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We investigated the genetic makeup of Lutz spruce, a natural hybrid between white and Sitka spruce on the Kenai Peninsula, Alaska. Microsatellites indicate 72% of individuals sampled had predominantly white spruce ancestry, whereas 14% had predominantly Sitka spruce ancestry; some individuals classified as white spruce had Sitka spruce plastid genotypes. As Picea mitochondria are maternally inherited and plastids are paternally inherited, it appears that white spruce was the ancestral seed parent of nearly all spruce on the western peninsula, whereas Sitka spruce alleles originated from pollen. Pollen records show that white spruce colonized the western peninsula ∼8500 YBP from glacial refugium, whereas Sitka spruce arrived on the eastern peninsula ∼4000 YBP after migrating up the Pacific coast. Our data suggest that Sitka spruce colonization west of the Kenai Mountains may have occurred not via seed dispersal but by long distance transport of wind-borne pollen and subsequent hybridization with established white spruce populations. Hybridization was an important mechanism that allowed Sitka spruce to expand the leading edge of its range in response to historical climate change. As the climate continues to warm, climate envelope modeling suggests Lutz spruce may ultimately displace white spruce on the western peninsula even as Sitka spruce is constrained to the eastern peninsula where it will continue to hybridize. [ABSTRACT FROM AUTHOR]

Climate change is increasing the frequency and intensity of drought events in many boreal forests. Trees are sessile organisms with a long generation time, which makes them vulnerable to fast climate change and hinders fast adaptations. Therefore, it is important to know how forests cope with drought stress and to explore the genetic basis of these reactions. We investigated three natural populations of white spruce (Picea glauca) in Alaska, located at one drought-limited and two cold-limited treelines with a paired plot design of one forest and one treeline plot. We obtained individual increment cores from 458 trees and climate data to assess dendrophenotypes, in particular the growth reaction to drought stress. To explore the genetic basis of these dendrophenotypes, we genotyped the individual trees at 3000 single nucleotide polymorphisms in candidate genes and performed genotype-phenotype association analysis using linear mixed models and Bayesian sparse linear mixed models. Growth reaction to drought stress differed in contrasting treeline populations. Therefore, the populations are likely to be unevenly affected by climate change. We identified 40 genes associated with dendrophenotypic traits that differed among the treeline populations. Most genes were identified in the drought-limited site, indicating comparatively strong selection pressure of drought-tolerant phenotypes. Contrasting patterns of drought-associated genes among sampled sites and in comparison to Canadian populations in a previous study suggest that drought adaptation acts on a local scale. Our results highlight genes that are associated with wood traits which in turn are critical for the establishment and persistence of future forests under climate change. [ABSTRACT FROM AUTHOR]

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]

Under climate change circumstances, increasing studies have reported the temporal instability of tree growth responses to climate, which poses a major challenge to linearly extrapolating past climate and future growth dynamics using tree‐ring data. Space‐for‐time substitution (SFTS) is a potential solution to this problem that is widely used in the dendrochronology field to project past or future temporal growth response trajectories from contemporary spatial patterns. However, the projected accuracy of the SFTS in the climate effects on tree growth remains uncertain. Here, we empirically test the SFTS method by comparing the effect of spatial and temporal climate variations on climate responses of white spruce (Picea glauca), which has a transcontinental range in North America. We first applied a response surface regression model to capture the variations in growth responses along the spatial climate gradients. The results showed that the relationships between growth and June temperature varied along spatial climate gradients in a predictable way. And their relationships varied mainly along with local temperate condition. Then, the projected correlation coefficients between growth and climate using SFTS were compared against the observed. We found that the growth response changes caused by spatial versus temporal climate variations showed opposite trends. Moreover, the projected correlation coefficients using the SFTS were significantly lower than the observed. This finding suggests that applying the SFTS to project the growth response of white spruce might lead to an overestimation of the degree of tree maladaptation in future climate scenarios. And the overestimation is likely to get weaker from Alaska and Yukon Territory in the west to Quebec in the east. Although this is only a case study of the SFTS method for projecting tree growth response, our findings suggest that direct application of the SFTS method may not be applicable to all regions and all tree species. [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]

