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Premise: Quaking aspen is a clonal tree species that has mixed ploidy, often with high relative abundance of both diploids and triploids but no haploids or tetraploids. Triploids typically have low fertility, leaving their occurrence apparently unlikely from an evolutionary perspective, unless they provide a "triploid bridge" to generating higher‐fitness tetraploids—which are not observed in this species. This study focused on how triploidy can be maintained in quaking aspen. Methods: A computational model was used to simulate gamete production, sexual reproduction, asexual reproduction, parent survival, and offspring survival in a population. All parameters were assumed to be cytotype‐dependent and environment‐independent. Sampling methods were used to identify parameter combinations consistent with observed cytotype frequencies. Results: Many processes and parameter values were sufficient to yield a moderate frequency of triploids, and very few were necessary. The most plausible route involved higher triploid survival at the parent or offspring stage and limited unreduced gamete production by either diploid or triploid parents. Triploid fertility was helpful but not necessary. Conclusions: The coexistence of diploids and triploids in quaking aspen is statistically likely and promoted by the existence of commonly observed, long‐lived triploid clones. However, other mechanisms not captured by the model related to environmental variation could also occur. Further empirical data or more complex but difficult‐to‐parameterize models are needed to gain further insight. [ABSTRACT FROM AUTHOR]

Globally, in remaining wildlands, tree densities and forested cover have increased in grasslands and open forests since European settlement. In the southern Rocky Mountains of Colorado, United States, we determined tree composition and tree cover from historical (years 1875 to 1896) surveys and compared them to current (2002 to 2011) tree composition and current (year 2016) forested land cover for 500,000 ha of the Routt National Forest. Additionally, we examined whether changes in precipitation occurred. Regarding composition, pine (primarily lodgepole pine; Pinus contorta) decreased from 65% to 32% of all trees, with increased subalpine fir (Abies lasiocarpa) from 0.5% to 23% of all trees, and quaking aspen (Populus tremuloides) from 13% to 30% of all trees. According to 80% of 5175 survey points not in forests, the historical landscape was very open, comprised of grasslands, mountain meadows, and other open ecosystems. In contrast, 75% of the current landscape is covered by forests. Change points in the Palmer Modified Drought Index were within historical limits, indicating that forestation was not related to a change in water availability. Based on historical surveys and accounts, we envisioned a historical landscape that was open but embedded with dense lodgepole pine clusters and spruce stands at high elevations, which has now become a predominantly forested landscape of dense forests, similar to global forestation patterns. [ABSTRACT FROM AUTHOR]

PremisePhloem tissue allows for sugar transport along the entirety of a plant and, thus, is one of the most important anatomical structures related to growth. It is thought that the sugar-conducting sieve tube may overwinter and that its cells persist multiple seasons in deciduous trees. One possible overwintering strategy is to build up callose on phloem sieve plates to temporarily cease their function. We tested the hypothesis that five deciduous tree species produce callose on their sieve plates on a seasonal basis.MethodsYoung shoots of five deciduous tree species were sampled periodically between April 2019 and February 2020 in Edmonton, Alberta, Canada. After enzymatic digestion of cytoplasmic constituents, cross sections were imaged using scanning electron microscopy to observe and quantify the level of callose deposition at monthly intervals, and sieve plate pore size was measured. Using a conductivity apparatus, we measured xylem native embolism during these sampling periods.ResultsContrary to past work on some of the same species, we found little evidence that sieve tubes overwinter by becoming occluded with callose. Instead, we found that most sieve plates remain open. Xylem embolism was minimal during the peak growing season, but increased over winter.ConclusionsMany species had been assumed to deposit callose on sieve plates over winter, though anatomical and phenological phloem data were sparse. Our data do not support this notion.

Globally, in remaining wildlands, tree densities and forested cover have increased in grasslands and open forests since European settlement. In the southern Rocky Mountains of Colorado, United States, we determined tree composition and tree cover from historical (years 1875 to 1896) surveys and compared them to current (2002 to 2011) tree composition and current (year 2016) forested land cover for 500,000 ha of the Routt National Forest. Additionally, we examined whether changes in precipitation occurred. Regarding composition, pine (primarily lodgepole pine; Pinus contorta) decreased from 65% to 32% of all trees, with increased subalpine fir (Abies lasiocarpa) from 0.5% to 23% of all trees, and quaking aspen (Populus tremuloides) from 13% to 30% of all trees. According to 80% of 5175 survey points not in forests, the historical landscape was very open, comprised of grasslands, mountain meadows, and other open ecosystems. In contrast, 75% of the current landscape is covered by forests. Change points in the Palmer Modified Drought Index were within historical limits, indicating that forestation was not related to a change in water availability. Based on historical surveys and accounts, we envisioned a historical landscape that was open but embedded with dense lodgepole pine clusters and spruce stands at high elevations, which has now become a predominantly forested landscape of dense forests, similar to global forestation patterns.

