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As they return to spawn and die in their natal streams, anadromous, semelparous fishes such as Pacific salmon import marine‐derived nutrients to otherwise nutrient‐poor freshwater and riparian ecosystems. Diverse organisms exploit this resource, and previous studies have indicated that riparian tree growth may be enhanced by such marine‐derived nutrients. However, these studies were largely inferential and did not account for all factors affecting tree growth. As an experimental test of the contribution of carcasses to tree growth, for 20 yr, we systematically deposited all sockeye salmon (Oncorhynchus nerka) carcasses (217,055 individual salmon) in the riparian zone on one bank of a 2‐km‐long stream in southwestern Alaska, reducing carcass accumulation on one bank and enhancing it on the other. After accounting for partial consumption and movement of carcasses by brown bears (Ursus arctos) and variation in salmon abundance and body size, we estimated that 267,620 kg of salmon were deposited on the enhanced bank and 45,200 kg on the depleted bank over the 20 yr, for a 5.9‐fold difference in total mass. In 2016, we sampled needles of 84 white spruce trees (Picea glauca) the dominant riparian tree species, for foliar nitrogen (N) content and stable isotope ratios (δ15N), and took core samples for annual growth increments. Stable isotope analysis indicated that marine‐derived N was incorporated into the new growth of the trees on the enhanced bank. Analysis of tree cores indicated that in the two decades prior to our enhancement experiment, trees on the south‐facing (subsequently the depleted) bank grew faster than those on the north‐facing (later enhanced) bank. This difference was reduced significantly during the two decades of fertilization, indicating an effect of the carcass transfer experiment against the background of other factors affecting tree growth. [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]

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]

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]

Increment cores from the boreal forest have long been used to reconstruct past climates. However, in recent years, numerous studies have revealed a deterioration of the correlation between temperature and tree growth that is commonly referred to as divergence. In the Brooks Range of northern Alaska, USA, studies of white spruce ( Picea glauca) revealed that trees in the west generally showed positive growth trends, while trees in the central and eastern Brooks Range showed mixed and negative trends during late 20th century warming. The growing season climate of the eastern Brooks Range is thought to be drier than the west. On this basis, divergent tree growth in the eastern Brooks Range has been attributed to drought stress. To investigate the hypothesis that drought-induced stomatal closure can explain divergence in the Brooks Range, we synthesized all of the Brooks Range white spruce data available in the International Tree Ring Data Bank ( ITRDB) and collected increment cores from our primary sites in each of four watersheds along a west-to-east gradient near the Arctic treeline. For cores from our sites, we measured ring widths and calculated carbon isotope discrimination (δ13C), intrinsic water-use efficiency ( iWUE), and needle intercellular CO2 concentration ( Ci) from δ13C in tree-ring alpha-cellulose. We hypothesized that trees exhibiting divergence would show a corresponding decline in δ13C, a decline in Ci, and a strong increase in iWUE. Consistent with the ITRDB data, trees at our western and central sites generally showed an increase in the strength of the temperature-growth correlation during late 20th century warming, while trees at our eastern site showed strong divergence. Divergent tree growth was not, however, associated with declining δ13C. Meanwhile, estimates of Ci showed a strong increase at all of our study sites, indicating that more substrate was available for photosynthesis in the early 21st than in the early 20th century. Our results, which are corroborated by measurements of xylem sap flux density, needle gas exchange, and measurements of growth and δ13C along moisture gradients within each watershed, suggest that drought-induced stomatal closure is probably not the cause of 20th century divergence in the Brooks Range. [ABSTRACT FROM AUTHOR]

The article provides information on the infanticides committed by the male red squirrels before a mast year. Topics discussed include their anticipation for White Spruce's production of cones, the death of pups after the female stops lactating, and the infanticide occuring frequently in species in which the male can monopolize the female species.

