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1. Boreal forest soils contain large stocks of soil carbon (C) that may be sensitive to changes in climate and disturbance. Destablization of boreal forest soil C through changes in C inputs, belowground C pools and/or wildfire could feedback to accelerate rising atmospheric CO2 concentration. Additionally, increasing frequency of severe fires may be changing the dominant forest types and reshaping aboveground C stocks. 2. Although controls on ecosystem C pools have received considerable attention, many studies have been limited to locations near the road system, leading to uncertainty in current and future C stocks across boreal Alaska. Here, we leveraged 545 randomly selected and spatially balanced Forest Inventory and Analysis (FIA) plots across ~13.5 million hectares in interior Alaska to examine the factors governing soil and live tree C pools. 3. Forest type mediated the effects of mean summer air temperature and the probability of near-surface permafrost on soil C. Meanwhile, forest type, stand age and aspect were the primary drivers of live tree C. Overall, plots with a known history of wildfire during the past 70 years did not have significantly different soil C stocks than plots without a known history of fire, likely due to the historical predominance of low severity fires. 4. Where wildfire likely initiated a transition to deciduous trees (19% of plots), live tree and soil C pools were reduced by 16% and 20%, respectively. Ecosystem C likely recovered over time, as maturing deciduous stands rapidly gained C in live trees. Deciduous stands without a known fire had comparatively very large live tree C stocks, suggesting a significant change in the distribution of ecosystem C following severe fire. 5. Synthesis. Our results highlight the nuanced interactions among wildfire, landscape position and forest type that will play important roles in shaping future boreal ecosystem C stocks. [ABSTRACT FROM AUTHOR]

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]

Climate change is expected to increase woody vegetation abundance in the Arctic, yet the magnitude, spatial pattern and pathways of change remain uncertain. We compared historical orthophotos photos (1952 and 1979) with high‐resolution satellite imagery (2015) to examine six decades of change in abundance of white spruce Picea glauca and tall shrubs (Salix spp., Alnus spp.) near the Agashashok River in northwest Alaska. We established ~3000 random points within our ~5500 ha study area for classification into nine cover types. To examine physiographic controls on tree abundance, we fit multinomial log‐linear models with predictors derived from a digital elevation model and with arctic tundra, alpine tundra and 'tree' as levels of a categorical response variable. Between 1952 and 2015, points classified as arctic and alpine tundra decreased by 31% and 15%, respectively. Meanwhile, tall shrubs increased by 86%, trees mixed with tall shrubs increased by 385% and forest increased by 84%. Tundra with tall shrubs rarely transitioned to forest. The best multinomial model explained 71% of variation in cover and included elevation, slope and an interaction between slope and 'northness'. Treeline was defined as the elevation where the probability of tree presence equaled that of tundra. Mean treeline elevation in 2015 was 202 m, corresponding with a June–August mean air temperature > 11°C, which is > 4°C warmer than the 6–7°C isotherm associated with global treeline elevations. Our results show dramatic increases in the abundance of trees and tall shrubs, question the universality of air temperature as a predictor of treeline elevation and suggest two mutually exclusive pathways of vegetation change, because tundra that gained tall shrubs rarely transitioned to forest. Conversion of tundra to tall shrubs and forest has important and potentially contrasting implications for carbon cycling, surface energy exchange and wildlife habitat in the Arctic. [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]

Long-term experiments provide a way to test presumed causes of successional or environmentally driven vegetation changes. Early-successional nitrogen (N)-fixing plants are widely thought to facilitate productivity and vegetation development on N-poor sites, thus accounting for observed vegetation patterns later in succession. We tested this facilitative impact on vegetation development in a 23-yr field experiment on an Interior Alaska ( USA) floodplain. On three replicate early-successional silt bars, we planted late-successional white spruce ( Picea glauca) seedlings in the presence and absence of planted seedlings of an early-successional N-fixing shrub, thinleaf alder ( Alnus incana). Alder initially facilitated survivorship and growth of white spruce. Within six years, however, after canopy closure, alder negatively affected spruce survivorship and growth. Our three replicate sites followed different successional trajectories. One site was eliminated by erosion and supported no vegetation development during our study. The other two sites, which differed in site moisture, diverged in vegetation composition. Structural equation modeling ( SEM) suggested that, in the drier of these two sites, alder inhibited spruce growth directly (presumably by competition) and indirectly through effects mediated by competition with other woody species. However, at the wetter site, alder had both positive and negative effects on spruce growth, with negative effects predominating. Snowshoe hares ( Lepus americanus) in alder thickets further reduced height growth of spruce in the wetter site. We conclude that net effects of alder on white spruce, the late-successional dominant, were primarily inhibitory and indirect, with the mechanisms depending on initial site moisture. Our results highlight the importance of long-term research showing that small differences among initial replicate sites can cause divergence in successional trajectories, consistent with individualistic distributions of species and communities along environmental gradients. This divergence was detectable only decades later. [ABSTRACT FROM AUTHOR]

