The carbon balance of tropical, temperate and boreal forests

Forest biomes are major reserves for terrestrial carbon, and major components of global primary productivity. The carbon balance of forests is determined by a number of component processes of carbon acquisition and carbon loss, and a small shift in the magnitude of these processes would have a large impact on the global carbon cycle. In this paper, we discuss the climatic influences on the carbon dynamics of boreal, temperate and tropical forests by presenting a new synthesis of micrometeorological, ecophysiological and forestry data, concentrating on three case-study sites. Historical changes in the carbon balance of each biome are also reviewed, and the evidence for a carbon sink in each forest biome and its likely behaviour under future global change are discussed. We conclude that there have been significant advances in determining the carbon balance of forests, but there are still critical uncertainties remaining, particularly in the behaviour of soil carbon stocks.     

Differential declines in Alaskan boreal forest vitality related to climate and competition

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.     

Large Impacts of Climatic Warming on Growth of Boreal Forests since 1960

Boreal forests are sensitive to climatic warming, because low temperatures hold back ecosystem processes, such as the mobilization of nitrogen in soils. A greening of the boreal landscape has been observed using remote sensing, and the seasonal amplitude of CO₂ in the northern hemisphere has increased, indicating warming effects on ecosystem productivity. However, field observations on responses of ecosystem productivity have been lacking on a large sub-biome scale. Here we report a significant increase in the annual growth of boreal forests in Finland in response to climatic warming, especially since 1990. This finding is obtained by linking meteorological records and forest inventory data on an area between 60° and 70° northern latitude. An additional increase in growth has occurred in response to changes in other drivers, such as forest management, nitrogen deposition and/or CO₂ concentration. A similar warming impact can be expected in the entire boreal zone, where warming takes place. Given the large size of the boreal biome – more than ten million km²– important climate feedbacks are at stake, such as the future carbon balance, transpiration and albedo.     

Plant Community Composition as a Predictor of Regional Soil Carbon Storage in Alaskan Boreal Black Spruce Ecosystems

The boreal forest is the largest terrestrial biome in North America and holds a large portion of the world’s reactive soil carbon. Therefore, understanding soil carbon accumulation on a landscape or regional scale across the boreal forest is useful for predicting future soil carbon storage. Here, we examined the relationship between floristic composition and ecosystem parameters, such as soil carbon pools, the carbon-to-nitrogen (C/N) ratio of live black spruce needles, and normalized basal area increment (NBAI) of trees in black spruce communities, the most widespread forest type in the boreal forest of Alaska. Variability in ecosystem properties among black spruce stands was as large as that which had previously been documented among all forest types in the central interior of Alaska; we found an eightfold range in NBAI and fivefold range in mineral soil carbon and nitrogen pools. Acidic black spruce communities had significantly more carbon in the organic soil horizon than did nonacidic black spruce communities, but did not differ in any other measured ecosystem parameter. We explained 48% of the variation in total soil carbon with a combination of plant community indices and abiotic and biotic factors. Plant community composition was at least as effective as any single environmental factor or stand characteristic in predicting soil C pools in Alaskan black spruce ecosystems. We conclude that among the community properties analyzed, the presence of key groups of species, overall species composition, and diversity of certain functional types, especially Sphagnum moss species, are important predictors of soil carbon sequestration in the black spruce forest type.     

The effect of biodiversity on tree productivity: from temperate to boreal forests

