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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.

The carbon isotope ratio (δ13 C) in tree rings is commonly used to derive estimates of the assimilation-to-stomatal conductance rate of trees, that is, intrinsic water-use efficiency (iWUE). Recent studies have observed increased iWUE in response to rising atmospheric CO2 concentrations (Ca ), in many different species, genera and biomes. However, increasing rates of iWUE vary widely from one study to another, likely because numerous covarying factors are involved. Here, we quantified changes in iWUE of two widely distributed boreal conifers using tree samples from a forest inventory network that were collected across a wide range of growing conditions (assessed using the site index, SI), developmental stages and stand histories. Using tree-ring isotopes analysis, we assessed the magnitude of increase in iWUE after accounting for the effects of tree size, stand age, nitrogen deposition, climate and SI. We also estimated how growth conditions have modulated tree physiological responses to rising Ca . We found that increases in tree size and stand age greatly influenced iWUE. The effect of Ca on iWUE was strongly reduced after accounting for these two variables. iWUE increased in response to Ca , mostly in trees growing on fertile stands, whereas iWUE remained almost unchanged on poor sites. Our results suggest that past studies could have overestimated the CO2 effect on iWUE, potentially leading to biased inferences about the future net carbon balance of the boreal forest. We also observed that this CO2 effect is weakening, which could affect the future capacity of trees to resist and recover from drought episodes.

In recent decades, many forest die-off events have been reported in relation to climate-change-induced episodes, such as droughts and heat waves. To understand how these extreme climatic events induce forest die-off, it is important to find a tool to standardize the climatic conditions experienced by different populations during a specific climatic event, taking into account the historic climatic conditions of the site where these populations live (bioclimatic niche). In this study, we used estimates of climatic suitability calculated from species distribution models (SDMs) for such purpose. We studied forest die-off across France during the 2003 heatwave that affected Western Europe, using 2,943 forest inventory plots dominated by 14 single tree species. Die-off severity was estimated by Normalized Difference Vegetation Index (NDVI) loss using Moderate-resolution Imaging Spectroradiometer remote sensor imagery. Climatic suitability at the local level during the historical 1979-2002 period (HCS), the episode time (2003; ECS) and suitability deviance during the historical period (HCS-SD) were calculated for each species by means of boosted regression tree models using the CHELSA climate database and occurrences extracted from European forest inventories. Low HCS-SD and high mean annual temperature explained the overall regional pattern of vulnerability to die-off across different monospecific forests. The combination of high historical and low episode climatic suitability also contributed significantly to overall forest die-off. Furthermore, we observed different species-specific relationships between die-off vulnerability and climatic suitability: Sub-Mediterranean and Mediterranean species tended to be vulnerable in historically more suitable localities (high HCS), whereas Euro-Siberian species presented greater vulnerability when the hot drought episode was more intense. We demonstrated that at regional scale, past climatic legacy plays an important role in explaining NDVI loss during the episode. Moreover, we demonstrated that SDMs-derived indexes, such as HCS, ECS and HCS-SD, could constitute a tool for standardizing the ways that populations and species experience climatic variability across time and space.

Carbon offsets are widely used by individuals, corporations, and governments to mitigate their greenhouse gas emissions on the assumption that offsets reflect equivalent climate benefits achieved elsewhere. These climate-equivalence claims depend on offsets providing real and additional climate benefits beyond what would have happened, counterfactually, without the offsets project. Here, we evaluate the design of California's prominent forest carbon offsets program and demonstrate that its climate-equivalence claims fall far short on the basis of directly observable evidence. By design, California's program awards large volumes of offset credits to forest projects with carbon stocks that exceed regional averages. This paradigm allows for adverse selection, which could occur if project developers preferentially select forests that are ecologically distinct from unrepresentative regional averages. By digitizing and analyzing comprehensive offset project records alongside detailed forest inventory data, we provide direct evidence that comparing projects against coarse regional carbon averages has led to systematic over-crediting of 30.0 million tCO2 e (90% CI: 20.5-38.6 million tCO2 e) or 29.4% of the credits we analyzed (90% CI: 20.1%-37.8%). These excess credits are worth an estimated $410 million (90% CI: $280-$528 million) at recent market prices. Rather than improve forest management to store additional carbon, California's forest offsets program creates incentives to generate offset credits that do not reflect real climate benefits.

