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Populus tremuloides Michx. is an iconic tree of the mountains of the western United States. In Utah, it very commonly suffers from white trunk rot caused by Phellinus tremulae (Bondartsev) Bondartsev & P.N.Borisov. The incidence of this disease is affected by tree and site characteristics, but the magnitude of these effects appears to be site-dependent. To minimize harvest wood loss in Utah, we determined the locally important factors that explain the wide variation in the incidence of sporocarps of Phellinus tremulae. To avoid confounding of factors, we utilized a multiple regression approach. We found that while the incidence of Phellinus tremulae sporocarps on quaking aspen was always low at high elevations, it was variable at low elevations. Our logistic regression model indicated that variation in the incidence of sporocarps at low elevations was attributable, in part, to variation in aspect, slope, environmental stress, and tree age and size. Based on these results, we recommend that harvesting at elevations below 2500 m be confined to younger trees or to sites on relatively steep, north-facing slopes. Because of site-dependency, the same general method could be used to establish harvesting criteria in other regions.

Recent concern regarding the potential decline of quaking aspen (Populus tremuloides Michx.) forests in the western United States has sparked concern over whether the species can be effectively regenerated. Using a retrospective approach, we quantified the response of regenerating aspen stems to an ordinary set of silvicultural treatments conducted over approximately the past decade in southern Utah, USA. A suite of variables describing stand structure and composition, stand vigor, physiographic factors, herbivore pressure, and treatment types were measured to predict the possible controls, as well as their relative importance, on aspen regeneration. Results suggested that aspen regeneration was most strongly related to browsing pressure, site preparation technique, and the presence of advance reproduction before treatment, which is a novel finding. Secondary predictors included elevation, site index, and overstory conditions, which are generally characteristics of stand vigor. Management recommendations based on our results should recognize the strong primary control that browsing pressure exerts on regeneration. First, the height of advance reproduction is inherently dependent on antecedent herbivory and also indicative of present browsing and should be assessed before treatment. Second, the most effective site preparation techniques, namely broadcast burning and browsing reduction, will directly reduce browsing pressure, assuming ungulate populations are not too large. Any management targeting timely and effective aspen regeneration should incorporate monitoring and (or) controlling browsing pressure, both before and after treatment.

Two ends of the fire regime spectrum are a frequent low-intensity fire regime and an infrequent high-intensity fire regime, but intermediate fire regimes combine high- and low-severity fire over space and time. We used fire-scar and tree-age data to reconstruct fire regime attributes of mixed-conifer and aspen forests in the North Rim area of Grand Canyon National Park, with a goal of estimating patch sizes of historical high-severity fire and comparing them with modern patch sizes. We used three methods based on (i) aspen groves, (ii) even-aged stands, and (iii) inverse distance weighting, to estimate occurrence and patch sizes of historical high-severity fire. Evidence of high-severity fire was common in the 1800s, and high-severity fire years were associated with drought. High-severity fire patch sizes likely ranged from 10−1 to 102 ha. However, the forest is quite young, and we cannot rule out a historical large high-severity fire that could have reinitiated much of the 1400 ha study area. Fire scars, which are indicative of low-severity fire, were also common. Historical fire was likely heterogeneous across the landscape. Maintaining heterogeneity of fire severity, size, and frequency would promote heterogeneity of forest structure and composition and resilience to future disturbances.

We analyzed a series of increment cores collected from 260 adult dominant or co-dominant quaking aspen (Populus tremuloides Michx.) trees from national forests across Colorado and southern Wyoming in 2009 and 2010. Half of the cores were collected from trees in stands with a high amount of crown dieback, and half were from lightly damaged stands. We define the level of stand damage based on stand survey data in which lightly damaged stands had average crown dieback of 16% and heavily damaged stands averaged 41%. Upon analysis, two-thirds of the cores collected did not exhibit radial growth correlated with region-wide patterns (e.g., climate) and were classified as having a low cohesive response. The site variable most predictive of whether a stand exhibited high cohesive response or low cohesive response was site elevation, followed by aspect, slope, and canopy closure. Sites with high cohesive response stands were more likely to have aspen bark beetle damage, white rot, and Cryptosphaeria canker. We did not detect relationships between tree growth and summer precipitation from 1900–2008, but there was a relationship between growth and annual precipitation. A growth model included maximum May and July temperatures, as well as the current and previous year’s annual precipitation.

In the southwestern United States, populations of quaking aspen (Populus tremuloides Michx.) are experiencing widespread mortality. Although environmental factors contributing to mortality have been well characterized, less is known about how genotype and particularly ploidy level affect susceptibility. We used five microsatellite markers to infer the ploidy level of 212 aspen stems in Kaibab National Forest, Arizona. Many multilocus genotypes showed three alleles at one or more loci, suggestive of frequent triploidy among our samples. Sites populated with putative triploids had higher mortality. In addition, heterozygosity was positively associated with mortality and crown dieback. Our results suggest that triploidy is a predisposing factor for aspen mortality in Kaibab National Forest.

