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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]

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]

Herbivores can modify the rate of shrub and treeline advance. Both direct and indirect effects of herbivory may simultaneously interact to affect the growth rates of plants at this ecotone. We investigated the effect of snowshoe hare herbivory on the height of white spruce at two treeline locations in Alaska, USA. White spruce is expanding its distribution both upwards in elevation and northward in latitude because of climate warming, and snowshoe hares are already present in areas likely to be colonized by spruce. We hypothesized that herbivory would result in browsed individuals having reduced height, suggesting herbivory is a direct, negative effect on spruce treeline advance. We found an interactive effect between browsing history and spruce age. When young (under 30 years old), individuals that were browsed tended to be taller than unbrowsed individuals. However, older seedlings (over 30 years old) that had been browsed were shorter than unbrowsed individuals of the same age. Hares suppress faster growing individuals that are initially taller by preferentially browsing them as they emerge above the winter snowpack. This reduced height, in combination with increased mortality associated with browsing, is predicted to slow the advance of both latitudinal and altitudinal treeline expansions and alter the structure of treeline forests. [ABSTRACT FROM AUTHOR]

Post-harvest regeneration failure of white spruce ( Picea glauca Moench [Voss]), has led to concerns of “de-coniferization” on productive site in the Alaskan boreal forest. Forest management in the region sought historically to increase spruce composition after harvest through silvicultural practices such as site preparation and assisted regeneration; however, successful reforestation requires the effects of these practices to persist over time and control non-target tree species. In order to identify the enduring effects of silvicultural regeneration practices, we sampled a large (26.7 ha) white spruce regeneration trial established immediately following a stand-replacing wildfire and subsequent salvage harvest in a productive upland forest. The original regeneration treatments followed a split-split plot experimental design on two landform types (LF), four ground scarification treatments (GST) plus a non-scarified control, and five artificial white spruce regeneration treatments (WSRT) plus a natural seedfall control (Densmore et al., 1999). Here we analyze the total biomass, stand density, and basal area for all tree species within each of the regeneration treatments 28 years post-establishment, and calculate seed dispersal distances. Our results show that compared to natural seedfall control plots, white spruce basal area was six times greater in planted seedling plots, and white spruce stem density (dbh ≥ 1.0 cm) was nearly three times greater in broadcast seeding plots. White spruce stem density from natural seedfall averaged 944 stems ha −1 , but was dependent on both topographic position and distance to wind-dispersed seed sources. Our results also indicate that GST had few significant effects on white spruce basal area or stem density. However, scarification nearly doubled Alaska birch ( Betula neoalaskana Sarg.) stem density and basal area compared to non-scarified control plots. Planted white spruce plots supported 19% less birch basal area, except in the most intensive scarification treatments in which birch basal area did not differ. Intensive scarification reduced quaking aspen ( Populus tremuloides Michx.) basal area by half on slope plots. Our results demonstrate that early regeneration practices profoundly influence stand development beyond the stem initiation stage, but pre-fire stand type, post-fire configuration of unburned seed sources, and topographical variation play a mediating role in determining species assemblages and competitive relationships. A fire-killed stand must be considered within its ecological and landscape context to determine the probable success of a management action such as salvage and tree regeneration. [ABSTRACT FROM AUTHOR]

Treelines in Alaska are advancing in elevation and latitude because of climate warming, which is expanding the habitat available for boreal wildlife species, including snowshoe hares ( Lepus americanus). Snowshoe hares are already present in tall shrub communities beyond treeline and are the main browser of white spruce ( Picea glauca), the dominant tree species at treeline in Alaska. We investigated the processes involved in a 'snowshoe hare filter' to white spruce establishment near treeline in Denali National Park, Alaska, USA. We modeled the pattern of spruce establishment from 1970 to 2009 and found that fewer spruce established during periods of high hare abundance. Multiple factors interact to influence browsing of spruce, including the hare cycle, snow depth and the characteristics of surrounding vegetation. Hares are abundant at treeline and may exclude spruce from otherwise optimal establishment sites, particularly floodplain locations with closed shrub canopies. The expansion of white spruce treeline in response to warming climate will be strongly modified by the spatial and temporal dynamics of the snowshoe hare filter. [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]

