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.

North American tree migration paced by climate in the West, lagging in the East.

Tree fecundity and recruitment have not yet been quantified at scales needed to anticipate biogeographic shifts in response to climate change. By separating their responses, this study shows coherence across species and communities, offering the strongest support to date that migration is in progress with regional limitations on rates. The southeastern continent emerges as a fecundity hotspot, but it is situated south of population centers where high seed production could contribute to poleward population spread. By contrast, seedling success is highest in the West and North, serving to partially offset limited seed production near poleward frontiers. The evidence of fecundity and recruitment control on tree migration can inform conservation planning for the expected long-term disequilibrium between climate and forest distribution.

Tracking 20 years of forest demographics in east Texas, USA, using national forest inventory data.

Forest resource reporting techniques primarily use the two most recent measurements for understanding forest change. Multiple remeasurements now exist within the US national forest inventory (NFI), providing an opportunity to examine long-term forest demographics. We leverage two decades of remeasurements to quantify live-dead wood demographics which can better inform estimates of resource changes in forest ecosystems. Our overall objective is to identify opportunities and gaps in tracking 20 years of forest demographics within the US NFI using east Texas as a pilot study region given its diversity of tree species, prevalence of managed conditions, frequency of disturbances, and relatively rapid change driven by a warm, humid climate. We examine growth and mortality rates, identify transitions to downed dead wood/litter and removal via harvest, and describe implications of these processes focusing on key species groups (i.e., loblolly pine, post oak, and water oak) and size classes (i.e., saplings, small and large trees). Growth and mortality rates fluctuated differently over time by species and stem sizes in response to large-scale disturbances, namely the 2011 drought in Texas. Tree-fall rates were highest in saplings and snag-fall rates trended higher in smaller trees. For removal rates, different stem sizes generally followed similar patterns within each species group. Forest demographics from the field-based US NFI are informative for identifying diffuse lagged mortality, species- and size-specific effects, and management effects. Moreover, researchers continually seek to employ ancillary data and develop new statistical methods to enhance understanding of forest resource changes from field-based inventories.

Carbon conundrums: Do United States' current carbon market baselines represent an undesirable ecological threshold?

Relative frequency distribution of observed annual mortality expressed in aboveground (AG) carbon (C) (Mg CO2 e ha-1 year-1 ) summarized across supersections by forest type [Hardwood (HW) vs. Softwood (SW)] and site class (Low vs. High) based on approximately 130,000 remeasured USDA Forest Service Forest Inventory and Analysis plots across the US. Top panel summarizes conditions in plots that do and do not meet the California Air Resources Board standards based on total basal area, whereas bottom panel summarizes conditions in plots falling inside and outside of optimum relative density levels. The latter represents a biophysically-informed approach accounting for changes in tree (and carbon) packing over forest development.

Estimating uncertainty in the volume and carbon storage of downed coarse woody debris.

Downed coarse woody debris, also known as coarse woody detritus or downed dead wood, is challenging to estimate for many reasons, including irregular shapes, multiple stages of decay, and the difficulty of identifying species. In addition, some properties are commonly not measured, such as wood density and carbon concentration. As a result, there have been few previous evaluations of uncertainty in estimates of downed coarse woody debris, which are necessary for analysis and interpretation of the data. To address this shortcoming, we quantified uncertainties in estimates of downed coarse woody debris volume and carbon storage using data collected from permanent forest inventory plots in the northeastern United States by the Forest Inventory and Analysis program of the USDA Forest Service. Quality assurance data collected from blind remeasurement audits were used to quantify error in diameter measurements, hollowness of logs, species identification, and decay class determination. Uncertainty estimates for density, collapse ratio, and carbon concentration were taken from the literature. Estimates of individual sources of uncertainty were combined using Monte Carlo methods. Volume estimates were more reliable than carbon storage, with an average 95% confidence interval of 15.9 m3 /ha across the 79 plots evaluated, which was less than the mean of 31.2 m3 /ha. Estimates of carbon storage (and mass) were more uncertain, due to poorly constrained estimates of the density of wood. For carbon storage, the average 95% confidence interval was 11.1 Mg C/ha, which was larger than the mean of 4.6 Mg C/ha. Accounting for the collapse of dead wood as it decomposes would improve estimates of both volume and carbon storage. On the other hand, our analyses suggest that consideration of the hollowness of downed coarse woody debris pieces could be eliminated in this region, with little effect. This study demonstrates how uncertainty analysis can be used to quantify confidence in estimates and to help identify where best to allocate resources to improve monitoring designs.

