The influences of conifer succession, physiographic conditions and herbivory on quaking aspen regeneration after fire

Fire suppression over the last century has increased conifer expansion and dominance in aspen-conifer forests, which appears to be a driving force behind aspen decline in some areas. The primary objective of this study was to examine how increasing conifer dominance affects aspen regeneration vigor following the return of fire. The influence of physiographic features and herbivory on aspen regeneration vigor were also examined. The study was conducted in the Sanford fire complex located in the Dixie National Forest in southern Utah, USA, where more than 31,000 hectares burned in the summer of 2002. Seven years after the burn (at 66 locations) we measured aspen regeneration density and height as response variables and former stand composition and density (the burned trees were still standing), soil characteristics, slope, aspect and presence or absence of herbivory as independent variables. Aspen regeneration (root suckering) densities ranged from <500 to 228,000 stems/hectare with an average of 37,000 stems/hectare. Post-fire aspen regeneration density was most strongly correlated with pre-fire stand successional status (as measured by stand composition and species abundance), with percent conifer abundance (R² = −0.55) and overstory aspen density (R² = + 0.50) being the most important. Average aspen suckering densities ranged from approximately 60,000 stems/hectare in what were relatively pure aspen stands (>90% aspen) to less than 5000 stems/hectare in stands where conifer abundance was greater than 90%. Soil C, N, and P showed positive correlations (R² = 0.07 to 0.17) with aspen regeneration vigor, while soil texture had a relatively weak influence on sucker regeneration. Aspen regeneration densities were 15% lower on north facing aspects compared to east, west and south facing aspects with slope steepness showing no correlation with regeneration vigor. Regeneration density was significantly lower (8%) at sites with evidence of herbivory versus sites where herbivory was absent. Overall, the aspen regeneration response in the Sanford fire complex was strong despite high wildlife densities, which may be related to disturbance size. Where the maintenance of aspen is desired in the landscape we recommend promoting fire when the percentage of overstory conifer stems is greater than 80% or overstory aspen density is less than 200 overstory stems/hectare.     

Effects and etiology of sudden aspen decline in southwestern Colorado,USA

Sudden aspen decline (SAD), affecting Populus tremuloides, was first observed in Colorado in 2004. By 2008 it affected at least 220,000 ha, an estimated 17% of the aspen cover type in the state. In southwestern Colorado, we examined site and stand features in paired healthy and damaged plots to assess the effects of SAD on aspen and to identify factors associated with decline. Root mortality increased significantly with recent crown loss. Consequently, density of regeneration did not increase as the overstory deteriorated, and regeneration that originated since 2002 decreased significantly in stands with moderate to severe SAD. However, mortality of regeneration did not increase with that of the overstory. Remeasurement of a subset of plots after 1–2 yrs showed significant increases in severity. Contrary to expectations, overstory age and diameter were not related to SAD severity as measured by recent crown loss or mortality. Severity of SAD was inversely, but weakly, related to basal area, stem slenderness, and site index, and positively related to upper slope positions. This is consistent with moisture stress as an underlying factor. To test the role of climate as an inciting factor for SAD, a landscape-scale climate model was used to compare moisture status of declining and healthy aspen at the height of the warm drought in water year 2002. Polygons identified as damaged aspen in the 2008 aerial survey had greater moisture deficits than healthy aspen in the 2002 water year. SAD has led to loss of aspen cover in some stands, and is occurring in areas where early loss of aspen due to climate change has been predicted. Further warm, dry growing seasons will likely lead to recurrence of SAD.     