The onset of lifetime reproduction is central to processes of plant adaptation and population changes. The relative contributions of size and age to the onset of reproduction are important to predicting that key event in individuals. To estimate respective size and age effects on first reproduction in a conifer, I observed white spruce (Picea glauca (Moench) Voss) trees in forest stands in Alaska, United States, over ranges of cumulative growth rates. Age was estimated by counting annular growth rings, size was indicated by tree height, and reproductive status was indicated by seed cones. Multiple logistic regression showed the relative contributions of tree height and age, adjusted for each other, to statistical predictions of reproductive maturity. Trees selected for having little competition for light first produced cones at heights between 1 and 5 m with little additional effect of age. Tree height contributed approximately 50% to regression predictions of reproductive maturity. In contrast, tree age contributed ≤1.5% and bore little relation to reproductive maturity other than as a component of growth rate. This is among the first reports to partial the effect sizes of tree size and age on the commencement of lifetime reproduction in a conifer. [ABSTRACT FROM AUTHOR]

Understanding interactions between competition and climate in relation to their effects on individual tree growth is crucial to the development of climate-sensitive growth models required for modelling boreal forest succession in a changing climate. We used data from permanent growth and yield sample plots in western Canada and Alaska to investigate the impact of competition within a regional gradient of climatic conditions for lodgepole pine (Pinus contorta Douglas ex Loudon), jack pine (Pinus banksiana Lamb.), trembling aspen (Populus tremuloides Michx.), balsam poplar (Populus balsamifera L.), white spruce (Picea glauca (Moench) Voss), and black spruce (Picea mariana (Mill.) Britton, Sterns & Poggenb.). We characterized the effects of competition (basal area of spruce–fir, deciduous, and pine trees larger than the focal tree) and climate (mean annual temperature and precipitation) and their interactions on basal area growth of individual trees using linear mixed-effects models. Our results indicated that intraspecific competition had stronger effects on growth than interspecific competition and climate. Moreover, significant interactions between intraspecific competition and climate suggest that an increase in intraspecific competition will lead to a reduction in tree growth for warmer regions (lodgepole pine, trembling aspen, balsam poplar, and white spruce) and wetter regions (jack pine). The manner in which interspecific competition altered tree growth responses to climate was variable, depending on tree species and competition type. These results indicate that the relationships between growth and climate may differ according to the degree of competition and the structure of the stand. [ABSTRACT FROM AUTHOR]

The negative growth response of North American boreal forest trees to warm summers is well documented and the constraint of competition on tree growth widely reported, but the potential interaction between climate and competition in the boreal forest is not well studied. Because competition may amplify or mute tree climate‐growth responses, understanding the role current forest structure plays in tree growth responses to climate is critical in assessing and managing future forest productivity in a warming climate. Using white spruce tree ring and carbon isotope data from a long‐term vegetation monitoring program in Denali National Park and Preserve, we investigated the hypotheses that (a) competition and site moisture characteristics mediate white spruce radial growth response to climate and (b) moisture limitation is the mechanism for reduced growth. We further examined the impact of large reproductive events (mast years) on white spruce radial growth and stomatal regulation. We found that competition and site moisture characteristics mediated white spruce climate‐growth response. The negative radial growth response to warm and dry early‐ to mid‐summer and dry late summer conditions intensified in high competition stands and in areas receiving high potential solar radiation. Discrimination against 13C was reduced in warm, dry summers and further diminished on south‐facing hillslopes and in high competition stands, but was unaffected by climate in open floodplain stands, supporting the hypothesis that competition for moisture limits growth. Finally, during mast years, we found a shift in current year's carbon resources from radial growth to reproduction, reduced 13C discrimination, and increased intrinsic water‐use efficiency. Our findings highlight the importance of temporally variable and confounded factors, such as forest structure and climate, on the observed climate‐growth response of white spruce. Thus, white spruce growth trends and productivity in a warming climate will likely depend on landscape position and current forest structure. Using tree ring and carbon isotopes data from Denali National Park and Preserve, we investigated whether competition and site moisture characteristics mediate white spruce radial growth response to climate. We found competition and solar radiation mediate the influence of climate on the radial growth. Our carbon isotope analysis suggests the mechanism behind these modified climate‐growth responses is increased competition for moisture in high basal area stands and dry sites. Overall, our results suggest dynamic growth responses to future climate change that are dependent on landscape position and stand competition and likely to result in feedbacks on forest structure. [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]

