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Complex interactions of deer herbivory, soil chemistry, and competing vegetation explain oak–hickory forest tree regeneration in central Pennsylvania, USA

The root causes of forest tree regeneration failure are difficult to resolve, although numerous studies show ungulate herbivory, soil conditions, and competition from undesirable vegetation as likely contributors. To better understand the relative importance of each issue, we conducted a 7-year manipulative experiment to assess the interactive effects of white-tailed deer ( Odocoileus virginianus ) herbivory, soil acidity, and competing vegetation on tree regeneration in oak–hickory forests of central Pennsylvania, USA. Outcomes depended on initial tree seedling abundance, and all three factors had significant interactions. At low initial seedling abundance, fencing resulted in the greatest increase, but all treatments had a positive effect on seedling growth and abundance. At higher initial seedling abundance, abundance failed to recover 7 years after herbicide treatment and soil pH was an important predictor. When soil pH was >4.6 from lime application, seedling growth and abundance in unfenced controls with high initial abundance was comparable to the fenced-only treatment. Competing vegetation, assumed to be a symptom of excessive, long-term deer herbivory, does not seem to be the primary factor limiting tree regeneration in our study area. Ameliorating acid deposition warrants greater consideration as a management action because it could provide long-lasting benefits compared to short-term fence installations.

Pennsylvania↗

Cross-site comparison of ecosystem nutrient cycling in hardwood and conifer forests on contrasting soils

In the northeastern United States, both hardwood and conifer forests have developed on sites with contrasting soils, allowing an examination of the effect of site and forest type on ecosystem nutrient cycling. We measured biomass production and nutrient fluxes in northern hardwood and conifer stands at three sites differing in soil fertility. We found that leaf, root, and wood concentrations of calcium (Ca), magnesium (Mg), and potassium reflected differences in soil base cation availability, while concentrations of nitrogen (N) and phosphorus (P) were more consistent across sites. Nutrient uptake was calculated as the sum of litterfall, net throughfall (throughfall minus precipitation), root turnover, and accumulation in perennial tissues (wood). We propose a novel metric of nutrient cycling, the nutrient retention fraction (NRF), defined as the proportion of annual nutrient uptake retained in biomass accretion. Because the NRF is unitless, it can be compared across nutrients; Ca and Mg had the highest NRF and P the lowest ( p = 0.05). Across sites and elements, NRFs were lower for conifers (5.0 ± 0.6%) than for hardwoods (7.2 ± 0.5%), associated with their lower productivity. Nutrient-use efficiency (biomass production divided by nutrient uptake) tended to be high where foliar concentrations indicated low availability of that nutrient. Nutrient retention of N and P was higher where availability of the other element was high, which could be a mechanism contributing to N and P co-limitation.

Canadian Journal of Forest Research↗

Assessing large-scale surveyor variability in the historic forest data of the original U.S. Public Land Survey

The U.S. General Land Office Public Land Survey (PLS) records are a valuable resource for studying pre-European settlement vegetation. However, these data were taken for legal, not ecological, purposes. In turn, the instructions the surveyors followed affected the data collected. For this reason, it has been suggested that the PLS data may not truly represent the surveyed landscapes. This study examined the PLS data of northern Wisconsin, U.S.A., to determine the extent of variability among surveyors. We statistically tested for differences among surveyors in recorded tree species, size, location, and distance from the survey point. While we cannot rule out effects from other influences (e.g., environmental factors), we found evidence suggesting some level of surveyor bias for four of five variables, including tree species and size. The PLS data remain one of the best records of pre-European settlement vegetation available. However, based on our findings, we recommend that projects using PLS records examine these data carefully. This assessment should include not only the choice of variables to be studied but also the spatial extent at which the data will be examined.

Canadian Journal of Forest Research↗

Soil respiration and photosynthetic uptake of carbon dioxide by ground-cover plants in four ages of jack pine forest

Soil carbon dioxide (CO2) emission (soil respiration), net CO2 exchange after photosynthetic uptake by ground-cover plants, and soil CO2 concentration versus depth below land surface were measured at four ages of jack pine (Pinus banksiana Lamb.) forest in central Saskatchewan. Soil respiration was smallest at a clear-cut site, largest in an 8-year-old stand, and decreased with stand age in 20-year-old and mature (60-75 years old) stands during May-September 1994 (12.1, 34.6, 31.5, and 24.9 mol C??m-2, respectively). Simulations of soil respiration at each stand based on continuously recorded soil temperature were within one standard deviation of measured flux for 48 of 52 measurement periods, but were 10%-30% less than linear interpolations of measured flux for the season. This was probably due to decreased soil respiration at night modeled by the temperature-flux relationships, but not documented by daytime chamber measurements. CO2 uptake by ground-cover plants ranged from 0 at the clear-cut site to 29, 25, and 9% of total growing season soil respiration at the 8-year, 20-year, and mature stands. CO2 concentrations were as great as 7150 ppmv in the upper 1 m of unsaturated zone and were proportional to measured soil respiration.

