Response of feather moss associated N2 fixation and litter decomposition to variations in simulated rainfall intensity and frequency

Feather mosses in boreal forests form a dense ground‐cover that is an important driver of both nutrient and carbon cycling. While moss growth is highly sensitive to moisture availability, little is known about how moss effects on nutrient and carbon cycling are affected by the dynamics of moisture input to the ecosystem. We experimentally investigated how rainfall regimes affected ecosystem processes driven by the dominant boreal feather moss Pleurozium schreberi by manipulating total moisture amount, frequency of moisture addition and moss presence/absence. Moisture treatments represented the range of rainfall conditions that occur in Swedish boreal forests as well as shifts in rainfall expected through climate change. We found that nitrogen (N) fixation by cyanobacteria in feather mosses (the main biological N input to boreal forests) was strongly influenced by both moisture amount and frequency, and their interaction; increased frequency had greater effects when amounts were higher. Within a given moisture amount, N fixation varied up to seven‐fold depending on how that amount was distributed temporally. We also found that mosses promoted vascular litter decomposition rates, concentrations of litter nutrients, and active soil microbial biomass, and reduced N release into soil solution. These effects were usually strongest under low moisture amount and/or frequency, and revealed a buffering effect of mosses on the decomposer subsystem under moisture limitation. These results highlight that both the amount and temporal distribution of rainfall, determine the effect of feather mosses on ecosystem N input and the decomposer subsystem. They also emphasize the role of feather mosses in mediating moisture effects on decomposer processes. Finally, our results suggest that projected shifts in precipitation in the Swedish boreal forest through climate change will result in increased moss growth and N₂ fixation but a reduced dependency of the decomposer subsystem on feather moss cover for moisture retention.     

The interactive effects of temperature and light on biological nitrogen fixation in boreal forests

Plant productivity is predicted to increase in northern latitudes as a result of climate warming; however, this may depend on whether biological nitrogen (N)-fixation also increases. We evaluated how the variation in temperature and light affects N-fixation by two boreal feather mosses, Pleurozium schreberi and Hylocomium splendens, which are the primary source of N-fixation in most boreal environments. We measured N-fixation rates 2 and 4 wk after exposure to a factorial combination of environments of normal, intermediate and high temperature (16.3, 22.0 and 30.3°C) and light (148.0, 295.7 and 517.3 μmol m(-2) s(-1)). Our results showed that P. schreberi achieved higher N-fixation rates relative to H. splendens in response to warming treatments, but that the highest warming treatment eventually caused N-fixation to decline for both species. Light strongly interacted with warming treatments, having positive effects at low or intermediate temperatures and damaging effects at high temperatures. These results suggest that climate warming may increase N-fixation in boreal forests, but that increased shading by the forest canopy or the occurrence of extreme temperature events could limit increases. They also suggest that P. schreberi may become a larger source of N in boreal forests relative to H. splendens as climate warming progresses.     

Ecosystem controls on nitrogen fixation in boreal feather moss communities

N fixation in feather moss carpets is maximized in late secondary successional boreal forests; however, there is limited understanding of the ecosystem factors that drive cyanobacterial N fixation in feather mosses with successional stage. We conducted a reciprocal transplant experiment to assess factors in both early and late succession that control N fixation in feather moss carpets dominated by Pleurozium schreberi. In 2003, intact microplots of moss carpets (30 cm × 30 cm × 10–20 cm deep) were excavated from three early secondary successional (41–101 years since last fire) forest sites and either replanted within the same stand or transplanted into one of three late successional (241–356 years since last fire) forest sites and the transverse was done for late successional layers of moss. Moss plots were monitored for changes in N-fixation rates by acetylene reduction (June 2003–September 2005) and changes in the presence of cyanobacteria on moss shoots by microscopy (2004). Forest nutrient status was measured using ionic resin capsules buried in the humus layer. Late successional forests exhibit high rates of N fixation and consistently high numbers of cyanobacteria on moss shoots, but low levels of available N. Conversely, early successional forests have higher N availability and have low rates of N fixation and limited presence of cyanobacteria on moss shoots. Transplantation of moss carpets resulted in a significant shift in presence and activity of cyanobacteria 1 year after initiation of the experiment responding to N fertility differences in early versus late successional forests.     