Growth of white spruce underplanted beneath spaced and unspaced aspen stands in northeastern B.C.—10 year results

Establishing white spruce (Picea glauca (Moench) Voss) by planting it under established aspen (Populus tremuloides Michx.), stands has substantial potential as a technique for regenerating boreal mixedwood stands. The presence of an aspen overstory serves to ameliorate frost and winter injury problems and suppresses understory vegetation that may compete with white spruce. In this study we examine the growth of white spruce during the first 10 years after being planted underneath a 39 year-old stand of trembling aspen following thinning and fertilization. Results indicate successful establishment and reasonable growth rates of white spruce planted under thinned and unthinned aspen stands, even with aspen basal area of 51 m² ha⁻¹. Thinning of overstory aspen to 1000 or 2000 stems ha⁻¹ did not increase light reaching seedlings, but did result in improvements in light above the shrub layer and in diameter and height growth of the underplanted seedlings. However, these increases in growth of underplanted spruce may not justify the expense of thinnings. Fertilization of these stands prior to planting had no effect on spruce growth. Growth of spruce underplanted at this site near Fort Nelson was similar to that at two other stands near Dawson Creek, B.C.     

Corrigendum to: Growth of white spruce underplanted beneath spaced and unspaced aspen stands in northeastern B.C.—10 year results

Establishing white spruce (Picea glauca (Moench) Voss) by planting it under established aspen (Populus tremuloides Michx.), stands has substantial potential as a technique for regenerating boreal mixedwood stands. The presence of an aspen overstory serves to ameliorate frost and winter injury problems and suppresses understory vegetation that may compete with white spruce. In this study we examine the growth of white spruce during the first 10 years after being planted underneath a 39-year-old stand of trembling aspen following thinning and fertilization. Results indicate successful establishment and reasonable growth rates of white spruce planted under thinned and unthinned aspen stands, even with aspen basal area of 51 m² ha⁻¹. Thinning of overstory aspen to 1000 or 2000 stems ha⁻¹ did not increase light reaching seedlings, but did result in improvements in light above the shrub layer and in diameter and height growth of the underplanted seedlings. However, these increases in growth of underplanted spruce may not justify the expense of thinnings. Fertilization of these stands prior to planting had no effect on spruce growth. Growth of spruce underplanted at this site near Fort Nelson was similar to that at two other stands near Dawson Creek, B.C.     

表达豇豆胰蛋白酶抑制剂转基因杨树的虫试试验

将转豇豆胰蛋白酶抑制剂基因的三倍体毛白杨回交杂种 [(毛新杨×毛白杨 )×毛白杨 ]]于春季萌发前截干 .用萌条上的树叶对 3种主要杨树害虫 :天幕毛虫 (Malacosomadisstria)、舞毒蛾 (Lymant riadisparL .)和杨雪毒蛾 (StilpnotiacandidaStaudinger)进行离体虫试试验 ,年底测量各转基因株系萌条树高和地径 ,以衡量其生长性状 .结果发现 ,与对照相比 ,转基因株系的树高和地径没有显著降低 ,说明外源基因的导入没有影响转基因杨树的速生特性 ,相反 ,TG0 4的树高和地径较对照有明显的提高 ,这可能是外源导入后促进转基因杨树生长的结果 ;转基因植株均能显著提高害虫的死亡率 ,降低幼虫的排粪量、蛹重、体重增加量和成虫的产卵量 ;其中 ,TG0 7、TG0 8和TG713个转基因株系的抗虫特性较为突出 ,表明CpTI基因在它们的树体内表达较为活跃和稳定     

Quantifying successional rates in western aspen woodlands: Current conditions,future predictions