Food acquisition and predation avoidance are key drivers of herbivore behaviour. We investigated the interaction of top-down (predator) and bottom-up (food, fire, thermal) effects by measuring the relationship between wolf (Canis lupus L., 1758) predation risk perceived by elk (Cervus elaphus L., 1758) and elk landscape use. We conducted fecal pellet and wolf scat surveys in three valleys with three wolf population levels (Saint Mary: low; Waterton: moderate; North Fork: high). In the North Fork, 90% of quaking aspen (Populus tremuloides Michx.) stands burned recently; the other valleys had no fire. We created predictive models of elk pellet density that incorporated bottom-up and top-down variables. All valleys had a high elk pellet density (≥10 per 100 m2). Wolf scat density was similar where there was no fire, but one order of magnitude greater in burned areas. Elk pellet density was lower in the North Fork, a predation-related response. In all valleys, site-specific elk density declined as impediments to detecting or escaping wolves increased, and elk avoided aspen, except for North Fork unburned areas. Models that best predicted elk density contained bottom-up and top-down effects. At local scales, high predation risk negatively influence elk occurrence, suggesting that even with minimal wolf exposure elk avoid risky sites.

High rates of triploidy have recently been described in quaking aspen (Populus tremuloides Michx.) of the Intermountain West, raising questions about the contributions of triploidy to stand persistence and dynamics. In this study, we investigated cytotype differences between diploid and triploid aspen clones using dendrochronological techniques. We used tree-ring data collected from stems within an aspen stand near Fish Lake, Utah, to test for differences in stem age, population structure, growth, and response to climate. This stand contains the well-known Pando clone, which is purported to be the largest organism documented on earth. Our results show that triploid aspen stems grew more rapidly than diploids, and that this difference was most pronounced early in stand development. Growth response to climate varied little between triploids and diploids, where wide rings were associated with cool, moist years, and narrow rings were associated with above-average growing season temperatures. Stand development processes and inherent genetic differences are mechanisms possibly controlling the observed differences in aspen ring width between triploids and diploids. Regardless of the mechanism, the results have specific management implications. Conventional regeneration methods involving coppicing and the associated intermediate treatments will promote asexually reproducing triploids, leading to static or reduced genetic diversity. Enhanced genetic diversity will be favored by management actions that explicitly account for (i) the potential existence of multiple cytotypes within a stand and (ii) the observed differences in growth rates between diploid and triploid individuals.

Tree mortality because of beetle outbreaks has become substantial and widespread in conifer forests in western North America. A number of environmental and physiological factors influence patterns of mortality. Tree diversity may reduce the severity and extent of insect damage to host trees by providing associational resistance, but the existence and importance of associational resistance varies by forest type and by tree and insect species. We assessed whether plot-level tree diversity contributed to survival of Engelmann spruce (Picea engelmannii Parry ex Engelm.) following a spruce beetle (Dendroctonus rufipennis Kirby) epidemic. Our study plots comprised 2 to 5 tree species including Engelmann spruce, subalpine fir (Abies lasiocarpa (Hook) Nutt.), Douglas-fir (Pseudostuga menziesii (Mirb.) Franco), quaking aspen (Populus tremuloides Michx.), and white fir (Abies concolor (Gordon & Glend.) Hildebr.). We used a model-selection analysis to compare the importance of tree diversity with other known factors that influence spruce survival. We found lower rates of spruce survival in stands where spruce was the dominant tree species (by percent of stand density index) and higher survival in stands where nonspruce conifers (Douglas-fir, subalpine, and white fir) were dominant. We also found that tree diversity (Shannon index) did not show a positive correlation to spruce survival and that there was no additional benefit derived from the presence of aspen, which has higher phylogenetic distance from Engelmann spruce than the other trees in this study. The relationship between diversity and survival is complicated by factors that naturally co-vary with diversity, such as elevation, aspect, and stand density of spruce. Our results best support an explanation that if associational resistance does increase spruce survival during a beetle epidemic, it is due to host or resource dilution, which may be an indirect effect of higher stand diversity.

In recent years, water stress has led to widespread growth declines and dieback of several North American tree species. In addition to its direct effects on tree physiology, water stress may compromise anti-herbivore defenses. We tested whether extrafloral (EF) sugar secretion rate in Populus tremuloides Michx. (quaking aspen) increases in response to defoliation and whether water stress impairs constitutive and inducible EF sugar secretion. We subjected P. tremuloides ramets of four genotypes to water restriction and defoliation in a factorial design and measured EF sugar secretion rates 2, 4, and 6 days after defoliation. The sugar secretion rate of defoliated ramets was significantly higher than that of undefoliated ramets 6 days after defoliation. Low water availability reduced the sugar secretion rate of one of the four aspen genotypes but did not prevent induction. Populus tremuloides genotypes secreted EF sugar at different average rates, suggesting genetic variation for this trait. The results indicate that EF nectar secretion is inducible in P. tremuloides, which could increase the effectiveness of indirect defense following herbivory. Genotypic differences in the response of P. tremuloides to water stress suggest that some clonal stands may be at a disadvantage when faced with compound stresses of drought and herbivory.