Mosses modify effects of warmer and wetter conditions on tree seedlings at the alpine treeline

Climate warming enables tree seedling establishment beyond the current alpine treeline, but to achieve this, seedlings have to establish within existing tundra vegetation. In tundra, mosses are a prominent feature, known to regulate soil temperature and moisture through their physical structure and associated water retention capacity. Moss presence and species identity might therefore modify the impact of increases in temperature and precipitation on tree seedling establishment at the arctic‐alpine treeline. We followed Betula pubescens and Pinus sylvestris seedling survival and growth during three growing seasons in the field. Tree seedlings were transplanted along a natural precipitation gradient at the subarctic‐alpine treeline in northern Sweden, into plots dominated by each of three common moss species and exposed to combinations of moss removal and experimental warming by open‐top chambers (OTCs). Independent of climate, the presence of feather moss, but not Sphagnum, strongly supressed survival of both tree species. Positive effects of warming and precipitation on survival and growth of B. pubescens seedlings occurred in the absence of mosses and as expected, this was partly dependent on moss species. P. sylvestris survival was greatest at high precipitation, and this effect was more pronounced in Sphagnum than in feather moss plots irrespective of whether the mosses had been removed or not. Moss presence did not reduce the effects of OTCs on soil temperature. Mosses therefore modified seedling response to climate through other mechanisms, such as altered competition or nutrient availability. We conclude that both moss presence and species identity pose a strong control on seedling establishment at the alpine treeline, and that in some cases mosses weaken climate‐change effects on seedling establishment. Changes in moss abundance and species composition therefore have the potential to hamper treeline expansion induced by climate warming.     

The impact of simulated chronic nitrogen deposition on the biomass and N2-fixation activity of two boreal feather moss–cyanobacteria associations

Bryophytes achieve substantial biomass and play several key functional roles in boreal forests that can influence how carbon (C) and nitrogen (N) cycling respond to atmospheric deposition of reactive nitrogen (N_r). They associate with cyanobacteria that fix atmospheric N₂, and downregulation of this process may offset anthropogenic N_r inputs to boreal systems. Bryophytes also promote soil C accumulation by thermally insulating soils, and changes in their biomass influence soil C dynamics. Using a unique large-scale (0.1 ha forested plots), long-term experiment (16 years) in northern Sweden where we simulated anthropogenic N_r deposition, we measured the biomass and N₂-fixation response of two bryophyte species, the feather mosses Hylocomium splendens and Pleurozium schreberi. Our data show that the biomass declined for both species; however, N₂-fixation rates per unit mass and per unit area declined only for H. splendens. The low and high treatments resulted in a 29% and 54% reduction in total feather moss biomass, and a 58% and 97% reduction in total N₂-fixation rate per unit area, respectively. These results help to quantify the sensitivity of feather moss biomass and N₂ fixation to chronic N_r deposition, which is relevant for modelling ecosystem C and N balances in boreal ecosystems.     

Nitrogen fixation in mixed <Emphasis Type="Italic">Hylocomium splendens</Emphasis> moss communities

