-2014, integrating over the legacy of disturbance events from 1875 to 2014. Major Taxa Studied: Trees. Methods: We constructed an empirical model of natural disturbance probability as a function of community traits and climate, based on observed disturbance rate and form across 77 protected forest landscapes distributed across three continents. Coupling this within a dynamic vegetation model simulating forest composition and structure, we generated estimates of stand-replacing disturbance return intervals in the absence of forestry for northern hemisphere temperate and boreal forests. We then applied this model to calculate forest stand age structure and carbon turnover rates. Results: Comparison with observed disturbance rates revealed human activities to have almost halved the median return interval of stand-replacing disturbances across temperate forest, with more moderate changes in the boreal region. The resulting forests are typically much younger, especially in northern Europe and south-eastern North America, resulting in a 32% reduction in vegetation carbon turnover time across temperate forests and a 7% reduction for boreal forests. Conclusions: The current northern hemisphere temperate forest age structure is dramatically out of equilibrium with its natural disturbance regimes. Shifts towards more nature-based approaches to forest policy and management should more explicitly consider the current disturbance surplus, as it substantially impacts carbon dynamics and litter (including deadwood) stocks.

Future supply of boreal forest ecosystem services is driven by management rather than by climate change.

Forests provide a wide variety of ecosystem services (ES) to society. The boreal biome is experiencing the highest rates of warming on the planet and increasing demand for forest products. To foresee how to maximize the adaptation of boreal forests to future warmer conditions and growing demands of forest products, we need a better understanding of the relative importance of forest management and climate change on the supply of ecosystem services. Here, using Finland as a boreal forest case study, we assessed the potential supply of a wide range of ES (timber, bilberry, cowberry, mushrooms, carbon storage, scenic beauty, species habitat availability and deadwood) given seven management regimes and four climate change scenarios. We used the forest simulator SIMO to project forest dynamics for 100 years into the future (2016-2116) and estimate the potential supply of each service using published models. Then, we tested the relative importance of management and climate change as drivers of the future supply of these services using generalized linear mixed models. Our results show that the effects of management on the future supply of these ES were, on average, 11 times higher than the effects of climate change across all services, but greatly differed among them (from 0.53 to 24 times higher for timber and cowberry, respectively). Notably, the importance of these drivers substantially differed among biogeographical zones within the boreal biome. The effects of climate change were 1.6 times higher in northern Finland than in southern Finland, whereas the effects of management were the opposite-they were three times higher in the south compared to the north. We conclude that new guidelines for adapting forests to global change should account for regional differences and the variation in the effects of climate change and management on different forest ES.

Large old trees increase growth under shifting climatic constraints: Aligning tree longevity and individual growth dynamics in primary mountain spruce forests.

In a world of accelerating changes in environmental conditions driving tree growth, tradeoffs between tree growth rate and longevity could curtail the abundance of large old trees (LOTs), with potentially dire consequences for biodiversity and carbon storage. However, the influence of tree-level tradeoffs on forest structure at landscape scales will also depend on disturbances, which shape tree size and age distribution, and on whether LOTs can benefit from improved growing conditions due to climate warming. We analyzed temporal and spatial variation in radial growth patterns from ~5000 Norway spruce (Picea abies [L.] H. Karst) live and dead trees from the Western Carpathian primary spruce forest stands. We applied mixed-linear modeling to quantify the importance of LOT growth histories and stand dynamics (i.e., competition and disturbance factors) on lifespan. Finally, we assessed regional synchronization in radial growth variability over the 20th century, and modeled the effects of stand dynamics and climate on LOTs recent growth trends. Tree age varied considerably among forest stands, implying an important role of disturbance as an age constraint. Slow juvenile growth and longer period of suppressed growth prolonged tree lifespan, while increasing disturbance severity and shorter time since last disturbance decreased it. The highest age was not achieved only by trees with continuous slow growth, but those with slow juvenile growth followed by subsequent growth releases. Growth trend analysis demonstrated an increase in absolute growth rates in response to climate warming, with late summer temperatures driving the recent growth trend. Contrary to our expectation that LOTs would eventually exhibit declining growth rates, the oldest LOTs (>400 years) continuously increase growth throughout their lives, indicating a high phenotypic plasticity of LOTs for increasing biomass, and a strong carbon sink role of primary spruce forests under rising temperatures, intensifying droughts, and increasing bark beetle outbreaks.

