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Numerous recent studies have argued that moisture limitation is leading to growth declines and mortality of black spruce (Picea mariana) and white spruce (Picea glauca) in the North American boreal forest. A parallel line of research suggests that increasingly common severe wildfires are altering successional pathways and leading to long‐term replacement of spruce forests with those dominated by paper birch (Betula papyrifera, Betula neoalaskana) and trembling aspen (Populus tremuloides). When both conifers and deciduous trees establish after fire, this biome shift hypothesis implicitly assumes that deciduous species will outcompete the conifers, owing to their more rapid vertical growth and because they might be less sensitive to warm and dry conditions.We established a research site in a white spruce‐paper birch forest on an east‐facing slope in interior Alaska and tested the hypothesis that Alaska paper birch are better adapted to warm and dry conditions than white spruce. Over 6 years (2013–2018), we made hourly measurements of microclimate and xylem sap flux of both species. We also collected increment cores and conducted climate‐growth analyses for both species.During our 6‐year study, growing seasons with low volumetric soil water content (SWC) were those that followed shallow winter snowpacks and had limited summer rainfall, not necessarily those with warm air temperature. Both species were sensitive to moisture limitation. The tree‐ring data revealed significant positive effects of cumulative water year precipitation on radial growth, with a steeper slope for paper birch than for white spruce. Radial growth of both species was also positively related to mean water year air temperature. Sap flux density declined progressively for white spruce over the range of observed SWC and abruptly for paper birch when SWC fell below ~15%.Synthesis. Our results show that, while paper birch might be less sensitive to mild drought than white spruce, it may be more sensitive to severe drought, raising questions about the ability of paper birch to outcompete co‐occurring white spruce in a drier climate. [ABSTRACT FROM AUTHOR]

Abstract: The boreal biome represents approximately one third of the world's forested area and plays an important role in global biogeochemical and energy cycles. Numerous studies in boreal Alaska have concluded that growth of black and white spruce is declining as a result of temperature‐induced drought stress. The combined evidence of declining spruce growth and changes in the fire regime that favor establishment of deciduous tree species has led some investigators to suggest the region may be transitioning from dominance by spruce to dominance by deciduous forests and/or grasslands. Although spruce growth trends have been extensively investigated, few studies have evaluated long‐term radial growth trends of the dominant deciduous species (Alaska paper birch and trembling aspen) and their sensitivity to moisture availability. We used a large and spatially extensive sample of tree cores from interior Alaska to compare long‐term growth trends among contrasting tree species (white and black spruce vs. birch and aspen). All species showed a growth peak in the mid‐1940s, although growth following the peak varied strongly across species. Following an initial decline from the peak, growth of white spruce showed little evidence of a trend, while black spruce and birch growth showed slight growth declines from ~1970 to present. Aspen growth was much more variable than the other species and showed a steep decline from ~1970 to present. Growth of birch, black and white spruce was sensitive to moisture availability throughout most of the tree‐ring chronologies, as evidenced by negative correlations with air temperature and positive correlations with precipitation. However, a positive correlation between previous July precipitation and aspen growth disappeared in recent decades, corresponding with a rise in the population of the aspen leaf miner (Phyllocnistis populiella), an herbivorous moth, which may have driven growth to a level not seen since the early 20th century. Our results provide important historical context for recent growth and raise questions regarding competitive interactions among the dominant tree species and exchanges of carbon and energy in the warming climate of interior Alaska. [ABSTRACT FROM AUTHOR]

Rising temperatures associated with climate change have been shown to negatively affect the photosynthetic rates of boreal forest tree saplings at their southern range limits. To quantify the responses of ectomycorrhizal ( EM) fungal communities associated with poorly performing hosts, we sampled the roots of Betula papyrifera and Abies balsamea saplings growing in the B4Warmed (Boreal Forest Warming at an Ecotone in Danger) experiment. EM fungi on the root systems of both hosts were compared from ambient and +3.4 °C air and soil warmed plots at two sites in northern Minnesota. EM fungal communities were assessed with high-throughput sequencing along with measures of plant photosynthesis, soil temperature, moisture, and nitrogen. Warming selectively altered EM fungal community composition at both the phylum and genus levels, but had no significant effect on EM fungal operational taxonomic unit ( OTU) diversity. Notably, warming strongly favored EM Ascomycetes and EM fungi with short-contact hyphal exploration types. Declining host photosynthetic rates were also significantly inversely correlated with EM Ascomycete and EM short-contact exploration type abundance, which may reflect a shift to less carbon demanding fungi due to lower photosynthetic capacity. Given the variation in EM host responses to warming, both within and between ecosystems, better understanding the link between host performance and EM fungal community structure will to clarify how climate change effects cascade belowground. [ABSTRACT FROM AUTHOR]