The pleurocarpus feather moss, Hylocomium splendens, is one of two co-dominant moss species in boreal forest ecosystems and one of the most common mosses on earth, yet little is known regarding its capacity to host cyanobacterial associates and thus contribute total ecosystem N. In these studies, we evaluated the N-fixation potential of the H. splendens–cyanobacteria association and contrasted the N-fixation activity with that of the putative N-fixing moss–cyanobacteria association of Pleurozium schreberi. Studies were conducted to: quantify N-fixation in H. splendens and P. schreberi in sites ranging from southern to northern Fennoscandia; assess N and P availability as drivers of N-fixation rates; contrast season-long N-fixation rates for both mosses; and characterize the cyanobacteria that colonize shoots of H. splendens. Nitrogen-fixation rates were generally low at southern latitudes and higher at northern latitudes (64–69°N) potentially related to anthropogenic N deposition across this gradient. Nitrogen fixation in H. splendens appeared to be less sensitive to N deposition than P. schreberi. The season-long assessment of N-fixation rates at a mixed feather moss site in northern Sweden showed that H. splendens fixed a substantial quantity of N, but about 50% less total N compared to the contribution from P. schreberi. In total, both species provided 1.6 kg fixed N ha⁻¹ year⁻¹. Interestingly, H. splendens demonstrated somewhat higher N-fixation rates at high fertility sites compared to P. schreberi. Nostoc spp. and Stigonema spp. were the primary cyanobacteria found to colonize H. splendens and P. schreberi. These results suggest that H. splendens with associated Nostoc or Stigonema communities contributes a significant quantity of N to boreal forest ecosystems, but the contribution is subordinate to that of P. schreberi at northern latitudes. Epiphytic cyanobacteria are likely a key factor determining the co-dominant presence of these two feather mosses across the boreal biome.     

Invited review: climate change impacts in polar regions: lessons from Antarctic moss bank archives

Mosses are the dominant plants in polar and boreal regions, areas which are experiencing rapid impacts of regional warming. Long‐term monitoring programmes provide some records of the rate of recent climate change, but moss peat banks contain an unrivalled temporal record of past climate change on terrestrial plant Antarctic systems. We summarise the current understanding of climatic proxies and determinants of moss growth for contrasting continental and maritime Antarctic regions, as informed by 13C and 18O signals in organic material. Rates of moss accumulation are more than three times higher in the maritime Antarctic than continental Antarctica with growing season length being a critical determinant of growth rate, and high carbon isotope discrimination values reflecting optimal hydration conditions. Correlation plots of 13C and 18O values show that species (Chorisodontium aciphyllum / Polytrichum strictum) and growth form (hummock / bank) are the major determinants of measured isotope ratios. The interplay between moss growth form, photosynthetic physiology, water status and isotope composition are compared with developments of secondary proxies, such as chlorophyll fluorescence. These approaches provide a framework to consider the potential impact of climate change on terrestrial Antarctic habitats as well as having implications for future studies of temperate, boreal and Arctic peatlands. There are many urgent ecological and environmental problems in the Arctic related to mosses in a changing climate, but the geographical ranges of species and life‐forms are difficult to track individually. Our goal was to translate what we have learned from the more simple systems in Antarctica, for application to Arctic habitats.     

Projecting the distribution of forests in New England in response to climate change

Aim To project the distribution of three major forest types in the northeastern USA in response to expected climate change. Location The New England region of the United States. Methods We modelled the potential distribution of boreal conifer, northern deciduous hardwood and mixed oak–hickory forests using the process‐based BIOME4 vegetation model parameterized for regional forests under historic and projected future climate conditions. Projections of future climate were derived from three general circulation models forced by three global warming scenarios that span the range of likely anthropogenic greenhouse gas emissions. Results Annual temperature in New England is projected to increase by 2.2–3.3 °C by 2041–70 and by 3.0–5.2 °C by 2071–99 with corresponding increases in precipitation of 4.7–9.5% and 6.4–11.4%, respectively. We project that regional warming will result in the loss of 71–100% of boreal conifer forest in New England by the late 21st century. The range of mixed oak–hickory forests will shift northward by 1.0–2.1 latitudinal degrees (c. 100–200 km) and will increase in area by 149–431% by the end of the 21st century. Northern deciduous hardwoods are expected to decrease in area by 26% and move upslope by 76 m on average. The upslope movement of the northern deciduous hardwoods and the increase in oak–hickory forests coincide with an approximate 556 m upslope retreat of the boreal conifer forest by 2071–99. In our simulations, rising atmospheric CO₂ concentrations reduce the losses of boreal conifer forest in New England from expected losses based on climatic change alone. Main conclusion Projected climate warming in the 21st century is likely to cause the extensive loss of boreal conifer forests, reduce the extent of northern hardwood deciduous forests, and result in large increases of mixed oak–hickory forest in New England.     