Drought then wildfire reveals a compound disturbance in a resprouting forest.

The frequency and intensity of forest disturbances, such as drought and fire, are increasing globally, with an increased likelihood of multiple disturbance events occurring in short succession. Disturbances layered over one another may influence the likelihood or intensity of subsequent events (a linked disturbance) or impact response and recovery trajectories (a compound disturbance), with substantial implications for ecological spatiotemporal vulnerability. This study evaluates evidence for disturbance interactions of drought followed by wildfire in a resprouting eucalypt-dominated forest (the Northern Jarrah Forest) in southwestern Australia. Sites were stratified by drought (high, low), from previous modeling and ground validation, and fire severity (high, moderate, unburnt), via remote sensing using the relative difference normalized burn ratio (RdNBR). Evidence of a linked disturbance was assessed via fine fuel consumption and fire severity. Compound disturbance effects were quantified at stand scale (canopy height, quadratic mean diameter, stem density) and stem scale (mortality). There was no evidence of prior drought influencing fine fuel consumption or fire severity and, hence, no evidence of a linked disturbance. However, compound disturbance effects were evident; stands previously affected by drought experienced smaller shifts in canopy height, quadratic mean diameter, and stem density than stands without prior drought impact. At the stem scale, size and fire severity were the strongest determinants of stem survival. Proportional resprouting height was greater in high drought sites than in low drought sites (p < 0.01), meaning, structurally, the low drought stands decreased in height more than the high drought stands. Thus, a legacy of the drought was evident after the wildfire. Although these resprouting eucalypt forests have been regarded as particularly resilient, this study illustrates how multiple disturbances can overwhelm the larger tree component and promote an abundance of smaller stems. We suggest that this is early evidence of a structural destabilization of these forests under a more fire-prone, hotter, and drier future climate.

Global tree-ring analysis reveals rapid decrease in tropical tree longevity with temperature.

Forests are the largest terrestrial biomass pool, with over half of this biomass stored in the highly productive tropical lowland forests. The future evolution of forest biomass depends critically on the response of tree longevity and growth rates to future climate. We present an analysis of the variation in tree longevity and growth rate using tree-ring data of 3,343 populations and 438 tree species and assess how climate controls growth and tree longevity across world biomes. Tropical trees grow, on average, two times faster compared to trees from temperate and boreal biomes and live significantly shorter, on average (186 ± 138 y compared to 322 ± 201 y outside the tropics). At the global scale, growth rates and longevity covary strongly with temperature. Within the warm tropical lowlands, where broadleaf species dominate the vegetation, we find consistent decreases in tree longevity with increasing aridity, as well as a pronounced reduction in longevity above mean annual temperatures of 25.4 °C. These independent effects of temperature and water availability on tree longevity in the tropics are consistent with theoretical predictions of increases in evaporative demands at the leaf level under a warmer and drier climate and could explain observed increases in tree mortality in tropical forests, including the Amazon, and shifts in forest composition in western Africa. Our results suggest that conditions supporting only lower tree longevity in the tropical lowlands are likely to expand under future drier and especially warmer climates.

Forest and woodland replacement patterns following drought-related mortality.

Forest vulnerability to drought is expected to increase under anthropogenic climate change, and drought-induced mortality and community dynamics following drought have major ecological and societal impacts. Here, we show that tree mortality concomitant with drought has led to short-term (mean 5 y, range 1 to 23 y after mortality) vegetation-type conversion in multiple biomes across the world (131 sites). Self-replacement of the dominant tree species was only prevalent in 21% of the examined cases and forests and woodlands shifted to nonwoody vegetation in 10% of them. The ultimate temporal persistence of such changes remains unknown but, given the key role of biological legacies in long-term ecological succession, this emerging picture of postdrought ecological trajectories highlights the potential for major ecosystem reorganization in the coming decades. Community changes were less pronounced under wetter postmortality conditions. Replacement was also influenced by management intensity, and postdrought shrub dominance was higher when pathogens acted as codrivers of tree mortality. Early change in community composition indicates that forests dominated by mesic species generally shifted toward more xeric communities, with replacing tree and shrub species exhibiting drier bioclimatic optima and distribution ranges. However, shifts toward more mesic communities also occurred and multiple pathways of forest replacement were observed for some species. Drought characteristics, species-specific environmental preferences, plant traits, and ecosystem legacies govern postdrought species turnover and subsequent ecological trajectories, with potential far-reaching implications for forest biodiversity and ecosystem services.