Aim An important issue regarding biodiversity concerns its influence on ecosystem functioning. Experimental work has led to the proposal of mechanisms such as niche complementarity. However, few attempts have been made to confirm these in natural systems, especially in forests. Furthermore, one of the most interesting unresolved questions is whether the effects of complementarity on ecosystem functioning (EF) decrease in favour of competitive exclusions over an increasing productivity gradient. Using records from permanent forest plots, we asked the following questions. (1) Is tree productivity positively related to diversity? (2) Does the effect of diversity increase in less productive forests? (3) What metric of diversity (e.g. functional or phylogenetic diversity) better relates to tree productivity? Location Temperate, mixed and boreal forests of eastern Canada. Methods Over 12,000 permanent forest plots, from temperate to boreal forests, were used to test our hypotheses in two steps. (1) Stepwise regressions were used to identify the best explanatory variables for tree productivity. (2) The selected climatic and environmental variables, as well as density and biodiversity indices, were included in a structural equation model where links (paths) between covarying variables are made explicit, making structural equation modelling the best tool to explore such complicated causal networks. Results This is the first large‐scale demonstration of a strong, positive and significant effect of biodiversity on tree productivity with control for climatic and environmental conditions. Important differences were noted between the two forest biomes investigated. Main conclusions We show for the first time that complementarity may be less important in temperate forests growing in a more stable and productive environment where competitive exclusion is the most probable outcome of species interactions, whereas in the more stressful environment of boreal forests, beneficial interactions between species may be more important. The present work is also a framework for the analysis of large datasets in biodiversity–ecosystem functioning (B‐EF) research.     

Stable carbon isotope analysis reveals widespread drought stress in boreal black spruce forests

Unprecedented rates of climate warming over the past century have resulted in increased forest stress and mortality worldwide. Decreased tree growth in association with increasing temperatures is generally accepted as a signal of temperature‐induced drought stress. However, variations in tree growth alone do not reveal the physiological mechanisms behind recent changes in tree growth. Examining stable carbon isotope composition of tree rings in addition to tree growth can provide a secondary line of evidence for physiological drought stress. In this study, we examined patterns of black spruce growth and carbon isotopic composition in tree rings in response to climate warming and drying in the boreal forest of interior Alaska. We examined trees at three nested scales: landscape, toposequence, and a subsample of trees within the toposequence. At each scale, we studied the potential effects of differences in microclimate and moisture availability by sampling on northern and southern aspects. We found that black spruce radial growth responded negatively to monthly metrics of temperature at all examined scales, and we examined ?¹³C responses on a subsample of trees as representative of the wider region. The negative ?¹³C responses to temperature reveal that black spruce trees are experiencing moisture stress on both northern and southern aspects. Contrary to our expectations, ?¹³C from trees on the northern aspect exhibited the strongest drought signal. Our results highlight the prominence of drought stress in the boreal forest of interior Alaska. We conclude that if temperatures continue to warm, we can expect drought‐induced productivity declines across large regions of the boreal forest, even for trees located in cool and moist landscape positions.     

Outbreaks by canopy-feeding geometrid moth cause state-dependent shifts in understorey plant communities

The increased spread of insect outbreaks is among the most severe impacts of climate warming predicted for northern boreal forest ecosystems. Compound disturbances by insect herbivores can cause sharp transitions between vegetation states with implications for ecosystem productivity and climate feedbacks. By analysing vegetation plots prior to and immediately after a severe and widespread outbreak by geometrid moths in the birch forest-tundra ecotone, we document a shift in forest understorey community composition in response to the moth outbreak. Prior to the moth outbreak, the plots divided into two oligotrophic and one eutrophic plant community. The moth outbreak caused a vegetation state shift in the two oligotrophic communities, but only minor changes in the eutrophic community. In the spatially most widespread communities, oligotrophic dwarf shrub birch forest, dominance by the allelopathic dwarf shrub Empetrum nigrum ssp. hermaphroditum, was effectively broken and replaced by a community dominated by the graminoid Avenella flexuosa, in a manner qualitatively similar to the effect of wild fires in E. nigrum communities in coniferous boreal forest further south. As dominance by E. nigrum is associated with retrogressive succession the observed vegetation state shift has widespread implications for ecosystem productivity on a regional scale. Our findings reveal that the impact of moth outbreaks on the northern boreal birch forest system is highly initial-state dependent, and that the widespread oligotrophic communities have a low resistance to such disturbances. This provides a case for the notion that climate impacts on arctic and northern boreal vegetation may take place most abruptly when conveyed by changed dynamics of irruptive herbivores.     