Herbivores have the capacity to modify plant community composition and ecosystem structure and function via browsing. For example, moose and snowshoe hare facilitate succession in Alaska’s boreal forest by preferentially browsing early successional species over late successional conifers. Snowshoe hares also eat conifers, including white spruce, and this browsing may affect the pattern of spruce establishment over time. We measured over 800 spruce at 18 locations along the Tanana River floodplain in interior Alaska, USA and demonstrated that the proportion of spruce browsed annually positively correlates with annual hare abundance. Nearly all seedlings sampled had been browsed. Further, we modeled the pattern of spruce establishment over the last 40 years and found that hare abundance, growing season temperature, early season snow depth, and flooding explain the majority of this pattern. This model demonstrated that less spruce established during periods of high hare abundance than during periods of low hare abundance. The extensive browsing of white spruce that occurs during periods of high hare abundance may further compound the negative effects of climate warming on spruce recruitment in these floodplain forests. [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]

Increasing temperatures have resulted in reduced growth and increased tree mortality across large areas of western North American forests. We use tree-ring isotope chronologies (δ13C and δ18O) from live and dead trees from four locations in south-central Alaska, USA, to test whether white spruce trees killed by recent spruce beetle ( Dendroctonus rufipennis Kirby) outbreaks showed evidence of drought stress prior to death. Trees that were killed were more sensitive to spring/summer temperature and/or precipitation than trees that survived. At two of our sites, we found greater correlations between the δ13C and δ18O chronologies and spring/summer temperatures in dead trees than in live trees, suggesting that trees that are more sensitive to temperature-induced drought stress are more likely to be killed. At one site, the difference between δ13C in live and dead trees was related to winter/spring precipitation, with dead trees showing stronger correlations between δ13C and precipitation, again suggesting increased water stress in dead trees. At all sites where δ18O was measured, δ18O chronologies showed the greatest difference in climate response between live and dead groups, with δ18O in live trees correlating more strongly with late winter precipitation than dead trees. Our results indicate that sites where trees are already sensitive to warm or dry early growing-season conditions experienced the most beetle-kill, which has important implications for forecasting future mortality events in Alaska. [ABSTRACT FROM AUTHOR]

Herbivores transform landscapes and affect succession via selective foraging that alters vegetation composition. In the boreal forest, mammalian herbivores, mainly moose, facilitate a shift toward the dominance of heavily defended species over time, such as white spruce. The effects of moose herbivory are intensified by the browsing of snowshoe hares. However, unlike moose, snowshoe hares also browse seedlings of white spruce. We quantified herbivory by snowshoe hares on white spruce along the Tanana River, interior Alaska, and assessed the effects on white spruce demography via two different herbivore exclosure experiments. We hypothesized that both experiments would show reduced plant density and height growth in the presence of hares. We found evidence of extensive browsing of white spruce seedlings by snowshoe hares, which negatively affected white spruce establishment along the Tanana River floodplain. Previous research has indicated that hares facilitate floodplain succession in the forward direction. However, hare herbivory can also retard succession by depressing white spruce establishment and increase the time individual trees spend in the understory. This stage-dependent effect on successional dynamics via a unique temporal variation in snowshoe hare herbivory may contribute to alternative successional trajectories in the boreal forest. [ABSTRACT FROM AUTHOR]