Fast but steady: An integrated leaf-stem-root trait syndrome for woody forest invaders.

Successful control and prevention of biological invasions depend on identifying traits of non-native species that promote fitness advantages in competition with native species. Here, we show that, among 76 native and non-native woody plants of deciduous forests of North America, invaders express a unique functional syndrome that combines high metabolic rate with robust leaves of longer lifespan and a greater duration of annual carbon gain, behaviours enabled by seasonally plastic xylem structure and rapid production of thin roots. This trait combination was absent in all native species examined and suggests the success of forest invaders is driven by a novel resource-use strategy. Furthermore, two traits alone-annual leaf duration and nuclear DNA content-separated native and invasive species with 93% accuracy, supporting the use of functional traits in invader risk assessments. A trait syndrome reflecting both fast growth capacity and understorey persistence may be a key driver of forest invasions.

Soils and topography control natural disturbance rates and thereby forest structure in a lowland tropical landscape.

Tree mortality is a major control over tropical forest carbon stocks globally but the strength of associations between abiotic drivers and tree mortality within forested landscapes is poorly understood. Here, we used repeat drone photogrammetry across 1500 ha of forest in Central Panama over 5 years to quantify spatial variation in canopy disturbance rates and its predictors. We identified 11,153 canopy disturbances greater than 25 m2 in area, including treefalls, large branchfalls and standing dead trees, affecting 1.9% of area per year. Soil type, forest age and topography explained up to 46%-67% of disturbance rate variation at spatial grains of 58-64 ha, with higher rates in older forests, steeper slopes and local depressions. Furthermore, disturbance rates predicted the proportion of low canopy area across the landscape, and mean canopy height in old growth forests. Thus abiotic factors drive variation in disturbance rates and thereby forest structure at landscape scales.

Size- and environment-driven seedling survival and growth are mediated by leaf functional traits.

Ecologists usually find that plant demography (e.g. survival and growth) changes along with plant size and environmental gradients, which suggests the effects of ontogeny-related processes and abiotic filtering. However, the role of functional traits underlying the size- and environment-demography relationships is usually overlooked. By measuring individual-level leaf traits of more than 2700 seedlings in a temperate forest, we evaluated how seedling traits mediated the size- and environment-demography relationships. We found leaves were larger for taller seedlings; leaf economics traits were more conservative in taller seedlings and under high-light and low-elevation conditions. Structural equation modelling showed that a higher survival probability for taller seedlings was indirectly driven by their larger leaf area. Although taller seedlings had lower growth rates, larger and more resource-conservative leaves could promote the growth of these tall seedlings. Environmental variables did not influence seedling survival and growth directly but did influence growth indirectly by mediating trait variation. Finally, species-specific variation in traits along with size and environments was associated with the species-specific variation in seedling survival and growth. Our study suggests that not only plant ontogeny- and environment-related ecological processes, but functional traits are also important intermediary agents underlying plant size- and environment-demography relationships.

Emerging infectious disease triggered a trophic cascade and enhanced recruitment of a masting tree.

