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Resilience and vulnerability of permafrost to climate change

The resilience and vulnerability of permafrost to climate change depends on complex interactions among topography, water, soil, vegetation, and snow, which allow permafrost to persist at mean annual air temperatures (MAATs) as high as +2 °C and degrade at MAATs as low as –20 °C. To assess these interactions, we compiled existing data and tested effects of varying conditions on mean annual surface temperatures (MASTs) and 2 m deep temperatures (MADTs) through modeling. Surface water had the largest effect, with water sediment temperatures being ~10 °C above MAAT. A 50% reduction in snow depth reduces MADT by 2 °C. Elevation changes between 200 and 800 m increases MAAT by up to 2.3 °C and snow depths by ~40%. Aspect caused only a ~1 °C difference in MAST. Covarying vegetation structure, organic matter thickness, soil moisture, and snow depth of terrestrial ecosystems, ranging from barren silt to white spruce (Picea glauca (Moench) Voss) forest to tussock shrub, affect MASTs by ~6 °C and MADTs by ~7 °C. Groundwater at 2–7 °C greatly affects lateral and internal permafrost thawing. Analyses show that vegetation succession provides strong negative feedbacks that make permafrost resilient to even large increases in air temperatures. Surface water, which is affected by topography and ground ice, provides even stronger negative feedbacks that make permafrost vulnerable to thawing even under cold temperatures.

Alaska

The role of mosses in ecosystem succession and function in Alaska’s boreal forest

Shifts in moss communities may affect the resilience of boreal ecosystems to a changing climate because of the role of moss species in regulating soil climate and biogeochemical cycling. Here, we use long-term data analysis and literature synthesis to examine the role of moss in ecosystem succession, productivity, and decomposition. In Alaskan forests, moss abundance showed a unimodal distribution with time since fire, peaking 30–70 years post-fire. We found no evidence of mosses compensating for low vascular productivity in low-fertility sites at large scales, although a trade-off between moss and vascular productivity was evident in intermediate-productivity sites. Mosses contributed 48% and 20% of wetland and upland productivity, respectively, but produced tissue that decomposed more slowly than both nonwoody and woody vascular tissues. Increasing fire frequency in Alaska is likely to favor feather moss proliferation and decrease Sphagnum abundance, which will reduce soil moisture retention and decrease peat accumulation, likely leading to deeper burning during wildfire and accelerated permafrost thaw. The roles of moss traits in regulating key aspects of boreal performance (ecosystem N supply, C sequestration, permafrost stability, and fire severity) represent critical areas for understanding the resilience of Alaska’s boreal forest region under changing climate and disturbance regimes.

Alaska

Resilience of Alaska’s boreal forest to climatic change

This paper assesses the resilience of Alaska’s boreal forest system to rapid climatic change. Recent warming is associated with reduced growth of dominant tree species, plant disease and insect outbreaks, warming and thawing of permafrost, drying of lakes, increased wildfire extent, increased postfire recruitment of deciduous trees, and reduced safety of hunters traveling on river ice. These changes have modified key structural features, feedbacks, and interactions in the boreal forest, including reduced effects of upland permafrost on regional hydrology, expansion of boreal forest into tundra, and amplification of climate warming because of reduced albedo (shorter winter season) and carbon release from wildfires. Other temperature-sensitive processes for which no trends have been detected include composition of plant and microbial communities, long-term landscape-scale change in carbon stocks, stream discharge, mammalian population dynamics, and river access and subsistence opportunities for rural indigenous communities. Projections of continued warming suggest that Alaska’s boreal forest will undergo significant functional and structural changes within the next few decades that are unprecedented in the last 6000 years. The impact of these social–ecological changes will depend in part on the extent of landscape reorganization between uplands and lowlands and on policies regulating subsistence opportunities for rural communities.