Societal Impact Statement: Syntheses clearly show that global warming is affecting ecosystems and biodiversity around the world. New methods and measures are needed to predict the climate resilience of plant species critical to ecosystem stability, to improve ecological management and to support habitat restoration and human well‐being. Widespread keystone species such as aspen are important targets in the study of resilience to future climate conditions because they play a crucial role in maintaining various ecosystem functions and may contain genetic material with untapped adaptive potential. Here, we present a new framework in support of climate‐resilient revegetation based on comprehensively understood patterns of genetic variation in aspen. Summary: Elucidating species' genetic makeup and seed germination plasticity is essential to inform tree conservation efforts in the face of climate change. Populus tremuloides Michx. (aspen) occurs across diverse landscapes and reaches from Alaska to central Mexico, thus representing an early‐successional model for ecological genomics. Within drought‐affected regions, aspen shows ploidy changes and/or shifts from sexual to clonal reproduction, and reduced diversity and dieback have already been observed.We genotyped over 1000 individuals, covering aspen's entire range, for approximately 44,000 single‐nucleotide polymorphisms (SNPs) to assess large‐scale and fine‐scale genetic structure, variability in reproductive type (sexual/clonal), polyploidy and genomic regions under selection. We developed and implemented a rapid and reliable analysis pipeline (FastPloidy) to assess the presence of polyploidy. To gain insights into plastic responses, we contrasted seed germination from western US and eastern Canadian natural populations under elevated temperature and water stress.Four major genetic clusters were identified range wide; a preponderance of triploids and clonemates was found within western and southern North American regions, respectively. Genomic regions involving approximately 1000 SNPs under selection were identified with association to temperature and precipitation variation. Under drought stress, western US genotypes exhibited significantly lower germination rates compared with those from eastern North America, a finding that was unrelated to differences in mutation load (ploidy).This study provided new insights into the adaptive evolution of a key indicator tree that provisions crucial ecosystem services across North America, but whose presence is steadily declining within its western distribution. We uncovered untapped adaptive potential across the species' range which can form the basis for climate‐resilient revegetation. [ABSTRACT FROM AUTHOR]

Spatial interactions between trees influence forest community succession. The objective of this study was to investigate how shifts in forest composition and proximity between tree species affect stand development over time in mixed forest systems. At six locations across the Fishlake National Forest, Utah, USA, in stands where facilitation has been documented previously, tree‐ring samples were collected from aspen and subalpine fir trees. Basal area increment was calculated to characterize the effects of the proximity of overstory trees on multidecadal growth responses of aspen and subalpine fir in aspen‐dominant and mixed aspen–conifer stands. Subalpine fir seedlings were established next to aspen (within 10 cm) when aspen was between 15 and 120 years old with a mean age of 60 years. Aspen and subalpine fir growth rates were reduced with increasing conifer abundance. Aspen trees growing next to a proximate subalpine fir tree had slower growth rates over time than aspen trees growing independently. Growth rates of subalpine fir in aspen‐dominated stands were similar when growing independently or near aspen trees. However, subalpine fir in conifer‐dominated stands maintained higher growth rates when growing next to an aspen tree than when growing independently. The data suggest that as stand competition increases with conifer abundance, the proximity of overstory trees increases competitive exclusion of aspen while having a beneficial growth effect on subalpine fir. These results underscore the importance of maintaining natural fire regimes in forest systems that keep competitive interactions in balance. [ABSTRACT FROM AUTHOR]

Abstract Populus tremuloides is the widest‐ranging tree species in North America and an ecologically important component of mesic forest ecosystems displaced by the Pleistocene glaciations. Using phylogeographic analyses of genome‐wide SNPs (34,796 SNPs, 183 individuals) and ecological niche modeling, we inferred population structure, ploidy levels, admixture, and Pleistocene range dynamics of P. tremuloides, and tested several historical biogeographical hypotheses. We found three genetic lineages located mainly in coastal–Cascades (cluster 1), east‐slope Cascades–Sierra Nevadas–Northern Rockies (cluster 2), and U.S. Rocky Mountains through southern Canadian (cluster 3) regions of the P. tremuloides range, with tree graph relationships of the form ((cluster 1, cluster 2), cluster 3). Populations consisted mainly of diploids (86%) but also small numbers of triploids (12%) and tetraploids (1%), and ploidy did not adversely affect our genetic inferences. The main vector of admixture was from cluster 3 into cluster 2, with the admixture zone trending northwest through the Rocky Mountains along a recognized phenotypic cline (Utah to Idaho). Clusters 1 and 2 provided strong support for the “stable‐edge hypothesis” that unglaciated southwestern populations persisted in situ since the last glaciation. By contrast, despite a lack of clinal genetic variation, cluster 3 exhibited “trailing‐edge” dynamics from niche suitability predictions signifying complete northward postglacial expansion. Results were also consistent with the “inland dispersal hypothesis” predicting postglacial assembly of Pacific Northwestern forest ecosystems, but rejected the hypothesis that Pacific‐coastal populations were colonized during outburst flooding from glacial Lake Missoula. Overall, congruent patterns between our phylogeographic and ecological niche modeling results and fossil pollen data demonstrate complex mixtures of stable‐edge, refugial locations, and postglacial expansion within P. tremuloides. These findings confirm and refine previous genetic studies, while strongly supporting a distinct Pacific‐coastal genetic lineage of quaking aspen.