Canadian Journal of Forest Research↗

Growth responses of subalpine fir to climatic variability in the Pacific Northwest

We studied regional variation in growth-limiting factors and responses to climatic variability in subalpine forests by analyzing growth patterns for 28 tree-ring growth chronologies from subalpine fir ( Abies lasiocarpa (Hook.) Nutt.) stands in the Cascade and Olympic Mountains (Washington and Oregon, U.S.A.). Factor analysis identified four distinct time series of common growth patterns; the dominant growth pattern at any site varied with annual precipitation and temperature (elevation). Throughout much of the region, growth is negatively correlated with winter precipitation and spring snowpack depth, indicating that growth is limited primarily by short growing seasons. On the driest and warmest sites, growth is negatively correlated with previous summer temperature, suggesting that low summer soil moisture limits growth. Growth patterns in two regions were sensitive to climatic variability associated with the Pacific Decadal Oscillation, apparently responding to low-frequency variation in spring snowpack and summer soil moisture (one negatively, one positively). This regional-scale analysis shows that subalpine fir growth in the Cascades and Olympics is limited by different climatic factors in different subregional climates. Climate–growth relationships are similar to those for a co-occurring species, mountain hemlock ( Tsuga mertensiana (Bong.) Carrière), suggesting broad biogeographic patterns of response to climatic variability and change by subalpine forest ecosystems in the Pacific Northwest.

Oregon, Washington↗

Vitality and chemistry of roots of red spruce in forest floors of stands with a gradient of soil Al/Ca ratios in the northeastern United States

Number of living root tips per branch, percent dead roots, percent mycorrhizae and mycorrhizal morphotype, response of woody roots to wounding and colonization by fungi, and concentrations of starch, soluble sugars, phenols, percent C and N and C/N ratio, and Al Ca, Fe, K, Mg, Mn, and P were measured for 2 consecutive years in roots of red spruce (Picea rubens Sarg.) in stands in the northeastern United States (nine in 1993 and two additional in 1994) dominated by red spruce and with a gradient of forest floor exchangeable Al/Ca ratios. Root vitality was measured for nonwoody and coarse woody roots; chemical variables were measured for nonwoody (<1 mm), fine woody (1 to <2 mm), and coarse woody (2 to <5 mm) roots. There were significant differences among sites for all variables, particularly in 1993, although few were related to the Al/Ca ratio gradient. Percent mycorrhizae decreased, while some morphotypes increased or decreased as the Al/Ca ratio increased. In nonwoody roots, N increased as the Al/Ca ratio increased. Most sampled trees appeared to be in good or fair health, suggesting that an adverse response of these root variables to high Al concentrations may be apparent only after a significant change in crown health.

Canadian Journal of Forest Research↗

Growth rate predicts mortality of Abies concolor in both burned and unburned stands

Tree mortality is often the result of both long-term and short-term stress. Growth rate, an indicator of long-term stress, is often used to estimate probability of death in unburned stands. In contrast, probability of death in burned stands is modeled as a function of short-term disturbance severity. We sought to narrow this conceptual gap by determining ( i ) whether growth rate, in addition to crown scorch, is a predictor of mortality in burned stands and ( ii ) whether a single, simple model could predict tree death in both burned and unburned stands. Observations of 2622 unburned and 688 burned Abies concolor (Gord. & Glend.) Lindl. (white fir) in the Sierra Nevada of California, U.S.A., indicated that growth rate was a significant predictor of mortality in the unburned stands, while both crown scorch and radial growth were significant predictors of mortality in the burned stands. Applying the burned stand model to unburned stands resulted in an overestimation of the unburned stand mortality rate. While failing to create a general model of tree death for A. concolor , our findings underscore the idea that similar processes may affect mortality in disturbed and undisturbed stands.