The climate,the fuel and the land use: Long‐term regional variability of biomass burning in boreal forests

The influence of different drivers on changes in North American and European boreal forests biomass burning (BB) during the Holocene was investigated based on the following hypotheses: land use was important only in the southernmost regions, while elsewhere climate was the main driver modulated by changes in fuel type. BB was reconstructed by means of 88 sedimentary charcoal records divided into six different site clusters. A statistical approach was used to explore the relative contribution of (a) pollen‐based mean July/summer temperature and mean annual precipitation reconstructions, (b) an independent model‐based scenario of past land use (LU), and (c) pollen‐based reconstructions of plant functional types (PFTs) on BB. Our hypotheses were tested with: (a) a west‐east northern boreal sector with changing climatic conditions and a homogeneous vegetation, and (b) a north‐south European boreal sector characterized by gradual variation in both climate and vegetation composition. The processes driving BB in boreal forests varied from one region to another during the Holocene. However, general trends in boreal biomass burning were primarily controlled by changes in climate (mean annual precipitation in Alaska, northern Quebec, and northern Fennoscandia, and mean July/summer temperature in central Canada and central Fennoscandia) and, secondarily, by fuel composition (BB positively correlated with the presence of boreal needleleaf evergreen trees in Alaska and in central and southern Fennoscandia). Land use played only a marginal role. A modification towards less flammable tree species (by promoting deciduous stands over fire‐prone conifers) could contribute to reduce circumboreal wildfire risk in future warmer periods.     

The effect of climate anomalies and human ignition factor on wildfires in Russian boreal forests

Over the last few years anomalies in temperature and precipitation in northern Russia have been regarded as manifestations of climate change. During the same period exceptional forest fire seasons have been reported, prompting many authors to suggest that these in turn are due to climate change. In this paper, we examine the number and areal extent of forest fires across boreal Russia for the period 2002-2005 within two forest categories: 'intact forests' and 'non-intact forests'. Results show a far lower density of fire events in intact forests (5-14 times less) and that those events tend to be in the first 10 km buffer zone inside intact forest areas. Results also show that, during exceptional climatic years (2002 and 2003), fire event density is twice that found during normal years (2004 and 2005) and average areal extent of fire events (burned area) in intact forests is 2.5 times larger than normal. These results suggest that a majority of the fire events in boreal Russia are of human origin and a maximum of one-third of their impact (areal extension) can be attributed to climate anomalies alone, the rest being due to the combined effect of human disturbances and climate anomalies.     

Precipitation frequency alters peatland ecosystem structure and CO2 exchange: Contrasting effects on moss,sedge, and shrub communities

Climate projections forecast a redistribution of seasonal precipitation for much of the globe into fewer, larger events spaced between longer dry periods, with negligible changes in seasonal rainfall totals. This intensification of the rainfall regime is expected to alter near‐surface water availability, which will affect plant performance and carbon uptake. This could be especially important in peatland systems, where large stores of carbon are tightly coupled to water surpluses limiting decomposition. Here, we examined the role of precipitation frequency on vegetation growth and carbon dioxide (CO₂) balances for communities dominated by a Sphagnum moss, a sedge, and an ericaceous shrub in a cool temperate poor fen. Field plots and laboratory monoliths received one of three rainfall frequency treatments, ranging from one event every three days to one event every 14 days, while total rain delivered in a two‐week cycle and the entire season to each treatment remained the same. Separating incident rain into fewer but larger events increased vascular cover in all peatland communities: vascular plant cover increased 6× in the moss‐dominated plots, nearly doubled in the sedge plots, and tripled in the shrub plots in Low‐Frequency relative to High‐Frequency treatments. Gross ecosystem productivity was lowest in moss communities receiving low‐frequency rain, but higher in sedge and shrub communities under the same conditions. Net ecosystem exchange followed this pattern: fewer events with longer dry periods increased CO₂ flux to the atmosphere from the moss while vascular plant‐dominated communities became more of a sink for CO₂. Results of this study suggest that changes to rainfall frequency already occurring and predicted to continue will lead to increased vascular plant cover in peatlands and will impact their carbon‐sink function.     