Although many studies have examined the phenological mismatches between interacting organisms, few have addressed the potential for mismatches between phenology and seasonal weather conditions. In the Arctic, rapid phenological changes in many taxa are occurring in association with earlier snowmelt. The timing of snowmelt is jointly affected by the size of the late winter snowpack and the temperature during the spring thaw. Increased winter snowpack results in delayed snowmelt, whereas higher air temperatures and faster snowmelt advance the timing of snowmelt. Where interannual variation in snowpack is substantial, changes in the timing of snowmelt can be largely uncoupled from changes in air temperature. Using detailed, long-term data on the flowering phenology of four arctic plant species from Zackenberg, Greenland, we investigate whether there is a phenological component to the temperature conditions experienced prior to and during flowering. In particular, we assess the role of timing of flowering in determining pre-flowering exposure to freezing temperatures and to the temperatures experienced prior to flowering. We then examine the implications of flowering phenology for flower abundance. Earlier snowmelt resulted in greater exposure to freezing conditions, suggesting an increased potential for a mismatch between the timing of flowering and seasonal weather conditions and an increased potential for negative consequences, such as freezing damage. We also found a parabolic relationship between the timing of flowering and the temperature experienced during flowering after taking interannual temperature effects into account. If timing of flowering advances to a cooler period of the growing season, this may moderate the effects of a general warming trend across years. Flower abundance was quadratically associated with the timing of flowering, such that both early and late flowering led to lower flower abundance than did intermediate flowering. Our results indicate that shifting the timing of flowering affects the temperature experienced during flower development and flowering beyond that imposed by interannual variations in climate. We also found that phenological timing may affect flower abundance, and hence, fitness. These findings suggest that plant population responses to future climate change will be shaped not only by extrinsic climate forcing, but also by species' phenological responses. [ABSTRACT FROM AUTHOR]

The position of the Arctic treeline, which is a key regulator of surface energy exchange and carbon cycling, is widely thought to be controlled by temperature. Here, we present evidence that soil nutrient availability, rather than temperature, may be the proximate control on growth of treeline trees at our study site in northwest Alaska. We examined constraints on growth and allocation of white spruce in three contrasting habitats. The habitats had similar aboveground climates, but soil temperature declined from the riverside terrace to the forest to the treeline. We identified six lines of evidence that conflict with the hypothesis of direct temperature control and/or point to the importance of soil nutrient availability. First, the magnitude of aboveground growth declined from the terrace to the forest to the treeline, along gradients of diminishing soil nitrogen (N) availability and needle N concentration. Second, peak rates of branch extension, main stem radial and fine-root growth were generally not coincident with seasonal air and soil temperature maxima. At the treeline, in particular, rates of aboveground and fine-root growth declined well before air and soil temperatures reached their seasonal peaks. Third, in contrast with the hypothesis of temperature-limited growth, growing season average net photosynthesis was positively related to the sum of normalized branch extension, main stem radial and fine-root growth across trees and sites. Fourth, needle nonstructural carbohydrate concentration was significantly higher on the terrace, where growth was greatest. Fifth, annual branch extension growth was positively related to snow depth, consistent with the hypothesis that deeper snow promotes microbial activity and greater soil nutrient availability. Finally, the tree ring record revealed a large growth increase during late 20th-century climate warming on the terrace, where soil N availability is relatively high. Meanwhile, trees in the forest and at the treeline showed progressively smaller growth increases. Our results suggest temperature effects on tree growth at our study sites may be mediated by soil nutrient availability, making responses to climate change more complex and our ability to interpret the tree ring record more challenging than previously thought. [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]