Alaska

Alaska’s changing fire regime - Implications for the vulnerability of its boreal forests

A synthesis was carried out to examine Alaska’s boreal forest fire regime. During the 2000s, an average of 767 000 ha·year –1 burned, 50% higher than in any previous decade since the 1940s. Over the past 60 years, there was a decrease in the number of lightning-ignited fires, an increase in extreme lightning-ignited fire events, an increase in human-ignited fires, and a decrease in the number of extreme human-ignited fire events. The fraction of area burned from human-ignited fires fell from 26% for the 1950s and 1960s to 5% for the 1990s and 2000s, a result from the change in fire policy that gave the highest suppression priorities to fire events that occurred near human settlements. The amount of area burned during late-season fires increased over the past two decades. Deeper burning of surface organic layers in black spruce (Picea mariana (Mill.) BSP) forests occurred during late-growing-season fires and on more well-drained sites. These trends all point to black spruce forests becoming increasingly vulnerable to the combined changes of key characteristics of Alaska’s fire regime, except on poorly drained sites, which are resistant to deep burning. The implications of these fire regime changes to the vulnerability and resilience of Alaska’s boreal forests and land and fire management are discussed.

Alaska

The role of soil drainage class in carbon dioxide exchange and decomposition in boreal black spruce (Picea mariana) forest stands

Black spruce (Picea mariana (Mill.) B.S.P.) forest stands range from well drained to poorly drained, typically contain large amounts of soil organic carbon (SOC), and are often underlain by permafrost. To better understand the role of soil drainage class in carbon dioxide (CO2) exchange and decomposition, we measured soil respiration and net CO2 fluxes, litter decomposition and litterfall rates, and SOC stocks above permafrost in three Alaska black spruce forest stands characterized as well drained (WD), moderately drained (MD), and poorly drained (PD). Soil respiration and net CO2 fluxes were not significantly different among sites, although the relation between soil respiration rate and temperature varied with site (Qw: WD > MD > PD). Annual estimated soil respiration, litter decomposition, and groundcover photosynthesis were greatest at PD. These results suggest that soil temperature and moisture conditions in shallow organic horizon soils at PD were more favorable for decomposition compared with the better drained sites. SOC stocks, however, increase from WD to MD to PD such that surface decomposition and C storage are diametric. Greater groundcover vegetation productivity, protection of deep SOC by permafrost and anoxic conditions, and differences in fire return interval and (or) severity at PD counteract the relatively high near-surface decomposition rates, resulting in high net C accumulation.

Canadian Journal of Forest Research

Tree species and soil nutrient profiles in old-growth forests of the Oregon Coast Range

Old-growth forests of the Pacific Northwest provide a unique opportunity to examine tree species – soil relationships in ecosystems that have developed without significant human disturbance. We characterized foliage, forest floor, and mineral soil nutrients associated with four canopy tree species (Douglas-fir ( Pseudotsuga menziesii (Mirbel) Franco), western hemlock ( Tsuga heterophylla (Raf.) Sarg.), western redcedar ( Thuja plicata Donn ex D. Don), and bigleaf maple ( Acer macrophyllum Pursh)) in eight old-growth forests of the Oregon Coast Range. The greatest forest floor accumulations of C, N, P, Ca, Mg, and K occurred under Douglas-fir, primarily due to greater forest floor mass. In mineral soil, western hemlock exhibited significantly lower Ca concentration and sum of cations (Ca + Mg + K) than bigleaf maple, with intermediate values for Douglas-fir and western redcedar. Bigleaf maple explained most species-based differences in foliar nutrients, displaying high concentrations of N, P, Ca, Mg, and K. Foliar P and N:P variations largely reflected soil P variation across sites. The four tree species that we examined exhibited a number of individualistic effects on soil nutrient levels that contribute to biogeochemical heterogeneity in these ecosystems. Where fire suppression and long-term succession favor dominance by highly shade-tolerant western hemlock, our results suggest a potential for declines in both soil Ca availability and soil biogeochemical heterogeneity in old-growth forests.