Organisms survive environmental variation by combining homeostatic regulation of critical states with allostatic variation of other traits, and species differences in these responses can contribute to coexistence in temporally variable environments.In this paper, we simultaneously record variation in three functional traits—body temperature (Tb), heart rate and activity—in relation to three forms of environmental variation—air temperature (Ta), photoperiod and experimentally manipulated resource levels—in free‐ranging snowshoe hares and North American red squirrels to characterize distinctions in homeotherm responses to the extreme conditions of northern boreal winters.Hares and squirrels differed in the level and precision of Tb regulation, but also in the allostatic pathways necessary to maintain thermal homeostasis. Hares demonstrated a stronger metabolic pathway (through heart rate variation reflective of the thermogenesis), while squirrels demonstrated a stronger behavioural pathway (through activity variation that minimizes cold exposure).As intermediate‐sized, winter‐active homeotherms, hares and squirrels share many functional attributes, yet, through the integrated monitoring of multiple functional traits in response to shared environmental variation, our study reveals many pairwise species differences in homeostatic and allostatic traits, that both define and are defined by the natural history, functional niches and coexistence of sympatric species. A free Plain Language Summary can be found within the Supporting Information of this article. [ABSTRACT FROM AUTHOR]

Many studies have focused on vegetation across forest edges to study impacts of edges created by human activities on forest structure and composition, or patterns of vegetation at inherent natural edges. Our objective was to create a database of plant‐related variables across different types of edges from various studies (mainly from across Canada, but also in Brazil and Belize) to facilitate edge research. We compiled data on vegetation along more than 300 transects perpendicular to forest edges adjacent to clear‐cuts, burned areas, bogs, lakes, barrens, insect disturbances, and riparian areas from 24 studies conducted over the past three decades. Data were compiled for more than 400 plant species and forest structure variables (e.g., trees, logs, canopy cover). All data were collected with a similar sampling design of quadrats along transects perpendicular to forest edges, but with varying numbers of transects and quadrats, and distances from the edge. The purpose for most of the studies was either to determine the distance of edge influence (edge width) or to explore the pattern of vegetation along the edge to interior gradient. We provide data tables for the cover of plant species and functional groups, the species and size of live and dead trees, the density of saplings, maximum height of functional groups and shrub species, and the cover of functional groups at different heights (vertical distribution of vegetation). The Forest Edge Research Network (FERN) database provides extensive data on many variables that can be used for further study including meta‐analyses and can assist in answering questions important to conservation efforts (e.g., how is distance of edge influence from created edges affected by different factors?). We plan to expand this database with subsequent studies from the authors and we invite others to contribute to make this a more global database. The data are released under a CC0 license. When using these data, we ask that you cite this data paper and any relevant publications listed in our metadata file. We also encourage you to contact the first author if you are planning to use or contribute to this database. [ABSTRACT FROM AUTHOR]

Summary: Nitrogen (N2)‐fixing moss microbial communities play key roles in nitrogen cycling of boreal forests. Forest type and leaf litter inputs regulate moss abundance, but how they control moss microbiomes and N2‐fixation remains understudied. We examined the impacts of forest type and broadleaf litter on microbial community composition and N2‐fixation rates of Hylocomium splendens and Pleurozium schreberi.We conducted a moss transplant and leaf litter manipulation experiment at three sites with paired paper birch (Betula neoalaskana) and black spruce (Picea mariana) stands in Alaska. We characterized bacterial communities using marker gene sequencing, determined N2‐fixation rates using stable isotopes (15N2) and measured environmental covariates.Mosses native to and transplanted into spruce stands supported generally higher N2‐fixation and distinct microbial communities compared to similar treatments in birch stands. High leaf litter inputs shifted microbial community composition for both moss species and reduced N2‐fixation rates for H. splendens, which had the highest rates. N2‐fixation was positively associated with several bacterial taxa, including cyanobacteria.The moss microbiome and environmental conditions controlled N2‐fixation at the stand and transplant scales. Predicted shifts from spruce‐ to deciduous‐dominated stands will interact with the relative abundances of mosses supporting different microbiomes and N2‐fixation rates, which could affect stand‐level N inputs. [ABSTRACT FROM AUTHOR]