Fine root foraging strategies in Norway spruce forests across a European climate gradient

Fine root acclimation to different environmental conditions is crucial for growth and sustainability of forest trees. Relatively small changes in fine root standing biomass (FRB), morphology or mycorrhizal symbiosis may result in a large change in forest carbon, nutrient and water cycles. We elucidated the changes in fine root traits and associated ectomycorrhizal (EcM) fungi in 12 Norway spruce stands across a climatic and N deposition gradient from subarctic‐boreal to temperate regions in Europe (68°N–48°N). We analysed the standing FRB and the ectomycorrhizal root tip biomass (EcMB, g m^?2) simultaneously with measurements of the EcM root morphological traits (e.g. mean root length, root tissue density (RTD), N% in EcM roots) and frequency of dominating EcM fungi in different stands in relation to climate, soil and site characteristics. Latitude and N deposition explained the greatest proportion of variation in fine root traits. EcMB per stand basal area (BA) increased exponentially with latitude: by about 12.7 kg m^?2 with an increase of 10° latitude from southern Germany to Estonia and southern Finland and by about 44.7 kg m^?2 with next latitudinal 10° from southern to northern Finland. Boreal Norway spruce forests had 4.5 to 11 times more EcM root tips per stand BA, and the tips were 2.1 times longer, with 1.5 times higher RTD and about 1/3 lower N concentration. There was 19% higher proportion of root tips colonized by long‐distance exploration type forming EcM fungi in the southern forests indicating importance of EcM symbiont foraging strategy in fine root nutrient acquisition. In the boreal zone, we predict ca. 50% decrease in EcMB per stand BA with an increase of 2 °C annual mean temperature. Different fine root foraging strategies in boreal and temperate forests highlight the importance of complex studies on respective regulatory mechanisms in changing climate.     

Precipitation regime controls bryosphere carbon cycling similarly across contrasting ecosystems

In arctic and boreal ecosystems, ground bryophytes play an important role in regulating carbon (C) exchange between vast belowground C stores and the atmosphere. Climate is changing particularly fast in these high-latitude regions, but it is unclear how altered precipitation regimes will affect C dynamics in the bryosphere (i.e. the ground moss layer including senesced moss, litter and associated biota) and the closely associated upper humus layer, and how these effects will vary across contrasting environmental conditions. Here, we set up a greenhouse experiment in which mesocosms were assembled containing samples of the bryosphere, dominated by the feather moss Hylocomium splendens, and the upper humus layer, that were collected from across a boreal forest chronosequence in northern Sweden which varies strongly in nutrient availability, productivity and soil biota. We tested the effect of variation in precipitation volume and frequency on CO₂ exchange and dissolved organic carbon (DOC) export, and on moss growth. As expected, reduced precipitation volume and frequency lowered net CO₂ efflux, DOC export and moss growth. However, by regulating moisture, the lower bryosphere and humus layers often mediated how precipitation volume and frequency interacted to drive C dynamics. For example, less frequent precipitation reduced moss growth only when precipitation volume was low. When volume was high, high moisture content of the humus layer helped avoid moss desiccation. Variation in precipitation regime affected C cycling consistently in samples collected across the chronosequence, despite large environmental variation along the sequence. This suggests that the bryosphere exerts a strong buffering effect on environmental variation at the forest floor, which leads to similar responses of C cycling to external perturbations across highly contrasting ecosystems. As such, our study indicates that projected increases in droughts and ground evapotranspiration in high-latitude regions resulting from climate change will consistently reduce C losses from moss-dominated ecosystems.     