Mast seeding involves the episodic and synchronous production of large seed crops by perennial plants. The predator satiation hypothesis proposes that mast seeding maximizes seed escape because seed predators consume a decreasing proportion of available seeds with increasing seed production. However, the seed escape benefits of masting depend not only on whether predators are satiated at high levels of seed production, but also on the shape of their functional response (type II vs. type III), and the actual proportion of available seeds that they consume at different levels of seed production. North American red squirrels (Tamiasciurus hudsonicus) are the primary vertebrate predator of white spruce (Picea glauca) mast seed crops in many boreal regions because they hoard unopened cones in underground locations, preempting the normal sequence of cone opening, seed dispersal, and seed germination. We document the functional response of cone-hoarding by red squirrels across three non-mast years and one mast year by estimating the number of cones present in the territories of individual red squirrels and the proportion of these cones that they hoarded each autumn. Even though red squirrels are not constrained by the ingestive and on-body (fat reserves) energy reserve limitations experienced by animals that consume seeds directly, most squirrels hoarded <10% of the cones present on their territories under mast conditions. Cone availability during non-mast years also reached levels that satiated the hoarding activity of red squirrels; however, this occurred only on the highest-quality territories. Squirrels switched to mushroom-hoarding when cone production was low and mushrooms were abundant. This resulted in type III functional response whereby the proportional harvest of cones was highest at levels of cone availability that were intermediate within non-mast years. Overall, more cones escaped squirrel cone-hoarding during a mast event than when cone production was low in non-mast years, which supports the predator satiation hypothesis. However, the highly variable seed escape in non-mast years may help to explain why all spruce cone production is not concentrated into fewer, larger, mast years. [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]

The northern Québec-Labrador tree lines are the most climatically stressed tree ecosystems of eastern North America. In particular, white spruce (Picea glauca) tree line populations distributed between 56° N and 58° N and 61° W and 66° W show contrasted responses to recent changes in climate according to their geographic position relative to the Labrador Sea. Along the coast, the northernmost latitudinal and altitudinal tree lines responded positively to warming over the last 50 years with invading spruce several tens of meters above the current tree line. In contrast, white spruce tree lines across the wind-exposed Labrador plateau are located much higher in altitude and have receded a few tens of meters beginning around AD 1740-1750 and have not yet recovered. Whereas no field evidence of recent fire and insect damage was found, all inland tree line stands were progressively damaged likely due to mechanical defoliation of wind-exposed trees. Massive tree death in the 19th century caused a reduction in the number of seed-bearing trees, and declining tree lines were not replenished by seedlings. Recent warming reported for northern latitudes has not been strong enough to change the regressive tree line trajectory in interior Labrador. However, white spruce expansion above coastal tree line in the northernmost forest site in Labrador is in line with current climatic trends. It is hypothesized that the species is still advancing toward its potential tree line higher in altitude due to delayed postglacial migration. The slow advance of white spruce in northernmost coastal Labrador is likely caused by the rugged topography of the Torngat-Kaumajet-Kiglapait mountains. [ABSTRACT FROM AUTHOR]

We used the mast-seeding tree Picea glauca (white spruce) to examine whether the timing of mast years relative to fire had a lasting effect on the density and timing of regeneration. We studied 17 fires that occurred in mast years and in years with low cone production between 1941 and 1994. Trees were carefully aged by crossdating procedures. Over the 59-yr period studied, there was significantly more regeneration after fires that occurred in mast years than after fires that occurred in years of low cone production. Spruce density was significantly lower after fires that occurred 1-3 years before a mast year than after fires during mast years. The cohort of trees that regenerated in the first mast year after a fire was critical to white spruce regeneration for fires that occurred 0-1 year before a mast year, but mast years occurring three or more years after a fire contributed few recruits. Our results suggest that masting is a key process that interacts' with fire to shape stand composition in boreal mixedwoods. For species like white spruce, for which establishment is linked to disturbance, masting may have a contingent, historical effect on succession and landscape structure. [ABSTRACT FROM AUTHOR]

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

Tests whether the presence of conspecifics or enhanced soil moisture increased the growth and survivorship of Symphoricarpos occidentalis and Picea glauca in the presence of the perrenial grass, Bromus inermis. Competition between the Symphoricarpos and Bromus; Intraspecific competition in Picea; Competitive effects of grasses on Picea survivorship.

Tests the hypothesis that acropetal dispersal by late second and early third-instar Zeiraphera canadensis larvae throughout the crown of white spruce is an adaptive response to within-plant variability in foliage development. Survival; Size and development rate of survivors; Intraspecific larval competition; Effect of formica ant predation.

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]