The aspen leaf miner, Phyllocnistis populiella Cham., has caused widespread and severe damage to aspen in the boreal forests of western North America for over a decade. We suppressed P. populiella on individual small aspen ramets using insecticide at two sites near Fairbanks, Alaska, annually for 7 years and compared plant performance with controls. Insecticide treatment successfully reduced leaf damage by P. populiella during most years and had little effect on herbivory by externally feeding invertebrates. By the end of the study, control ramets had suffered a reduction in height and girth relative to treated ramets and to the original, pretreatment size. Control ramets produced smaller leaves during some years and, after 7 years, produced fewer total shoots and leaves than ramets sprayed with insecticide. Treatment did not affect mortality, but at the warmer of the two sites, ramets sustaining ambient levels of leaf mining were significantly more likely to die back to basal sprouts than those treated with insecticide. We conclude that a decade of P. populiella outbreak has caused strongly negative effects on aspen development and the production of aboveground tissues.

A conventional view of regeneration ecology of quaking aspen ( Populus tremuloides Michx.) in western North American holds that reproduction is strictly vegetative and, except on some marginal sites, only successful following high-severity disturbance. This view has strongly influenced silvicultural treatment of western aspen and has led to low expectations concerning genetic diversity of stands and landscapes. However, recent discoveries are fundamentally altering our understanding of western aspen regeneration ecology and genetics. For example, there are clearly multiple pathways of aspen regeneration and stand development. Research on a variety of fronts indicates that seedling establishment is common enough to be ecologically important and that genetic diversity is substantially greater than previously thought. We review conventional understanding of western aspen and put this into the context of silvicultural practice. We then review recent developments in aspen research and assess the silvicultural implications of these insights.

Characteristics of edges affect the behavior of species that are active in and near edges. Forest canopies may provide edge-like habitat for bats, though bat response to edge orientation has not been well examined. We sampled bat activity in quaking aspen ( Populus tremuloides Michx.) forest canopies and edges in Heber Valley, Utah, during summer 2009 using Anabat detectors. Categorization and regression tree (CART) analysis of echolocation characteristics (e.g., frequency, duration) identified two guilds based on characteristic frequency (i.e., high- and low-frequency guilds). We used linear regression to compare characteristics of canopy and edge vegetation (e.g., tree height, diameter at breast height) to bat activity levels. Activity levels of high-frequency bats did not respond differentially to edge vegetation; low-frequency bat activity seemed to respond to canopy height. Activity levels of high-frequency bats were significantly greater than low-frequency bats in both edges and canopies. We detected significantly more bat activity in forest edges than in forest canopies, indicating the importance of edges to bats in forests.

Mature aspen (genus Populus L..) and maple (genus Acer L.) forests appear to be high-quality breeding habitat for Yellow-bellied Sapsuckers ( Sphyrapicus varius (L., 1766)). To explore some of the mechanisms that influence quality of the breeding habitat, we measured demographic and breeding resources in four forest stands each of 95-year-old quaking aspen ( Populus tremuloides Michx.) and 182-year-old sugar maple ( Acer saccharum Marsh.) in Algonquin Provincial Park, Ontario, from 2007 to 2009. Population density was higher, egg laying earlier, clutches larger, hatching success greater, and per capita and population fledgling production higher in maple stands compared with aspen stands. Longer nest-building and prelaying stages in aspen stands delayed egg laying by 5 days relative to maple stands. The delay in egg laying in aspen stands may have been caused by lower quality sap resources, which then resulted in lower productivity. Adults delivered smaller food loads to nests in aspen stands than in maple stands, which may have increased nestling mortality in aspen stands. Although per capita fledgling production was lower in aspen stands than in maple stands, Yellow-bellied Sapsuckers in both forests were able to replace themselves, suggesting that mature deciduous and mixed-deciduous forests, in general, are high-quality breeding habitat for this sapsucker.

Extrafloral nectaries (EFNs) are secretory glands most commonly linked to defensive mutualisms. Both a plant’s need for defense and the strength of defense provided by mutualists will vary with plant condition and local insect community. Thus, the benefit of EFNs may vary spatially and temporally. However, little attention has been paid to natural variation in the presence and abundance of EFNs within and among individuals of the same species. Quaking aspen, Populus tremuloides Michx., bear EFNs on a subset of their leaves. Here, we describe patterns of EFN expression on shoots within ramets, among ramets, and among putative clones in interior Alaska. We also examine the relationship between EFN presence and herbivory by both the very abundant aspen leaf miner, Phyllocnistis populiella Chambers, and less common chewing herbivores. The proportion of leaves bearing EFNs varied from 33% to 87% among distinct aspen stands. Within stands, short (1–2 m height) ramets had higher EFN frequency than their taller (>4 m) neighbors. Patterns of herbivory also differed between short and tall ramets. Compared with leaves without EFNs, those with EFNs suffered less mining damage on short ramets but slightly higher damage on tall ramets. Tall ramets suffered more chewing damage than short ramets, but this damage was unrelated to the presence of EFNs. Our results suggest that variable EFN expression may be explained by variation in the benefits of EFNs. Leaves with EFNs on short ramets benefit through reduction in leaf mining, but this benefit does not extend to tall ramets or other forms of herbivory.

Determinate growth involves cessation of leaf formation prior to the end of the growing season, whereas indeterminate growth involves leaf formation for as long as environmental conditions allow. Quaking aspen ( Populus tremuloides Michx.) has been described as having both types of growth. This variation is partially due to canopy bud position, but environmental factors may also contribute. In this study, buds from the same canopy position of aspen trees in northern Colorado, USA, exhibited less neoformation with increasing elevation (R2= 0.78). Evidence suggests that this is primarily a response to the characteristically cooler temperatures and shorter growing seasons of higher elevations.