Changes in growing season climate are often the foci of research exploring forest response to climate change. By contrast, little is known about tree growth response to projected declines in winter snowpack and increases in soil freezing in seasonally snow-covered forest ecosystems, despite extensive documentation of the importance of winter climate in mediating ecological processes. We conducted a 5-year snow-removal experiment whereby snow was removed for the first 4-5 weeks of winter in a northern hardwood forest at the Hubbard Brook Experimental Forest in New Hampshire, USA. Our results indicate that adverse impacts of reduced snowpack and increased soil freezing on the physiology of Acer saccharum (sugar maple), a dominant species across northern temperate forests, are accompanied by a 40 ± 3% reduction in aboveground woody biomass increment, averaged across the 6 years following the start of the experiment. Further, we find no indication of growth recovery 1 year after cessation of the experiment. Based on these findings, we integrate spatial modeling of snowpack depth with forest inventory data to develop a spatially explicit, regional-scale assessment of the vulnerability of forest aboveground growth to projected declines in snowpack depth and increased soil frost. These analyses indicate that nearly 65% of sugar maple basal area in the northeastern United States resides in areas that typically experience insulating snowpack. However, under the RCP 4.5 and 8.5 emissions scenarios, we project a 49%-95% reduction in forest area experiencing insulating snowpack by the year 2099 in the northeastern United States, leaving large areas of northern forest vulnerable to these changes in winter climate, particularly along the northern edge of the region. Our study demonstrates that research focusing on growing season climate alone overestimates the stimulatory effect of warming temperatures on tree and forest growth in seasonally snow-covered forests.

Symbiotic nitrogen (N)-fixing trees can drive N and carbon cycling and thus are critical components of future climate projections. Despite detailed understanding of how climate influences N-fixation enzyme activity and physiology, comparatively little is known about how climate influences N-fixing tree abundance. Here, we used forest inventory data from the USA and Mexico (>125,000 plots) along with climate data to address two questions: (1) How does the abundance distribution of N-fixing trees (rhizobial, actinorhizal, and both types together) vary with mean annual temperature (MAT) and precipitation (MAP)? (2) How will changing climate shift the abundance distribution of N-fixing trees? We found that rhizobial N-fixing trees were nearly absent below 15°C MAT, but above 15°C MAT, they increased in abundance as temperature rose. We found no evidence for a hump-shaped response to temperature throughout the range of our data. Rhizobial trees were more abundant in dry than in wet ecosystems. By contrast, actinorhizal trees peaked in abundance at 5-10°C MAT and were least abundant in areas with intermediate precipitation. Next, we used a climate-envelope approach to project how N-fixing tree relative abundance might change in the future. The climate-envelope projection showed that rhizobial N-fixing trees will likely become more abundant in many areas by 2080, particularly in the southern USA and western Mexico, due primarily to rising temperatures. Projections for actinorhizal N-fixing trees were more nuanced due to their nonmonotonic dependence on temperature and precipitation. Overall, the dominant trend is that warming will increase N-fixing tree abundance in much of the USA and Mexico, with large increases up to 40° North latitude. The quantitative link we provide between climate and N-fixing tree abundance can help improve the representation of symbiotic N fixation in Earth System Models.

Rising global temperatures are suggested to be drivers of shifts in tree species ranges. The resulting changes in community composition may negatively impact forest ecosystem function. However, long-term shifts in tree species ranges remain poorly documented. We test for shifts in the northern range limits of 16 temperate tree species in Quebec, Canada, using forest inventory data spanning three decades, 15° of longitude and 7° of latitude. Range shifts were correlated with climate warming and dispersal traits to understand potential mechanisms underlying changes. Shifts were calculated as the change in the 95th percentile of latitudinal occurrence between two inventory periods (1970–1978, 2000–2012) and for two life stages: saplings and adults. We also examined sapling and adult range offsets within each inventory, and changes in the offset through time. Tree species ranges shifted predominantly northward, although species responses varied. As expected shifts were greater for tree saplings, 0.34 km yr−1, than for adults, 0.13 km yr−1. Range limits were generally further north for adults compared to saplings, but the difference diminished through time, consistent with patterns observed for range shifts within each life stage. This suggests caution should be exercised when interpreting geographic range offsets between life stages as evidence of range shifts in the absence of temporal data. Species latitudinal velocities were on average