Since the mid 1970s, Interior Alaska white spruce trees experienced markedly lower growth than during the 19th and early 20th centuries. This observation raises the question of forest persistence on certain sites of lowland central and eastern Alaska. We analyzed white spruce growth across a 36-site network (540 trees) on three major river floodplains in boreal Alaska along a longitudinal gradient from eastern Interior to the southwest tree limit to test for the presence of tree growth patterns and climate sensitivities. Chronologies are compared for temperature sensitivity at both stand and individual tree levels, using data from Bethel, McGrath, and Fairbanks NWS stations during the common period of 1952–2001. Cross-dated stand-level ring width chronologies indicate three regions of common signal in tree growth across the gradient. Temperature sensitivity of stand- and individual-tree chronologies is spatially coherent. Most downriver chronologies correlate positively with spring mean monthly temperatures (MMT) at Bethel, mid- and upriver chronologies correlate negatively with MMT of May and previous year July at either McGrath or Fairbanks, and an area in between is a mixed population of positive and negative responders. In downriver positive responders, recent increases from suboptimal cool temperatures accelerated tree growth, while in mid- and upriver negative responders, recent increases from optimal or above-optimal temperatures decreased growth. Fairbanks negative responders are also negatively correlated with a 200-yr index of recorded and reconstructed Fairbanks summer temperatures, and recent sustained record high summer temperatures are associated with the lowest relative growth. Until the 1940s, absolute growth rate of negative responders was greater than positive responders, but from the 1970s the positive responders grew more. These results explain why northern ring width samples can display opposite temperature sensitivity and contribute to understanding recent “divergence” or loss of temperature sensitivity in a changing climate. We find that July MMT and annual precipitation at Fairbanks are now outside the limits that previously characterized the North American distribution of white spruce, and are near the reported physiological limits of the species. Our results of the spatial and temporal change of white spruce temperature sensitivity provide strong empirical evidence of previously proposed early stage biome shift in boreal Alaska due to clear climatic causes. Already, western Alaska, previously extending to tree limit, has become the optimum climate region for the species. With modest additional warming widespread tree death will be unavoidable on warmer lowland interior sites, where persistence of white spruce is unlikely. [ABSTRACT FROM AUTHOR]

Mast-seeding conifers such as Picea glauca exhibit synchronous production of large seed crops over wide areas, suggesting climate factors as possible triggers for episodic high seed production. Rapidly changing climatic conditions may thus alter the tempo and spatial pattern of masting of dominant species with potentially far-reaching ecological consequences. Understanding the future reproductive dynamics of ecosystems including boreal forests, which may be dominated by mast-seeding species, requires identifying the specific cues that drive variation in reproductive output across landscape gradients and among years. Here we used annual data collected at three sites spanning an elevation gradient in interior Alaska, USA between 1986 and 2011 to produce the first quantitative models for climate controls over both seedfall and seed viability in P. glauca, a dominant boreal conifer. We identified positive associations between seedfall and increased summer precipitation and decreased summer warmth in all years except for the year prior to seedfall. Seed viability showed a contrasting response, with positive correlations to summer warmth in all years analyzed except for one, and an especially positive response to warm and wet conditions in the seedfall year. Finally, we found substantial reductions in reproductive potential of P. glauca at high elevation due to significantly reduced seed viability there. Our results indicate that major variation in the reproductive potential of this species may occur in different landscape positions in response to warming, with decreasing reproductive success in areas prone to drought stress contrasted with increasing success in higher elevation areas currently limited by cool summer temperatures. [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]

Throughout interior Alaska (USA), a gradual warming trend in mean monthly temperatures occurred over the last few decades (∼2-4°°C). The accompanying increases in woody vegetation at many alpine treeline (hereafter treeline) locations provided an opportunity to examine how biotic and abiotic local site conditions interact to control tree establishment patterns during warming. We devised a landscape ecological approach to investigate these relationships at an undisturbed treeline in the Alaska Range. We identified treeline changes between 1953 (aerial photography) and 2005 (satellite imagery) in a geographic information system (GIS) and linked them with corresponding local site conditions derived from digital terrain data, ancillary climate data, and distance to 1953 trees. Logistic regressions enabled us to rank the importance of local site conditions in controlling tree establishment. We discovered a spatial transition in the importance of tree establishment controls. The biotic variable (proximity to 1953 trees) was the most important tree establishment predictor below the upper tree limit, providing evidence of response lags with the abiotic setting and suggesting that tree establishment is rarely in equilibrium with the physical environment or responding directly to warming. Elevation and winter sun exposure were important predictors of tree establishment at the upper tree limit, but proximity to trees persisted as an important tertiary predictor, indicating that tree establishment may achieve equilibrium with the physical environment. However, even here, influences from the biotic variable may obscure unequivocal correlations with the abiotic setting (including temperature). Future treeline expansion will likely be patchy and challenging to predict without considering the spatial variability of influences from biotic and abiotic local site conditions. [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]