Quaking aspen (Populus tremuloides Michx.) comprises only a small fraction of western USA forests, yet contributes significant biological diversity and is considered by many to be the most important deciduous forest type in western North America. There is currently a high level of concern in the western United States as many seral aspen populations are declining in vigor due to drought, ungulate browsing, and lack of disturbance. It is also highly uncertain if aspen will successfully accommodate future climate warming via migration through seedling establishment, which has been assumed to be extremely rare. In recent years, fundamental assumptions concerning aspen clonal age, regeneration, and genetic diversity have been challenged, and these findings have important implications for management and persistence of aspen in western USA forests. In this study, we compared regeneration dynamics of aspen revitalization strategies (conifer removal and prescribed fire) to unplanned wildfires of low, moderate, and high severity in the Sierra Nevada, and related multiple components of pre-fire stand composition to post-fire aspen regeneration. To better understand the viability of aspen migration to accommodate future climate warming, we examined recent events of aspen seedling establishment. We found substantial evidence that greater disturbance severity yields increased aspen sprout density and growth rates, and that live conifer and/or dead aspen basal area in a stand before a fire reduces post fire sprout density. Additionally, we found evidence that aspen seedling establishment is more common than has been assumed, and represents a viable means for aspen migration. Future climate changes will present both challenges and opportunities for aspen. Increased temperatures and drought will stress existing populations, but increased high severity fire in forested areas, may provide opportunity for successful aspen migration and genet establishment. In addition to revitalizing existing aspen stands, future management goals should include the establishment of new stands in more suitable habitat.

In recent decades, a paradigm shift in forest management and associated policies has led to greater emphasis on harvest practices that retain mature, overstory trees in forest stands that would otherwise be clear‐cut. While it is often assumed that the maintenance of compositional and structural complexity, such as that achieved through retention forestry approaches, will also mitigate negative impacts to functional diversity, empirical evidence of this relationship is sparse. We examined the effects of an aggregated retention system on taxonomic and functional diversity in a regenerating aspen‐dominated forest. Sampling was conducted along transects arranged to capture the transition from harvested (regenerating) forest to mature, unharvested forest (both intact forest stands and 0.1 ha retention aggregates). We then assessed the magnitude and distance of edge effects on multiple indices of taxonomic and functional diversity as well as functional identity. Twelve years after harvest, the distance and magnitude of edge effects on functional and taxonomic diversity did not differ between the two unharvested patch sizes (intact vs. aggregate); however, intact forest exhibited greater resistance to edge effects and greater depth of edge influence into harvested areas for some traits compared to aggregates. Analyses relying on functional traits were generally applicable across sites within a highly variable forest type, and our results demonstrate the promise of using functional traits to assess management impacts on plant diversity across a landscape. Aggregates maintained some functional attributes associated with interior forest and influenced adjacent regeneration. However, trends in some traits (i.e., shade tolerance and seed mass), particularly in the seedling layer, suggest aggregates of this size provide primarily edge habitat. [ABSTRACT FROM AUTHOR]

Populus tremuloides is the widest‐ranging tree species in North America and an ecologically important component of mesic forest ecosystems displaced by the Pleistocene glaciations. Using phylogeographic analyses of genome‐wide SNPs (34,796 SNPs, 183 individuals) and ecological niche modeling, we inferred population structure, ploidy levels, admixture, and Pleistocene range dynamics of P. tremuloides, and tested several historical biogeographical hypotheses. We found three genetic lineages located mainly in coastal–Cascades (cluster 1), east‐slope Cascades–Sierra Nevadas–Northern Rockies (cluster 2), and U.S. Rocky Mountains through southern Canadian (cluster 3) regions of the P. tremuloides range, with tree graph relationships of the form ((cluster 1, cluster 2), cluster 3). Populations consisted mainly of diploids (86%) but also small numbers of triploids (12%) and tetraploids (1%), and ploidy did not adversely affect our genetic inferences. The main vector of admixture was from cluster 3 into cluster 2, with the admixture zone trending northwest through the Rocky Mountains along a recognized phenotypic cline (Utah to Idaho). Clusters 1 and 2 provided strong support for the "stable‐edge hypothesis" that unglaciated southwestern populations persisted in situ since the last glaciation. By contrast, despite a lack of clinal genetic variation, cluster 3 exhibited "trailing‐edge" dynamics from niche suitability predictions signifying complete northward postglacial expansion. Results were also consistent with the "inland dispersal hypothesis" predicting postglacial assembly of Pacific Northwestern forest ecosystems, but rejected the hypothesis that Pacific‐coastal populations were colonized during outburst flooding from glacial Lake Missoula. Overall, congruent patterns between our phylogeographic and ecological niche modeling results and fossil pollen data demonstrate complex mixtures of stable‐edge, refugial locations, and postglacial expansion within P. tremuloides. These findings confirm and refine previous genetic studies, while strongly supporting a distinct Pacific‐coastal genetic lineage of quaking aspen. [ABSTRACT FROM AUTHOR]

