Forest density intensifies the importance of snowpack to growth in water-limited pine forests.

Warming climate and resulting declines in seasonal snowpack have been associated with drought stress and tree mortality in seasonally snow-covered watersheds worldwide. Meanwhile, increasing forest density has further exacerbated drought stress due to intensified tree-tree competition. Using a uniquely detailed data set of population-level forest growth (n = 2,495 sampled trees), we examined how inter-annual variability in growth relates to snow volume across a range of forest densities (e.g., competitive environments) in sites spanning a broad aridity gradient across the United States. Forest growth was positively related to snowpack in water-limited forests located at low latitude, and this relationship was intensified by forest density. However, forest growth was negatively related to snowpack in a higher latitude more energy-limited forest, and this relationship did not interact with forest density. Future reductions in snowpack may have contrasting consequences, as growth may respond positively in energy-limited forests and negatively in water-limited forests; however, these declines may be mitigated by reducing stand density through forest thinning.

Retention forestry influences understory diversity and functional identity.

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.

Assessing the ecological impacts of biomass harvesting along a disturbance severity gradient.

Disturbance is a central driver of forest development and ecosystem processes with variable effects within and across ecosystems. Despite the high levels of variation in disturbance severity often observed in forests following natural and anthropogenic disturbance, studies quantifying disturbance impacts often rely on categorical classifications, thus limiting opportunities to examine potential gradients in ecosystem response to a given disturbance or management regime. Given the potential increases in disturbance severity associated with global change, as well as shifts in management regimes related to procurement of biofuel feedstocks, there is an increasing need to quantitatively describe disturbance severity and associated responses of forest development, soil processes, and structural conditions. This study took advantage of two replicated large-scale studies of forest biomass harvesting in Populus tremuloides and Pinus bansksiana forests, respectively, to develop and test the utility of a continuous, quantitative, disturbance severity index (DSI) for describing postharvest response of plant communities and nutrient pools to different levels of biomass removal and legacy retention (i.e., live trees and coarse woody material). There was a high degree of variability in DSI within categorical treatments associated with different levels of legacy retention and regression models using DSI as a predictor explained a portion of the variation (>50%) for many of the ecosystem- and community-level responses to biomass harvesting examined. Nutrient losses associated with biomass harvesting were positively related to disturbance severity, particularly in P. tremuloides forests, with postharvest nutrient availability generally declining along the gradient of impacts. Consistent with expectations from ecological theory, species richness and diversity of woody plant communities were greatest at intermediate disturbance severities and regeneration densities of dominant trees species were most abundant at highest levels of disturbance. Although categorical benchmarks will continue to be the primary way through which management guidelines are conveyed to practitioners, evaluation of their effectiveness at sustaining ecosystem functioning should be through continuous analyses, such as the DSI approach used in this study, to allow for the more precise identification of thresholds that ensure a range of desirable outcomes exist across managed landscapes.

Forested versus herbaceous wetlands: Can management mitigate ecohydrologic regime shifts from invasive emerald ash borer?

Wetlands self-organize through reciprocal controls between vegetation and hydrology, but external disturbance may disrupt these feedbacks with consequent changes to ecosystem state. Imminent and widespread emerald ash borer (EAB) infestation throughout North American forested wetlands has raised concern over possible ecosystem state shifts (i.e., wetter, more herbaceous systems) and loss of forest function, calling for informed landscape-scale management strategies. In response, we employed a large-scale manipulative study to assess the ecohydrologic response of black ash wetlands to three alternative EAB management strategies: 1) a do-nothing approach (i.e., simulated EAB infestation via tree girdling), 2) a preemptive, complete harvesting approach (i.e., clearcut), and 3) an overstory replacement approach via group selection. We analyzed six years of daily water table and evapotranspiration (ET) dynamics in six blocks comprising black ash wetlands (controls) and management strategy treatments to quantify potential for hydrologic change and subsequent recovery. In both the do-nothing approach and complete harvesting approach, we found persistent changes in hydrologic regime defined by shallower water tables and lower ET rates coupled with increased herbaceous vegetation growth, indicating ecosystem state shifts driven by vegetation-water table interactions. The do-nothing approach showed the least hydrologic recovery after five years, which we attribute to reduction in overstory transpiration as well as greater shade (via standing dead trees) that reduces open water evaporation and herbaceous layer transpiration compared to complete harvesting. We found no evidence of ecohydrologic disturbance in the overstory replacement approach, highlighting its potential as a management strategy to preserve forested wetland habitat if periodically executed over time before EAB infestation. Although the scale of potential disturbance is daunting, our findings provide a baseline assessment for forest managers to develop preemptive mitigation strategies to address the threat of EAB to ecological functions in black ash wetlands.

A cross-continental comparison of plant and beetle responses to retention of forest patches during timber harvest.