Persistent and pervasive compositional shifts of western boreal forest plots in Canada

Species compositional shifts have important consequences to biodiversity and ecosystem function and services to humanity. In boreal forests, compositional shifts from late‐successional conifers to early‐successional conifers and deciduous broadleaves have been postulated based on increased fire frequency associated with climate change truncating stand age‐dependent succession. However, little is known about how climate change has affected forest composition in the background between successive catastrophic fires in boreal forests. Using 1797 permanent sample plots from western boreal forests of Canada measured from 1958 to 2013, we show that after accounting for stand age‐dependent succession, the relative abundances of early‐successional deciduous broadleaves and early‐successional conifers have increased at the expense of late‐successional conifers with climate change. These background compositional shifts are persistent temporally, consistent across all forest stand ages and pervasive spatially across the region. Rising atmospheric CO₂ promoted early‐successional conifers and deciduous broadleaves, and warming increased early‐successional conifers at the expense of late‐successional conifers, but compositional shifts were not associated with climate moisture index. Our results emphasize the importance of climate change on background compositional shifts in the boreal forest and suggest further compositional shifts as rising CO₂ and warming will continue in the 21st century.     

Anthropogenic nitrogen deposition in boreal forests has a minor impact on the global carbon cycle

It is proposed that increases in anthropogenic reactive nitrogen (N_r) deposition may cause temperate and boreal forests to sequester a globally significant quantity of carbon (C); however, long‐term data from boreal forests describing how C sequestration responds to realistic levels of chronic N_r deposition are scarce. Using a long‐term (14‐year) stand‐scale (0.1 ha) N addition experiment (three levels: 0, 12.5, and 50 kg N ha⁻¹ yr⁻¹) in the boreal zone of northern Sweden, we evaluated how chronic N additions altered N uptake and biomass of understory communities, and whether changes in understory communities explained N uptake and C sequestration by trees. We hypothesized that understory communities (i.e. mosses and shrubs) serve as important sinks for low‐level N additions, with the strength of these sinks weakening as chronic N addition rates increase, due to shifts in species composition. We further hypothesized that trees would exhibit nonlinear increases in N acquisition, and subsequent C sequestration as N addition rates increased, due to a weakening understory N sink. Our data showed that understory biomass was reduced by 50% in response to the high N addition treatment, mainly due to reduced moss biomass. A ¹⁵N labeling experiment showed that feather mosses acquired the largest fraction of applied label, with this fraction decreasing as the chronic N addition level increased. Contrary to our hypothesis, the proportion of label taken up by trees was equal (ca. 8%) across all three N addition treatments. The relationship between N addition and C sequestration in all vegetation pools combined was linear, and had a slope of 16 kg C kg⁻¹ N. While canopy retention of N_r deposition may cause C sequestration rates to be slightly different than this estimate, our data suggest that a minor quantity of annual anthropogenic CO₂ emissions are sequestered into boreal forests as a result of N_r deposition.     

Climate change drives substantial decline of understorey species richness and abundance in Norway spruce forests during 32 years of vegetation monitoring

Questions

Observations in permanent forest vegetation plots in Norway and elsewhere indicate that complex changes have taken place over the period 1988–2020. These observations are summarised in the “climate-induced understorey change (CIUC)” hypothesis, i.e. that the understorey vegetation of old-growth boreal forests in Norway undergoes significant long-term changes and that these changes are consistent with the ongoing climate change as an important driver. Seven testable predictions were derived from the CIUC hypothesis.

Location

Norway.

Methods

Vegetation has been monitored in a total of 458 permanently marked plots, each 1 m², in nine old-growth forest sites dominated by Picea abies at intervals of 5–8 years over the 32-year study period. For each of the 52 combinations of site and year, we obtained response variables for the abundance of single species, abundance and species density of taxonomic–ecological species groups and two size classes of cryptogams, and site species richness. All of these variables were subjected to linear regression modelling with site and year as predictors.