Scavenging by vertebrates can have important impacts on food web stability and persistence, and can alter the distribution of nutrients throughout the landscape. However, scavenging communities have been understudied in most regions around the globe, and we lack understanding of the biotic drivers of vertebrate scavenging dynamics.In this paper, we examined how changes in prey density and carrion biomass caused by population cycles of a primary prey species, the snowshoe hare Lepus americanus, influence scavenging communities in the northern boreal forest. We further examined the impact of habitat and temperature on scavenging dynamics.We monitored the persistence time, time until first scavenger, and number of species scavenging experimentally‐placed hare carcasses over four consecutive years in the southwestern Yukon. We simultaneously monitored hare density and carrion biomass to examine their influence relative to temperature, habitat, and seasonal effects. For the primary scavengers, we developed species‐specific scavenging models to determine variation on the effects of these factors across species, and determine which species may be driving temporal patterns in the entire community.We found that the efficiency of the scavenging community was affected by hare density, with carcass persistence decreasing when snowshoe hare densities declined, mainly due to increased scavenging rates by Canada lynx Lynx canadensis. However, prey density did not influence the number of species scavenging a given carcass, suggesting prey abundance affects carrion recycling but not necessarily the number of connections in the food web. In addition, scavenging rates increased in warmer temperatures, and there were strong seasonal effects on the richness of the vertebrate scavenging community.Our results demonstrate that vertebrate scavenging communities are sensitive to changes in species' demography and environmental change, and that future assessments of food web dynamics should consider links established through scavenging. [ABSTRACT FROM AUTHOR]

As boreal forests rapidly warm due to anthropogenic climate change, long‐term baseline community data are needed to effectively characterize the corresponding ecological changes that are occurring in these forests. The combined seasonal dynamics (SEADYN) and annual dynamics (ANNDYN) data set, which documents the vegetative changes in boreal forests during the snow‐free period, is one such source of baseline community data. These data were collected by George H. La Roi and colleagues in Alberta, Canada from 1980 to 2015 within permanent sampling plots established in the Hondo‐Slave Lake area (eight stands; 1980–2015) in central Alberta and the Athabasca Oil Sands (AOS) region (17 stands; 1981–1984) near Fort McMurray in northeastern Alberta. Various data were collected, with temporal and spatial coverage differing by data set. These data sets include, but are not limited to, cover of each identified vascular plant and bryoid (moss, liverwort, and lichen) species; forest mensuration; forest litter production; and soil temperature and moisture. Notably, permanent sampling plots were set up as a grid, which will facilitate analyses of spatial relations. These data can be used to analyze long‐term changes in seasonal dynamics and succession within boreal forest communities and serve as a baseline for comparison with future forest conditions in unmanaged, managed, and reclaimed forests. Data are released under a CC‐BY license; please cite this data paper when using the data for analyses. [ABSTRACT FROM AUTHOR]

Aim: Previous analyses of remotely sensed data detected the multimodality of the tree‐cover distribution of the boreal forest, and identified areas with potentially alternative tree‐cover states. This paper aims at investigating the causes of multimodality and multistability of the boreal forest, their influence on the asymmetric tree species distribution between Eurasia and North America, and whether multistability could be associated with recent greening trends in leaf area index (LAI) and normalized difference vegetation index (NDVI). Location: Eurasian and North American boreal forests. Time period: 2000–2010. Major taxa studied: Boreal forest plant functional types. Methods: We employ a conceptual model based on tree species competition to simulate the sensitivity of tree cover to stochastic disturbances and to changes in environmental factors. We include different plant functional types based on survival adaptations, and force the model with remotely sensed environmental data. We analyse the model as a dynamical system. We use metrics from statistics and information theory to compare the detection of alternative tree‐cover states and greening trends in LAI and NDVI. Results: We find that multimodality and multistability can emerge through competition between different plant functional types. Additionally, our model is able to reproduce the asymmetry in tree species distribution between Eurasia and North America. Moreover, changes in permafrost distribution can be associated with phenomenological bifurcation points of the model. Finally, we find that the detection of multistable areas is not affected by recent vegetation trends, whereas shifts between alternative states could have affected the greening trends. Main conclusions: Tree‐cover multistability in the boreal region can emerge through competition between species subject to periodic disturbances. Changes in permafrost thaw and distribution could be responsible for the asymmetry in tree species distribution between North America and Eurasia. Climate change and permafrost degradation could cause shifts in tree‐cover states and dominant species. Recent vegetation greening trends in multistable areas could have been affected by shifts between alternative states. [ABSTRACT FROM AUTHOR]