Abstract: White spruce (Picea glauca (Moench) Voss) is a dominant species in late-successional ecosystems along the Tanana River, interior Alaska, and the most important commercial timber species in these boreal floodplain forests. Whereas white spruce commonly seed in on young terraces in early primary succession, the species does not become a conspicuous component of the vegetation until after 60–80 years. To address what abiotic and/or biotic factors may explain the paucity of spruce in earlier stages of succession, we examined germination and growth of planted white spruce seedlings across an environmental gradient that included variation in soil physico-chemical properties in the presence and absence of mammal browsing. The effect of browsing pressure over the first four years after planting was most noticeable on the older terraces. Likewise, direct effects of hare browsing on spruce seedling mortality were only manifested at the oldest sites. Spruce germination and survival was inversely proportional to soil cation concentrations, which was largely controlled by temperature-driven evapotranspiration. High light intensities and high air temperatures significantly reduced seedling growth, whereas variation in soil moisture only explained a significant amount of variation in seedling survival. Temperatures within the needle clusters on terminal shoots reached values that adversely affect photosynthesis (>32°C) on multiple occasions over the growing season. We conclude that the direct (temperature) and indirect (soil chemistry) effects of high insolation are major factors constraining spruce performance on early successional terraces, and that these effects can be significantly exacerbated by mammal browsing on associated deciduous vegetation. [Copyright &y& Elsevier]

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]

Abstract: Between 1987 and 2000, a spruce beetle (Dendroctonus rufipennis) outbreak infested 1.19 million ha of spruce (Picea spp.) forests in Alaska, killing most of the large diameter trees. We evaluated whether these forests would recover to their pre-outbreak density, and determined the site conditions on which spruce germinated and survived following the spruce beetle outbreak in forests of the Anchor River watershed, Kenai Peninsula, Alaska. White spruce (Picea glauca) and Lutz''s spruce (Picea × lutzii), a hybrid between white and Sitka spruce (Picea sitchensis), dominate the study area. We measured the pre- and post-outbreak density of spruce in 108 3m×80m plots across the study area by recording all live trees and all dead trees >1.5m tall in each plot. To determine the fine scale site conditions on which spruce germinated and survived, we measured ground surface and substrate characteristics within 20cm circular plots around a subset of post-outbreak spruce seedlings. The density of post-outbreak spruce (855/ha) was adequate to restock the stands to their pre-outbreak densities (643/ha) for trees >1.5m tall. We could not accurately estimate recovery for pre-outbreak spruce seedlings because dead seedlings may have decayed in the 5–18 years since the beetle outbreak occurred. At the fine scale, spruce that germinated post-outbreak grew on a wide variety of substrates including downed log, stump, mesic organic mat, peat, hummocks and mineral soil. They exhibited a strong preference for downed logs (53%) and stumps (4%), and most (91%) of the downed logs and stumps that spruce rooted on were heavily decayed. This preference for heavily decayed logs and stumps was especially evident given that their combined mean cover was only 2% in the 3m×80m plots. Within the 3m×80m plots, spruce seedling survival was negatively correlated with bluejoint (Calamagrostis canadensis) litter cover. [Copyright &y& Elsevier]