Results

Mean annual temperature, growing-season length and the number of days with precipitation were higher in the study period than in the preceding ca. 30-year period, resulting in increasingly favourable conditions for bryophyte growth. Site species richness decreased by 13% over the 32-year study period. On average, group abundance of vascular plants decreased by 24% (decrease in forbs: 38%). Patterns of group abundance change differed among cryptogam groups: although peat-moss abundance increased by 39%, the abundance of mosses, hepatics and lichens decreased by 13%, 49% and 67%, respectively. Group abundance of small cryptogams decreased by 61%, whereas a 13% increase was found for large cryptogams. Of 61 single species tested for abundance change, a significant decrease was found for 43 species, whereas a significant increase was found only for 6 species.

Conclusions

The major patterns of change in species richness, group species density and group abundance observed over the 32-year study period accord with most predictions from the CIUC hypothesis and are interpreted as direct and indirect responses to climate change, partly mediated through changes in the population dynamics of microtine rodents. The more favourable climate for bryophyte growth explains the observed increase for a few large bryophyte species, whereas the decrease observed for small mosses and hepatics is interpreted as an indirect amensalistic effect, brought about by shading and burial in mats of larger species and accelerated by reduced fine-scale disturbance by microtine rodents. Indirect effects of a thicker moss mat most likely drive the vascular plant decline although long-term effects of tree-stand dynamics and former logging cannot be completely ruled out. Our results suggest that the ongoing climate change has extensive, cascading effects on boreal forest ecosystems. The importance of long time-series of permanent vegetation plots for detecting and understanding the effects of climate change on boreal forests is emphasised.     

Climate warming feedback from mountain birch forest expansion: reduced albedo dominates carbon uptake

Expanding high‐elevation and high‐latitude forest has contrasting climate feedbacks through carbon sequestration (cooling) and reduced surface reflectance (warming), which are yet poorly quantified. Here, we present an empirically based projection of mountain birch forest expansion in south‐central Norway under climate change and absence of land use. Climate effects of carbon sequestration and albedo change are compared using four emission metrics. Forest expansion was modeled for a projected 2.6 °C increase in summer temperature in 2100, with associated reduced snow cover. We find that the current (year 2000) forest line of the region is circa 100 m lower than its climatic potential due to land‐use history. In the future scenarios, forest cover increased from 12% to 27% between 2000 and 2100, resulting in a 59% increase in biomass carbon storage and an albedo change from 0.46 to 0.30. Forest expansion in 2100 was behind its climatic potential, forest migration rates being the primary limiting factor. In 2100, the warming caused by lower albedo from expanding forest was 10 to 17 times stronger than the cooling effect from carbon sequestration for all emission metrics considered. Reduced snow cover further exacerbated the net warming feedback. The warming effect is considerably stronger than previously reported for boreal forest cover, because of the typically low biomass density in mountain forests and the large changes in albedo of snow‐covered tundra areas. The positive climate feedback of high‐latitude and high‐elevation expanding forests with seasonal snow cover exceeds those of afforestation at lower elevation, and calls for further attention of both modelers and empiricists. The inclusion and upscaling of these climate feedbacks from mountain forests into global models is warranted to assess the potential global impacts.     

The origin of soil organic matter controls its composition and bioreactivity across a mesic boreal forest latitudinal gradient