Large-diameter, tall-stature, and big-crown trees are the main stand structures of forests, generally contributing a large fraction of aboveground biomass, and hence play an important role in climate change mitigation strategies. Here, we hypothesized that the effects of large-diameter, tall-stature, and big-crown trees overrule the effects of species richness and remaining trees attributes on aboveground biomass in tropical forests (i.e., we term the "big-sized trees hypothesis"). Specifically, we assessed the importance of: (a) the "top 1% big-sized trees effect" relative to species richness; (b) the "99% remaining trees effect" relative to species richness; and (c) the "top 1% big-sized trees effect" relative to the "99% remaining trees effect" and species richness on aboveground biomass. Using environmental factor and forest inventory datasets from 712 tropical forest plots in Hainan Island of southern China, we tested several structural equation models for disentangling the relative effects of big-sized trees, remaining trees attributes, and species richness on aboveground biomass, while considering for the full (indirect effects only) and partial (direct and indirect effects) mediation effects of climatic and soil conditions, as well as interactions between species richness and trees attributes. We found that top 1% big-sized trees attributes strongly increased aboveground biomass (i.e., explained 55%-70% of the accounted variation) compared to species richness (2%-18%) and 99% remaining trees attributes (6%-10%). In addition, species richness increased aboveground biomass indirectly via increasing big-sized trees but via decreasing remaining trees. Hence, we show that the "big-sized trees effect" overrides the effects of remaining trees attributes and species richness on aboveground biomass in tropical forests. This study also indicates that big-sized trees may be more susceptible to atmospheric drought. We argue that the effects of big-sized trees on species richness and aboveground biomass should be tested for better understanding of the ecological mechanisms underlying forest functioning.

Rapid warming and changes in water availability at high latitudes alter resource abundance, tree competition, and disturbance regimes. While these changes are expected to disrupt the functioning of boreal forests, their ultimate implications for forest composition are uncertain. In particular, recent site-level studies of the Alaskan boreal forest have reported both increases and decreases in productivity over the past few decades. Here, we test the idea that variations in Alaskan forest growth and mortality rates are contingent on species composition. Using forest inventory measurements and climate data from plots located throughout interior and south-central Alaska, we show significant growth and mortality responses associated with competition, midsummer vapor pressure deficit, and increased growing season length. The governing climate and competition processes differed substantially across species. Surprisingly, the most dramatic climate response occurred in the drought tolerant angiosperm species, trembling aspen, and linked high midsummer vapor pressure deficits to decreased growth and increased insect-related mortality. Given that species composition in the Alaskan and western Canadian boreal forests is projected to shift toward early-successional angiosperm species due to fire regime, these results underscore the potential for a reduction in boreal productivity stemming from increases in midsummer evaporative demand.

The role of tree mortality in the global carbon balance is complicated by strong spatial and temporal heterogeneity that arises from the stochastic nature of carbon loss through disturbance. Characterizing spatio-temporal variation in mortality (including disturbance) and its effects on forest and carbon dynamics is thus essential to understanding the current global forest carbon sink, and to predicting how it will change in future. We analyzed forest inventory data from the eastern United States to estimate plot-level variation in mortality (relative to a long-term background rate for individual trees) for nine distinct forest regions. Disturbances that produced at least a fourfold increase in tree mortality over an approximately 5 year interval were observed in 1–5% of plots in each forest region. The frequency of disturbance was lowest in the northeast, and increased southwards along the Atlantic and Gulf coasts as fire and hurricane disturbances became progressively more common. Across the central and northern parts of the region, natural disturbances appeared to reflect a diffuse combination of wind, insects, disease, and ice storms. By linking estimated covariation in tree growth and mortality over time with a data-constrained forest dynamics model, we simulated the implications of stochastic variation in mortality for long-term aboveground biomass changes across the eastern United States. A geographic gradient in disturbance frequency induced notable differences in biomass dynamics between the least- and most-disturbed regions, with variation in mortality causing the latter to undergo considerably stronger fluctuations in aboveground stand biomass over time. Moreover, regional simulations showed that a given long-term increase in mean mortality rates would support greater aboveground biomass when expressed through disturbance effects compared with background mortality, particularly for early-successional species. The effects of increased tree mortality on carbon stocks and forest composition may thus depend partly on whether future mortality increases are chronic or episodic in nature.