Understanding how severe disturbances and their interactions affect forests is key to projecting ecological change under a warming climate. Substantial increases in some biotic disturbances, such as bark beetle outbreaks, in temperate forest ecosystemsmay compromise recovery to a forest vegetation type (i.e., physiognomic recovery or resilience), especially if subsequent biotic disturbances (e.g., herbivory) alter recovery mechanisms. From 2005 to 2017, severe outbreaks (>90% mortality) of spruce bark beetles (SB, Dendroctonus rufipennis) affected Engelmann spruce (Picea engelmannii) across 325,000 ha of spruce and subalpine fir (Abies lasiocarpa) forest in the southern Rocky Mountains, USA. Concurrently, an outbreak of western balsam bark beetle (WBBB, Dryocoetes confuses) infested subalpine fir across at least 47,000 of these hectares. We explored the capacity of 105 stands affected by one or two bark beetle outbreaks and browsing of juvenile trees by ungulates to return to a forest vegetation type in the context of pre‐outbreak forest conditions and topography. Nine initial forest trajectories (i.e., at least several decades) were identified from four pre‐outbreak forest types affected by three biotic disturbances that occurred at different spatial scales and severities. Most stands (86%) contained surviving nonhost adult trees in the main canopy (fir and aspen [Populus tremuloides]) and many surviving juveniles of all species, implying that they are currently on a trajectory for physiognomic recovery. Stands composed exclusively of large‐diameter spruce were affected by a severe SB outbreak and were most vulnerable to a transition to a low‐density forest, below regional stocking levels (<370 trees/ha). Greater pre‐outbreak stand structural complexity and species diversity were key traits of stands with a higher potential for physiognomic recovery. However, all multispecies stands shifted in relative composition of the main canopy to nonhost species, suggesting low potential for compositional recovery over the next several decades. Most post‐outbreak stands (86%) exceeded regional stocking levels with trees taller than the browse zone (<2 m). As such, ungulate browsing on over half of all juveniles will primarily affect the rate of infilling of the forest canopy and preferential browsing of more palatable species will influence the composition of the future forest canopy. [ABSTRACT FROM AUTHOR]

Ploidy level in plants may influence ecological functioning, demography and response to climate change. However, measuring ploidy level typically requires intensive cell or molecular methods.We map ploidy level variation in quaking aspen, a dominant North American tree species that can be diploid or triploid and that grows in spatially extensive clones. We identify the predictors and spatial scale of ploidy level variation using a combination of genetic and ground‐based and airborne remote sensing methods.We show that ground‐based leaf spectra and airborne canopy spectra can both classify aspen by ploidy level with a precision‐recall harmonic mean of 0.75–0.95 and Cohen's kappa of c. 0.6–0.9. Ground‐based bark spectra cannot classify ploidy level better than chance. We also found that diploids are more common on higher elevation and steeper sites in a network of forest plots in Colorado, and that ploidy level distribution varies at subkilometer spatial scales.Synthesis. Our proof‐of‐concept study shows that remote sensing of ploidy level could become feasible in this tree species. Mapping ploidy level across landscapes could provide insights into the genetic basis of species' responses to climate change. [ABSTRACT FROM AUTHOR]

Highlights • Review of current knowledge related to aspen seed ecology and early establishment. • Establishment requirements and approaches for seed-based aspen restoration. • Aspen seedling stock quality and manipulation of aspen seedling characteristics. • Management approaches to enhance the restoration of aspen on the landscape. • Passive and active seedling based aspen regeneration and restoration techniques. Abstract Quaking or trembling aspen (Populus tremuloides Michx.) is a foundational tree species, which is native, common, and broadly distributed in North America. The ecology of aspen has been extensively studied throughout its range, but both research and forest management practices have focused primarily on its ability to regenerate asexually via root suckering. The seed-based reproductive ecology of aspen has received comparatively little attention, and information on the underlying processes, mechanisms, and requirements of seed regeneration tends to be scattered, somewhat anecdotal, or based only on localized research efforts. Here we review and explore some of the variables that influence the sexual reproduction and early establishment of aspen. We focus this review on western North America, where trembling aspen plays a dominant ecological role and may be disproportionately impacted by climate change. This synthesis presents existing information and identifies critical knowledge gaps in our understanding of seed –based aspen regeneration, in particular as it relates to flowering and seed production, as well as germination, first year growth, and survival of aspen seedlings. This information is discussed further in the context of aspen ecology and its application in both passive and active management approaches to aspen seedling regeneration and restoration. [ABSTRACT FROM AUTHOR]