Canadian Journal of Forest Research↗

Large wood recruitment and redistribution in headwater streams in the southern Oregon Coast Range, U.S.A

Large wood recruitment and redistribution mechanisms were investigated in a 3.9 km 2 basin with an old-growth Pseudotsuga menziesii (Mirb.) Franco and Tsuga heterophylla (Raf.) Sarg. forest, located in the southern Coast Range of Oregon. Stream size and topographic setting strongly influenced processes that delivered wood to the channel network. In small colluvial channels draining steep hillslopes, processes associated with slope instability dominated large wood recruitment. In the larger alluvial channel, windthrow was the dominant recruitment process from the local riparian area. Consequently, colluvial channels received wood from further upslope than the alluvial channel. Input and redistribution processes influenced piece location relative to the direction of flow and thus, affected the functional role of wood. Wood recruited directly from local hillslopes and riparian areas was typically positioned adjacent to the channel or spanned its full width, and trapped sediment and wood in transport. In contrast, wood that had been fluvially redistributed was commonly located in mid-channel positions and was associated with scouring of the streambed and banks. Debris flows were a unique mechanism for creating large accumulations of wood in small streams that lacked the capacity for abundant fluvial transport of wood, and for transporting wood that was longer than the bank-full width of the channel.

Oregon↗

An experimental demonstration of stem damage as a predictor of fire-caused mortality for ponderosa pine

We subjected 159 small ponderosa pine (Pinus ponderosa Dougl. ex P. & C. Laws.) to treatments designed to test the relative importance of stem damage as a predictor of postfire mortality. The treatments consisted of a group with the basal bark artificially thinned, a second group with fuels removed from the base of the stem, and an untreated control. Following prescribed burning, crown scorch severity was equivalent among the groups. Postfire mortality was significantly less frequent in the fuels removal group than in the bark removal and control groups. No model of mortality for the fuels removal group was possible, because dead trees constituted <4% of subject trees. Mortality in the bark removal group was best predicted by crown scorch and stem scorch severity, whereas death in the control group was predicted by crown scorch severity and bark thickness. The relative lack of mortality in the fuels removal group and the increased sensitivity to stem damage in the bark removal group suggest that stem damage is a critical determinant of postfire mortality for small ponderosa pine.

Canadian Journal of Forest Research↗

Does coring contribute to tree mortality?

We assess the potential of increment coring, a common method for measuring tree ages and growth, to contribute to mortality. We used up to 21 years of annual censuses from two cored and two uncored permanent plots in the Sierra Nevada of California, to detect changes in mortality rates 12 years following coring for individuals >5 cm DBH from two coniferous species, Abies concolor (Gordon & Glend.) Lindl. (white fir) and Abies magnifica A. Murr. (red fir). Using a randomized before-after control impact (BACI) design, we found no differences in mortality rates following coring for 825 cored and 525 uncored A. concolor and 104 cored and 66 uncored A. magnifica . These results support the view that collecting tree cores can be considered nondestructive sampling, but we emphasize that our 12-year postcoring records are short compared with the maximum life-span of these trees and that other species in different environments may prove to be more sensitive to coring.

California↗

Simulating the dynamics of linear forests in Great Plains agroecosystems under changing climates

Most forest growth models are not suitable for the highly fragmented, linear (or linearly shaped) forests in the Great Plains agroecosystems (e.g., windbreaks, riparian forest buffers), where such forests are a minor but ecologically important component of the land mosaics. This study used SEEDSCAPE, a recently modified gap model designed for cultivated land mosaics in the Great Plains, to simulate the effects of climate change on the dynamics of such linear forests. We simulated the dynamics of windbreaks with different initial planting species richness and widths (light changes as the selected resulting factor) using current climate data and nested regional circulation models (RegCMs). Results indicated that (i) it took 70-80 simulation years for the linear forests to reach a steady state under both normal (present-day) and warming climates; (ii) warming climates would reduce total aboveground tree biomass and the spatial variation in biomass, but increase dominance in the linear forests, especially in the upland forests; (iii) linear forests with higher planting species richness and smaller width produced higher aboveground tree biomass per unit area; and (iv) the same species performed very differently with different climate scenarios, initial planting diversity, and forest widths. Although the model still needs further improvements (e.g., the effects of understory species should be included), the model can serve as a useful tool in modeling the succession of linear forests in human-dominated land mosaics under changing climates and may also have significant practical implications in other systems. ?? 2004 NRC Canada.