Substantial reorganization of China's tropical and subtropical forests: based on the permanent plots

There is evidence that climate change induced tree mortalities in boreal and temperate forests and increased forest turnover rates (both mortality and recruitment rates) in Amazon forests. However, no study has examined China's tropical and subtropical evergreen broadleaved forests (TEBF) that cover >26% of China's terrestrial land. The sustainability of this biome is vital to the maintenance of local ecosystem services (e.g., carbon sequestration, biodiversity conservation, climatic regulation), many of which may influence patterns of atmospheric circulation and composition at regional to global scales. Here, we analyze time‐series data collected from thirteen permanent plots within China's unmanaged TEBF to study whether and how this biome has changed over recent decades. We find that the numbers of individuals and species for shrub and small tree have increased since 1978, whereas the numbers of individuals and species for tree have decreased over this same time period. The shift in species composition is accompanied by a decrease in the mean diameter at breast height (DBH) for all individuals combined. China's TEBF may thereby be transitioning from cohorts of fewer and larger individuals to ones of more and smaller individuals, which shows a unique change pattern differing from the documented. Regional‐scale drying is likely responsible for the biome's reorganization. This biome‐wide reconstitution would deeply impact the regimes of carbon sequestration and biodiversity conservation and have implications for the sustainability of economic development in the area.     

Global patterns in the vulnerability of ecosystems to vegetation shifts due to climate change

Aim Climate change threatens to shift vegetation, disrupting ecosystems and damaging human well‐being. Field observations in boreal, temperate and tropical ecosystems have detected biome changes in the 20th century, yet a lack of spatial data on vulnerability hinders organizations that manage natural resources from identifying priority areas for adaptation measures. We explore potential methods to identify areas vulnerable to vegetation shifts and potential refugia. Location Global vegetation biomes. Methods We examined nine combinations of three sets of potential indicators of the vulnerability of ecosystems to biome change: (1) observed changes of 20th‐century climate, (2) projected 21st‐century vegetation changes using the MC1 dynamic global vegetation model under three Intergovernmental Panel on Climate Change (IPCC) emissions scenarios, and (3) overlap of results from (1) and (2). Estimating probability density functions for climate observations and confidence levels for vegetation projections, we classified areas into vulnerability classes based on IPCC treatment of uncertainty. Results One‐tenth to one‐half of global land may be highly (confidence 0.80–0.95) to very highly (confidence ≥ 0.95) vulnerable. Temperate mixed forest, boreal conifer and tundra and alpine biomes show the highest vulnerability, often due to potential changes in wildfire. Tropical evergreen broadleaf forest and desert biomes show the lowest vulnerability. Main conclusions Spatial analyses of observed climate and projected vegetation indicate widespread vulnerability of ecosystems to biome change. A mismatch between vulnerability patterns and the geographic priorities of natural resource organizations suggests the need to adapt management plans. Approximately a billion people live in the areas classified as vulnerable.     

Tree line identification from pollen data: beyond the limit?