Quaking aspen (Populus tremuloides Michx.) comprises only a small fraction of western USA forests, yet contributes significant biological diversity and is considered by many to be the most important deciduous forest type in western North America. There is currently a high level of concern in the western United States as many seral aspen populations are declining in vigor due to drought, ungulate browsing, and lack of disturbance. It is also highly uncertain if aspen will successfully accommodate future climate warming via migration through seedling establishment, which has been assumed to be extremely rare. In recent years, fundamental assumptions concerning aspen clonal age, regeneration, and genetic diversity have been challenged, and these findings have important implications for management and persistence of aspen in western USA forests. In this study, we compared regeneration dynamics of aspen revitalization strategies (conifer removal and prescribed fire) to unplanned wildfires of low, moderate, and high severity in the Sierra Nevada, and related multiple components of pre‐fire stand composition to post‐fire aspen regeneration. To better understand the viability of aspen migration to accommodate future climate warming, we examined recent events of aspen seedling establishment. We found substantial evidence that greater disturbance severity yields increased aspen sprout density and growth rates, and that live conifer and/or dead aspen basal area in a stand before a fire reduces post fire sprout density. Additionally, we found evidence that aspen seedling establishment is more common than has been assumed, and represents a viable means for aspen migration. Future climate changes will present both challenges and opportunities for aspen. Increased temperatures and drought will stress existing populations, but increased high severity fire in forested areas, may provide opportunity for successful aspen migration and genet establishment. In addition to revitalizing existing aspen stands, future management goals should include the establishment of new stands in more suitable habitat.

Timing of herbivory or selection of specific plant tissues (mode of herbivory) by different ungulate herbivore species are likely to have important influences on plant defense strategies. In this study, we devised two different modes of simulated herbivory, representing a selective ungulate feeding strategy (defoliation: leaf tissue removal only) and a bulk feeding strategy (clipping: leaves, twigs and meristems taken together). We applied these contrasting herbivory treatments to juvenile aspen suckers (Populus tremuloides Michx.) regenerating underneath aspen stands in early summer (June), late summer (August) or at both times to determine the effects of herbivory mode, timing and frequency on regenerating aspen. In response to the simulated herbivory treatments, we measured traits related to three plant defense strategies: tolerance (aboveground biomass and stem diameter), resistance (foliar phenolic glycosides) and vertical escape (sucker height and average leader length). There was no evidence that mode, timing or frequency of simulated herbivory induced or repressed phenolic glycoside production. Early summer herbivory was more detrimental than late summer herbivory on aspen tolerance and escape. Repeat herbivory in late summer did not amplify the negative effects of early summer herbivory. Clipping and defoliation tended to have similar effects on tolerance but clipping was more detrimental than defoliation on vertical escape. These results suggest that different ungulate herbivore species may have disparate impacts on the plant communities by selecting different tissues of the same plant, or browsing the plant at different times in the growing season. [ABSTRACT FROM AUTHOR]

Soil moisture varies within landscapes in response to vegetative, physiographic, and climatic drivers, which makes quantifying soil moisture over time and space difficult. Nevertheless, understanding soil moisture dynamics for different ecosystems is critical, as the amount of water in a soil determines a myriad ecosystem services and processes such as net primary productivity, runoff, microbial decomposition, and soil fertility. We investigated the patterns and variability in in situ soil moisture measurements converted to plant-available water across time and space under different vegetative cover types and topographic positions at the Marcell Experimental Forest (Minnesota, USA). From 0- to 228.6-cm soil depth, plant-available water was significantly higher under the hardwoods (12%), followed by the aspen (8%) and red pine (5%) cover types. Across the same soil depth, toeslopes were wetter (mean plant-available water = 10%) than ridges and backslopes (mean plant-available water was 8%), although these differences were not statistically significant ( p < .05). Using a mixed model of fixed and random effects, we found that cover type, soil texture, and time were related to plant-available water and that topography was not significantly related to plant-available water within this low-relief landscape. Additionally, during the 3-year monitoring period, red pine and quaking aspen sites experienced plant-available water levels that may be considered limiting to plant growth and function. Given that increasing temperatures and more erratic precipitation patterns associated with climate change may result in decreased soil moisture in this region, these species may be sensitive and vulnerable to future shifts in climate. [ABSTRACT FROM AUTHOR]

Recent emphasis on increasing structural complexity and species diversity reflective of natural ecosystems through the use of retention harvesting approaches is coinciding with increased demand for forest-derived bioenergy feedstocks, largely sourced through the removal of harvest residues associated with whole-tree harvest. Uncertainties about the consequences of such approaches prompted us to examine the combined impacts of aggregated overstory retention and harvest residue retention on the composition and density of regeneration following biomass harvests on four operational-scale (40 ha) study areas dominated by Populus tremuloides Michx. in northern Minnesota. Whole-tree harvest had no statistically significant effects on initial (2-year) regeneration densities, including root suckers, sprouts, and seedlings relative to conventional, stem-only harvest. The density of shrub stems was also unaffected by harvest residue retention. Despite having a lower mean leaf area index than intact forest controls, aggregates maintained comparable densities of the four most common tree species, individually, as well as all tree species combined. The composition of regeneration within aggregates differed from surrounding harvested areas as expected, but this increase in complexity at the stand scale was achieved without diminishing P. tremuloides densities in the edge area (0-5 m) surrounding aggregates 2 years after harvest. These initial findings suggest even small aggregates of overstory reserves may achieve basic objectives related to structural complexity and sustaining shade-tolerant tree species in harvested units without compromising regeneration objectives for less tolerant species. [ABSTRACT FROM AUTHOR]