The impacts of increasing drought on forest dynamics, structure, and biodiversity in the United States.

We synthesize insights from current understanding of drought impacts at stand-to-biogeographic scales, including management options, and we identify challenges to be addressed with new research. Large stand-level shifts underway in western forests already are showing the importance of interactions involving drought, insects, and fire. Diebacks, changes in composition and structure, and shifting range limits are widely observed. In the eastern US, the effects of increasing drought are becoming better understood at the level of individual trees, but this knowledge cannot yet be confidently translated to predictions of changing structure and diversity of forest stands. While eastern forests have not experienced the types of changes seen in western forests in recent decades, they too are vulnerable to drought and could experience significant changes with increased severity, frequency, or duration in drought. Throughout the continental United States, the combination of projected large climate-induced shifts in suitable habitat from modeling studies and limited potential for the rapid migration of tree populations suggests that changing tree and forest biogeography could substantially lag habitat shifts already underway. Forest management practices can partially ameliorate drought impacts through reductions in stand density, selection of drought-tolerant species and genotypes, artificial regeneration, and the development of multistructured stands. However, silvicultural treatments also could exacerbate drought impacts unless implemented with careful attention to site and stand characteristics. Gaps in our understanding should motivate new research on the effects of interactions involving climate and other species at the stand scale and how interactions and multiple responses are represented in models. This assessment indicates that, without a stronger empirical basis for drought impacts at the stand scale, more complex models may provide limited guidance.

Prevalence and strength of density-dependent tree recruitment.

Density dependence could maintain diversity in forests, but studies continue to disagree on its role. Part of the disagreement results from the fact that different studies have evaluated different responses (survival, recruitment, or growth) of different stages (seeds, seedlings, or adults) to different inputs (density of seedlings, density or distance to adults). Most studies are conducted on a single site and thus are difficult to generalize. Using USDA Forest Service's Forest Inventory and Analysis data, we analyzed over a million seedling-to-sapling recruitment observations of 50 species from the eastern United States, controlling for the effects of climate. We focused on the per-seedling recruitment rate, because it is most likely to promote diversity and to be identified in observational or experimental data. To understand the prevalence of density dependence, we quantified the number of species with significant positive or negative effects. To understand the strength of density dependence, we determined the magnitude of effects among con- and heterospecifics, and how it changes with overall species abundance. We found that density dependence is pervasive among the 50 species, as the majority of them have significant effects and mostly negative. Density-dependence effects are stronger from conspecific than heterospecfic adult neighbors, consistent with the predictions of the Janzen-Connell hypothesis. Contrary to recent reports, density-dependence effects are more negative for common than rare species, suggesting disproportionately stronger population regulation in common species. We conclude that density dependence is pervasive, and it is strongest from conspecific neighbors of common species. Our analysis provides direct evidence that density dependence reaulates opulation dynamics of tree species in eastern U.S. forests.

Decomposing biodiversity data using the Latent Dirichlet Allocation model, a probabilistic multivariate statistical method.

We propose a novel multivariate method to analyse biodiversity data based on the Latent Dirichlet Allocation (LDA) model. LDA, a probabilistic model, reduces assemblages to sets of distinct component communities. It produces easily interpretable results, can represent abrupt and gradual changes in composition, accommodates missing data and allows for coherent estimates of uncertainty. We illustrate our method using tree data for the eastern United States and from a tropical successional chronosequence. The model is able to detect pervasive declines in the oak community in Minnesota and Indiana, potentially due to fire suppression, increased growing season precipitation and herbivory. The chronosequence analysis is able to delineate clear successional trends in species composition, while also revealing that site-specific factors significantly impact these successional trajectories. The proposed method provides a means to decompose and track the dynamics of species assemblages along temporal and spatial gradients, including effects of global change and forest disturbances.

Dual impacts of climate change: forest migration and turnover through life history.

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.

More than the sum of the parts: forest climate response from joint species distribution models.