Carbon release from thawing permafrost soils could significantly exacerbate global warming as the active-layer deepens, exposing more carbon to decay. Plant community and soil properties provide a major control on this by influencing the maximum depth of thaw each summer (active-layer thickness; ALT), but a quantitative understanding of the relative importance of plant and soil characteristics, and their interactions in determine ALTs, is currently lacking. To address this, we undertook an extensive survey of multiple vegetation and edaphic characteristics and ALTs across multiple plots in four field sites within boreal forest in the discontinuous permafrost zone ( NWT, Canada). Our sites included mature black spruce, burned black spruce and paper birch, allowing us to determine vegetation and edaphic drivers that emerge as the most important and broadly applicable across these key vegetation and disturbance gradients, as well as providing insight into site-specific differences. Across sites, the most important vegetation characteristics limiting thaw (shallower ALTs) were tree leaf area index ( LAI), moss layer thickness and understory LAI in that order. Thicker soil organic layers also reduced ALTs, though were less influential than moss thickness. Surface moisture (0-6 cm) promoted increased ALTs, whereas deeper soil moisture (11-16 cm) acted to modify the impact of the vegetation, in particular increasing the importance of understory or tree canopy shading in reducing thaw. These direct and indirect effects of moisture indicate that future changes in precipitation and evapotranspiration may have large influences on ALTs. Our work also suggests that forest fires cause greater ALTs by simultaneously decreasing multiple ecosystem characteristics which otherwise protect permafrost. Given that vegetation and edaphic characteristics have such clear and large influences on ALTs, our data provide a key benchmark against which to evaluate process models used to predict future impacts of climate warming on permafrost degradation and subsequent feedback to climate. [ABSTRACT FROM AUTHOR]

Wood harvesting in boreal forests typically consists of sequential harvesting operations within a rotation: a few thinnings and a final felling. The aim of this paper is to model differentiated relative global warming potential ( GWP) coefficients for stemwood use from different thinnings and final fellings, and correction factors for long-lived wood products, potentially applicable in life cycle assessment studies. All thinnings and final fellings influence the development of forest carbon stocks. The climate impact of a single harvesting operation is generated in comparison with no harvesting, thus encountering a methodological problem on how to handle the subsequent operations. The dynamic forest stand simulator MOTTI was applied in the modelling of evolution of forest carbon stocks at landscape level in Southern Finland. The landscape-level approach for climate impact assessment gave results similar to some stand-level approaches presented in previous literature that included the same forest C pools and also studied the impacts relative to the no-harvest situation. The climate impacts of stemwood use decreased over time. For energy use, the impacts were higher or similar in the short term and 0-50% lower in the midterm in comparison with an identical amount of fossil CO2. The impacts were to some extent (approximately 20-40%) lower for wood from intermediate thinnings than for wood from final fellings or first thinnings. However, the study reveals that product lifetime has higher relative influence on the climate impacts of wood-based value chains than whether the stemwood originates from thinnings or final fellings. Although the evolution of future C stocks in unmanaged boreal forests is uncertain, a sensitivity analysis suggests that landscape-level model results for climate impacts would not be sensitive to the assumptions made on the future evolution of C stocks in unmanaged forest. Energy use of boreal stemwood seems to be far from climate neutral. [ABSTRACT FROM AUTHOR]

Tree growth at northern treelines is generally temperature‐limited due to cold and short growing seasons. However, temperature‐induced drought stress was repeatedly reported for certain regions of the boreal forest in northwestern North America, provoked by a significant increase in temperature and possibly reinforced by a regime shift of the pacific decadal oscillation (PDO). The aim of this study is to better understand physiological growth reactions of white spruce, a dominant species of the North American boreal forest, to PDO regime shifts using quantitative wood anatomy and traditional tree‐ring width (TRW) analysis. We investigated white spruce growth at latitudinal treeline across a >1,000 km gradient in northwestern North America. Functionally important xylem anatomical traits (lumen area, cell‐wall thickness, cell number) and TRW were correlated with the drought‐sensitive standardized precipitation–evapotranspiration index of the growing season. Correlations were computed separately for complete phases of the PDO in the 20th century, representing alternating warm/dry (1925–1946), cool/wet (1947–1976) and again warm/dry (1977–1998) climate regimes. Xylem anatomical traits revealed water‐limiting conditions in both warm/dry PDO regimes, while no or spatially contrasting associations were found for the cool/wet regime, indicating a moisture‐driven shift in growth‐limiting factors between PDO periods. TRW reflected only the last shift of 1976/1977, suggesting different climate thresholds and a higher sensitivity to moisture availability of xylem anatomical traits compared to TRW. This high sensitivity of xylem anatomical traits permits to identify first signs of moisture‐driven growth in treeline white spruce at an early stage, suggesting quantitative wood anatomy being a powerful tool to study climate change effects in the northwestern North American treeline ecotone. Projected temperature increase might challenge growth performance of white spruce as a key component of the North American boreal forest biome in the future, when drier conditions are likely to occur with higher frequency and intensity. [ABSTRACT FROM AUTHOR]