Canadian Journal of Forest Research↗

Woody debris along an upland chronosequence in boreal Manitoba and its impact on long-term carbon storage

This study investigated the role of fire-killed woody debris as a source of soil carbon in black spruce (Picea mariana (Mill.) BSP) stands in Manitoba, Canada. We measured the amount of standing dead and downed woody debris along an upland chronosequence, including wood partially and completely covered by moss growth. Such woody debris is rarely included in measurement protocols and composed up to 26% of the total amount of woody debris in older stands, suggesting that it is important to measure all types of woody debris in ecosystems where burial by organic matter is possible. Based on these data and existing net primary production (NPP) values, we used a mass-balance model to assess the potential impact of fire-killed wood on long-term carbon storage at this site. The amount of carbon stored in deeper soil organic layers, which persists over millennia, was used to represent this long-term carbon. We estimate that between 10% and 60% of the deep-soil carbon is derived from wood biomass. Sensitivity analyses suggest that this estimate is most affected by the fire return interval, decay rate of wood, amount of NPP, and decay rate of the char (postfire) carbon pool. Landscape variations in these terms could account for large differences in deep-soil carbon. The model was less sensitive to fire consumption rates and to rates at which standing dead becomes woody debris. All model runs, however, suggest that woody debris plays an important role in long-term carbon storage for this area. ?? 2005 NRC Canada.

Canadian Journal of Forest Research↗

Assessment of calcium status in Maine forests: Review and future projection

Forest harvesting and acidic deposition can cause substantial decreases in the calcium (Ca) inventory of forest soils if such losses are not replenished through mineral weathering, atmospheric deposition, or fertilization. The net balance between losses and gains defines the forest Ca status. Site-specific studies have measured Ca pools and fluxes in Maine forests, but no synthesis has been published. In this paper, I review the literature on forest Ca and assess the current status and potential future trends. Forest soils in Maine are currently at lesser risk of Ca depletion compared with many forest soils in the central and southeastern United States, because levels of acidic deposition and rates of Ca accumulation in trees are lower in Maine. The rate of Ca accumulation in trees is reduced in Maine as a result of lower growth rates and a higher proportion of conifer trees that require less Ca than hardwoods. However, field-scale biogeochemical studies in Maine and New Hampshire, and regional estimates of harvest removals and soil inventories coupled with low weathering estimates, indicate that Ca depletion is a realistic concern in Maine. The synthesis of site-specific and regional data for Maine in conjunction with the depletion measured directly in surrounding areas indicates that the Ca status of many forest soils in Maine is likely declining. Ca status could decrease further in the future if forest growth rates increase in response to climate trends and recovery from insect-induced mortality and excessive harvesting in recent years. Proposed climate change induced reductions in spruce and fir and increases in hardwoods would also increase the risk of Ca depletion. ?? 2005 NRC.

Canadian Journal of Forest Research↗

Characterizing flow regimes for floodplain forest conservation: An assessment of factors affecting sapling growth and survivorship on three cold desert rivers

I analyzed annual height growth and survivorship of Fremont cottonwood ( Populus fremontii S. Watson) saplings on three floodplains in Colorado and Utah to assess responses to interannual variation in flow regime and summer precipitation. Mammal exclosures, supplemented with an insecticide treatment at one site, were used to assess flow regime – herbivore interactions. Multiple regression analyses on data collected over 7–11 years indicated that growth of continuously injury-free saplings was positively related to either peak discharge or the maximum 30-day discharge but was not related to interannual decline in the late-summer river stage (Δ W MIN ) or precipitation. Growth was fastest where Δ W MIN was smallest and depth to the late-summer water table moderate (≤1.5 m). Survivorship increased with Δ W MIN where the water table was at shallow depths. Herbivory reduced long-term height growth and survivorship by up to 60% and 50%, respectively. The results support the concept that flow history and environmental context determine whether a particular flow will have a net positive or negative influence on growth and survivorship and suggest that the flow regime that best promotes sapling growth and survival along managed rivers features a short spring flood pulse and constant base flow, with no interannual variation in the hydrograph. Because environmental contexts vary, interannual variation may be necessary for best overall stand performance.

Canadian Journal of Forest Research↗

Factors affecting distribution of wood, detritus, and sediment in headwater streams draining managed young-growth red alder - Conifer forests in southeast Alaska

Factors (riparian stand condition, management regimes, and channel properties) affecting distributions of wood, detritus (leaves and branches), and sediment were examined in headwater streams draining young-growth red alder (Alnus rubra Bong.) - conifer riparian forests (<40 years old) in southeast Alaska. More riparian red alder were found along streams affected by both timber harvesting and mass movement than in streams affected by timber harvesting alone. Young-growth stands produced little large wood material (diameter ???10 cm) and had little effect on altering the size distribution of functional large wood in channels, although more alder wood pieces were found in streams with greater numbers of riparian alder trees. Legacy wood pieces (>40 years old) remained in channels and provided sites for sediment and organic matter storage. Despite various alder-conifer mixtures and past harvesting effects, the abundance of large wood, fine wood, and detritus accumulations significantly decreased with increasing channel bank-full width (0.5-3.5 m) along relatively short channel distances (up to 700 m). Changes in wood, detritus, and sediment accumulations together with changes in riparian stand characteristics create spatial and temporal variability of in-channel conditions in headwater systems. A component of alder within young-growth riparian forests may benefit both wood production and biological recovery in disturbed headwater stream channels. ?? 2006 NRC.