The perceived threat of climate change is often evaluated from species distribution models that are fitted to many species independently and then added together. This approach ignores the fact that species are jointly distributed and limit one another. Species respond to the same underlying climatic variables, and the abundance of any one species can be constrained by competition; a large increase in one is inevitably linked to declines of others. Omitting this basic relationship explains why responses modeled independently do not agree with the species richness or basal areas of actual forests. We introduce a joint species distribution modeling approach (JSDM), which is unique in three ways, and apply it to forests of eastern North America. First, it accommodates the joint distribution of species. Second, this joint distribution includes both abundance and presence-absence data. We solve the common issue of large numbers of zeros in abundance data by accommodating zeros in both stem counts and basal area data, i.e., a new approach to zero inflation. Finally, inverse prediction can be applied to the joint distribution of predictions to integrate the role of climate risks across all species and identify geographic areas where communities will change most (in terms of changes in abundance) with climate change. Application to forests in the eastern United States shows that climate can have greatest impact in the Northeast, due to temperature, and in the Upper Midwest, due to temperature and precipitation. Thus, these are the regions experiencing the fastest warming and are also identified as most responsive at this scale.

Sapling recruitment as an indicator of carbon resiliency in forests of the northern USA.

Tree regeneration shapes forest carbon dynamics by determining long-term forest composition and structure, which suggests that threats to natural regeneration may diminish the capacity of forests to replace live tree carbon transferred to the atmosphere or other pools through tree mortality. Yet, the potential implications of tree regeneration patterns for future carbon dynamics have been sparsely studied. We used forest inventory plots to investigate whether the composition of existing tree regeneration is consistent with aboveground carbon stock loss, replacement, or gain for forests across the northeastern and midwestern USA, leveraging a recently developed method to predict the likelihood of sapling recruitment from seedling abundance tallied within six seedling height classes. A comparison of carbon stock predictions from tree and seedling composition suggested that 29% of plots were poised to lose carbon based on seedling composition, 55% were poised for replacement of carbon stocks (<5 Mg ha-1 difference) and 16% were poised to gain carbon. Forests predicted to lose carbon tended to be on steeper slopes, at lower latitudes, and in rolling upland environments. Although plots predicted to gain and lose carbon had similar stand ages, carbon loss plots had greater current carbon stocks. Synthesis and applications. Our results demonstrate the utility of considering tree regeneration through the lens of carbon replacement to develop effective management strategies to secure long-term carbon storage and resilience in the context of global change. Forests poised to lose C due to climate change and other stressors could be prioritized for regeneration strategies that enhance long-term carbon resilience and stewardship.

Mapping multi-dimensional variability in water stress strategies across temperate forests.

Increasing water stress is emerging as a global phenomenon, and is anticipated to have a marked impact on forest function. The role of tree functional strategies is pivotal in regulating forest fitness and their ability to cope with water stress. However, how the functional strategies found at the tree or species level scale up to characterise forest communities and their variation across regions is not yet well-established. By combining eight water-stress-related functional traits with forest inventory data from the USA and Europe, we investigated the community-level trait coordination and the biogeographic patterns of trait associations for woody plants, and analysed the relationships between the trait associations and climate factors. We find that the trait associations at the community level are consistent with those found at the species level. Traits associated with acquisitive-conservative strategies forms one dimension of variation, while leaf turgor loss point, associated with stomatal water regulation strategy, loads along a second dimension. Surprisingly, spatial patterns of community-level trait association are better explained by temperature than by aridity, suggesting a temperature-driven adaptation. These findings provide a basis to build predictions of forest response under water stress, with particular potential to improve simulations of tree mortality and forest biomass accumulation in a changing climate.

Trends over time in tree and seedling phylogenetic diversity indicate regional differences in forest biodiversity change.

Changing climate conditions may impact the short-term ability of forest tree species to regenerate in many locations. In the longer term, tree species may be unable to persist in some locations while they become established in new places. Over both time frames, forest tree biodiversity may change in unexpected ways. Using repeated inventory measurements five years apart from more than 7000 forested plots in the eastern United States, we tested three hypotheses: phylogenetic diversity is substantially different from species richness as a measure of biodiversity; forest communities have undergone recent changes in phylogenetic diversity that differ by size class, region, and seed dispersal strategy; and these patterns are consistent with expected early effects of climate change. Specifically, the magnitude of diversity change across broad regions should be greater among seedlings than in trees, should be associated with latitude and elevation, and should be greater among species with high dispersal capacity. Our analyses demonstrated that phylogenetic diversity and species richness are decoupled at small and medium scales and are imperfectly associated at large scales. This suggests that it is appropriate to apply indicators of biodiversity change based on phylogenetic diversity, which account for evolutionary relationships among species and may better represent community functional diversity. Our results also detected broadscale patterns of forest biodiversity change that are consistent with expected early effects of climate change. First, the statistically significant increase over time in seedling diversity in the South suggests that conditions there have become more favorable for the reproduction and dispersal of a wider variety of species, whereas the significant decrease in northern seedling diversity indicates that northern conditions have become less favorable. Second, we found weak correlations between seedling diversity change and latitude in both zones, with stronger relationships apparent in some ecoregions. Finally, we detected broadscale seedling diversity increases among species with longer-distance dispersal capacity, even in the northern zone, where overall seedling diversity declined. The statistical power and geographic extent of such analyses will increase as data become available over larger areas and as plot measurements are repeated at regular intervals over a longer period of time.