Stands of quaking aspen (Populus tremuloides) rank among the most biologically diverse plant communities across the intermountain region of western North America. Marked declines of aspen have occurred in recent decades, likely due to a combination of effects from changes in fire regimes, herbivory, climate (e.g. drought), and interspecific competition with conifer species. However, it is poorly understood how the effects of these factors are manifested at a landscape scale over decadal time periods. Analysis of field data combined with topographic information collected across the 500,000 ha Owyhee Plateau in southwestern Idaho revealed that aspen in the area occur in three different biophysical settings; First, aspen stands exist at high altitudes on south-facing slopes where local conifer species are not likely to occur because of limiting temperature or precipitation levels under current climate conditions. In these areas aspen is the potential vegetation type rather than conifers. Second, aspen grow on anomalously wet microsites (e.g. near springs), and third, aspen grow within upland mixed aspen/conifer stands, which are experiencing rapid rates of conifer establishment. Based on a paired t-test (α = 0.05) we conclude that stands growing on wet microsites show significantly slower successional rates of conifer establishment relative to upland aspen stands. We developed a conceptual state-and-transition model for upland aspen/conifer stands occurring across a range of topographic positions. We then parameterized the model using extensive field data in the vegetation dynamics computer simulation model Vegetation Dynamics Development Tool (VDDT), and examined the current and future aspen distribution under varying fire regimes. Model results indicate that average fire return intervals of 50–70 years are desirable for maintenance of aspen in upland areas where conifers are present. Under the current fire regime in the area many upland aspen/conifer stands will likely be lost within 80–200 years. Thresholds for the effect of conifer encroachment and browsing on aspen regeneration identified through this research are similar to those described by others across the West. We therefore suggest that the results presented for the Owyhee Plateau are likely applicable to semi-arid aspen woodlands across the American West where succession to conifers is a cause of aspen decline.     

Forest fragmentation and duration of forest tent caterpillar (Malacosoma disstria Hübner) outbreaks in northern hardwood forests

In northern forests dominated by aspen (Populus spp.), the duration of outbreaks of forest tent caterpillar (Malacosoma disstria Hübner) has been reported to increase with forest fragmentation. This relationship has not been tested in other forest types affected by this widespread native defoliator. From 2002 to 2007, a large-scale outbreak of this insect in the northeastern United States defoliated millions of hectares, with sugar maple (Acer saccharum Marsh.) the primary host. We used digital defoliation maps generated from aerial surveys and national land cover data to assess the effect of fragmentation on outbreak duration in areas of NY, MA, VT, and NH. We found that outbreak duration increased with forest cover and decreased with the forest edge, in opposition to the pattern previously reported for aspen-dominated forests in Canada. This pattern was significant from plot sizes ranging from 100 m to 1000 m in radius. The relationship between FTC and its natural enemies, which was postulated to underlie the effect of fragmentation on outbreaks in aspen forests, may be affected differently in northern hardwood forests, or other factors may be more important in determining outbreak duration in this forest type.     

Early stand production of hybrid poplar and white spruce in mixed and monospecific plantations in eastern Maine

Forest plantations in the northeastern United States comprise a small proportion of the total forest area. Most plantations are typically softwood dominated and managed for sawlog and pulpwood production, while high-yield hardwood plantations for bioenergy feedstocks have not been as widely investigated. The objective of this study was to compare the biomass production of planted white spruce (Picea glauca (Moench) Voss) and hybrid poplar (Populus spp.) plantations (four clones) in monoculture, and in mixture of the two on a typical reforestation site in Maine. Three years following planting, hybrid poplar height and ground line diameter growth rates began to diverge among clones, and by 6?years, the Populus nigra?×?Populus maximowiczii (NM6) clone clearly outperformed three Populus deltoides?×?Populus nigra clones (D51, DN10 and DN70) both in pure stands and in mixtures with white spruce. In mixture, we found the yield of white spruce to decline as the yield of hybrid poplar increased. Overall, yields of white spruce monocultures were comparable to those reported in eastern Canada, while the hybrid poplar biomass yields were substantially lower than those reported from studies on abandoned agricultural lands, likely due to the harsher soil conditions at our site. The dominance of rocky and poorly drained sites (like the one tested in this study) across Maine will likely limit the feasibility of widespread hybrid poplar plantations, and thus constrains their potential use as a bioenergy feedstock.     

Spring frost and decay fungi are implicated in suppressing aspen re-growth following partial cleaning in juvenile stands

? Aspen (Populus tremuloides Michx.) regenerates at high densities following manual cleaning.

? Ten-year-old stands located near Lac La Biche and Peace River, Alberta were manually cleaned to three densities (0, 500 or 1 500 stems ha^?1) at three times (bud set, dormancy or bud flush) to test the hypothesis that maintaining residual aspen reduces regeneration.