Oregon

Changes in faunal and vegetation communities along a soil calcium gradient in northern hardwood forests

Depletion of Ca from forest soils due to acidic deposition has had potentially pervasive effects on forest communities, but these impacts remain largely unknown. Because snails, salamanders, and plants play essential roles in the Ca cycle of northern hardwood forests, we hypothesized that their community diversity, abundance, and structure would vary with differences in biotic Ca availability. To test this hypothesis, we sampled 12 upland hardwood forests representing a soil Ca gradient in the Adirondack Mountains, New York (USA), where chronic deposition has resulted in acidified soils but where areas of well-buffered soils remain Ca rich due to parent materials. Along the gradient of increasing soil [Ca 2+ ], we observed increasing trends in snail community richness and abundance, live biomass of redback salamanders (Plethodon cinereus (Green, 1818)), and canopy tree basal area. Salamander communities were dominated by mountain dusky salamanders (Desmognathus ochrophaeus Cope, 1859) at Ca-poor sites and changed continuously along the Ca gradient to become dominated by redback salamanders at the Ca-rich sites. Several known calciphilic species of snails and plants were found only at the highest-Ca sites. Our results indicated that Ca availability, which is shaped by geology and acidic deposition inputs, influences northern hardwood forest ecosystems at multiple trophic levels, although the underlying mechanisms require further study.

New York

The effect of size and competition on tree growth rate in old-growth coniferous forests

Tree growth and competition play central roles in forest dynamics. Yet models of competition often neglect important variation in species-specific responses. Furthermore, functions used to model changes in growth rate with size do not always allow for potential complexity. Using a large data set from old-growth forests in California, models were parameterized relating growth rate to tree size and competition for four common species. Several functions relating growth rate to size were tested. Competition models included parameters for tree size, competitor size, and competitor distance. Competitive strength was allowed to vary by species. The best ranked models (using Akaike’s information criterion) explained between 18% and 40% of the variance in growth rate, with each species showing a strong response to competition. Models indicated that relationships between competition and growth varied substantially among species. The results also suggested that the relationship between growth rate and tree size can be complex and that how we model it can affect not only our ability to detect that complexity but also whether we obtain misleading results. In this case, for three of four species, the best model captured an apparent and unexpected decline in potential growth rate for the smallest trees in the data set.

Canadian Journal of Forest Research

Litter dynamics in two Sierran mixed conifer forests. I. Litterfall and decomposition rates

Litterfall was measured for 4 years and leaf litter decomposition rates were studied for 3.6 years in two mixed conifer forest (giant sequoia-fir and fir-pine) in the southern Sierra Nevada of California. The giant sequoia-fir forest (GS site) was dominated by giant sequoia ( Sequoiadendron giganteum (Lindl.) Buchh.), white fir ( Abies concolor Lindl. & Gord.), and sugar pine ( Pinus lambertiana Dougl.). The fir-pine forest (FP site) was dominated by white fir, sugar pine, and incense cedar ( Calocedrus decurrens (Torr.) Florin). Litterfall, including large woody debris <15.2 cm in diameter, at the GS site averaged 6364 kg•ha -1 •year -1 compared with 4355 kg•ha -1 •year -1 at the FP site (3.4:1). In the GS site, leaf litter decomposition after 3.6 years was slowest for giant sequoia (28.2% mass loss), followed by sugar pine (34.3%) and white fie (45.1%). In the FP site, mass loss was slowest for sugar pine (40.0%), followed by white fir (45.1%), while incense cedar showed the greatest mass loss (56.9%) after 3.6 years. High litterfall rates of large woody debris (i.e., 2.5-15.2 cm diameter) and slow rates of leaf litter decomposition in the giant sequoia-fir forest type may result in higher litter accumulation rates than in the fir-pine type. Leaf litter times to 95% decay for the GS and FP sites were 30 and 27 years, respectively, if the initial 0.7-year period (a short period of rapid mass decay) was ignored in the calculation. A mass balance approach for total litterfall (<15.2 cm diameter) decomposition yielded lower decay constants than did the litterbag study and therefore longer times to 95% decay (57 years for the GS site and 62 years for the FP site).