Aim A critical concern for boreal ecosystems centres on broad-scale responses to warming, i.e. declining growth and mortality, or enhanced growth and greater productivity. However, few studies have synthesized tree growth along biogeographic gradients to address this issue. This study develops a broader understanding of how growth of a dominant conifer has responded to recent warming near the western forest margin of Alaska. Location Alaska, United States. Methods Thirty Picea glauca sites in south-west Alaska (1216 trees ≥ 4 cm dbh) were evaluated for growth differences from the southern boreal forest to the western forest-tundra margin across low-elevation forests and woodlands, and altitudinal tree line. Regional climate records were used to evaluate (1) whether tree growth near western tree line showed greater sensitivity to temperature and/or precipitation than southern boreal sites, (2) if the climate-growth response varied through time, across ecoregions and ecosystems, with spruce beetle disturbance, and by tree age, and (3) if there was a temperature threshold that limited growth. Results Positive growth trends since the 1980s in many open stands were consistent with the predicted expansion of western and altitudinal tree line. However, growth levelled off with temperatures ≥ 13°C at all but altitudinal tree line. An increasingly positive effect of precipitation on growth occurred after 1985, particularly at boreal tree line and closed forest sites. Closed-canopy forests showed lower rates of growth, greater spruce beetle activity and less potential for resiliency to warming than other ecosystem types. Main Conclusions Warming has led to markedly different growth responses according to ecosystem type and ecoregion near the forest-tundra margin. Altitudinal tree line sites showed consistently positive growth with warmer temperatures in recent decades, whereas low-elevation forests had reduced growth and greater beetle mortality. Strong positive correlations between growth and summer-fall precipitation since the mid-1980s suggest that precipitation will become an increasingly important factor with further warming. [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 late-Holocene shift from Picea glauca (white spruce) to Picea mariana (black spruce) forests marked the establishment of modern boreal forests in Alaska. To understand the patterns and drivers of this vegetational change and the associated late-Holocene environmental dynamics, we analyzed radiocarbon-dated sediments from Grizzly Lake for chironomids, diatoms, pollen, macrofossils, charcoal, element composition, particle size, and magnetic properties for the period 4100–1800 cal BP. Chironomid assemblages reveal two episodes of decreased July temperature, at ca. 3300–3150 (ca −1 °C) and 2900–2550 cal BP (ca −2 °C). These episodes coincided with climate change elsewhere in the Northern Hemisphere, atmospheric reorganization, and low solar activity. Diatom-inferred lake levels dropped by ca. 5 m at 3200 cal BP, suggesting dry conditions during the period 3200–1800 cal BP. P. glauca declined and P. mariana expanded at ca. 3200 cal BP; this vegetational change was linked to diatom-inferred low lake levels and thus decreased moisture availability. Forest cover declined at 3300–3100, 2800–2500 and 2300–2100 cal BP and soil erosion as inferred from increased values of Al, K, Si, Ti, and Ca intensified, when solar irradiance was low. Plant taxa adapted to disturbance and cold climate (e.g. Alnus viridis , shrub Betula , Epilobium ) expanded during these periods of reduced forest cover. This open vegetation type was associated with high fire activity that peaked at 2800 cal BP, when climatic conditions were particularly cold and dry. Forest recovery lagged behind subsequent climate warming (≤+3 °C) by ca. 75–225 years. Our multiproxy data set suggests that P. glauca was dominant under warm-moist climatic conditions, whereas P. mariana prevailed under cold-dry and warm-dry conditions. This pattern implies that climatic warming, as anticipated for this century, may promote P. glauca expansions, if moisture availability will be sufficiently high, while P. mariana may expand under dry conditions, possibly exacerbating climate impacts on the fire regime. [ABSTRACT FROM AUTHOR]