Summary: Plant‐associated mycobiomes in extreme habitats are understudied and poorly understood.We analysed Illumina‐generated ITS1 sequences from the needle mycobiome of white spruce (Picea glauca) at the northern treeline in Alaska (USA). Sequences were obtained from the same DNA that was used for tree genotyping. In the present study, fungal metabarcoding and tree microsatellite data were compared for the first time.In general, neighbouring trees shared more fungal taxa with each other than trees growing in further distance. Mycobiomes correlated strongly with phenological host traits and local habitat characteristics contrasting a dense forest stand with an open treeline site. Genetic similarity between trees did not influence fungal composition and no significant correlation existed between needle mycobiome and tree genotype.Our results suggest the pronounced influence of local habitat conditions and phenotypic tree traits on needle‐inhabiting fungi. By contrast, the tree genetic identity cannot be benchmarked as a dominant driver for needle‐inhabiting mycobiomes, at least not for white spruce in this extreme environment. [ABSTRACT FROM AUTHOR]

The prevailing hypothesis states that treeline positions are defined by the direct effects of cold temperatures on cell division and tree growth. Meanwhile, photosynthesis is thought to be relatively unrestricted in treeline trees. Support for this hypothesis comes from the global correlation between temperature and treeline position, the observation that many treelines have advanced in response to recent warming and the frequent use of increment cores from treeline trees to reconstruct past climates. However, studies of the physiology of trees at the Arctic treeline are rare, and this hypothesis remains largely untested., To improve our understanding of the relationships between temperature and performance of white spruce near the Arctic treeline, we made measurements of needle gas exchange, needle nutrition and soil nutrient availability over 2 years in three contrasting habitats: riverside terrace, hillslope forest and treeline. The sites had similar above-ground microclimates, but very different soil conditions. Soils were warm and dry on the terrace, cool and moist in the forest and cold and seasonally wet the treeline., Photosynthesis, needle nitrogen (N) concentration and soil N availability declined from the terrace to the forest to the treeline. Low N availability at the treeline was likely a consequence of limited microbial activity in the cold and seasonally wet soils. Soils at the treeline were colder than the terrace during the growing season and colder than the forest in winter, when the treeline maintains a shallow snowpack., Our results highlight the potential for an indirect effect of temperature on the growth of trees at the Arctic treeline and suggest that treeline responses to changes in climate may be more complex than previously thought. [ABSTRACT FROM AUTHOR]

Interactions between vegetation and fire have the potential to overshadow direct effects of climate change on fire regimes in boreal forests of North America. We develop methods to compare sediment-charcoal records with fire regimes simulated by an ecological model, ALFRESCO (Alaskan Frame-based Ecosystem Code) and apply these methods to evaluate potential causes of a mid-Holocene fire-regime shift in boreal forests of the south-central Brooks Range, Alaska, USA. Fire-return intervals (FRIs, number of years between fires) are estimated over the past 7000 calibrated 14C years (7-0 kyr BP [before present]) from short-term variations in charcoal accumulation rates (CHARs) at three lakes, and an index of area burned is inferred from long-term CHARs at these sites. ALFRESCO simulations of FRIs and annual area burned are based on prescribed vegetation and climate for 7-5 kyr BP and 5-0 kyr BP, inferred from pollen and stomata records and qualitative paleoclimate proxies. Two sets of experiments examine potential causes of increased burning between 7-5 and 5-0 kyr BP. (I) Static-vegetation scenarios: white spruce dominates with static mean temperature and total precipitation of the growing season for 7-0 kyr BP or with decreased temperature and/or increased precipitation for 5-0 kyr BP. (2) Changed-vegetation scenarios: black spruce dominates 5-0 kyr BP, with static temperature and precipitation or decreased temperature and/or increased precipitation. Median FRIs decreased between 7-5 and 5-0 kyr BP in empirical data and changed-vegetation scenarios but remained relatively constant in static-vegetation scenarios. Median empirical and simulated FRIs are not statistically different for 7-5 kyr BP and for two changed-vegetation scenarios (temperature decrease, precipitation increase) for 5-0 kyr BP. In these scenarios, cooler temperatures or increased precipitation dampened the effect of increased landscape flammability resulting from the increase in black spruce. CHAR records and all changed-vegetation scenarios indicate long-term increases in area burned between 7-5 and 5-0 kyr BP. The similarity of CHAR and ALFRESCO results demonstrates the compatibility of these independent data sets for investigating ecological mechanisms causing past fire-regime changes. The finding that vegetation flammability was a major driver of Holocene fire regimes is consistent with other investigations that suggest that landscape fuel characteristics will mediate the direct effects of future climate change on boreal fire regimes. [ABSTRACT FROM AUTHOR]