Analysis of the potential genetic variation in quaking aspen (Populus tremuloides Michx.) wood extractives was performed by sampling nine natural clones at breast height. Significant clonal differences were found not only in the levels of total acetone extractives but also in the levels of sterols–triterpenes, steryl esters–waxes, and triglycerides, all of which are known to contribute to pitch formation in pulping and papermaking. In addition, extractive compounds known to elicit toxic effects in aquatic organisms showed statistically significant differences between natural aspen clones. In some cases (e.g., flavonoids), these compounds formed a part of the defense system in the living tree and, thus, were affected in part by the presence of decayed wood.

Low soil temperatures, common during the growing season in northern forests, have the potential to impede plant growth. In this study, water uptake, water relations, and growth characteristics were examined in aspen (Populus tremuloides) and white spruce (Picea glauca) seedlings that were inoculated with ectomycorrhizal fungi and grown at 20°C daytime air temperatures and low soil temperatures of 4°C and 8°C. Mycorrhizal associations had little effect on root and shoot biomass at both soil temperatures. Root hydraulic conductance (Kr) was higher in both mycorrhizal plant species compared to nonmycorrhizal plants, but there was no soil temperature effect on Kr in either species. Mycorrhizae also increased shoot water potential (Ψw) in Populus tremuloides but had no effect on Ψw in Picea glauca. The increases in Kr and Ψw were not reflected by changes in stomatal conductance (gs) and transpiration rates (E), suggesting that the reduction of water flow in seedlings exposed to low soil temperature was not likely the factor limiting gs in both plant species.Key words: boreal forest, root hydraulic conductance, root growth, stomatal conductance, water uptake.

Forest biomass, rates of production, and carbon dynamics are a function of climate, plant species present, and the structure of the soil organic and mineral layers. Inventory data from the U.S. Forest Service (USFS) Inventory Analysis Unit was used to develop estimates of the land area represented by the major overstory species at various age-classes. The CENTURY model was then used to develop an estimate of carbon dynamics throughout the age sequence of forest development for the major ecosystem types. The estimated boreal forest area in Alaska, based on USFS inventory data is 17 244 098 ha. The total aboveground biomass within the Alaska boreal forest was estimated to be 815 330 000 Mg. The CENTURY model estimated maximum net ecosystem production (NEP) at 137, 88, 152, 99, and 65 g·m–2·year–1 for quaking aspen (Populus tremuloides Michx.), paper birch (Betula papyrifera Marsh.), balsam poplar (Populus balsamifera L.), white spruce (Picea glauca (Moench) Voss), and black spruce (Picea mariana (Mill.) BSP) forest stands, respectively. These values were predicted at stand ages of 80, 60, 41, 68, and 100 years, respectively. The minimum values of NEP for aspen, paper birch, balsam poplar, white spruce, and black spruce were –171, –166, –240, –300, and –61 g·m–2·year–1 at the ages of 1, 1, 1, 1, and 12, respectively. NEP became positive at the ages of 14, 19, 16, 13, and 34 for aspen, birch, balsam poplar, white spruce, and black spruce ecosystems, respectively. A 5°C increase in mean annual temperature resulted in a higher amount of predicted production and decomposition in all ecosystems, resulting in an increase of NEP. We estimate that the current vegetation absorbs approximately 9.65 Tg of carbon per year within the boreal forest of the state. If there is a 5°C increase in the mean annual temperature with no change in precipitation we estimated that NEP for the boreal forest in Alaska would increase to 16.95 Tg of carbon per year.

The response of net ecosystem productivity (NEP) and evaporation in a boreal aspen (Populus tremuloides Michx.) forest and a black spruce (Picea mariana (Mill.) BSP) forest in Canada was compared using a newly developed realistic model of surface-atmosphere exchanges of carbon dioxide (CO2), water vapor, and energy as well as eddy covariance flux measurements made over a 6-year period (1994-1999). The model was developed by incorporating a process-based two-leaf (sunlit and shaded) canopy conductance and photosynthesis submodel in the Canadian Land Surface Scheme (CLASS). A simple submodel of autotrophic and heterotrophic respiration was combined with the photosynthesis model to simulate NEP. The model performed well in simulating half-hourly, daily, and monthly mean CO2 exchange and evaporation values in both deciduous and coniferous forests. Modeled and measured results showed a linear relationship between CO2 uptake and evaporation, and for each kilogram of water transpired, approximately 3 g of carbon (C) were photosynthesized by both ecosystems. The model results confirmed that the aspen forest was a weak to moderate C sink with considerable interannual variability in C uptake. In the growing season, the C uptake capacity of the aspen forest was over twice that of the black spruce forest. Warm springs enhanced NEP in both forests; however, high mid-summer temperatures appear to have significantly reduced NEP at the black spruce forest as a result of increased respiration. The model suggests that the black spruce forest is a weak C sink in cool years and a weak C source in warm years. These results show that the C balance of these two forests is sensitive to seasonal and interannual climatic variability and stresses the importance of continuous long-term flux measurement to confirm modeling results.