Early establishment of competing vegetation often presents an obstacle to the success of planted white spruce ( Picea glauca (Moench) Voss) seedlings. We followed growth and development of white spruce and associated vegetation for up to 17 years in Alaska's boreal forests to quantify roles of overtopping plant cover in suppressing conifers. The three study areas represented a range of site conditions of varying productivity and species of competing cover, different site preparation and release treatments, and different bare-root and container white spruce stock types. Herbaceous overtopping peaked early after planting and decreased as white spruce were able to outgrow competitors. Overtopping by shrubs and hardwoods, especially aspen ( Populus tremuloides Michx.) and resin birch ( Betula neoalaskana Sarg.) peaked somewhat later than herbaceous overtopping and decreased over time for most sites and treatments. In a model that combined all sites, vegetation management treatments, and years, overtopping and previous year's volume explained approximately 85% of the variation in volume growth. Increasing the size of planting stock helped reduce overtopping, hence suppression, even in treatments dominated by hardwood species. Results suggested that control of overtopping was essential for maximum growth and long-term or increasing levels of overtopping severely suppressed white spruce seedling growth. [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]

White spruce (Picea glauca [Moench] Voss) is a valuable commercial species found in interior and southceniral Alaska. Numerous regional and local volume tables or equations exist; however, no statewide model exists or has been tested for accuracy. There is a demand for an accurate model to determine the cubic-foot volume of white spruce trees in Alaska. Multiple models were developed for white spruce to estimate total and merchantable cubic-foot volume to a 2-, 4-, and 6-in, top. These multiple-entry (diameter and height) models were developed for both inside and outside bark volume from a 6-in, stump. The models were tested on a regional basis at various geographic locations and were shown to be highly accurate. The Alaska models chosen have R2 at or near 0.99 and mean square error from 0 to 0.16 for all models. These models are shown to be superior to other white spruce models in Alaska. [ABSTRACT FROM AUTHOR]

Ongoing warming at high latitudes is expected to lead to large changes in the structure and function of boreal forests. Our objective in this research is to determine the climatic controls over the growth of white spruce ( Picea glauca (Moench) Voss) at the warmest driest margins of its range in interior Alaska. We then use those relationships to determine the climate variables most likely to limit future growth. We collected tree cores from white spruce trees growing on steep, south-facing river bluffs at five sites in interior Alaska, and analyzed the relationship between ring widths and climate using boosted regression trees. Precipitation and temperature of the previous growing season are important controls over growth at most sites: trees grow best in the coolest, wettest years. We identify clear thresholds in growth response to a number of variables, including both temperature and precipitation variables. General circulation model (GCM) projections of future climate in this region suggest that optimum climatic conditions for white spruce growth will become increasingly rare in the future. This is likely to cause short-term declines in productivity and, over the longer term, probably lead to a contraction of white spruce to the cooler, moister parts of its range in Alaska. [ABSTRACT FROM AUTHOR]