Warmer climates have been associated with reduced bioreactivity of soil organic matter (SOM) typically attributed to increased diagenesis; the combined biological and physiochemical transformation of SOM. In addition, cross‐site studies have indicated that ecosystem regime shifts, associated with long‐term climate warming, can affect SOM properties through changes in vegetation and plant litter production thereby altering the composition of soil inputs. The relative importance of these two controls, diagenesis and inputs, on SOM properties as ecosystems experience climate warming, however, remains poorly understood. To address this issue we characterized the elemental, chemical (nuclear magnetic resonance spectroscopy and total hydrolysable amino acids analysis), and isotopic composition of plant litter and SOM across a well‐constrained mesic boreal forest latitudinal transect in Atlantic Canada. Results across forest sites within each of three climate regions indicated that (1) climate history and diagenesis affect distinct parameters of SOM chemistry, (2) increases in SOM bioreactivity with latitude were associated with elevated proportions of carbohydrates relative to plant waxes and lignin, and (3) despite the common forest type across regions, differences in SOM chemistry by climate region were associated with chemically distinct litter inputs and not different degrees of diagenesis. The observed climate effects on vascular plant litter chemistry, however, explained only part of the regional differences in SOM chemistry, most notably the higher protein content of SOM from warmer regions. Greater proportions of lignin and aliphatic compounds and smaller proportions of carbohydrates in warmer sites' soils were explained by the higher proportion of vascular plant relative to moss litter in the warmer relative to cooler forests. These results indicate that climate change induced decreases in the proportion of moss inputs not only impacts SOM chemistry but also increases the resistance of SOM to decomposition, thus significantly altering SOM cycling in these boreal forest soils.     

The climate sensitivity of carbon,timber, and species richness covaries with forest age in boreal–temperate North America

Climate change threatens the provisioning of forest ecosystem services and biodiversity (ESB). The climate sensitivity of ESB may vary with forest development from young to old‐growth conditions as structure and composition shift over time and space. This study addresses knowledge gaps hindering implementation of adaptive forest management strategies to sustain ESB. We focused on a number of ESB indicators to (a) analyze associations among carbon storage, timber growth rate, and species richness along a forest development gradient; (b) test the sensitivity of these associations to climatic changes; and (c) identify hotspots of climate sensitivity across the boreal–temperate forests of eastern North America. From pre‐existing databases and literature, we compiled a unique dataset of 18,507 forest plots. We used a full Bayesian framework to quantify responses of nine ESB indicators. The Bayesian models were used to assess the sensitivity of these indicators and their associations to projected increases in temperature and precipitation. We found the strongest association among the investigated ESB indicators in old forests (>170 years). These forests simultaneously support high levels of carbon storage, timber growth, and species richness. Older forests also exhibit low climate sensitivity of associations among ESB indicators as compared to younger forests. While regions with a currently low combined ESB performance benefitted from climate change, regions with a high ESB performance were particularly vulnerable to climate change. In particular, climate sensitivity was highest east and southeast of the Great Lakes, signaling potential priority areas for adaptive management. Our findings suggest that strategies aimed at enhancing the representation of older forest conditions at landscape scales will help sustain ESB in a changing world.     

Photosynthetic traits of Sphagnum and feather moss species in undrained,drained and rewetted boreal spruce swamp forests

In restored peatlands, recovery of carbon assimilation by peat‐forming plants is a prerequisite for the recovery of ecosystem functioning. Restoration by rewetting may affect moss photosynthesis and respiration directly and/or through species successional turnover. To quantify the importance of the direct effects and the effects mediated by species change in boreal spruce swamp forests, we used a dual approach: (i) we measured successional changes in moss communities at 36 sites (nine undrained, nine drained, 18 rewetted) and (ii) photosynthetic properties of the dominant Sphagnum and feather mosses at nine of these sites (three undrained, three drained, three rewetted). Drainage and rewetting affected moss carbon assimilation mainly through species successional turnover. The species differed along a light‐adaptation gradient, which separated shade‐adapted feather mosses from Sphagnum mosses and Sphagnum girgensohnii from other Sphagna, and a productivity and moisture gradient, which separated Sphagnum riparium and Sphagnum girgensohnii from the less productive S. angustifolium, S. magellanicum and S. russowii. Undrained and drained sites harbored conservative, low‐production species: hummock‐Sphagna and feather mosses, respectively. Ditch creation and rewetting produced niches for species with opportunistic strategies and high carbon assimilation. The direct effects also caused higher photosynthetic productivity in ditches and in rewetted sites than in undrained and drained main sites.