Tree species are expected to track warming climate by shifting their ranges to higher latitudes or elevations, but current evidence of latitudinal range shifts for suites of species is largely indirect. In response to global warming, offspring of trees are predicted to have ranges extend beyond adults at leading edges and the opposite relationship at trailing edges. Large-scale forest inventory data provide an opportunity to compare present latitudes of seedlings and adult trees at their range limits. Using the USDA Forest Service’s Forest Inventory and Analysis data, we directly compared seedling and tree 5th and 95th percentile latitudes for 92 species in 30 longitudinal bands for 43 334 plots across the eastern United States. We further compared these latitudes with 20th century temperature and precipitation change and functional traits, including seed size and seed spread rate. Results suggest that 58.7% of the tree species examined show the pattern expected for a population undergoing range contraction, rather than expansion, at both northern and southern boundaries. Fewer species show a pattern consistent with a northward shift (20.7%) and fewer still with a southward shift (16.3%). Only 4.3% are consistent with expansion at both range limits. When compared with the 20th century climate changes that have occurred at the range boundaries themselves, there is no consistent evidence that population spread is greatest in areas where climate has changed most; nor are patterns related to seed size or dispersal characteristics. The fact that the majority of seedling extreme latitudes are less than those for adult trees may emphasize the lack of evidence for climate-mediated migration, and should increase concerns for the risks posed by climate change.

Substantial uncertainty surrounds how forest ecosystems will respond to the simultaneous impacts of multiple global change drivers. Long-term forest dynamics are sensitive to changes in tree mortality rates; however, we lack an understanding of the relative importance of the factors that affect tree mortality across different spatial and temporal scales. We used the US Forest Service Forest Inventory and Analysis database to evaluate the drivers of tree mortality for eastern temperate forest at the individual-level across spatial scales from tree to landscape to region. We investigated 13 covariates in four categories: climate, air pollutants, topography, and stand characteristics. Overall, we found that tree mortality was most sensitive to stand characteristics and air pollutants. Different functional groups also varied considerably in their sensitivity to environmental drivers. This research highlights the importance of considering the interactions among multiple global change agents in shaping forest ecosystems.

Stem radial growth responds to environmental conditions, and has been widely used as a proxy to study long-term patterns of tree growth and to assess the impact of environmental changes on growth patterns. In this study, we use a tree ring dataset from the Catalan Ecological and Forest Inventory to study the temporal variability of Scots pine (Pinus sylvestris L.) stem growth during the 20th century across a relatively large region (Catalonia, NE Spain) close to the southern limit of the distribution of the species. Basal area increment (BAI) was modelled as a function of tree size and environmental variables by means of mixed effects models. Our results showed an overall increase of 84% in Scots pine BAI during the 20th century, consistent with most previous studies for temperate forests. This trend was associated with increased atmospheric CO2 concentrations and, possibly, with a general increase in nutrient availability, and we interpreted it as a fertilization effect. Over the same time period, there was also a marked increase in temperature across the study region (0.19 °C per decade on average). This warming had a negative impact on radial growth, particularly at the drier sites, but its magnitude was not enough to counteract the fertilization effect. In fact, the substantial warming observed during the 20th century in the study area did not result in a clear pattern of increased summer drought stress because of the large variability in precipitation, which did not show any clear time trend. But the situation may change in the future if temperatures continue to rise and/or precipitation becomes scarcer. Such a change could potentially reverse the temporal trend in growth, particularly at the driest sites, and is suggested in our data by the relative constancy of radial growth after ca. 1975, coinciding with the warmer period. If this situation is representative of other relatively dry, temperate forests, the implications for the regional carbon balance would be substantial.

The amount and spatial distribution of forest biomass in the Amazon basin is a major source of uncertainty in estimating the flux of carbon released from land-cover and land-use change. Direct measurements of aboveground live biomass (AGLB) are limited to small areas of forest inventory plots and site-specific allometric equations that cannot be readily generalized for the entire basin. Furthermore, there is no spaceborne remote sensing instrument that can measure tropical forest biomass directly. To determine the spatial distribution of forest biomass of the Amazon basin, we report a method based on remote sensing metrics representing various forest structural parameters and environmental variables, and more than 500 plot measurements of forest biomass distributed over the basin. A decision tree approach was used to develop the spatial distribution of AGLB for seven distinct biomass classes of lowland old-growth forests with more than 80% accuracy. AGLB for other vegetation types, such as the woody and herbaceous savanna and secondary forests, was directly estimated with a regression based on satellite data. Results show that AGLB is highest in Central Amazonia and in regions to the east and north, including the Guyanas. Biomass is generally above 300Mgha(sup 1) here except in areas of intense logging or open floodplains. In Western Amazonia, from the lowlands of Peru, Ecuador, and Colombia to the Andean mountains, biomass ranges from 150 to 300Mgha(sup 1). Most transitional and seasonal forests at the southern and northwestern edges of the basin have biomass ranging from 100 to 200Mgha(sup 1). The AGLB distribution has a significant correlation with the length of the dry season. We estimate that the total carbon in forest biomass of the Amazon basin, including the dead and below ground biomass, is 86 PgC with +/- 20% uncertainty.