To better understand the regeneration ecology of northern red oak (Quercus rubra L.) in mesic forests, we compared its seedling growth and physiology with those of two other hardwoods, differing in successional status, along a gradient in light availability ranging from forest understories (2.6% of full light) to small clearings (69% of full light). Oak's relative growth rate (RGR) closely resembled that of shade-tolerant sugar maple (Acer saccharum Marsh.), and the positive response of both to increasing light was modest, especially beyond a relative light availability of 15%. Intolerant trembling aspen (Populus tremuloides Michx.) outgrew the others when relative light availability exceeded 5%, and its RGR increased more or less linearly with increasing light. However, there was a rank reversal of RGR in deep shade, where maple and oak had a higher RGR than aspen. This reversal was mirrored by seedling demography, as aspen survival was comparatively high in small clearings but negligible in deep shade. Aspen's low RGR and poor survival in low light were associated with a high rate of shoot dark respiration and minimal allocation to starch reserves. Aspen's high RGR in openings was attributed primarily to a high photosynthetic capacity per unit leaf mass. Thus, differential growth and survival among species could be explained in part by trade-offs in attributes that promoted rapid growth in high light at the expense of a favorable carbon balance in low light, or vice versa. Oak's suite of traits facilitated a positive carbon balance in perhaps all but the darkest understories. We suggest that the lack of persistence of oak seedlings in many understory environments may center around factors (e.g., vulnerability to biotic stresses) not directly related to carbon gain.

The early colonization of newly created coarse woody material (CWM) by beetles was studied in aspen mixedwood forests at two locations in north-central Alberta. Healthy trembling aspen (Populus tremuloides Michx.) trees, in old (>100 years) and mature (40–80 years) stands, were cut to provide three types of CWM: stumps, bolts on the ground (logs), and bolts suspended above the ground to simulate snags. Over 2 years, 1049 Coleoptera, representing 49 taxa, were collected. Faunal structure differed little between the two locations. Species diversity was higher in old than in mature stands, and higher in stumps and logs than in suspended bolts; however, these "snags" tended to have higher abundance when compared with stumps and logs. Overall beetle abundance and the catch of wood-boring beetles was significantly higher in the first year post-treatment, mainly because of the ambrosia beetle (Trypodendron retusum (LeConte)) and one of its predators, Rhizophagus remotus LeConte; however, beetle diversity was higher in the second year, suggesting that early wood-boring species may "precondition" the wood for a number of succeeding species. The high turnover rate of taxa and spatial or temporal variation in faunal structure suggests that effort focused on habitat classification of CWM will facilitate management to conserve saproxylic faunal diversity.

A generalized logistic model of individual tree mortality was developed for trembling aspen (Populus tremuloides Michx.), white spruce (Picea glauca (Moench) Voss), and lodgepole pine (Pinus contorta Dougl. ex Loud. var. latifolia Engelm.) in Alberta boreal mixedwood forests based on an empirical data base of permanent sample plots. The model is suitable for observations from unequal remeasurement intervals. The maximum likelihood estimation was used to fit the model, the likelihood ratio test was combined with our understanding of mortality process to select the important variables, and the Hosmer-Lemeshow goodness-of-fit test was conducted to evaluate the fit. The fitted model predicts the survival probability of an individual tree based on the tree diameter at breast height, annual diameter increment, stand basal area, species composition, and site productivity.

Fallen logs of trembling aspen (Populus tremuloides Michx.) and white spruce (Picea glauca (Moench) Voss) at various stages of decomposition were sampled from undisturbed and 1-, 14-, and 28-year-old post-fire and post-harvest sites in northern Alberta and studied for differences in the associated microfungus communities. Wood samples were plated directly onto each of six different media and, from these, fungal species were identified and enumerated over a 24-month period. Approximately 10 000 isolates were obtained, representing 292 species of filamentous microfungi, including 41 ascomycetes, 29 zygomycetes, and 222 mitosporic fungi. The most commonly isolated species were Trichoderma viride (Gray) Pers., Rhinocladiella atrovirens Nannf., Penicillium pinophilum Hedgcock, and Mortierella ramanniana (Moller) Linnem. Cluster analysis and ordination of microfungus communities in logs showed that the tree species of the log had the greatest influence on the species composition of communities. Fungus community composition was also correlated with the stage of decomposition. Species richness was highest in logs from undisturbed sites and lowest in logs from the most recently disturbed sites. Species diversity (Shannon-Weaver) was only slightly higher at undisturbed sites than at disturbed sites. The most significant environmental factor was log moisture, which increased proportionately with stage of decomposition and was significantly correlated with climatic factors.Key words: ecology, fungi, ordination, succession, species richness, diversity.