Abstract: From 1989 to 2003, a widespread outbreak of spruce beetles (Dendroctonus rufipennis) in the Copper River Basin, Alaska, infested over 275,000ha of forests in the region. During 1997 and 1998, we measured forest vegetation structure and composition on one hundred and thirty-six 20-m×20-m plots to assess both the immediate stand and landscape level effects of the spruce beetle infestation. A photo-interpreted vegetation and infestation map was produced using color-infrared aerial photography at a scale of 1:40,000. We used linear regression to quantify the effects of the outbreak on forest structure and composition. White spruce (Picea glauca) canopy cover and basal area of medium-to-large trees [≥15cm diameter-at-breast height (1.3m, dbh)] were reduced linearly as the number of trees attacked by spruce beetles increased. Black spruce (Picea mariana) and small diameter white spruce (<15cm dbh) were infrequently attacked and killed by spruce beetles. This selective attack of mature white spruce reduced structural complexity of stands to earlier stages of succession and caused mixed tree species stands to lose their white spruce and become more homogeneous in overstory composition. Using the resulting regressions, we developed a transition matrix to describe changes in vegetation types under varying levels of spruce beetle infestations, and applied the model to the vegetation map. Prior to the outbreak, our study area was composed primarily of stands of mixed white and black spruce (29% of area) and pure white spruce (25%). However, the selective attack on white spruce caused many of these stands to transition to black spruce dominated stands (73% increase in area) or shrublands (26% increase in area). The post-infestation landscape was thereby composed of more even distributions of shrubland and white, black, and mixed spruce communities (17–22% of study area). Changes in the cover and composition of understory vegetation were less evident in this study. However, stands with the highest mortality due to spruce beetles had the lowest densities of white spruce seedlings suggesting a longer forest regeneration time without an increase in seedling germination, growth, or survival. [Copyright &y& Elsevier]

Abstract: The northern treeline is generally limited by available warmth. However, in recent years, more and more studies have identified drought stress as an additional limiting factor for tree growth in northern boreal forests and at treelines. Three growth responses to warming have been identified: increase in growth, decrease in growth, and nonsignificant correlation of tree growth with climate. Here we investigate the effect of drought stress on radial growth of white spruce at northern treelines along a longitudinal gradient spanning the entire Brooks Range in Alaska. We systematically sampled 687 white spruce at seven treeline sites. Where possible, we sampled three site types at a given site: high-density forest, low-density forest, and floodplain forest. We investigated the relationship of site and site type to tree growth responses. In the western part of our study area, we found very high numbers of trees responding with increase in growth to recent warming; while in the eastern part, trees responding with decrease in growth to recent warming are predominant. Within a given site, more trees reacting positively to warming grow on site types characterized by low tree density. These patterns coincide with precipitation decreases from west to east and local water availability gradients, therefore pointing to drought stress as the controlling factor for the distribution of trees responding with increase or decrease in growth to recent warming. Compared to 20th century climate, we project a 25–50% basal area (BA) increase in the western region for the 21st century due to climate warming as projected by five general circulation models, 4–11% in the central region and decreases (+1 to −11%) in the eastern region. The overall net change in projected 21st century BA increase at each site seems to be controlled by the relative proportion of responder groups. If these are similar, differences in the magnitude of increase versus decrease in growth control BA projections for that site. This study highlights the importance of regional-scale investigations of biosphere–climate interactions, since our results indicate a substantial gain in aboveground biomass as a result of future warming only in the western regions; while in the eastern regions, climate warming will decrease overall wood production and therefore carbon uptake potential. [Copyright &y& Elsevier]

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]

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]

The Spatial Alaskan Forest Ecosystem Dynamics (SAFED) model was validated across four of the most common vegetation types found in interior Alaska. The vegetation types were an alder (Alnus spp.) - balsam poplar (Populus balsamifera L.) site (FP2), an old-growth balsam poplar and white spruce (Picea glauca (Moench) Voss) site (FP3), a mixed deciduous (primarily birch (Betula papyrifera Marsh.) and aspen (Populus tremuloides Michx.)) and white spruce site (UP2), and a mature white spruce site (UP3). The FP site types are common on the floodplain along the Tanana River and the UP site types are common in the uplands in interior Alaska. SAFED is based on nitrogen productivity for vegetation growth, litter fall quantity and quality, and microbial efficiency for forest floor decomposition. The state factors (climate, topography, and disturbance) are used to describe a broad-scale classification of the landscape to define basic limitations for the driving variables. Climate and ecosystem-level disturbances are handled as restricted stochastic processes. The model has been programed in a spatial framework as an ARC/INFO AML within the GRID package. The current version of the model has been validated as functional from an individual tree basis (1-m2 cell size) in a number of forest types found in interior Alaska. The growth, litter fall, and forest floordecomposition were compared with data from the sites. An estimate ofyearly carbon balance for the four sites was calculated. [ABSTRACT FROM AUTHOR]