This paper calibrates climate controls over radial growth of floodplain white spruce ( (Moench) Voss) and examines whether growth in these populations responds similarly to climate as upland trees in Interior Alaska. Floodplain white spruce trees hold previously unrecognized potential for long-term climate reconstruction because they are the source of driftwood that becomes frozen in coastal deposits, where archeological timbers and beach logs represent well-preserved datable material. We compared ring width chronologies for 135 trees in six stands on the Yukon Flats and Tanana River with temperature and precipitation at Fairbanks from 1912-2001. Our sample contains a stable common signal representing a strong negative relationship between summer temperature and tree growth. We developed a floodplain temperature index (FPTI), which explains half of the variability of the composite chronology, and a supplemental precipitation index (SPI) based on correlation of monthly precipitation with the residual of the temperature-based prediction of growth. We then combined FPTI and SPI into a climate favorability index (CFI) in which above-normal precipitation partially compensates for temperature-induced drought reduction of growth and vice versa. CFI and growth have been particularly low since 1969. Our results provide a basis for building longer chronologies based on archeological wood and for projecting future growth. [ABSTRACT FROM AUTHOR]

To identify suitable methods for reforestation, we evaluated the interacting effects of past disturbance, stock types, and site preparation treatments on white spruce ( Picea glauca (Moench) Voss) seedling survival and growth across a range of sites in Alaska. Replicated experiments were established in five regions. At each site, two complete installations differed in time since disturbance: 'new' units were harvested immediately before spring planting and 'old' units were harvested at least 3 years before planting. We compared mechanical scarification before planting, broadcast herbicide application during the fall before planting, and no site preparation with 1-year-old container-grown seedlings from two sources, 2-year-old bare-root transplants from two sources, and 3-year-old bare-root transplants. Seedlings were followed for 11 years on most sites. Based on meta-analyses, seedling survival increased 10% with herbicide application and 15% with mechanical scarification compared with no site preparation. Scarification and herbicide application increased seedling height by about 28% and 35%, respectively, and increased seedling volume by about 86% and 195%, respectively, compared with no site preparation. Soil temperature did not differ among site preparation methods after the first 7 years. Results suggest that white spruce stands may be successfully restored through a combination of vegetation control and use of quality planting stock. [ABSTRACT FROM AUTHOR]

Although a recent study of white spruce using chloroplast DNA uncovered the presence of a glacial refuge in Alaska, chloroplast failed to provide information on the number or specific localities of refugia. Recent studies have demonstrated the utility of nuclear microsatellites to refine insights into postglacial histories. The greater relative rate of mutation may allow finer scale resolution of historic dynamics, including the number, location, and sizes of refugia. Genetic data were acquired from screening 6 microsatellite loci on approximately 14 trees from each of 22 populations located across the central and western boreal forests of Canada and Alaska. Our studies combining microsatellites with Bayesian analyses of population structure in white spruce support the phylogeographic patterns uncovered using chloroplast, separating Alaskan from non-Alaskan regions. Results also support the idea that north-central Alaska served as a glacial refugium during the last glacial maximum. Additionally, the relationship between the degree of genetic differentiation and geographic distance indicated that gene flow played a more important role in structuring non-Alaskan populations, whereas drift played a more important role in structuring Alaskan populations (RST’s for non-Alaskan populations 0.029 ± 0.007 and 0.083 ± 0.012 for Alaskan populations). Microsatellites also substantiate the bidirectional patterns of gene flow previously uncovered using chloroplast DNA but indicate much greater movement and mixing. Results from our Bayesian analyses also suggest the existence of additional cryptic refugia. However, the locations have been obscured by high gene flow (RST averaging 0.057 ± 0.004). [ABSTRACT FROM PUBLISHER]

This paper integrates dendrochronological, demographic, and experimental perspectives to improve understanding of the response of white spruce (Picea glauca (Moench) Voss) tree growth to climatic variability in interior Alaska. The dendrochronological analyses indicate that climate warming has led to widespread declines in white spruce growth throughout interior Alaska that have become more prevalent during the 20th century. Similarly, demographic studies show that white spruce tree growth is substantially limited by soil moisture availability in both mid- and late-successional stands. Interannual variability in tree growth among stands within a landscape exhibits greater synchrony than does growth of trees that occupy different landscapes, which agrees with dendrochronological findings that the responses depend on landscape position and prevailing climate. In contrast, the results from 18 years of a summer moisture limitation experiment showed that growth in midsuccessional upland stands was unaffected by moisture limitation and that moisture limitation decreased white spruce growth in floodplain stands where it was expected that growth would be less vulnerable because of tree access to river water. Taken together, the evidence from the different perspectives analyzed in this study clearly indicates that white spruce tree growth in interior Alaska is vulnerable to the effects of warming on plant water balance. [ABSTRACT FROM AUTHOR]