Quaking aspen (Populus tremuloides Michx.) seedlings were grown in nutrient solutions to investigate their responses to a range of solution Ca levels (10-1000 µM) similar to those found in aspen stands of upper Lakes States. Growth increased significantly with increasing level of solution Ca. The level of solution Ca associated with 90% of maximum elongation (critical level) was 61 µM for shoots and 88 µM for roots. Critical Ca levels for biomass probably were lower. Concentrations of Ca in leaves and roots increased significantly with increasing solution Ca. Elongation of shoots and roots was also strongly and positively related to Ca concentrations in leaves and roots. Critical Ca concentrations (oven-dry mass) for shoot elongation were 0.46% in leaf tissue and 0.12% in root tissue, while critical Ca concentrations for root elongation were 0.54% in leaf tissue and 0.13% in root tissue. Solution Ca may have induced deficiencies of other elements, but direct Ca deficiency was the primary cause of growth reduction. These critical Ca levels in solutions or tissues cannot be used to diagnose Ca deficiency in aspen forests until it is known how other soil and plant factors affect the Ca requirement of aspen.

Ring widths from 1458 trees from Massachusetts, New York, and New Jersey were measured to determine the influence of gypsy moth (Lymantria dispar L.) defoliation on both hosts (primarily oaks (Quercus spp.)) and nonhosts. Previous year's defoliation negatively influenced radial increment in all oak species. Defoliation also negatively affected radial growth of pitch pine (Pinus rigida Mill.), an intermediate host. There was little or no effect of defoliation on increment of trembling aspen (Populus tremuloides Michx.) and bigtooth aspen (Populus grandidentata Michx.), but both are preferred hosts. Defoliation of host trees in mixed stands resulted in increased increment in the nonhosts tulip-tree (Liriodendron tulipifera L.) and ash (Fraxinus spp.) in the year following defoliation. The effect of defoliation was also evident in the relative production of earlywood and latewood with a pronounced dominance of earlywood production in host trees during the same year as defoliation and often in the following year.

Rates of mass loss of foliar litter of trembling aspen (Populus tremuloides Michx.) were measured for 4 years after a single fertilization with N at 200 kg N/ha or a mix of N, P, K. Ca, Mg, S, and B. Foliar litter from fertilized plots had higher N concentrations than litter from control plots; litter from the nutrient-mix plots had higher concentrations of N, P, and Ca. The average rate of mass loss of aspen foliar litter was 60% after 4 years. The higher concentrations of nutrients in litter from fertilized plots did not affect the rate of decomposition. There was some difference in rates of decomposition of litter incubated in plots of the three treatments. Mass loss was significantly slower in the nutrient-mix plots after 12 and 18 months compared with control plots. This experiment provided no evidence that fertilization of boreal aspen forests will increase rates of litter decomposition.

The effects of different amounts of residual canopy on stand development of quaking aspen (Populus tremuloides Michx.) were examined in a chronosequence of 32 stands spanning 6-10 years since harvest. Residual canopy covers ranged from 0 to 65%, and residual basal areas ranged from 0 to 14.4 m2/ha. Aspen regeneration densities ranged from 7130 to 43 672 stems/ha. Regeneration stem density was affected primarily by residual canopy cover (R2 = 0.27, P = 0.0001) and secondarily by stand age (R2 = 0.09, P = 0.004). Aspen density decreased significantly with increasing residual canopy cover for 7-year-old and 8-year-old regeneration. Residual canopy cover did not significantly affect aspen density in 9-year-old regeneration (R2 = 0.02, P = 0.579) but was negatively related to total height of 9-year-old codominant aspens (R2 = 0.49, P = 0.002). Canopy cover was a more accurate representation of the amount of shade the regeneration received than the density or basal area of residual trees. However, the low value of the coefficient of determination from a multiple-regression model indicates that considerable variation in stem densities and height was unexplained by residual canopy cover, even though it was the best predictor of the variables measured.

We examined the effects of woody substrate and stand age on diversity and relative abundance of bryophytes, lichens, and fungi in trembling aspen (Populus tremuloides Michx.) mixedwood boreal forests in Alberta. Point sampling was carried out on surfaces of downed woody material in young (23-26 years), mature (51-63 years), and old (122-146 years) aspen mixedwood stands. Downed woody material was categorized into one of seven decay stages. Diversity and relative abundance of bryophytes, lichens, and fungi were related to decay stage of the log and to stand age. Old aspen mixedwood stands had the highest nonvascular species richness and the greatest diversity of woody substrates in each of the decay stages. The seven decay stages shared many species. Species of bryophytes, lichens, and fungi on particular decay stages were different across the age-classes, indicating that time, as well as structural attributes, were important in determining species assemblages. These results suggest that both the presence of old aspen mixedwood stands and downed wood in different stages of decay are important to maintaining assemblages of bryophytes, lichens, and fungi.Key words: bryophytes, lichens, decay, aspen mixedwood, downed woody material.

This research evaluated the direct and interactive effects of atmospheric CO2 and soil NO3 - availability on growth and biomass partitioning of quaking aspen (Populus tremuloides Michx.), red oak (Quercus rubra L.), and sugar maple (Acer saccharum Marsh.). In the split split plot experimental design, NO3 - availability (low and high) and tree species were nested in two levels of atmospheric CO2 (ambient, 355 μL/L; elevated, 650 μL/L). Seedlings were grown for 57 days in environmental control rooms. Increased CO2 and NO3 - availability positively and (mostly) independently influenced total growth and relative growth rates. Moderate to weak interactions between CO2 and NO3 - for several growth parameters (e.g., leaf production, shoot length, root collar diameter) in some species indicated an enhanced response to CO2 enrichment under conditions of high NO3 - availability. Interactive effects were most pronounced in aspen. Seedling growth and allocation responses to CO2 and NO3 - were frequently species specific and associated with successional status. For example, proportional increases in growth in response to elevated CO2 were greatest for sugar maple and least for quaking aspen, whereas the converse was true with respect to response to high NO3 - availability. This research indicates that the impact of enriched CO2 atmospheres on forest communities will be influenced by both nutrient availability and unique species characteristics.