? At Lac La Biche up to 98% of the aspen regeneration died in the partially-cleaned plots compared to 67% at Peace River five years post-treatment. A spring frost in the second growing season at Lac La Biche is hypothesized to be the inciting factor predisposing the stump sprouts to infection by decay fungi such as Armillaria root rot, resulting in reduced density and height of the aspen regeneration at Lac La Biche relative to Peace River. Drought and ungulate herbivory provided additional stresses. The high mortality of aspen regeneration at Lac La Biche shifted the understory regeneration from aspen to balsam poplar (Populus balsamifera L.).

? These results indicate that maintaining 1 500 stems ha^?1 of residual aspen will not effectively control the re-sprouting of aspen; however, the vulnerability of aspen regeneration to spring frost and other stressors can nearly eradicate the re-growth of aspen.

     

Stand development patterns as a consequence of the mortality in Austrocedrus chilensis forests

The forests of Austrocedrus chilensis (D. Don) Pic. Sern. et Bizarri in Argentina suffer decline and mortality throughout their natural distribution known as ‘mal del ciprés’. While several aspects of this, to date, temporally unpredictable process of overstory tree mortality have been the focus of detailed studies, there has been little research on tree growth and stand dynamics in symptomatic forests nor stand development patterns and prediction of future stand structure. We studied 12 stands in northern Patagonia (Province of Río Negro) using stand reconstruction studies to examine the changes in stand structure over time as a consequence of overstory mortality and the implications of these structural changes on the establishment and growth of the residual overstory. Dendrochronological analyses were used to reconstruct stand establishment and structure over time, and to study past diameter growth patterns. Mortality in A. chilensis forests was variable in time among stands. As expected, overstory mortality led to the successful establishment of trees in the understory in all stands; however, the response of residual overstory trees was variable. Understory establishment was low in some stands and high in others depending on the density of the overstory. While overstory trees in almost all stands released after the onset of the mortality, the pattern was not distinctive, varying in time, number and magnitude. In some stands, growth releases occurred after single or multiple tree deaths suggesting a relationship between processes, while in others this was not the case. Even when the patterns of recruitment in the understory and the overstory response varied greatly among stands, when examined together, some general patterns emerged. This study is the first to intensively explore the dynamics of A. chilensis forests affected by ‘mal del ciprés’. Additionally, this study showed that arbitrarily categorizing disturbances as discrete or chronic masks the true process of release of growing space and the resulting stand dynamics.     

Susceptibility of northern British Columbia forests to spruce budworm defoliation

Stand susceptibility to defoliation by spruce budworm, Choristoneura fumiferana (Clem.), was examined in the Fort Nelson area of the Prince George Forest Region of British Columbia. In a retrospective study, defoliation maps of the study area were overlaid onto British Columbia Ministry of Forests cover type maps using a geographic information system. Analysis of the combined data identified forest characteristics associated with increased susceptibility to defoliation by spruce budworm. These were stands where the leading species was white spruce (Picea glauca (Moench) Voss), or where spruce was associated with aspen (Populus tremuloides Michx. and P. balsamifera L.) in mixed stands. Susceptibility to defoliation also was related to site quality, level of crown closure and stand age. Spruce stands on medium quality sites (site index 15 to 25 m, at reference breast height age 50 years) were more susceptible than stands on both poor- and high-quality sites. When spruce was mixed with aspen, stands on higher quality sites were more susceptible to budworm attack than poor sites. Open stands, where crown closure was <50%, were more susceptible to attack by spruce budworm than closed canopy stands. Older stands (120–199 years) were more susceptible to budworm attack than younger stands (40–110 years). In defoliated plots monitored for 6 years, tree mortality and top-kill reached a maximum of 30.4 and 47.2%, respectively. The losses varied with level of defoliation and were reduced by applications of the biological pesticide Bacillus thuringiensis.     