Aim The boreal tree line is a prominent biogeographic feature, the position of which reflects climatic conditions. Pollen is the key sensor used to reconstruct past tree line patterns. Our aims in this study were to investigate pollen–vegetation relationships at the boreal tree line and to assess the success of a modified version of the biomization method that incorporates pollen productivity and dispersal in distinguishing the tree line. Location Northern Canada (307 sites) and Alaska (316 sites). Methods The REVEALS method for estimating regional vegetation composition from pollen data was simplified to provide correction factors to account for differential production and dispersal of pollen among taxa. The REVEALS‐based correction factors were used to adapt the biomization method and applied as a set of experiments to pollen data from lake sediments and moss polsters from the boreal tree line. Proportions of forest and tundra predicted from modern pollen samples along two longitudinal transects were compared with those derived from a vegetation map by: (1) a tally of ‘correct’ versus ‘incorrect’ assignments using vegetation in the relevant map pixels, and (2) a comparison of the shape and position of north–south forest‐cover curves generated from all transect pixels and from pollen data. Possible causes of bias in the misclassifications were assessed. Results Correcting for pollen productivity alone gave fewest misclassifications and the closest estimate of the modern mapped tree line position (Canada, + 300 km; Alaska, + 10 km). In Canada success rates were c. 40–70% and all experiments over‐predicted forest cover. Most corrections improved results over uncorrected biomization; using only lakes improved success rates to c. 80%. In Alaska success rates were 70–80% and classification errors were more evenly distributed; there was little improvement over uncorrected biomization. Main conclusions Corrected biomization should improve broad‐scale reconstructions of spatial patterns in forest/non‐forest vegetation mosaics and across climate‐sensitive ecotones. The Canadian example shows this is particularly the case in regions affected by taxa with extremely high pollen productivity (such as Pinus). Improved representation of actual vegetation distribution is most likely if pollen data from lake sediments are used because the REVEALS algorithm is based on the pollen dynamics of lake‐based systems.     

Relationships Among pH,Minerals, and Carbon in Soils from Tundra to Boreal Forest Across Alaska

Tundra and boreal forests in northern high latitudes contain significant amounts of carbon (C) in the soil, indicating the importance of clarifying controls on soil C dynamics in the region and their feedback effects on climate systems. In northern Alaska, variations in soil C processes are closely associated with variations in soil acidity within ecosystems; however, the reason for this association remains unclear. In this study, we demonstrate that it results from weathering and subsequent changes in soil geochemical characteristics, including minerals and adsorptive organic C. We sampled soils from 12 sites in Alaska along a 600-km transect from the Arctic Ocean to interior Alaska, spanning the biomes of tundra, tundra–boreal forest ecotone, and boreal forest. Mineral soil analyses revealed that soils with low pH have fewer base cations, more aluminum/iron minerals, and lower base saturation, indicating that weathering is a major function of these geochemical characteristics in the broad area over northern Alaska. Adsorbed organic C in soil presented strong correlations with Al and Fe minerals, soil pH, and soil total C and represented approximately 30–55% of total soil C, suggesting that soil C accumulation in the Alaskan ecosystems is strongly controlled by weathering-related changes in geochemical characteristics. An adsorption test supported these observations and illustrated a greater capacity for acidic soil to adsorb organic C. These findings demonstrate that variations in weathering-associated characteristics have a strong influence on the regional variation in C dynamics and biogeochemical consequences in the Alaskan ecosystems.     

A biome classification of China based on plant functional types and the BIOME3 model

A biome classification for China was established based on plant functional types (PFTs) using the BIOME3 model to include 16 biomes. In the eastern part of China, the PFTs of trees determine mostly the physiognomy of landscape. Biomes range from boreal deciduous coniferous forest/woodland, boreal mixed forest/woodland, temperate mixed forest, temperate broad-leaved deciduous forest, warm-temperate broad-leaved evergreen/mixed forest, warm-temperate/cool-temperate evergreen coniferous forest, xeric woodland/scrub, to tropical seasonal and rain forest, and tropical deciduous forest from north to south. In the northern and western part of China, grass is the dominant PFT. From northeast to west and southwest the biomes range from moist savannas, tall grassland, short grassland, dry savannas, arid shrubland/steppe, desert, to alpine tundra/ice/polar desert. Comparisons between the classification introduced here and the four classifications which were established over the past two decades, i.e. the vegetation classification, the vegetation division, the physical ecoregion, and the initial biome classification have showed that the different aims of biome classifications have resulted in different biome schemes each with its own unique characteristics and disadvantages for global change study. The new biome classification relies not only on climatic variables, but also on soil factor, vegetation functional variables, ecophysiological parameters and competition among the PFTs. It is a comprehensive classification that using multivariables better expresses the vegetation distribution and can be compared with world biome classifications. It can be easily used in the response study of Chinese biomes to global change, regionally and globally.     