Growth of red pine (Pinusresinosa Ait.) and quaking aspen (Populustremuloides Michx.) target trees with varying degrees of competition by neighboring aspen was measured in a 7-year-old clearcut in northeastern Minnesota. Competitive conditions were measured using indices of resource availability (percent open sky) and neighborhood stand density. Three-year diameter growth of target trees was reduced by more than 50% in both species because of competition from neighboring quaking aspen. The last 2 year's height growth was also reduced by approximately 30% for both species. The relationship between growth and percent open sky was convex for red pine and concave for quaking aspen, potentially indicating a greater sensitivity in aspen to any reduction below optimal resource availability. Competition reduced crown size in both species, but self-pruning occurred only in quaking aspen. Needle and leaf size were not affected by competition, but red pine needle density was lower in highly competitive environments. Red pine had a higher specific gravity when growing under competition, partially offsetting the reduction in wood volume.

Forages in boreal ecosystems are often deficient in sodium for mammalian herbivores. Moreover, consumption of various plant secondary metabolites has been associated with negative sodium balance in mammals. Neither of these issues has been investigated in birds, which differ from mammals in their ion-exchange processes and postrenal absorption of urine. Our objectives were to determine if ruffed grouse (Bonasa umbellus) can maintain sodium balance on quaking aspen (Populus tremuloides) flower buds, an important winter food, and to determine if the buds' primary plant secondary metabolite (coniferyl benzoate) further compromises a bird's sodium balance. Captive ruffed grouse were fed either aspen buds (0.063 mg∙g−1 sodium) or a formulated diet having different concentrations of coniferyl benzoate in no-choice feeding trials. Sodium excretion did not change in response to coniferyl benzoate intake or acid load from detoxication processes; however, birds were marginally in negative sodium balance (P = 0.035; −5.06 ± 2.05 mg∙kg−1∙d−1) when feeding on aspen buds. Sodium levels in the feces from free-ranging grouse (0.050 ± 0.0 mg∙g−1) and in their winter foods (0.065 mg∙g−1) indicated that these birds likely maintained sodium balance. We estimated that free-ranging ruffed grouse may need as little as 7 mg∙kg−1∙d−1 of sodium to maintain sodium balance, which is lower than the minimum sodium requirements for poultry and mammals.

The increasing commercial demand for quaking aspen (Populustremuloides Michx.) has sparked renewed interest in aspen silvics and management. While aspen generally regenerates vigorously by root suckering following clear-cutting, recent reports of unsuccessful regeneration suggest a linkage to harvesting activities. Accordingly we evaluated the effects of harvest season and equipment trafficking on aspen regeneration in a glaciolacustrine landform in northern Minnesota. Regeneration vigor, as measured by stem density, stem height, and crown closure was greater in winter-harvested areas. Increased equipment trafficking significantly reduced regeneration in stands harvested during the summer. We detected no significant increases in soil density associated with equipment trafficking; however, reduced regeneration was correlated with increased amounts of rutting and soil scarification. For a given level of trafficking, aspen regeneration increased along the annual chronosequence of harvesting later and later in the summer.

Variation at 10 polymorphic isozyme loci was examined for nine populations of quaking aspen (Populustremuloides Michx.) in Minnesota. Mean observed and expected estimates of isozyme heterozygosity were 0.217 and 0.220, respectively, with no significant differences in allele frequencies among populations and no significant deviations from genotype frequencies expected under Hardy–Weinberg equilibrium. These results suggest relatively high levels of gene flow among populations and a mating system involving little inbreeding.

A pollen mixture from three male quaking aspen (Populus tremuloides Michx.) trees was irradiated at exposures of 484, 968, 1453, 1937, 3874, 7747, and 15 494 R and used to control-pollinate cut branches from three female trees. The pollen radiosensitivity was determined from various seed yield and seed quality factors. The pollen LD50 exposure varied with the end point evaluated, ranging from 255 R for number of 50-mesh seed per catkin to 8800 R for total seeds per catkin. The mean LD50 for nine seed yield and seed quality end points was 3995 R. A significant stimulatory response in seed yield was noted at low pollen irradiation levels, particularly at the 484-R exposure. The LD100 was approached but not reached at 15 494 R. Irradiated quaking aspen pollen may be useful in breeding experiments.

Soils on 53 burned sites in the upper peninsula of Michigan were examined for fire-induced water repellency. The development of water repellency was found to be fire related with over 40% of the burned soils showing at least some water-repellent properties. Most water-repellent layers occurred in the upper 5 cm of mineral soil and were related to burn intensity. Laboratory burning experiments with 10 hardwood and conifer litters showed that white pine, red pine, and quaking aspen litter produced water repellency in underlying mineral soil. Repellency was also found on unburned sites, particularly under aspen. Water-repellent soils were widely distributed, but their nonwettable properties generally decreased rapidly over time. Fire-induced water repellency does not appear to present a major long-term management problem on most soils in this region. However, on certain burned sites, water repellency may influence seedling survival and subsequent stand establishment.