Predicting downed woody material carbon stocks in forests of the conterminous United States.

Downed woody material (DWM) is a unique part of the forest carbon cycle serving as a pool between living biomass and subsequent atmospheric emission or transference to other forest pools. Thus, DWM is an individually defined pool in national greenhouse gas inventories. The diversity of DWM carbon drivers (e.g., decay, tree mortality, or wildfire) and associated high spatial variability make this a difficult-to-predict component of forest ecosystems. Using the now fully established nationwide inventory of DWM across the United States (US), we developed models, which substantially improved predictions of stand-level DWM carbon density relative to the current national-reporting model ('previous' model, here). The previous model was developed from published DWM carbon densities prior to the NFI DWM inventory. Those predictions were tested using NFI DWM carbon densities resulting in a poor fit to the data (coefficient of determination, or R2 = 0.03). We present new random forest (RF) and stochastic gradient boosted (SGB) regression models to prediction DWM carbon density on all NFI plots and spatially on all forest land pixels. We evaluated various biotic and abiotic regression predictors, and the most important were standing dead trees, long-term annual precipitation, and long-term maximum summer temperature. A RF model scored best for expanding predictions to NFI plots (R2 = 0.31), while an SGB model was identified for DWM carbon predictions based on purely spatial data (i.e., NFI-plot-independent, with R2 = 0.23). The new RF model predicts conterminous US DWM carbon stocks to be 15% lower than the previous model and 2% higher than NFI data expanded according to inventory design-based inference. The new NFI data-driven models not only improve the predictions of DWM carbon density on all plots, they also provide flexibility in extending these predictions beyond the NFI to make spatially explicit and spatially continuous estimates of DWM carbon on all forest land in the US.

Co-limitation towards lower latitudes shapes global forest diversity gradients.

The latitudinal diversity gradient (LDG) is one of the most recognized global patterns of species richness exhibited across a wide range of taxa. Numerous hypotheses have been proposed in the past two centuries to explain LDG, but rigorous tests of the drivers of LDGs have been limited by a lack of high-quality global species richness data. Here we produce a high-resolution (0.025° × 0.025°) map of local tree species richness using a global forest inventory database with individual tree information and local biophysical characteristics from ~1.3 million sample plots. We then quantify drivers of local tree species richness patterns across latitudes. Generally, annual mean temperature was a dominant predictor of tree species richness, which is most consistent with the metabolic theory of biodiversity (MTB). However, MTB underestimated LDG in the tropics, where high species richness was also moderated by topographic, soil and anthropogenic factors operating at local scales. Given that local landscape variables operate synergistically with bioclimatic factors in shaping the global LDG pattern, we suggest that MTB be extended to account for co-limitation by subordinate drivers.

Similar tree species richness-productivity response but differing effects on carbon stocks and timber production in eastern US and continental Spain.

Unimodal response of tree species richness to increases in aboveground productivity is evident in grasslands but to a lesser extent in forests, where confounding factors (e.g., abiotic factors and management regimes) may alter the response and compromise the delivery of ecosystem services. We hypothesize that unimodal response of biomass accumulation through increased species richness leads to greater tree above ground carbon (AGC) stocks and thus climate regulation but not necessarily higher timber volume production for human consumption across portions of North American and European forests. We first evaluated the biodiversity-productivity pattern and assessed if the addition of potential confounding variables altered the response. Afterwards, we integrated direct and indirect effects of species richness and confounding factors in the modelling of aboveground carbon stock and timber volume. We confirm an increase in carbon stocks concomitant with an increase in tree species richness up to an optimum biomass value in both regions. Tree species richness had a marginal effect on both aboveground carbon stocks and timber volume with a trade-off in the eastern US. Biomass accumulation is lower in tree plantations than in natural forests, although volume increased with species richness. Naturally-regenerated forests needed as much as double the number of tree species than plantations to reach the same carbon stocks. Distinct ecosystem services (AGC and timber volume) showed unique pathways of achieving their maximum provisioning. As increasing forest resilience to global change requires a fundamental understanding of how tree species combine with changing climatic conditions to drive the provisioning of various ecosystem services, further examination of this study's findings across additional biogeographical regions may lead the way to unraveling such dynamics and empowering adaptive management.