Regeneration of Populus nine years after variable retention harvest in boreal mixedwood forests

Aspen and balsam poplar regeneration from root suckers were assessed in boreal mixedwood forests nine years after logging in a variable retention experiment (EMEND Project—Ecosystem Management Emulating Natural Disturbance) located north of Peace River, Alberta, Canada. Five levels of retention of mature trees (2%, 10%, 20%, 50% or 75% of the original basal area) were applied in stands dominated by aspen, white spruce or mixtures of the two species. Basal area of aspen (or that of aspen plus balsam poplar combined) prior to logging strongly influenced sucker density of aspen (or aspen + balsam poplar combined) and in some cases their growth. Nine years after harvest there was a decline in sucker density and volume ha⁻¹ with increasing retention levels of aspen (or both poplars combined); sucker density declined by 50% when only 20% of the original basal area was left in the stand. Retaining mature spruce trees in the stand had little influence on the number of suckers but did affect their total volume ha⁻¹. Thus, we suggest that by knowing stand aspen and balsam poplar density prior to logging and varying levels of retention of aspen and balsam poplar or conifers at harvest, the density of Populus regeneration can be predicted by managers, thereby allowing them to create a range of mixedwood conditions.     

Leaf area dynamics of a boreal black spruce fire chronosequence

Specific leaf area (SLA) and leaf area index (LAI) were estimated using site-specific allometric equations for a boreal black spruce (Picea mariana (Mill.) BSP) fire chronosequence in northern Manitoba, Canada. Stands ranged from 3 to 131 years in age and had soils that were categorized as well or poorly drained. The goals of the study were to: (i) measure SLA for the dominant tree and understory species of boreal black spruce-dominated stands, and examine the effect of various biophysical conditions on SLA; and (ii) examine leaf area dynamics of both understory and overstory for well- and poorly drained stands in the chronosequence. Overall, average SLA values for black spruce (n = 215), jack pine (Pinus banksiana Lamb., n = 72) and trembling aspen (Populus tremuloides Michx., n = 27) were 5.82 +/- 1.91, 5.76 +/- 1.91 and 17.42 +/- 2.21 m2 x kg-1, respectively. Foliage age, stand age, vertical position in the canopy and soil drainage had significant effects on SLA. Black spruce dominated overstory LAI in the older stands. Well-drained stands had significantly higher overstory LAI (P < 0.001), but lower understory LAI (P = 0.022), than poorly drained stands. Overstory LAI was negligible in the recent (3-12 years old) burn sites and highest in the 70-year-old burn site (6.8 and 3.0 in the well- and poorly drained stands, respectively), declining significantly (by 30-50%) from this peak in the oldest stands. Understory leaf area represented a significant portion (> 40%) of total leaf area in all stands except the oldest.     

Structure and composition changes following restoration treatments of longleaf pine forests on the Gulf Coastal Plain of Alabama

Longleaf pine (Pinus palustris Mill.) forests of the Gulf Coastal Plain historically burned every 2–4 years with low intensity fires, which maintained open stands with herbaceous dominated understories. During the early and mid 20th century however, reduced fire frequency allowed fuel to accumulate and hardwoods to increase in the midstory and overstory layers, while woody shrubs gained understory dominance. In 2001, a research study was installed in southern Alabama to develop management options that could be used to reduce fuel loads and restore the ecosystem. As part of a nationwide fire and fire surrogates study, treatments included a control (no fire or other disturbance), prescribed burning only, thinning of selected trees, thinning plus prescribed burning, and herbicide plus prescribed burning. After two cycles of prescribed burning, applied biennially during the growing season, there were positive changes in ecosystem composition. Although thinning treatments produced revenue, while reducing midstory hardwoods and encouraging growth of a grassy understory, burning was needed to discourage regrowth of the hardwood midstory and woody understory. Herbicide application followed by burning gave the quickest changes in understory composition, but repeated applications of fire eventually produced the same results at the end of this 8-year study. Burning was found to be a critical component of any restoration treatment for longleaf communities of this region with positive changes in overstory, midstory and understory layers after just three or four burns applied every 2 or 3 years.     