Modeling analysis of primary controls on net ecosystem productivity of seven boreal and temperate coniferous forests across a continental transect

Process‐based models are effective tools to synthesize and/or extrapolate measured carbon (C) exchanges from individual sites to large scales. In this study, we used a C‐ and nitrogen (N)‐cycle coupled ecosystem model named CN‐CLASS (Carbon Nitrogen‐Canadian Land Surface Scheme) to study the role of primary climatic controls and site‐specific C stocks on the net ecosystem productivity (NEP) of seven intermediate‐aged to mature coniferous forest sites across an east–west continental transect in Canada. The model was parameterized using a common set of parameters, except for two used in empirical canopy conductance–assimilation, and leaf area–sapwood relationships, and then validated using observed eddy covariance flux data. Leaf Rubisco‐N dynamics that are associated with soil–plant N cycling, and depend on canopy temperature, enabled the model to simulate site‐specific gross ecosystem productivity (GEP) reasonably well for all seven sites. Overall GEP simulations had relatively smaller differences compared with observations vs. ecosystem respiration (RE), which was the sum of many plant and soil components with larger variability and/or uncertainty associated with them. Both observed and simulated data showed that, on an annual basis, boreal forest sites were either carbon‐neutral or a weak C sink, ranging from 30 to 180 g C m^?2 yr^?1; while temperate forests were either a medium or strong C sink, ranging from 150 to 500 g C m^?2 yr^?1, depending on forest age and climatic regime. Model sensitivity tests illustrated that air temperature, among climate variables, and aboveground biomass, among major C stocks, were dominant factors impacting annual NEP. Vegetation biomass effects on annual GEP, RE and NEP showed similar patterns of variability at four boreal and three temperate forests. Air temperature showed different impacts on GEP and RE, and the response varied considerably from site to site. Higher solar radiation enhanced GEP, while precipitation differences had a minor effect. Magnitude of forest litter content and soil organic matter (SOM) affected RE. SOM also affected GEP, but only at low levels of SOM, because of low N mineralization that limited soil nutrient (N) availability. The results of this study will help to evaluate the impact of future climatic changes and/or forest C stock variations on C uptake and loss in forest ecosystems growing in diverse environments.     

Range Expansion of Moose in Arctic Alaska Linked to Warming and Increased Shrub Habitat

Twentieth century warming has increased vegetation productivity and shrub cover across northern tundra and treeline regions, but effects on terrestrial wildlife have not been demonstrated on a comparable scale. During this period, Alaskan moose (Alces alces gigas) extended their range from the boreal forest into tundra riparian shrub habitat; similar extensions have been observed in Canada (A. a. andersoni) and Eurasia (A. a. alces). Northern moose distribution is thought to be limited by forage availability above the snow in late winter, so the observed increase in shrub habitat could be causing the northward moose establishment, but a previous hypothesis suggested that hunting cessation triggered moose establishment. Here, we use recent changes in shrub cover and empirical relationships between shrub height and growing season temperature to estimate available moose habitat in Arctic Alaska c. 1860. We estimate that riparian shrubs were approximately 1.1 m tall c. 1860, greatly reducing the available forage above the snowpack, compared to 2 m tall in 2009. We believe that increases in riparian shrub habitat after 1860 allowed moose to colonize tundra regions of Alaska hundreds of kilometers north and west of previous distribution limits. The northern shift in the distribution of moose, like that of snowshoe hares, has been in response to the spread of their shrub habitat in the Arctic, but at the same time, herbivores have likely had pronounced impacts on the structure and function of these shrub communities. These northward range shifts are a bellwether for other boreal species and their associated predators.     