There are several mechanisms that allow plants to temporarily escape from top-down control. One of them is trophic cascades triggered by top predators or pathogens. Another is satiation of consumers by mast seeding. These two mechanisms have traditionally been studied in separation. However, their combined action may have a greater effect on plant release than either process alone. In 2015, an outbreak of a disease (African swine fever, ASF) caused a crash in wild boar (Sus scrofa) abundance in Bialowieza Primeval Forest. Wild boar are important consumers of acorns and are difficult to satiate relative to less mobile granivores. We hypothesized that the joint action of the ASF outbreak and masting would enhance regeneration of oaks (Quercus robur). Data from ungulate exclosures demonstrated that ASF led to reduction in acorn predation. Tree seedling data indicated that oak recruitment increased twofold relative to pre-epidemic period. Our results showed that perturbations caused by wildlife disease travel through food webs and influence forest dynamics. The outbreak of ASF acted synergistically with masting and removed herbivore top-down control of oaks by mobile consumers. This illustrates that the ASF epidemic that currently occurs across Europe can have broad effects on forest dynamics.

Deciphering the fingerprint of disturbance on the three-dimensional structure of the world's forests.

Canopy gaps and the processes that generate them play an integral role in shaping the structure and dynamics of forests. However, it is only with recent advances in remote sensing technologies such as airborne laser scanning that studying canopy gaps at scale has become a reality. Consequently, we still lack an understanding of how the size distribution and spatial organization of canopy gaps varies among forests ecosystems, nor have we determined whether these emergent properties can be reconciled with existing theories of forest dynamics. Here, I outline a roadmap for integrating remote sensing with field data and individual-based models to build a comprehensive picture of how environmental constraints and disturbance regimes shape the three-dimensional structure of the world's forests.

Warming increased bark beetle-induced tree mortality by 30% during an extreme drought in California.

Quantifying the responses of forest disturbances to climate warming is critical to our understanding of carbon cycles and energy balances of the Earth system. The impact of warming on bark beetle outbreaks is complex as multiple drivers of these events may respond differently to warming. Using a novel model of bark beetle biology and host tree interactions, we assessed how contemporary warming affected western pine beetle (Dendroctonus brevicomis) populations and mortality of its host, ponderosa pine (Pinus ponderosa), during an extreme drought in the Sierra Nevada, California, United States. When compared with the field data, our model captured the western pine beetle flight timing and rates of ponderosa pine mortality observed during the drought. In assessing the influence of temperature on western pine beetles, we found that contemporary warming increased the development rate of the western pine beetle and decreased the overwinter mortality rate of western pine beetle larvae leading to increased population growth during periods of lowered tree defense. We attribute a 29.9% (95% CI: 29.4%-30.2%) increase in ponderosa pine mortality during drought directly to increases in western pine beetle voltinism (i.e., associated with increased development rates of western pine beetle) and, to a much lesser extent, reductions in overwintering mortality. These findings, along with other studies, suggest each degree (°C) increase in temperature may have increased the number of ponderosa pine killed by upwards of 35%-40% °C-1 if the effects of compromised tree defenses (15%-20%) and increased western pine beetle populations (20%) are additive. Due to the warming ability to considerably increase mortality through the mechanism of bark beetle populations, models need to consider climate's influence on both host tree stress and the bark beetle population dynamics when determining future levels of tree mortality.

Demographic composition, not demographic diversity, predicts biomass and turnover across temperate and tropical forests.

The growth and survival of individual trees determine the physical structure of a forest with important consequences for forest function. However, given the diversity of tree species and forest biomes, quantifying the multitude of demographic strategies within and across forests and the way that they translate into forest structure and function remains a significant challenge. Here, we quantify the demographic rates of 1961 tree species from temperate and tropical forests and evaluate how demographic diversity (DD) and demographic composition (DC) differ across forests, and how these differences in demography relate to species richness, aboveground biomass (AGB), and carbon residence time. We find wide variation in DD and DC across forest plots, patterns that are not explained by species richness or climate variables alone. There is no evidence that DD has an effect on either AGB or carbon residence time. Rather, the DC of forests, specifically the relative abundance of large statured species, predicted both biomass and carbon residence time. Our results demonstrate the distinct DCs of globally distributed forests, reflecting biogeography, recent history, and current plot conditions. Linking the DC of forests to resilience or vulnerability to climate change, will improve the precision and accuracy of predictions of future forest composition, structure, and function.

Small and slow is safe: On the drought tolerance of tropical tree species.