Long-term response of planted conifers, natural regeneration, and vegetation to harvesting, scalping, and weeding on a boreal mixedwood site

This study reports 14th-year response of a boreal mixedwood stand to different harvest intensities (uncut, 50% partial cut with and without removal of residuals after 3 years, and clearcut), spot site preparation treatments (none and scalped), and chemical weeding frequencies (none, single, and multiple) in northeastern Ontario. The response variables include the survival and growth of planted white spruce (Picea glauca [Moench] Voss) and jack pine (Pinus banksiana Lamb.), height and density of natural regeneration and shrubs, and cover of shrubs and non-woody vegetation. Harvesting and weeding generally improved survival and growth of planted trees, although white spruce survival did not significantly differ among the three weeding frequencies. Harvesting tended to increase heights of hardwood (mostly trembling aspen (Populus tremuloides Michx.)) and conifer (largely balsam fir (Abies balsamea (L.) Mill.).) natural regeneration, cover and density of shrubs, and cover of herbs, lichens, and ferns. Chemical weeding reduced height, density and cover of shrubs, height and density of hardwood regeneration, and fern cover, but increased moss and lichen cover. Spot scalping did not significantly affect planted seedling, natural regeneration, or the vegetation.Maximum survival and growth of planted white spruce and jack pine were achieved using a combination of clearcutting and multiple weeding. However, partial cutting followed by a single weeding produced acceptable survival and reasonable growth of planted trees, particularly for white spruce. Partial canopy removal alone substantially reduced the amount of hardwood regeneration, relative to clearcutting, but did not adequately suppress understory shrubs. Significant improvement in seedling growth following multiple weedings was evident primarily in the complete canopy removal treatments: 50% partial cut with removal of residuals after 3 years and clearcut. While the effects of harvesting and weeding on planted crop trees found in the 5th-year assessments generally persisted at year 14, survival decreased, likely due to light competition from developing hardwood and shrubs.     

Influence of browsing damage and overstory cover on regeneration of American beech and sugar maple nine years following understory herbicide release in central Maine

Northern hardwood stands, notably those with American beech (Fagus grandifolia Ehrh), sugar maple (Acer saccharum Marsh.), and yellow birch (Betula alleghaniensis Britton), are abundant across the forested landscapes of northeastern USA and southeastern Canada. Recent studies have reported an increasing dominance of American beech in the understory and midstory of these forests. Beech is a commercially less desirable tree species due to its association with beech-bark disease, and because it commonly interferes with the regeneration of other more desirable tree species. We examined hardwood regeneration characteristics nine years after application of a 3 × 4 factorial combination of glyphosate herbicide (0.56, 1.12, and 1.68 kg ha^?1) and surfactant concentrations (0.0, 0.25, 0.5, and 1.0% v v^?1) to release sugar maple regeneration from beech-dominated understories using three stands that received shelterwood seed cutting in central Maine. Measurements nine years after treatment showed that glyphosate rate increased both absolute (AD) and relative density (RD) of sugar maple regeneration, but not its height (HT). In contrast, beech AD, RD, and HT were all significantly reduced with increasing glyphosate rate. Post-release browsing by ungulates and a high residual overstory basal area resulted in reduced sugar maple HT. Our results indicated that glyphosate herbicide applied in stands that have been recently shelterwood seed cut can significantly increase the abundance of sugar maple regeneration. However, subsequent browsing damage combined with the negative influence of the residual overstory cover can limit the longer-term benefit of understory herbicide treatments. Subsequent removal of the overstory and browsing-control measures may be needed to promote sugar maple regeneration over beech in similar northern hardwood stands.     

The effect of multi-nutrient fertilization on understory vegetation nutrient concentrations in inland Northwest conifer stands

This study looked at the effects that multi-nutrient fertilization had on understory vegetation nutrient concentrations at four conifer forested locations in the inland Northwest. Multi-nutrient fertilization of conifer stands cannot only enhance the overstory species in the inland Northwest but also the understory vegetation. Determination of nutrient concentration response to fertilization treatments can provide managers the ability to better manipulate their forests for grazing and wildlife habitat. We grouped the understory vegetation into three general life forms: forbs, grasses and grass-likes, and shrubs. Multi-nutrient fertilization had little effect on nitrogen concentration across all life forms. Potassium and sulfur generally increased in concentration. Micronutrients as a whole showed less variability in response to multi-nutrient fertilization. Boron, copper, molybdenum, and zinc generally showed increases in concentrations across all life forms. We were able to conduct analyses on a selected number of understory vegetation species. Individual species showed variability in nutrient concentration response to multi-nutrient fertilization. Wildlife habitat and grazing quality were both increased and decreased following multi-nutrient fertilization. Increases in nutrient concentrations will provide more nutritious vegetation to these animals and vice versa for decreases in concentrations.     