Canadian Journal of Forest Research

Litter dynamics in two Sierran mixed conifer forests. II. Nutrient release in decomposing leaf litter

The factors influencing leaf litter decomposition and nutrient release patterns were investigated for 3.6 years in two mixed conifer forests in the southern Sierra Nevada of California. The giant sequoia–fir forest was dominated by giant sequoia ( Sequoiadendrongiganteum (Lindl.) Buchh.), white fir ( Abiesconcolor Lindl. & Gord.), and sugar pine ( Pinuslambertiana Dougl.). The fir–pine forest was dominated by white fir, sugar pine, and incense cedar ( Calocedrusdecurrens (Torr.) Florin). Initial concentrations of nutrients and percent lignin, cellulose, and acid detergent fiber vary considerably in freshly abscised leaf litter of the studied species. Giant sequoia had the highest concentration of lignin (20.3%) and the lowest concentration of nitrogen (0.52%), while incense cedar had the lowest concentration of lignin (9.6%) and second lowest concentration of nitrogen (0.63%). Long-term (3.6 years) foliage decomposition rates were best correlated with initial lignin/N ( r 2 = 0.94, p < 0.05), lignin concentration ( r 2 = 0.92, p < 0.05), and acid detergent fiber concentration ( r 2 = 0.80, p < 0.05). Patterns of nutrient release were highly variable. Giant sequoia immobilized N and P, incense cedar immobilized N and to a lesser extent P, while sugar pine immobilized Ca. Strong linear or negative exponential relationships existed between initial concentrations of N, P, K, and Ca and percent original mass remaining of those nutrients after 3.6 years. This suggests efficient retention of these nutrients in the litter layer of these ecosystems. Nitrogen concentrations steadily increase in decomposing leaf litter, effectively reducing the C/N ratios from an initial range of 68–96 to 27–45 after 3.6 years.

Canadian Journal of Forest Research

Spatial patterns of giant sequoia ( Sequoiadendron giganteum ) in two sequoia groves in Sequoia National Park, California

Although Muir Grove and Castle Creek Grove are similar in area, elevation, and number of giant sequoias, various spatial pattern analysis techniques showed that they had dissimilar spatial patterns for similar-sized trees. Two-dimensional and transect two-term local quadrat variance techniques detected general trends in the spatial patterns of different-sized trees, detected multiple-scale patterns within individual size classes, and provided information on the scale and intensity of patches of individual size classes of trees in Muir and Castle Creek groves. In Muir Grove, midsized sequoias (1.5 to 2.4 m DBH classes) had major pattern scales 350–450 m in diameter, whereas the same-sized trees in Castle Creek Grove had pattern scales >1000 m in diameter. Many size classes of trees had minor patches superimposed on larger scale patterns in both groves. There may be different recruitment patterns in core (i.e., central) areas compared with peripheral areas of sequoia groves; core areas of both groves had more small live sequoias and dead sequoias than peripheral areas of the groves. Higher densities of sequoias and, perhaps, more rapid turnover of individuals in core areas may indicate (i) differences in disturbance histories and favorability of microsites in the core and peripheral areas of groves; (ii) different responses to disturbance due to shifts in the species composition of the stand and thus, the relative influences of intra- to inter-specific competition; or (iii) slower growth or lower survivorship rates in marginal habitat (i.e., peripheral areas).