Ecotones are transition zones that form, in forests, where distinct forest types meet across a climatic gradient. In mountains, ecotones are compressed and act as potential harbingers of species shifts that accompany climate change. As the climate warms in New England, USA, high-elevation boreal forests are expected to recede upslope, with northern hardwood species moving up behind. Yet recent empirical studies present conflicting findings on this dynamic, reporting both rapid upward ecotonal shifts and concurrent increases in boreal species within the region. These discrepancies may result from the limited spatial extent of observations. We developed a method to model and map the montane forest ecotone using Landsat imagery to observe change at scales not possible for plot-based studies, covering mountain peaks over 39 000 km(2) . Our results show that ecotones shifted downward or stayed stable on most mountains between 1991 and 2010, but also shifted upward in some cases (13-15% slopes). On average, upper ecotone boundaries moved down -1.5 m yr(-1) in the Green Mountains, VT, and -1.3 m yr(-1) in the White Mountains, NH. These changes agree with remeasured forest inventory data from Hubbard Brook Experimental Forest, NH, and suggest that processes of boreal forest recovery from prior red spruce decline, or human land use and disturbance, may swamp out any signal of climate-mediated migration in this ecosystem. This approach represents a powerful framework for evaluating similar ecotonal dynamics in other mountainous regions of the globe.

Assessments made over the past few decades have suggested that boreal forests may act as a sink for atmospheric carbon dioxide. However, the fate of the newly accumulated carbon in the living forest biomass is not well understood, and the estimates of carbon sinks vary greatly from one assessment to another. Analysis of remote sensing data has indicated that the carbon sinks in the Russian forests are larger than what has been estimated from forest inventories. In this study, we show that over the past four decades, the allometric relationships among various plant parts have changed in the Russian forests. To this end, we employ two approaches: (1) analysis of the database, which contains 3196 sample plots; and (2) application of developed models to forest inventory data. Within the forests as a whole, when assessed at the continental scale, we detect a pronounced increase in the share of green parts (leaves and needles). However, there is a large geographical variation. The shift has been largest within the European Russia, where summer temperatures and precipitation have increased. In the Northern Taiga of Siberia, where the climate has become warmer but drier, the fraction of the green parts has decreased while the fractions of aboveground wood and roots have increased. These changes are consistent with experiments and mathematical models that predict a shift of carbon allocation to transpiring foliage with increasing temperature and lower allocation with increasing soil drought. In light of this, our results are a possible demonstration of the acclimation of trees to ongoing warming and changes in the surface water balance. Independent of the nature of the observed changes in allometric ratios, the increase in the share of green parts may have caused a misinterpretation of the satellite data and a systematic overestimation by remote sensing methods of the carbon sink for living biomass of the Russian forest.

Temperate forest ecosystems have recently been identified as an important net sink in the global carbon budget. The factors responsible for the strength of the sinks and their permanence, however, are less evident. In this paper, we quantify the present carbon sequestration in Thuringian managed coniferous forests. We quantify the effects of indirect human-induced environmental changes (increasing temperature, increasing atmospheric CO2 concentration and nitrogen fertilization), during the last century using BIOME-BGC, as well as the legacy effect of the current age-class distribution (forest inventories and BIOME-BGC). We focused on coniferous forests because these forests represent a large area of central European forests and detailed forest inventories were available. The model indicates that environmental changes induced an increase in biomass C accumulation for all age classes during the last 20 years (1982‐2001). Young and old stands had the highest changes in the biomass C accumulation during this period. During the last century mature stands (older than 80 years) turned from being almost carbon neutral to carbon sinks. In high elevations nitrogen deposition explained most of the increase of net ecosystem production (NEP) of forests. CO2 fertilization was the main factor increasing NEP of forests in the middle and low elevations. According to the model, at present, total biomass C accumulation in coniferous forests of Thuringia was estimated at 1.51tCha � 1 yr � 1 with an averaged annual NEP of 1.42tCha � 1 yr � 1 and total net biome production of 1.03tCha � 1 yr � 1 (accounting for harvest). The annual averaged biomass carbon balance (BCB: biomass accumulation rateharvest) was 1.12tCha � 1 yr � 1 (not including soil respiration), and was close to BCB from forest inventories (1.15tCha � 1 yr � 1 ). Indirect human impact resulted in 33% increase in modeled biomass carbon accumulation in coniferous forests in Thuringia during the last century. From the forest inventory data we estimated the legacy effect of the age-class distribution to account for 17% of the inventory-based sink. Isolating the environmental change effects showed that these effects can be large in a long-term, managed conifer forest.