Retention harvesting, an approach that intentionally retains legacy features such as mature overstory trees, provides options for achieving ecological objectives. At the same time, retained overstory trees may compete with the nearby recovering understory for resources, and much remains to be learned about potential trade-offs with regeneration objectives, particularly over extended time periods. We assessed the influence of aggregated retention (reserved mature overstory and understory patches) versus intact forest on structure and productivity (standing biomass) of the adjacent woody understory and regeneration 12 years after harvest in northern Minnesota, USA. Each site was dominated by Populus tremuloides Michx., a species that regenerates prolifically via root sprouts following disturbance. Overall, fewer differences than expected occurred between the effects of intact forest and aggregated retention on regeneration, despite the small size (0.1 ha) of aggregates. Instead, harvest status and distance from harvest edge had a greater influence on structure and standing woody biomass. Proximity to aggregates reduced large sapling biomass (all species, combined) relative to open conditions, but only up to 5 m into harvested areas. This suggests the trade-off for achieving productivity objectives might be minimal if managers use retention aggregates in this region to achieve ecological objectives and meet management guidelines. [ABSTRACT FROM AUTHOR]

The continual emergence of pathogen resistance is a recurring challenge and pushes for the development of antimicrobial compounds. Here, we investigated compounds from quaking aspen trees (Populus tremuloides) as potential antimicrobial agents. Several extractions using different solvents were realized, and corresponding antimicrobial activity was tested against eight microorganisms. Results revealed that polar extraction solvents including water, ethanol and methanol gave the best extraction yields (>15.07%). Minimal inhibition concentration (MIC) and minimal bactericidal/fungicidal concentration (MBC/MFC) demonstrated that water extracts had the best antimicrobial activity by a weak to moderate inhibition of growth of all eight tested microorganisms in addition to having a bactericidal effect on three of them. The quaking aspen methanol extract also displayed antimicrobial activity but to a lower level than the water extract. Ultra-performance liquid chromatography quadrupole time-of flight mass spectrometry (UPLC-QTOF-MS) analysis led to the identification of 92 compounds, mainly polyphenols in both extracts, with 22 molecules previously known for their antimicrobial properties. According to the relative abundance, 4-hydroxybenzaldehyde (5.44% in methanol extract) and kaempferol (5.03% in water extract) were the most abundant antimicrobial compounds. Among antimicrobial molecules identified, nine were from the flavonoid family. The results of our study demonstrate the interest of using quaking aspen as source of antimicrobial compounds.

• High variability in aspen stands makes fire activity in aspen difficult to predict. • Pure stands with herbaceous understories are more likely to reduce fire behavior. • Aspen can burn easily and intensely in windy, dry, or certain seasonal conditions. • Specific management guidelines for aspen fire risk reduction treatments are needed. Quaking aspen (Populus tremuloides Michx.) stands have historically been referred to as "firebreak" forest types that can reduce fire activity, but high-intensity and high-severity fires have been observed to burn through aspen stands. Clearly, fire activity in aspen is highly variable, which may be due to the wide variation in aspen stand composition and structure and because the species occurs across wide geographic, environmental, and climatic gradients. In the western U.S., there is growing interest in promoting aspen stands within wildland-urban interface communities to reduce fire risk, but studies that refer to the low flammability of aspen stands rely on limited citations. If promoting aspen to reduce fire risk is a desirable forest management practice, consolidating the available literature is necessary to understand when, where, and how management might achieve this goal. Here, we synthesized literature and conducted a survey of forest and fire managers to assess current understanding of how fire interacts with aspen stands, as well as to examine possible factors that influence fire occurrence, behavior, and severity in aspen communities. We found evidence that the presence of aspen reduces fire occurrence, fire behavior, and fire severity, but this effect is dependent on many factors, including the percentage of aspen vs conifers in the overstory, load and type of understory fuels, weather, and season. We did not find any quantitative management guidelines on how to create, maintain, or use aspen stands to reduce fire risk. The large gap between "common knowledge" and empirical evidence regarding aspen's ability to inhibit fire requires further research. [ABSTRACT FROM AUTHOR]