Most wilderness areas still lack accurate distribution information on tree species. We met this need with a predictive GIS modeling approach, using freely available digital data and computer programs to efficiently obtain high-quality species distribution maps. Here we present a digital map with the predicted distribution of white spruce ( Picea glauca) in Alaska (4 km resolution, accuracy over 90%). Our presented concept represents a role-model for predicting tree species distribution for remote areas world-wide. Although this model intends to be accurate in making predictions rather than to give detailed biological mechanistic explanations, it can also be used as a baseline for further research and testable hypothesis on the importance of the environmental variables used to build a generalizable model. Further, we emphasize that work like presented here is a pre-condition for assessing human impacts and impacts of climate change on species distribution in a quantitative and transparent fashion, allowing for improved sustainable decision-making world-wide. [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]

We describe a new tree-ring width data set of 14 white spruce chronologies for the Seward Peninsula (SP), Alaska, based on living and subfossil wood dating from 1358 to 2001AD. A composite chronology derived from these data correlates positively and significantly with summer temperatures at Nome from 1910 to 1970, after which there is some loss of positive temperature response. There is inferred cooling during periods within the Little Ice Age (LIA) from the early to middle 1600s and late 1700s to middle 1800s; and warming from the middle 1600s to early 1700s. We also present a larger composite data set covering 978-2001AD, utilizing the SP ring-width data in combination with archaeological wood measurements and other recent collections from northwestern Alaska. The Regional Curve Standardization (RCS) method was employed to maximize potential low-frequency information in this data set. The RCS chronology shows intervals of persistent above-average growth around the time of the Medieval Warm Period (MWP) early in the millennium, which are comparable to growth levels in recent centuries. There is a more sustained cold interval during the LIA inferred from the RCS record as compared to the SP ring-width series. The chronologies correlate significantly with Bering and Chukchi Sea sea surface temperatures and with the Pacific Decadal Oscillation index. These atmosphere-ocean linkages probably account for the differences between these records and large-scale reconstructions of Arctic and Northern Hemisphere temperatures based largely on continental interior proxy data. [ABSTRACT FROM AUTHOR]

Northern and high-latitude alpine treelines are generally thought to be limited by available warmth. Most studies of tree-growth–climate interaction at treeline as well as climate reconstructions using dendrochronology report positive growth response of treeline trees to warmer temperatures. However, population-wide responses of treeline trees to climate remain largely unexamined. We systematically sampled 1558 white spruce at 13 treeline sites in the Brooks Range and Alaska Range. Our findings of both positive and negative growth responses to climate warming at treeline challenge the widespread assumption that arctic treeline trees grow better with warming climate. High mean temperatures in July decreased the growth of 40% of white spruce at treeline areas in Alaska, whereas warm springs enhance growth of additional 36% of trees and 24% show no significant correlation with climate. Even though these opposing growth responses are present in all sampled sites, their relative proportion varies between sites and there is no overall clear relationship between growth response and landscape position within a site. Growth increases and decreases appear in our sample above specific temperature index values (temperature thresholds), which occurred more frequently in the late 20th century. Contrary to previous findings, temperature explained more variability in radial growth after 1950. Without accounting for these opposite responses and temperature thresholds, climate reconstructions based on ring width will miscalibrate past climate, and biogeochemical and dynamic vegetation models will overestimate carbon uptake and treeline advance under future warming scenarios. [ABSTRACT FROM AUTHOR]