Understanding how evolutionary history and the coordination between trait trade-off axes shape the drought tolerance of trees is crucial to predict forest dynamics under climate change. Here, we compiled traits related to drought tolerance and the fast-slow and stature-recruitment trade-off axes in 601 tropical woody species to explore their covariations and phylogenetic signals. We found that xylem resistance to embolism (P50) determines the risk of hydraulic failure, while the functional significance of leaf turgor loss point (TLP) relies on its coordination with water use strategies. P50 and TLP exhibit weak phylogenetic signals and substantial variation within genera. TLP is closely associated with the fast-slow trait axis: slow species maintain leaf functioning under higher water stress. P50 is associated with both the fast-slow and stature-recruitment trait axes: slow and small species exhibit more resistant xylem. Lower leaf phosphorus concentration is associated with more resistant xylem, which suggests a (nutrient and drought) stress-tolerance syndrome in the tropics. Overall, our results imply that (1) drought tolerance is under strong selective pressure in tropical forests, and TLP and P50 result from the repeated evolutionary adaptation of closely related taxa, and (2) drought tolerance is coordinated with the ecological strategies governing tropical forest demography. These findings provide a physiological basis to interpret the drought-induced shift toward slow-growing, smaller, denser-wooded trees observed in the tropics, with implications for forest restoration programmes.

Trajectories of wildfire behavior under climate change. Can forest management mitigate the increasing hazard?

Mediterranean forests and fire regimes are closely intertwined. Global change is likely to alter both forest dynamics and wildfire activity, ultimately threatening the provision of ecosystem services and posing greater risks to society. In this paper we evaluate future wildfire behavior by coupling climate projections with simulation models of forest dynamics and wildfire hazard. To do so, we explore different forest management scenarios reflecting different narratives related to EU forestry (promotion of carbon stocks, reduction of water vulnerability, biomass production and business-as-usual) under the RCP 4.5 and RCP 8.5 climate pathways in the period 2020-2100. We used as a study model pure submediterranean Pinus nigra forests of central Catalonia (NE Spain). Forest dynamics were simulated from the 3rd National Forest Inventory (143 stands) using SORTIE-nd software based on climate projections under RCPs 4.5 and 8.5. The climate products were also used to estimate fuel moisture conditions (both live and dead) and wind speed. Fuel parameters and fire behavior were then simulated, selecting crown fire initiation potential and rate of spread as key indicators. The results revealed consistent trade-offs between forest dynamics, climate and wildfire. Despite the clear influence exerted by climate, forest management modulates fire behavior, resulting in different trends depending on the climatic pathway. In general, the maintenance of current practices would result in the highest rates of crown fire activity, while management for water vulnerability reduction is postulated as the best alternative to surmount the increasingly hazardous conditions envisaged in RCP 8.5.

Ecological legacies of past human activities in Amazonian forests.

In Amazonia, human activities that occurred hundreds of years ago in the pre-European era can leave long-lasting effects on the forests - termed ecological legacies. These legacies include the intentional or nonintentional enrichment or depletion of certain species. The persistence of these legacies through time varies by species, and creates complex long-term trajectories of post-disturbance succession that affect ecosystem processes for hundreds of years. Most of our knowledge of Amazonian biodiversity and carbon storage comes from a series of several hundred forest plots, and we only know the disturbance history of four of them. More empirical data are needed to determine the degree to which past human activities and their ecological legacies affect our current understanding of Amazonian forest ecology.

Increased mortality of tropical tree seedlings during the extreme 2015-16 El Niño.

As extreme climate events are predicted to become more frequent because of global climate change, understanding their impacts on natural systems is crucial. Tropical forests are vulnerable to droughts associated with extreme El Niño events. However, little is known about how tropical seedling communities respond to El Niño-related droughts, even though patterns of seedling survival shape future forest structure and diversity. Using long-term data from eight tropical moist forests spanning a rainfall gradient in central Panama, we show that community-wide seedling mortality increased by 11% during the extreme 2015-16 El Niño, with mortality increasing most in drought-sensitive species and in wetter forests. These results indicate that severe El Niño-related droughts influence understory dynamics in tropical forests, with effects varying both within and across sites. Our findings suggest that predicted increases in the frequency of extreme El Niño events will alter tropical plant communities through their effects on early life stages.