Climatic conditions exert important control over the growth, productivity, and distribution of forests, and characterizing these relationships is essential for understanding how forest ecosystems will respond to climate change. We used dendrochronological methods to develop climate-growth relationships for two dominant species, Populus tremuloides (quaking aspen) and Pinus resinosa (red pine), in the upper Great Lakes region to understand how climate and water availability influence annual forest productivity. Trees were sampled along a topographic gradient at the Marcell Experimental Forest (Minnesota, USA) to assess growth response to variations in temperature and different water availability metrics (precipitation, potential evapotranspiration (PET), cumulative moisture index (CMI), and soil water storage). Climatic variables were able to explain 33-58% of the variation in annual growth (as measured by ring-width increment) for quaking aspen and 37-74% of the variation for red pine. Climate-growth relationships were influenced by topography for quaking aspen but not for red pine. Annual ring growth for quaking aspen decreased with June CMI on ridges, decreased with temperature in the November prior to the growing season on sideslopes, and decreased with June PET on toeslopes. Red pine growth increased with increasing July PET across all topographic positions. These results indicate the sensitivity of both quaking aspen and red pine to local climate and show several vulnerabilities of these species to shifts in water supply and temperature because of climate change. Copyright © 2015 John Wiley & Sons, Ltd. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Mixed-severity fire regimes may be the most extensive yet poorly understood fire regimes of western North America. Understanding their long-term spatiotemporal dynamics is central to debates regarding altered fire regimes and the need for restoration in the context of changing climate and nearly a century of active fire suppression. However, the complexity of fire patterns and forest stand and landscape structures characteristic of mixed-severity regimes poses a substantial challenge to understanding their long-term dynamics. In this study, we develop analysis methods aimed at understanding the fire-driven forest dynamics of mixed-severity systems and apply them in the San Juan Mountains of southwestern Colorado. We sampled fire scars, stand structure, and >4300 tree ages across two 1340-ha landscapes (Williams Creek and Squaretop Mountain) that span the environmental gradient of montane mixed-conifer and aspen forests. New approaches were applied to identify pulses of tree recruitment, evaluate climate and fire as potential drivers of synchronous recruitment pulses, and combine fire scar and recruitment data to reconstruct fires. The reconstructions provided detailed fire history for each stand, which in turn was used to develop a fire-severity metric, compare fire frequency and severity by forest type, and develop a simulation procedure to evaluate the degree to which tree regeneration depended on fire by species within each forest type. Twenty fires were reconstructed since 1685 at Williams Creek and 13 fires since 1748 at Squaretop Mountain. Patterns of fire severity varied within each fire and over successive events, including high-severity patches of hundreds of hectares in both study areas. Dry mixed-conifer forests experienced relatively short fire intervals (mean 21 years) and low fire severity, and regeneration of the main conifer species was largely dependent on open conditions sustained over successive fires. Moist mixed-conifer forests experienced longer fire intervals (mean 32 years) and a broader range of severities, and fire-caused canopy openings were important for initiating pulses of tree recruitment. Most (83%) aspen stands included two or more post-fire cohorts. The methods presented here can be adapted to other mixed-severity systems to better understand their long-term spatial and temporal dynamics and develop restoration priorities. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Content: Changing aspen distribution in response to climate change and fire is a major focus of biodiversity conservation, yet little is known about the potential response of aspen to these two driving forces along topoclimatic gradients. Objective: This study is set to evaluate how aspen distribution might shift in response to different climate-fire scenarios in a semi-arid montane landscape, and quantify the influence of fire regime along topoclimatic gradients. Methods: We used a novel integration of a forest landscape succession and disturbance model (LANDIS-II) with a fine-scale climatic water deficit approach to simulate dynamics of aspen and associated conifer and shrub species over the next 150 years under various climate-fire scenarios. Results: Simulations suggest that many aspen stands could persist without fire for centuries under current climate conditions. However, a simulated 2-5 °C increase in temperature caused a substantial reduction of aspen coverage at lower elevations and a modest increase at upper elevations, leading to an overall reduction of aspen range at the landscape level. Increasing fire activity may favor aspen increase at its upper elevation limits adjacent to coniferous forest, but may also favor reduction of aspen at lower elevation limits adjacent to xeric shrubland. Conclusions: Our study highlights the importance of incorporating fine-scale terrain effects on climatic water deficit and ecohydrology when modeling species distribution response to climate change. This modeling study suggests that climate mitigation and adaptation strategies that use fire would benefit from consideration of spatial context at landscape scales. [ABSTRACT FROM AUTHOR]