Release of coarse woody detritus-related carbon: a synthesis across forest biomes.

BACKGROUND: Recent increases in forest tree mortality should increase the abundance coarse woody detritus (CWD) and ultimately lead to increased atmospheric carbon dioxide. However, the time course of carbon release from CWD is not well understood. We compiled CWD decomposition rate-constants (i.e., k) to examine how tree species, piece diameter, position (i.e., standing versus downed), canopy openness, and macroclimate influenced k. To illustrate their implications we modeled the effect of species and position on estimates of decomposition-related carbon flux. We examined a subset of currently used models to determine if their structure accounted for these factors. RESULTS: Globally k of downed CWD varied at least 244-fold with interspecies variation at individual sites up to 76-fold. While k generally decreased with increasing piece diameter, under open canopies the opposite occurred. Standing CWD sometimes exhibited little decomposition, but sometimes had k values up to 3 times faster than downed CWD. There was a clear response of k to mean annual temperature of ≈ 2.6 times per 10 â„ƒ; however, there was considerable variation for a given mean annual temperature related to species, diameter, and position. A key feature of carbon release from CWD after disturbance was the "evolution" of the ecosystem-level k value as positions and species mixtures of the remaining CWD changed. Variations in decomposition caused by disturbance (e.g., changes in species, positions, sizes, and microclimate) had the potential to cause net carbon fluxes to the atmosphere to be highly nonlinear. While several models currently being used for carbon accounting and assessing land-use/climate change would potentially capture some of these post disturbance changes in fluxes and carbon balances, many would not. CONCLUSIONS: While much has been learned in the last 5 decades about CWD decomposition, to fully understand the time course of carbon release from increased mortality and other aspects of global change a new phase of global CWD research that is more systematic, experimental, and replicated needs to be initiated. If our findings are to be fully applied in modeling, an approach acknowledging how the rate of carbon release evolves over time should be implemented.

Estimating dead wood during national forest inventories: a review of inventory methodologies and suggestions for harmonization.

Efforts to assess forest ecosystem carbon stocks, biodiversity, and fire hazards have spurred the need for comprehensive assessments of forest ecosystem dead wood (DW) components around the world. Currently, information regarding the prevalence, status, and methods of DW inventories occurring in the world's forested landscapes is scattered. The goal of this study is to describe the status, DW components measured, sample methods employed, and DW component thresholds used by national forest inventories that currently inventory DW around the world. Study results indicate that most countries do not inventory forest DW. Globally, we estimate that about 13% of countries inventory DW using a diversity of sample methods and DW component definitions. A common feature among DW inventories was that most countries had only just begun DW inventories and employ very low sample intensities. There are major hurdles to harmonizing national forest inventories of DW: differences in population definitions, lack of clarity on sample protocols/estimation procedures, and sparse availability of inventory data/reports. Increasing database/estimation flexibility, developing common dimensional thresholds of DW components, publishing inventory procedures/protocols, releasing inventory data/reports to international peer review, and increasing communication (e.g., workshops) among countries inventorying DW are suggestions forwarded by this study to increase DW inventory harmonization.

The downed and dead wood inventory of forests in the United States.

The quantity and condition of downed dead wood (DDW) is emerging as a major factor governing forest ecosystem processes such as carbon cycling, fire behavior, and tree regeneration. Despite this, systematic inventories of DDW are sparse if not absent across major forest biomes. The Forest Inventory and Analysis program of the United States (US) Forest Service has conducted an annual DDW inventory on all coterminous US forest land since 2002 (~1 plot per 38,850 ha), with a sample intensification occurring since 2012 (~1 plot per 19,425 ha). The data are organized according to DDW components and by sampling information which can all be linked to a multitude of auxiliary information in the national database. As the sampling of DDW is conducted using field efficient line-intersect approaches, several assumptions are adopted during population estimation that serve to identify critical knowledge gaps. The plot- and population-level DDW datasets and estimates provide the first insights into an understudied but critical ecosystem component of temperate forests of North America with global application.