Maximum latewood density and δ 13C discrimination of Interior Alaska white spruce were used to reconstruct summer (May through August) temperature at Fairbanks for the period 1800–1996, one of the first high-resolution reconstructions for this region. This combination of latewood density and δ 13C discrimination explains 59.9% of the variance in summer temperature during the period of record 1906–1996. The 200-yr. reconstruction is characterized by 7 decadal-scale regimes. Regime changes are indicated at 1816, 1834, 1879, 1916, 1937, and 1974, are abrupt, and appear to be the result of synoptic scale climate changes. The mean of summer temperature for the period of reconstruction (1800–1996) was 13.49 °C. During the period of instrument record (1903–1996) the mean of summer temperature was 13.31 °C for both the reconstruction and the recorded data. The coldest interval was 1916–1937 (12.62 ° C) and the warmest was 1974–1996 (14.23 °C) for the recorded data. The reconstruction differs from records of northern hemisphere temperatures over this period, especially because of Interior Alaska warm periods reconstructed from 1834 to 1851 (14.24 °C) and from 1862 to 1879 (14.19 °C) and because of the cool period in the early part of the 20th century (1917–1974). We show additional tree ring data that support our reconstruction of these warm periods. Alternate hypotheses involving autogenic effect of tree growth on the site, altered tree sensitivity, or novel combinations of temperature and precipitation were explored and while they cannot be ruled out as contributors to the anomalously warm 19th century reconstruction, they were not supported by available data. White spruce radial growth is highly correlated with reconstructed summer temperature, and temperature appears to be a reliable index of carbon uptake in this system. [ABSTRACT FROM AUTHOR]

Presents multi-proxy tree-ring data from twenty productive stands of white spruce in the interior of Alaska. Evidence that radial tree growth has decreased with increasing temperature; Effect of temperature-induced drought; Suggestion that drought stress may have been an important factor limiting carbon intake in North American forests; Implications for the future capacity of northern latitudes to sequester carbon.

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]

Needle hardiness of introduced yellow pine, Pinus banksiana Lamb., lodgepole pine, P. contorta Dougl. and native white spruce, Picea glauca (Moench) Voss, were assessed by the effective prefreezing temperature method. Yellow pine needles were less hardy than lodgepole pine or white spruce needles in Alaska on each date measured. Although hardiness decreased in springtime in all species, decreases in hardiness in yellow pine began before temperatures were above -20°C, apparently in response to day length, while decreases in hardiness in lodgepole pine and white spruce began only when mean temperatures were above 0°C. Hardiness was increased by decreasing the water content of yellow pine and spruce needles. However, only the latter increased its field hardiness by decreased water contents, and only to a small degree. Large decreases in phospholipid occurred during the dehardening period, indicating the presence of major membrane-associated changes. However, changes in hardiness did not closely parallel those in phospholipid; hardiness decreased before phospholipid did in spruce and after phospholipid did in lodgepole pine. In yellow pine, changes in hardiness were more closely related to changes in phospholipid content. Decreases in phospholipid appeared to be correlated with the day length in all species. [ABSTRACT FROM AUTHOR]

A white spruce seedling, Picea glauca (Moench) Voss, was found at the northern edge of the Brooks Range in Alaska, more than 50 km north of the latitudinal tree line. The seedling, 19 cm tall and about nine years old, was growing at the side of the Dalton Highway to Prudhoe Bay. It most likely sprouted from a seed transported across the Brooks Range on a vehicle and has survived on the well-drained gravel road berm, where site conditions are more favorable for germination and survival than in the surrounding tundra. This spruce has survived for about a decade under current climatic conditions. Even with a warming climate, natural seed dispersal is severely hampered by the rugged topographic barrier of the Brooks Range. Considering the amount of vehicle traffic on the Dalton Highway, however, it is likely that more pioneering spruce seedlings will turn up along this corridor. Once over the Brooks Range, a spruce population can potentially develop and expand. [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]