Local distributions of black spruce (Picea mariana) and white spruce (Picea glauca) are largely determined by edaphic and topographic factors in the interior of Alaska, with black spruce dominant on moist permafrost sites and white spruce dominant on drier upland sites. Given the recent evidence for climate warming and permafrost degradation, the distribution of white spruce is expected to expand, but the transition from black to white spruce may be dispersal limited: unlike the semi-serotinous black spruce, postfire regeneration of white spruce relies on seed dispersal from unburned areas. To determine the relative roles of dispersal, establishment, and growth in recruitment of white and black spruce, we studied postfire spruce regeneration in a 21-year-old burn across a white spruce-black spruce transition in the interior of Alaska. Although prefire spatial distributions of adults of the two species were well separated along the topographic sequence from upland to floodplain sites, the spatial distributions of recruits overlapped considerably. Even >700 m away from its seed source, white spruce sapling density on typical black spruce sites was high enough to form fully stocked stands. In contrast, black spruce regeneration was sparse on typical white spruce upland sites. Establishment rates of both species, estimated from a statistical model, were highest in mossy, wet depressions, which tended to have a thick residual postfire organic layer (∼10 cm). On all site types, height growth rates inferred from age-height relationships were comparable for recruits of both species. On typical black spruce sites ⩾300 m into the burn, white spruce was younger (and, therefore, shorter) than black spruce due to the timing of masting events following the fire. There was no indication that dispersal, establishment, or edaphic constraints on juvenile growth limit white spruce's capacity to invade typical black spruce stands during the recruitment stage in our study area. It is unlikely that white spruce recruits would persist to the adult stage if the permafrost returned to the original prefire levels during future postfire succession. However, if permafrost continues to degrade under climate warming, transition to a white spruce-dominated landscape could be rapid. [ABSTRACT FROM AUTHOR]

Investigates competition and microenvironmental modification in an Alaskan postfire boreal forest. Establishment of seedlings by white spruce and paper birch in patches dominated by horsetail and bluestem; Reduction of survivorship, growth and nitrogen accumulation in the presence of aboveground vegetation.

The treeline in Alaska is usually mapped as the limit of white spruce Picea glauca (Moench) Voss along the south slope of the Brooks Range and in western Alaska and as the limit of Sitka spruce Picea sitchensis (Bong.) Can. on Kodiak Island and in the Alaska Peninsula. In some localities black spruce Picea mariana (Mill.) B.S.P. and paper birch Bemia papyrifera Marsh, fonn components of the treeline community with white spruce. Of special interest are the groves of balsam poplar Populus balsamifera L. occurring on the north slope of the Brooks Range and to the west and southwest of the spruce treeline. There is some question as to whether trees survived in unglaciated portions of Alaska during the last glacial period. Spruce trees became prevalent about 10000 yr ago and may still be expanding to their climatic limits. In some areas of Alaska the treeline was more expanded during the Hypsithermal period than it is at present, but in other regions evidence for such an expansion is lacking. Treetine in some areas seems to have been stable for the last several centuries, but is has been expanding rapidly for the last 40 yr in entral and western Alaska and for a longer period on Kodiak Island. [ABSTRACT FROM AUTHOR]

Trees growing near the Arctic treeline have long been used to reconstruct past climates. However, recent studies have shown deterioration of historically strong positive correlations between air temperature and tree growth (known as "divergence"). Divergence has important implications for confidence in paleoclimate reconstructions and ecosystem–atmosphere carbon exchange. Studies in the Brooks Range of northern Alaska showed that white spruce in the west increased growth in response to late 20th century warming, whereas those in the east failed to show a growth increase. In an earlier study across four watersheds in the Brooks Range, we tested and rejected the hypothesis that divergence in the easternmost watershed reflects moisture limitation of growth. Here, using 16 sites distributed across the same four watersheds, we tested an alternative hypothesis, that greater nutrient limitation in the east may have impeded positive growth responses to warming. Climate comparison across the four Brooks Range study watersheds revealed that, although the easternmost watershed generally had a drier growing‐season climate, the most consistent difference was that winter air temperature and both winter and summer soil temperatures were much colder in the central and eastern watersheds. Soil nutrient availability, foliar nutrient concentrations, and tree growth were all generally lower in the central and eastern than in the western watersheds. Foliar phosphorus concentration was the best predictor of spatial variation in branch extension growth—a finding that is somewhat inconsistent with the theory that forest productivity on young, glacially derived soils should be strongly nitrogen limited. Experimental fertilization yielded the greatest growth increase in the eastern, an intermediate response in the central, and the smallest growth increase in the western watershed, generally mirroring trends in soil temperature, soil nutrient availability, foliar nutrient concentrations, and growth of control trees. Our results confirm that growth in the easternmost watershed is more nutrient limited and suggest that phosphorus limitation may be at least as important as nitrogen limitation of growth. We hypothesize that cold soil effects on tree access to nutrients might explain divergence in the eastern Brooks Range and elsewhere near the Arctic treeline, particularly in areas with cold winters and widespread permafrost. [ABSTRACT FROM AUTHOR]