Timber harvest can adversely affect forest biota. Recent research and application suggest that retention of mature forest elements (retention forestry), including unharvested patches (or aggregates) within larger harvested units, can benefit biodiversity compared to clearcutting. However, it is unclear whether these benefits can be generalized among the diverse taxa and biomes in which retention forestry is practiced. Lack of comparability in methods for sampling and analyzing responses to timber harvest and edge creation presents a challenge to synthesis. We used a consistent methodology (similarly spaced plots or traps along transects) to investigate responses of vascular plants and ground-active beetles to aggregated retention at replicate sites in each of four temperate and boreal forest types on three continents: Douglas-fir forests in Washington, USA; aspen forests in Minnesota, USA; spruce forests in Sweden; and wet eucalypt forests in Tasmania, Australia. We assessed (1) differences in local (plot-scale) species richness and composition between mature (intact) and regenerating (previously harvested) forest; (2) the lifeboating function of aggregates (capacity to retain species of unharvested forest); and whether intact forests and aggregates (3) are susceptible to edge effects and (4) influence the adjacent regenerating forest. Intact and harvested forests differed in composition but not richness of plants and beetles. The magnitude of this difference was generally similar among regions, but there was considerable heterogeneity of composition within and among replicate sites. Aggregates within harvest units were effective at lifeboating for both plant and beetle communities. Edge effects were uncommon even within the aggregates. In contrast, effects of forest influence on adjacent harvested areas were common and as strong for aggregates as for larger blocks of intact forest. Our results provide strong support for the widespread application of aggregated retention in boreal and temperate forests. The consistency of pattern in four very different regions of the world suggests that, for forest plants and beetles, responses to aggregated retention are likely to apply more widely. Our results suggest that through strategic placement of aggregates, it is possible to maintain the natural heterogeneity and biodiversity of mature forests managed for multiple objectives.

Biomass growth response to spatial pattern of variable­retention harvesting in a northern Minnesota pine ecosystem.

Variable-retention harvesting (VRH) is an approach for sustaining complex structure in managed forests. A criticism of VRH is that ecological benefits may come at a cost of reduced growth of regeneration, due to competition with residual trees. However, the spatial pattern of retention, i.e., dispersed or aggregated, in VRH systems can be manipulated to minimize suppression of regeneration, and resource limitation to regeneration might be mitigated by reduction of woody shrubs. Continued growth of the residual cohort will compensate for growth reduction of regeneration, although this may differ with retention pattern. We examined aboveground whole-stand biomass growth of trees in a VRH experiment in Pinus resinosa forest in Minnesota, USA. Treatments included dispersed retention, aggregated retention, and an uncut control, as well as a shrub treatment (reduced density or ambient). We addressed the following hypotheses: (1) biomass growth of a cohort of planted pine seedlings will be highest with aggregated rather than dispersed retention, (2) biomass growth of the planted seedlings will increase with shrub reduction, and (3) biomass growth of the residual overstory will be higher with dispersed rather than aggregated retention. Aboveground biomass growth of the planted pines ranged from 0.4 kg·ha−1·yr−1 in the overstory-control­ambient-shrub treatment to 23 kg·ha−1·yr−1 in the aggregated-retention­shrub-reduction treatment. The difference between the control and the retention treatments was significant (P < 0.0001), but not between dispersed and aggregated retention (P = 0.97). Thus, our first hypothesis was not supported. In all treatments, biomass growth was significantly higher (>100% increase) with shrub reduction (P = 0.001), supporting our second hypothesis. Biomass growth of residual trees ranged from 2404 kg·ha−1·yr−1 in the uncut-control­ambient-shrub treatment to 1043 kg·ha−1·yr−1 in the aggregated-retention­shrub-reduction treatment. Differences were significant between the control and retention treatments (P = 0.003), and marginally higher with dispersed vs. aggregated retention (P = 0.09), lending support to our third hypothesis. Our results suggest that managers have flexibility in application of VRH and can expect similar stand-level biomass growth of planted regeneration regardless of retention pattern, but somewhat higher stand-level biomass growth of retained trees with dispersed retention.

Winning and losing tree species of reassembly in Minnesota's mixed and broadleaf forests.

We examined reassembly of winning and losing tree species, species traits including shade and fire tolerance, and associated disturbance filters and forest ecosystem types due to rapid forest change in the Great Lakes region since 1850. We identified winning and losing species by changes in composition, distribution, and site factors between historical and current surveys in Minnesota's mixed and broadleaf forests. In the Laurentian Mixed Forest, shade-intolerant aspen replaced shade-intolerant tamarack as the most dominant tree species. Fire-tolerant white pine and jack pine decreased, whereas shade-tolerant ashes, maples, and white cedar increased. In the Eastern Broadleaf Forest, fire-tolerant white oaks and red oaks decreased, while shade-tolerant ashes, American basswood, and maples increased. Tamarack, pines, and oaks have become restricted to sites with either wetter or sandier and drier soils due to increases in aspen and shade-tolerant, fire-sensitive species on mesic sites. The proportion of shade-tolerant species increased in both regions, but selective harvest reduced the applicability of functional groups alone to specify winners and losers. Harvest and existing forestry practices supported aspen dominance in mixed forests, although without aspen forestry and with fire suppression, mixed forests will transition to a greater composition of shade-tolerant species, converging to forests similar to broadleaf forests. A functional group framework provided a perspective of winning and losing species and traits, selective filters, and forest ecosystems that can be generalized to other regions, regardless of species identity.