Volatile organic compounds (VOCs) emitted by woody vegetation influence global climate forcing and the formation of tropospheric ozone. We use data from over 250 000 re-surveyed forest plots in the eastern US to estimate emission rates for the two most important biogenic VOCs (isoprene and monoterpenes) in the 1980s and 1990s, and then compare these estimates to give a decadal change in emission rate. Over much of the region, particularly the southeast, we estimate that there were large changes in biogenic VOC emissions: half of the grid cells (1°× 1°) had decadal changes in emission rate outside the range −2.3% to +16.8% for isoprene, and outside the range 0.2–17.1% for monoterpenes. For an average grid cell the estimated decadal change in heatwave biogenic VOC emissions (usually an increase) was three times greater than the decadal change in heatwave anthropogenic VOC emissions (usually a decrease, caused by legislation). Leaf-area increases in forests, caused by anthropogenic disturbance, were the most important process increasing biogenic VOC emissions. However, in the southeast, which had the largest estimated changes, there were substantial effects of ecological succession (which decreased monoterpene emissions and had location-specific effects on isoprene emissions), harvesting (which decreased monoterpene emissions and increased isoprene emissions) and plantation management (which increased isoprene emissions, and decreased monoterpene emissions in some states but increased monoterpene emissions in others). In any given region, changes in a very few tree species caused most of the changes in emissions: the rapid changes in the southeast were caused almost entirely by increases in sweetgum (Liquidambar styraciflua) and a few pine species. Therefore, in these regions, a more detailed ecological understanding of just a few species could greatly improve our understanding of the relationship between natural ecological processes, forest management, and biogenic VOC emissions.

The carbon balance of the world's terrestrial ecosystems is uncertain. Both top-down (atmospheric) and bottom-up (forest inventory and land-use change) approaches have been used to calculate the sign and magnitude of a net terrestrial flux. Different methods often include different processes, however, and comparisons can be misleading. Differences are not necessarily the result of uncertainties or errors, but often result from incomplete accounting inherent in some of the methods. Recent estimates are reviewed here. Overall, a northern mid-latitude carbon sink of approximately 2 Pg C yr−1 appears robust, although the mechanisms responsible are uncertain. Several lines of evidence point to environmentally enhanced rates of carbon accumulation. Other lines suggest that recovery from past disturbances is largely responsible for the sink. The tropics appear to be a small net source of carbon or nearly neutral, and the same uncertainties of mechanism exist. In addition, studies in the tropics do not permit an unequivocal choice between two alternatives: large emissions of carbon from deforestation offset by large sinks in undisturbed forests, or moderate emissions from land-use change with essentially no change in the carbon balance in undisturbed forests. Resolution of these uncertainties is most likely to result from spatially detailed historical reconstructions of land-use change and disturbance in selected northern mid-latitude regions where such data are available, and from systematic monitoring of changes in the area of tropical forests with satellite data of high spatial resolution collected over the last decades and into the future.

Altitudinal treelines are typically temperature limited such that increasing temperatures linked to global climate change are causing upslope shifts of treelines worldwide. While such elevational increases are readily predicted based on shifting isotherms, at the regional level the realized response is often much more complex, with topography and local environmental conditions playing an important modifying role. Here, we used repeated aerial photographs in combination with forest inventory data to investigate changes in treeline position in the Central Mountain Range of Taiwan over the last 60 years. A highly spatially variable upslope advance of treeline was identified in which topography is a major driver of both treeline form and advance. The changes in treeline position that we observed occurred alongside substantial increases in forest density, and lead to a large increase in overall forest area. These changes will have a significant impact on carbon stocking in the high altitude zone, while the concomitant decrease in alpine grassland area is likely to have negative implications for alpine species. The complex and spatially variable changes that we report highlight the necessity for considering local factors such as topography when attempting to predict species distributional responses to warming climate.