California

Recent growth increases in old-growth longleaf pine

Longleaf pine ( Pinus palustris Mill.) tree-ring data were obtained from an old-growth stand located in Thomas County, Georgia. The tree-ring chronology from the pine stand is composed of a collection of cores extracted from 26 trees ranging in age from approximately 100 to 400 years. These cores were prepared, dated, and measured, and the resulting data were examined with dendrochronological and statistical techniques. Beginning in approximately 1950 and continuing to the present, annual increments of all age classes examined in this study have increased, resulting in an average annual ring increment approximately 40% greater in 1987 than in 1950. When compared with expected annual increment, the increase for 100- to 150-year-old trees is approximately 45%, while the increase for 200- to 400-year-old trees is approximately 35%. In terms of stand-level aboveground biomass accumulation, the increased growth has resulted in approximately 5% more biomass than expected. The increased growth cannot be explained by disturbance; stand history; or trends in precipitation, temperature, or Palmer drought severity index over the last 57 years. Increased atmospheric CO 2 is a possible explanation for initiation of the observed trend, while SO x and NO x may be augmenting continuation of this phenomenon.

Georgia

Effect of timber harvest on soil carbon storage at Blodgett Experimental Forest, California

Four plots from a mixed conifer forest were similarly cleared, burned, and replanted at various times over 17 years; a plot logged 79 years before sampling was used as a control. The plots had similar slope (2 to 15%, midslope position), aspect (south to southeast), and soil type (Holland series: mesic Haploxeralf; a Gray Brown Luvisol in the Canadian classification system). Twenty sites at each plot were sampled volumetrically by horizon to 20 cm below the organic–mineral soil boundary. Samples were analyzed for bulk density, organic C, and total N. There was an initial loss (15%) of organic C from the soil within 1 to 7 years, likely the result of oxidation (burning and decomposition) and erosion. For 17 years of forest regrowth, the soil continued to lose C (another 15%), probably owing to decomposition of slash material and possibly erosion, despite the slight accumulation of new litter and roots. After 80 years of regrowth, rates of carbon accumulation exceeded rates of loss, but carbon storage had declined and was not likely to recover to preharvest levels. Timber harvest and site preparation dramatically altered soil C and N distribution, in which C/N ratios after site preparation were initially high throughout the upper 20 cm. Subsequently, C/N ratios became lower with depth and with recovery age. Although stocks of C and N varied considerably among the plots and did not change consistently as a function of recovery age, the C/N ratios did vary systematically with recovery age. We hypothesize that the amount of C ultimately stored in the soil at steady state depends largely on N reserves and potentials, which appear to vary with erosion, intensity of burning, and site treatment.

California

Density, ages, and growth rates in old-growth and young-growth forests in coastal Oregon

We studied the ages and diameter growth rates of trees in former Douglas-fir (Pseudotsuga menziesii (Mirb.)Franco) old-growth stands on 10 sites and compared them with young-growth stands (50-70 years old, regenerated after timber harvest) in the Coast Range of western Oregon. The diameters and diameter growth rates for the first 100 years of trees in the old-growth stands were significantly greater than those in the young-growth stands. Growth rates in the old stands were comparable with those from long-term studies of young stands in which density is about 100-120 trees/ha; often young-growth stand density is well over 500 trees/ha. Ages of large trees in the old stands ranged from 100 to 420 years; ages in young stands varied by only about 5 to 10 years. Apparently, regeneration of old-growth stands on these sites occurred over a prolonged period, and trees grew at low density with little self-thinning; in contrast, after timber harvest, young stands may develop with high density of trees with similar ages and considerable self-thinning. The results suggest that thinning may be needed in dense young stands where the management objective is to speed development of old-growth characteristics.