Sapling-sized quaking aspen (Populits tremuloides) in five natural clones were inoculated with mass and single ascospore cultures of Hypoxylon mammatum. Fungus isolates varied in ability to produce cankers. 'Conidial-type' isolates generally produced few cankers, while 'mycelial-type' isolates produced cankers on 87.7% of inoculated branches. Isolates from a given ascus inoculated into a clone generally produced cankers of similar length after 2 months. However, some significant differences in canker length did occur. Clones varied in susceptibility to canker enlargement. No single isolate produced the largest cankers on all clones.

Field plantings of Populustremuloides Michx. cuttings in two areas with elevated levels of ambient ozone (New York and Michigan) were evaluated for differences in relative ozone sensitivity among five natural populations from areas that differed in air quality. At least fivefold differences in leaf injury were measured among population means in each planting, and the means were negatively correlated with ozone concentrations for each area. The relative sensitivity rankings among populations and among clones within populations agreed closely with rankings derived from laboratory fumigations of the same plant material. The hypothesis that ozone-sensitive clones of this species are less common in relatively polluted parts than they are in pristine parts of the eastern United States is therefore further supported by field verification of laboratory results. Data from a 10-year-old aspen planting in New York showing impaired growth of ozone-sensitive clones suggest that sensitive individuals can be eliminated from populations by the mechanism of competition.

Yearly applications of N, P, and K fertilizer for a 6-year period to a young, postfire aspen forest, resulted in substantial increases in tree growth primarily in response to nitrogen. The main effect of N was to increase, by at least a factor of two, the stand leaf area index, and bole and total aboveground tree mass. Improved nitrogen nutrition appeared to reduce the efficiency of production per unit of nitrogen incorporated into tree crowns.

Soil chemical properties were studied after a wildfire in stands of white spruce (Piceaglauca (Moench) Voss), black spruce (Piceamariana (Mill.) B.S.P.), paper birch (Betulapapyrifera Marsh.), and quaking aspen (Populustremuloides Michx.). Samples of the forest floor and surface 5 cm of mineral soil were collected from burned sites and unburned controls and analyzed soon after the fire. With the exception of soil pH, effects of the fire on soil chemistry differed among the four forest types. Generally, amounts of exchangeable K, Ca, and Mg did not appreciably increase in the forest floor and surface mineral soil except in heavily burned areas in white spruce and black spruce. Fire reduced amounts of N by about 50% in white spruce, aspen, and birch forest floors. In black spruce, quantities of N were slightly higher in heavily burned locations. Forest floor C:N ratios were substantially lower in heavily burned locations in white spruce and black spruce than in unburned controls. Burning did not have a marked influence on supplies of available P in the forest floor, except in heavily burned black spruce, where average amounts were 12.50 g/m2 versus only 0.46 g/m2 in the control. Burning caused more moderate gains in available P in surface mineral soils under aspen and white spruce. We concluded that fire caused marked short-term changes in soil chemistry in the four forest types. How long these changes will persist is unknown.

Soil and site relationships were studied in 32 mature quaking aspen (Populustremuloides Michx.) communities in north central Wisconsin. Multiple linear regression techniques indicated that soil texture, available water-holding capacity, water-table depth, and stand exposure were the most important factors controlling site index. Two models developed from combinations of these factors accounted for 62% of the variation in site index. Values for soil calcium, magnesium, potassium, and phosphorus were corrected for bulk density and summed to 12-, 24-, 36-, and 60-in. depths. When included in additional models, only magnesium to a 60-in. depth was found to have additional effect on aspen growth over and above that implicit in available water-holding capacity. The models were validated using 10 additional mature aspen stands and then applied to data from 10 stands in early stages of deterioration.

Following a uniform thinning, a young bigtooth aspen (Populusgrandidentata Michx.), quaking aspen (Populustremuloides Michx.), paper birch (Betulapapyrifera Marsh.), and red maple (Acerrubrum L.) stand was treated with nitrogen (N), phosphorus (P), and lime, singly and combined. Nitrogen increased the growth of all species. Lime and P also tended to increase the growth of bigtooth aspen and paper birch. Nitrogen increased mortality and lime reduced mortality. Quaking aspen suffered proportionally greater mortality than other species. The results suggest that both N and lime will be required for an optimum response of aspen and birch growing on acid spodosols in the Northeast, but the cause of and a means to avoid increased mortality caused by fertilizer needs to be determined before large-scale N fertilizer treatments are adopted.

Quaking aspen branches bearing female inflorescences were forced, artificially pollinated, and grown in water in the laboratory at 25 °C. The pistillate flowers developed normally to produce normal seeds in about 2 weeks. Sectioned material, which had been fixed at different times before and after pollination, revealed that the megagametophyte was in the four-nucleate stage at receptivity of the stigmas, and that it developed through the stages that Nagaraj (1952) described for natural material. Fertilization probably occurred between 72 and 120 h after pollination. Unfertilized ovules within ovaries containing one or more fertilized ovules developed normally for 1 week after pollination, but they began to degenerate soon thereafter.