Climate change is anticipated to alter plant species distributions. Regional context, notably the spatial complexity of climatic gradients, may influence species migration potential. While high-elevation species may benefit from steep climate gradients in mountain regions, their persistence may be threatened by limited suitable habitat as land area decreases with elevation. To untangle these apparently contradictory predictions for mountainous regions, we evaluated the climatic suitability of four coniferous forest tree species of the western United States based on species distribution modeling (SDM) and examined changes in climatically suitable areas under predicted climate change. We used forest structural information relating to tree species dominance, productivity, and demography from an extensive forest inventory system to assess the strength of inferences made with a SDM approach. We found that tree species dominance, productivity, and recruitment were highest where climatic suitability (i.e., probability of species occurrence under certain climate conditions) was high, supporting the use of predicted climatic suitability in examining species risk to climate change. By predicting changes in climatic suitability over the next century, we found that climatic suitability will likely decline, both in areas currently occupied by each tree species and in nearby unoccupied areas to which species might migrate in the future. These trends were most dramatic for high elevation species. Climatic changes predicted over the next century will dramatically reduce climatically suitable areas for high-elevation tree species while a lower elevation species, Pinus ponderosa, will be well positioned to shift upslope across the region. Reductions in suitable area for high-elevation species imply that even unlimited migration would be insufficient to offset predicted habitat loss, underscoring the vulnerability of these high-elevation species to climatic changes.

Warmer conditions over the past two decades have contributed to rapid expansion of bark beetle outbreaks killing millions of trees over a large fraction of western United States (US) forests. These outbreaks reduce plant productivity by killing trees and transfer carbon from live to dead pools where carbon is slowly emitted to the atmosphere via heterotrophic respiration which subsequently feeds back to climate change. Recent studies have begun to examine the local impacts of bark beetle outbreaks in individual stands, but the full regional carbon consequences remain undocumented for the western US. In this study, we quantify the regional carbon impacts of the bark beetle outbreaks taking place in western US forests. The work relies on a combination of postdisturbance forest regrowth trajectories derived from forest inventory data and a process-based carbon cycle model tracking decomposition, as well as aerial detection survey (ADS) data documenting the regional extent and severity of recent outbreaks. We find that biomass killed by bark beetle attacks across beetle-affected areas in western US forests from 2000 to 2009 ranges from 5 to 15 Tg C yr(-1) and caused a reduction of net ecosystem productivity (NEP) of about 6.1-9.3 Tg C y(-1) by 2009. Uncertainties result largely from a lack of detailed surveys of the extent and severity of outbreaks, calling out a need for improved characterization across western US forests. The carbon flux legacy of 2000-2009 outbreaks will continue decades into the future (e.g., 2040-2060) as committed emissions from heterotrophic respiration of beetle-killed biomass are balanced by forest regrowth and accumulation.

Tree species are predicted to track future climate by shifting their geographic distributions, but climate-mediated migrations are not apparent in a recent continental-scale analysis. To better understand the mechanisms of a possible migration lag, we analyzed relative recruitment patterns by comparing juvenile and adult tree abundances in climate space. One would expect relative recruitment to be higher in cold and dry climates as a result of tree migration with juveniles located further poleward than adults. Alternatively, relative recruitment could be higher in warm and wet climates as a result of higher tree population turnover with increased temperature and precipitation. Using the USDA Forest Service’s Forest Inventory and Analysis data at regional scales, we jointly modeled juvenile and adult abundance distributions for 65 tree species in climate space of the eastern United States. We directly compared the optimal climate conditions for juveniles and adults, identified the climates where each species has high relative recruitment, and synthesized relative recruitment patterns across species. Results suggest that for 77% and 83% of the tree species, juveniles have higher optimal temperature and optimal precipitation, respectively, than adults. Across species, the relative recruitment pattern is dominated by relatively more abundant juveniles than adults in warm and wet climates. These different abundance-climate responses through life history are consistent with faster population turnover and inconsistent with the geographic trend of large-scale tree migration. Taken together, this juvenile–adult analysis suggests that tree species might respond to climate change by having faster turnover as dynamics accelerate with longer growing seasons and higher temperatures, before there is evidence of poleward migration at biogeographic scales.