Canadian Journal of Forest Research

Response of high-elevation forests in the Olympic Mountains to climatic change

The gap model ZELIG was used to examine the effects of increased temperature (2°C) and altered precipitation on high-elevation ecosystems of the Olympic Mountains, Washington, U.S.A. Changes in tree species distribution and abundance, as well as stand biomass, were examined on north and south aspects in the dry northeast (NE) and wet southwest (SW) regions of the Olympics for ( i ) warmer, ( ii ) warmer and 20% wetter, and ( iii ) warmer and 20% drier climatic-change scenarios. Dominant tree species shift upwards 300-600 m in elevation in the SW, with subalpine meadows and Tsuga mertensiana (Bong.) Carr. forests being replaced by Abies amabilis (Dougl.) Forbes forests at higher elevations and A. amabilis forests being replaced by Tsuga heterophylla (Raf.) Sarg. forests at lower elevations. In the NE, drought-tolerant species become dominant approximately 200 m lower than present, with A. lasiocarpa dominating the north aspect and Pinus contorta Dougl. ex Loud. the south aspect. Biomass increases in the SW and generally decreases in the NE, depending on aspect and precipitation regime. This study suggests that species and site-specific responses at mesoscale (e.g., wet vs. dry climatic regime) and microscale (e.g., north vs. south aspect) resolutions must be characterized to quantify the variation in potential effects of climatic change on forest vegetation in mountainous regions.

Canadian Journal of Forest Research

Methods for evaluating crown area profiles of forest stands

Canopy architectures of five structurally complex forest stands and three structurally simple forest stands in southwest Oregon and the Willamette Valley, Oregon, were evaluated and quantified through crown area profiles. Mixed conifer and mixed conifer hardwood stands across a range of sites were sampled for crown widths and heights. Crown width and shape equations were derived and used to quantify the stand crown area at incremental heights above the forest floor. Crown area profiles describe the spatial arrangement of aboveground forest vegetation and the total pore spaces between crowns. Plot by plot profiles were combined to produce vertical and horizontal displays of the stand crown area distribution. In complex stands, the forest space was moderately occupied by crowns from the forest floor up to heights over 30 m, producing uniform distributions of between-crown porosity. The structurally complex stands had between-crown porosity values of 70% to 90% for more than 23 vertical metres of canopy, and they had total between-crown porosities of 86% to 91%. The structurally simple stands had between-crown porosity values of 70% to 90% for less than 8 vertical metres of canopy, and they had total between-crown porosities of 69% to 85%. Variances in crown area indicate that variation in horizontal crown area (within heights) was larger in complex stands than in simple stands, but vertical crown areas (between heights) varied less in complex stands. The study provides a basis for discriminating between canopy architectures and for quantifying the porosity of forest canopies.

Canadian Journal of Forest Research

Salmonberry and salal annual aerial stem production: The maintenance of shrub cover in forest stands

Annual sprouting of aerial stems and ramets enables populations of salmonberry (Rubus spectabilis Pursh), salal (Gaultheria shallon Pursh), and probably other forest shrubs to maintain dense covers (>20 000 stems/ha). We studied annual stem production of salmonberry on cut (all stems cut within 15 cm of the ground) and uncut (stems were not treated) plots for 8 years and salal for 5 years in the understories of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), alder, and riparian stands, as well as clearcuts, which are all common stand types in western Oregon. Mean salmonberry stem production on uncut plots ranged from 4.7 stemsA?ma??2A?yeara??1 (95% CI 2.9a??7.4) in alder stands and clearcuts to 1.6 stemsA?ma??2A?yeara??1 (95% CI 1.0a??2.6) in conifer stands. Mean salal production was greater, ranging from 58 stemsA?ma??2A?yeara??1 (95% CI 25a??135) to 8.6 stemsA?ma??2A?yeara??1 (95% CI 3.7a??20.1) on uncut plots in clearcuts and unthinned Douglas-fir stands, respectively. Annual production of both species was somewhat greater on cut plots. Most stems produced in early spring die by December, but enough are recruited to replace mortality of older stems. Stem density was maintained for 8 years for salmonberry and 5 years for salal on both cut and uncut plots. Based on length of rhizomes and bud density we estimate that only 1a??5% of the buds in the rhizomes are needed to support this annual stem production. Although these species sprout vigorously after their aerial stems are killed, disturbance is not necessary for maintaining a dense cover. It appears that, once established, salal, salmonberry, and probably other clonal forest shrubs can maintain a dense cover that can interfere with establishment of trees and other shrubs in canopy gaps or other openings.

Canadian Journal of Forest Research