Testing for changes in biomass dynamics in large‐scale forest datasets

Tropical forest responses to climate and atmospheric change are critical to the future of the global carbon budget. Recent studies have reported increases in estimated above‐ground biomass (EAGB) stocks, productivity, and mortality in old‐growth tropical forests. These increases could reflect a shift in forest functioning due to global change and/or long‐lasting recovery from past disturbance. We introduce a novel approach to disentangle the relative contributions of these mechanisms by decomposing changes in whole‐plot biomass fluxes into contributions from changes in the distribution of gap‐successional stages and changes in fluxes for a given stage. Using 30 years of forest dynamic data at Barro Colorado Island, Panama, we investigated temporal variation in EAGB fluxes as a function of initial EAGB (EAGB_i) in 10 × 10 m quadrats. Productivity and mortality fluxes both increased strongly with initial quadrat EAGB. The distribution of EAGB (and thus EAGB_i) across quadrats hardly varied over 30 years (and seven censuses). EAGB fluxes as a function of EAGB_i varied largely and significantly among census intervals, with notably higher productivity in 1985–1990 associated with recovery from the 1982–1983 El Niño event. Variation in whole‐plot fluxes among census intervals was explained overwhelmingly by variation in fluxes as a function of EAGB_i, with essentially no contribution from changes in EAGB_i distributions. The high observed temporal variation in productivity and mortality suggests that this forest is very sensitive to climate variability. There was no consistent long‐term trend in productivity, mortality, or biomass in this forest over 30 years, although the temporal variability in productivity and mortality was so strong that it could well mask a substantial trend. Accurate prediction of future tropical forest carbon budgets will require accounting for disturbance‐recovery dynamics and understanding temporal variability in productivity and mortality.     

Boreal Forest Floor Greenhouse Gas Emissions Across a Pleurozium schreberi-Dominated,Wildfire-Disturbed Chronosequence

Ecosystems - The boreal forest is a globally critical biome for carbon cycling. Its forests are shaped by wildfire events that affect ecosystem properties and climate feedbacks including greenhouse...     

Tundra and boreal forest of interior Alaska during terminal MIS 6 and MIS 5e

Two sites within the boreal forest of interior Alaska shed light on the climate and vegetation of terminal marine isotope stage (MIS) 6 (ca. 140–130 kyr ago) and MIS 5e (125–116 kyr ago). The Birch Creek and Koyukuk localities are river-cut exposures with sediments dating from the penultimate glaciation (at least) to the present. Plant macrofossils, pollen, and beetles were analyzed at these sites. Terminal MIS 6-aged samples indicate a cooler than modern climate and the presence of shrub tundra. During MIS 5e, boreal forest grew at the sites and temperatures were similar to modern times. However, the forest may also have been more mesic than today, as indicated by relatively abundant ferns. Winters may have been warmer than today, as suggested by beetle-based climatic reconstructions as well as the presence of two extralimital taxa that today live in regions where winter temperatures are up to 15 °C warmer than at the site localities.     

Impacts of climate change on natural forest productivity – evidence since the middle of the 20th century

Changes to forest production drivers (light, water, temperature, and site nutrient) over the last 55 years have been documented in peer‐reviewed literature. The main objective of this paper is to review documented evidence of the impacts of climate change trends on forest productivity since the middle of the 20th century. We first present a concise overview of the climate controls of forest production, provide evidence of how the main controls have changed in the last 55 years, followed by a core section outlining our findings of observed and documented impacts on forest productivity and a brief discussion of the complications of interpreting trends in net primary production (NPP). At finer spatial scales, a trend is difficult to decipher, but globally, based on both satellite and ground‐based data, climatic changes seemed to have a generally positive impact on forest productivity when water was not limiting. Of the 49 papers reporting forest production levels we reviewed, 37 showed a positive growth trend, five a negative trend, three reported both a positive and a negative trend for different time periods, one reported a positive and no trend for different geographic areas, and two reported no trend. Forests occupy ≈52% of the Earth's land surface and tend to occupy more temperature and radiation‐limited environments. Less than 7% of forests are in strongly water‐limited systems. The combined and interacting effects of temperature, radiation, and precipitation changes with the positive effect of CO₂, the negative effects of O₃ and other pollutants, and the presently positive effects of N will not be elucidated with experimental manipulation of one or a few factors at a time. Assessments of the greening of the biosphere depend on both accurate measurements of rates (net ecosystem exchange, NPP), how much is stored at the ecosystem level (net ecosystem production) and quantification of disturbances rates on final net biome production.     