Landscape assessment of a stable aspen community in southern Utah,USA

Recent reports of rapid die-off of aspen (Populus tremuloides), coupled with vigorous debate over long-term reduction of aspen cover in western North America, has prompted considerable research given the importance of this forest type for economic and non-economic interests. Despite this interest, indicators of aspen conditions are poorly understood, and there is a lack of systematic monitoring of stable aspen landscapes. Stable aspen are defined here as being predominantly aspen overstorey (>80% basal area) with little or no conifer regeneration. We examined a putative stable aspen landscape in southern Utah and addressed (1) stand structure and (2) indicators of decline. We sampled 83 aspen-dominated stands within a 275 km² landscape using established forest health monitoring protocols. Eighty-four percent of sample stands on Cedar Mountain exhibited stable aspen characteristics. Principal findings include: (1) a relatively uniform age of adults within the study area; (2) approximately 10% crown dieback on half of the plots sampled; (3) roughly 50% of the study plots had greater than 50% of the trees with damage to the bole; (4) about 25% of the adult basal area was dead; and (5) over half the plots had few sub-canopy individuals and/or limited regeneration. Physiographic variables including elevation, slope, and aspect were generally not strong indicators of aspen condition, typically explaining less than 15% of the variation in basal area, mortality, dieback, or damage. Healthy stands were rarely observed in the most drought prone locations, though the inverse was not necessarily true; healthy and unhealthy stands were found in more mesic settings. Principal components analysis identified two clusters of plots that differed considerably in regeneration; however, no other variables differed between these groupings. We suggest exogenous factors such as land-use history or altered disturbance regimes and endogenous factors such as soils and geology influence aspen condition on this landscape. Further research is necessary to test these hypotheses.     

Understory vegetation in old-growth and second-growth Tsuga canadensis forests in western Massachusetts

We compared the understory communities (herbs, shrubs, and tree seedlings and saplings) of old-growth and second-growth eastern hemlock forests (Tsuga canadensis) in western Massachusetts, USA. Second-growth hemlock forests originated following clear-cut logging in the late 1800s and were 108–136 years old at the time of sampling. Old-growth hemlock forests contained total ground cover of herbaceous and shrub species that was approximately 4 times greater than in second-growth forests (4.02 ± 0.41%/m² versus 1.06 ± 0.47%/m²) and supported greater overall species richness and diversity. In addition, seedling and sapling densities were greater in old-growth stands compared to second-growth stands and the composition of these layers was positively correlated with overstory species composition (Mantel tests, r > 0.26, P < 0.05) highlighting the strong positive neighborhood effects in these systems. Ordination of study site understory species composition identified a strong gradient in community composition from second-growth to old-growth stands. Vector overlays of environmental and forest structural variables indicated that these gradients were related to differences in overstory tree density, nitrogen availability, and coarse woody debris characteristics among hemlock stands. These relationships suggest that differences in resource availability (e.g., light, moisture, and nutrients) and microhabitat heterogeneity between old-growth and second-growth stands were likely driving these compositional patterns. Interestingly, several common forest understory plants, including Aralia nudicaulis, Dryopteris intermedia, and Viburnum alnifolium, were significant indicator species for old-growth hemlock stands, highlighting the lasting legacy of past land use on the reestablishment and growth of these common species within second-growth areas. The return of old-growth understory conditions to these second-growth areas will largely be dependent on disturbance and self-thinning mediated changes in overstory structure, resource availability, and microhabitat heterogeneity.