To investigate the extent and causes of recent quaking aspen (Populus tremuloides) recruitment in northern Yellowstone National Park, we measured browsing intensity and height of young aspen in 87 randomly selected aspen stands in 2012, and compared our results to similar data collected in 1997-1998. We also examined the relationship between aspen recovery and the distribution of Rocky Mountain elk (Cervus elaphus) and bison (Bison bison) on the Yellowstone northern ungulate winter range, using ungulate fecal pile densities and annual elk count data. In 1998, 90% of young aspen were browsed and none were taller than 200 cm, the height at which aspen begin to escape from elk browsing. In 2012, only 37% in the east and 63% in the west portions of the winter range were browsed, and 65% of stands in the east had young aspen taller than 200 cm. Heights of young aspen were inversely related to browsing intensity, with the least browsing and greatest heights in the eastern portion of the range, corresponding with recent changes in elk density and distribution. In contrast with historical elk distribution (1930s-1990s), the greatest densities of elk recently (2006-2012) have been north of the park boundary (~5 elk/km²), and in the western part of the range (2-4 elk/km²), with relatively few elk in the eastern portion of the range (<2 elk/km2), even in mild winters. This redistribution of elk and decrease in density inside the park, and overall reduction in elk numbers, explain why many aspen stands have begun to recover. Increased predation pressure following the reintroduction of gray wolves (Canis lupus) in 1995-1996 played a role in these changing elk population dynamics, interacting with other influences including increased predation by bears (Ursus spp.), competition with an expanding bison population, and shifting patterns of human land use and hunting outside the park. The resulting new aspen recruitment is evidence of a landscape-scale trophic cascade in which a resurgent large carnivore community, combined with other ecological changes, has benefited aspen through effects on ungulate prey. [ABSTRACT FROM AUTHOR]

Because forest stand structure, age, and productivity can mediate the impacts of climate on quaking aspen (Populus tremuloides) mortality, ignoring stand-scale factors limits inference on the drivers of recent sudden aspen decline. Using the proportion of aspen trees that were dead as an index of recent mortality at 841 forest inventory plots, we examined the relationship of this mortality index to forest structure and climate in the Rocky Mountains and Intermountain Western United States. We found that forest structure explained most of the patterns in mortality indices, but that variation in growing-season vapor pressure deficit and winter precipitation over the last 20 years was important. Mortality index sensitivity to precipitation was highest in forests where aspen exhibited high densities, relative basal areas, quadratic mean diameters, and productivities, whereas sensitivity to vapor pressure deficit was highest in young forest stands. These results indicate that the effects of drought on mortality may be mediated by forest stand development, competition with encroaching conifers, and physiological vulnerabilities of large trees to drought. By examining mortality index responses to both forest structure and climate, we show that forest succession cannot be ignored in studies attempting to understand the causes and consequences of sudden aspen decline. [ABSTRACT FROM AUTHOR]

Widespread dieback of aspen forests, sometimes called sudden aspen decline, has been observed throughout much of western North America, with the highest mortality rates in the southwestern United States. Recent aspen mortality has been linked to drought stress and elevated temperatures characteristic of conditions expected under climate change, but the role of individual aspen tree growth patterns in contributing to recent tree mortality is less well known. We used tree-ring data to investigate the relationship between an individual aspen tree's lifetime growth patterns and mortality. Surviving aspen trees had consistently higher average growth rates for at least 100 years than dead trees. Contrary to observations from late successional species, slow initial growth rates were not associated with a longer lifespan in aspen. Aspen trees that died had slower lifetime growth and slower growth at various stages of their lives than those that survived. Differences in average diameter growth between live and dead trees were significant ( α = 0.05) across all time periods tested. Our best logistical model of aspen mortality indicates that younger aspen trees with lower recent growth rates and higher frequencies of abrupt growth declines had an increased risk of mortality. Our findings highlight the need for species-specific mortality functions in forest succession models. Size-dependent mortality functions suitable for late successional species may not be appropriate for species with different life history strategies. For some early successional species, like aspen, slow growth at various stages of the tree's life is associated with increased mortality risk. [ABSTRACT FROM AUTHOR]

Heavy and repeated ungulate browsing on reproductive suckers has limited quaking aspen [Populus tremuloides Michx.) regeneration on many Western landscapes. However, little is known about the specific effects of season and intensity of browsing. The objectives of this study were to determine the effects of season and intensity of clipping (simulated browsing) on density and growth characteristics of suckers. Three randomly selected stands were clear-felled in mid-July 2005 and fenced. Simulated browsing intensities of 0, 20, 40, and 60% removal of the current year's growth on aspen suckers were randomly applied in early, mid, and late summers of 2006 and 2007 on permanently demarcated quadrats. Sucker density and sucker height were monitored in each quadrat. Changes in annual sucker numbers for the ends of 2006 and 2007 were compared with declines found that reflected both winter and summer (2007 only) mortality. Mortality was not found for suckers clipped in early summer compared with mortalities of 48 and 46% for mid and late summer clipped suckers, respectively. However, even at the highest mortality, there were still ample numbers (∼25,000/ha) of suckers for stand regeneration. At the end of the study, sucker height was reduced by all summer treatment intensities. After adjustment for the controls, clipping (of current year's growth) at 20 and 40% in mid and late summer resulted in suckers that were 47 and 41 cm tall, respectively. However, clipping at 60% resulted in suckers that were only 20 cm tall. Application of information found in this study to new areas under similar circumstances suggests that browsing of terminal leaders at intensities ⩽40% in midsummer for 1 or 2 years would not have a substantial impact on aspen regeneration. [ABSTRACT FROM AUTHOR]