Susceptibility of burned black spruce (Picea mariana) forests to non-native plant invasions in interior Alaska

As climate rapidly warms at high-latitudes, the boreal forest faces the simultaneous threats of increasing invasive plant abundances and increasing area burned by wildfire. Highly flammable and widespread black spruce (Picea mariana) forest represents a boreal habitat that may be increasingly susceptible to non-native plant invasion. This study assess the role of burn severity, site moisture and time elapsed since burning in determining the invisibility of black spruce forests. We conducted field surveys for presence of non-native plants at 99 burned black spruce forest sites burned in 2004 in three regions of interior Alaska that spanned a gradient of burn severities and site moisture levels, and a chronosequence of sites in a single region that had burned in 1987, 1994, and 1999. We also conducted a greenhouse experiment where we grew invasive plants in vegetation and soil cores taken from a subset of these sites. In both our field survey and the greenhouse experiment, regional differences in soils and vegetation between burn complexes outweighed local burn severity or site moisture in determining the invasibility of burned black spruce sites. In the greenhouse experiments using cores from the 2004 burns, we found that the invasive focal species grew better in cores with soil and vegetation properties characteristic of low severity burns. Invasive plant growth in the greenhouse was greater in cores from the chronosequence burns with higher soil water holding capacity or lower native vascular biomass. We concluded that there are differences in susceptibility to non-native plant invasions between different regions of boreal Alaska based on native species regeneration. Re-establishment of native ground cover vegetation, including rapidly colonizing bryophytes, appear to offer burned areas a level of resistance to invasive plant establishment.     

Carbon cycling and storage in world forests: biome patterns related to forest age

Forest age, which is affected by stand‐replacing ecosystem disturbances (such as forest fires, harvesting, or insects), plays a distinguishing role in determining the distribution of carbon (C) pools and fluxes in different forested ecosystems. In this synthesis, net primary productivity (NPP), net ecosystem productivity (NEP), and five pools of C (living biomass, coarse woody debris, organic soil horizons, soil, and total ecosystem) are summarized by age class for tropical, temperate, and boreal forest biomes. Estimates of variability in NPP, NEP, and C pools are provided for each biome‐age class combination and the sources of variability are discussed. Aggregated biome‐level estimates of NPP and NEP were higher in intermediate‐aged forests (e.g., 30–120 years), while older forests (e.g., >120 years) were generally less productive. The mean NEP in the youngest forests (0–10 years) was negative (source to the atmosphere) in both boreal and temperate biomes (?0.1 and –1.9 Mg C ha^?1 yr^?1, respectively). Forest age is a highly significant source of variability in NEP at the biome scale; for example, mean temperate forest NEP was ?1.9, 4.5, 2.4, 1.9 and 1.7 Mg C ha^?1 yr^?1 across five age classes (0–10, 11–30, 31–70, 71–120, 121–200 years, respectively). In general, median NPP and NEP are strongly correlated (R²=0.83) across all biomes and age classes, with the exception of the youngest temperate forests. Using the information gained from calculating the summary statistics for NPP and NEP, we calculated heterotrophic soil respiration (R_h) for each age class in each biome. The mean R_h was high in the youngest temperate age class (9.7 Mg C ha^?1 yr^?1) and declined with age, implying that forest ecosystem respiration peaks when forests are young, not old. With notable exceptions, carbon pool sizes increased with age in all biomes, including soil C. Age trends in C cycling and storage are very apparent in all three biomes and it is clear that a better understanding of how forest age and disturbance history interact will greatly improve our fundamental knowledge of the terrestrial C cycle.