Canadian Journal of Forest Research↗

Role of burning season on initial understory vegetation response to prescribed fire in a mixed conifer forest

Although the majority of fires in the western United States historically occurred during the late summer or early fall when fuels were dry and plants were dormant or nearly so, early-season prescribed burns are often ignited when fuels are still moist and plants are actively growing. The purpose of this study was to determine if burn season influences postfire vegetation recovery. Replicated early-season burn, late-season burn, and unburned control units were established in a mixed conifer forest, and understory vegetation was evaluated before and after treatment. Vegetation generally recovered rapidly after prescribed burning. However, late-season burns resulted in a temporary but significant drop in cover and a decline in species richness at the 1 m 2 scale in the following year. For two of the several taxa that were negatively affected by burning, the reduction in frequency was greater after late-season than early-season burns. Early-season burns may have moderated the effect of fire by consuming less fuel and lessening the amount of soil heating. Our results suggest that, when burned under high fuel loading conditions, many plant species respond more strongly to differences in fire intensity and severity than to timing of the burn relative to stage of plant growth. ?? 2007 NRC.

Canadian Journal of Forest Research↗

The relationship between tree growth patterns and likelihood of mortality: A study of two tree species in the Sierra Nevada

We examined mortality of Abies concolor (Gord. & Glend.) Lindl. (white fir) and Pinus lambertiana Dougl. (sugar pine) by developing logistic models using three growth indices obtained from tree rings: average growth, growth trend, and count of abrupt growth declines. For P. lambertiana , models with average growth, growth trend, and count of abrupt declines improved overall prediction (78.6% dead trees correctly classified, 83.7% live trees correctly classified) compared with a model with average recent growth alone (69.6% dead trees correctly classified, 67.3% live trees correctly classified). For A. concolor , counts of abrupt declines and longer time intervals improved overall classification (trees with DBH ≥20 cm: 78.9% dead trees correctly classified and 76.7% live trees correctly classified vs. 64.9% dead trees correctly classified and 77.9% live trees correctly classified; trees with DBH <20 cm: 71.6% dead trees correctly classified and 71.0% live trees correctly classified vs. 67.2% dead trees correctly classified and 66.7% live trees correctly classified). In general, count of abrupt declines improved live-tree classification. External validation of A. concolor models showed that they functioned well at stands not used in model development, and the development of size-specific models demonstrated important differences in mortality risk between understory and canopy trees. Population-level mortality-risk models were developed for A. concolor and generated realistic mortality rates at two sites. Our results support the contention that a more comprehensive use of the growth record yields a more robust assessment of mortality risk.

California↗

Differential impacts of the southern pine beetle, Dendroctonus frontalis, on Pinus palustris and Pinus taeda

Patterns of host use by herbivore pests can have serious consequences for natural and managed ecosystems but are often poorly understood. Here, we provide the first quantification of large differential impacts of the southern pine beetle, Dendroctonus frontalis Zimmermann, on loblolly pine, Pinus taeda L., and longleaf pine, Pinus palustris P. Mill., and evaluate putative mechanisms for the disparity. Spatially extensive survey data from recent epidemics indicate that, per square kilometre, stands of loblolly versus longleaf pine in four forests (380-1273 km2) sustained 3-18 times more local infestations and 3-116 times more tree mortality. Differences were not attributable to size or age structure of pine stands. Using pheromone-baited traps, we found no differences in the abundance of dispersing D. frontalis or its predator Thanasimus dubius Fabricius between loblolly and longleaf stands. Trapping triggered numerous attacks on trees, but the pine species did not differ in the probability of attack initiation or in the surface area of bark attacked by growing aggregations. We found no evidence for postaggregation mechanisms of discrimination or differential success on the two hosts, suggesting that early colonizers discriminate between host species before a pheromone plume is present. ?? 2007 NRC.

Canadian Journal of Forest Research↗