Effects of thinning on drought vulnerability and climate response in north temperate forest ecosystems.

Reducing tree densities through silvicultural thinning has been widely advocated as a strategy for enhancing resistance and resilience to drought, yet few empirical evaluations of this approach exist. We examined detailed dendrochronological data from a long-term (> 50 years) replicated thinning experiment to determine if density reductions conferred greater resistance and/or resilience to droughts, assessed by the magnitude of stand-level growth reductions. Our results suggest that thinning generally enhanced drought resistance and resilience; however, this relationship showed a pronounced reversal over time in stands maintained at lower tree densities. Specifically, lower-density stands exhibited greater resistance and resilience at younger ages (49 years), yet exhibited lower resistance and resilience at older ages (76 years), relative to higher-density stands. We attribute this reversal to significantly greater tree sizes attained within the lower-density stands through stand development, which in turn increased tree-level water demand during the later droughts. Results from response-function analyses indicate that thinning altered growth-climate relationships, such that higher-density stands were more sensitive to growing-season precipitation relative to lower-density stands. These results confirm the potential of density management to moderate drought impacts on growth, and they highlight the importance of accounting for stand structure when predicting climate-change impacts to forests.

Pseudoabsence generation strategies for species distribution models.

BACKGROUND: Species distribution models require selection of species, study extent and spatial unit, statistical methods, variables, and assessment metrics. If absence data are not available, another important consideration is pseudoabsence generation. Different strategies for pseudoabsence generation can produce varying spatial representation of species. METHODOLOGY: We considered model outcomes from four different strategies for generating pseudoabsences. We generating pseudoabsences randomly by 1) selection from the entire study extent, 2) a two-step process of selection first from the entire study extent, followed by selection for pseudoabsences from areas with predicted probability <25%, 3) selection from plots surveyed without detection of species presence, 4) a two-step process of selection first for pseudoabsences from plots surveyed without detection of species presence, followed by selection for pseudoabsences from the areas with predicted probability <25%. We used Random Forests as our statistical method and sixteen predictor variables to model tree species with at least 150 records from Forest Inventory and Analysis surveys in the Laurentian Mixed Forest province of Minnesota. CONCLUSIONS: Pseudoabsence generation strategy completely affected the area predicted as present for species distribution models and may be one of the most influential determinants of models. All the pseudoabsence strategies produced mean AUC values of at least 0.87. More importantly than accuracy metrics, the two-step strategies over-predicted species presence, due to too much environmental distance between the pseudoabsences and recorded presences, whereas models based on random pseudoabsences under-predicted species presence, due to too little environmental distance between the pseudoabsences and recorded presences. Models using pseudoabsences from surveyed plots produced a balance between areas with high and low predicted probabilities and the strongest relationship between density and area with predicted probabilities ≥75%. Because of imperfect accuracy assessment, the best assessment currently may be evaluation of whether the species has been sufficiently but not excessively predicted to occur.

Carbon stocks across a chronosequence of thinned and unmanaged red pine (Pinus resinosa) stands.

Forests function as a major global C sink, and forest management strategies that maximize C stocks offer one possible means of mitigating the impacts of increasing anthropogenic CO2 emissions. We studied the effects of thinning, a common management technique in many forest types, on age-related trends in C stocks using a chronosequence of thinned and unmanaged red pine (Pinus resinosa) stands ranging from 9 to 306 years old. Live tree C stocks increased with age to a maximum near the middle of the chronosequence in unmanaged stands, and increased across the entire chronosequence in thinned stands. C in live understory vegetation and C in the mineral soil each declined rapidly with age in young stands but changed relatively little in middle-aged to older stands regardless of management. Forest floor C stocks increased with age in unmanaged stands, but forest floor C decreased with age after the onset of thinning around age 40 in thinned stands. Deadwood C was highly variable, but decreased with age in thinned stands. Total ecosystem C increased with stand age until approaching an asymptote around age 150. The increase in total ecosystem C was paralleled by an age-related increase in total aboveground C, but relatively little change in total belowground C. Thinning had surprisingly little impact on total ecosystem C stocks, but it did modestly alter age-related trends in total ecosystem C allocation between aboveground and belowground pools. In addition to characterizing the subtle differences in C dynamics between thinned and unmanaged stands, these results suggest that C accrual in red pine stands continues well beyond the 60-100 year management rotations typical for this system. Management plans that incorporate longer rotations and thinning in some stands could play an important role in maximizing C stocks in red pine forests while meeting other objectives including timber extraction, biodiversity conservation, restoration, and fuel reduction goals.

Evaluating the effects of alternative forest management plans under various physiographic settings using historical records as a reference.

Using historical General Land Office record as a reference, this study employed a landscape-scale disturbance and succession model to estimate the future cumulative effects of six alternative management plans on the tree species composition for various physiographic settings for the Mark Twain National Forest in Missouri. The results indicate that over a 200-year horizon, the relative abundance of black oak and pine species groups will decrease and the relative abundance of the white oak species group will increase, regardless of management strategy. General Land Office witness tree records provide a measure of tree species composition in the period from 1800 to 1850, prior to the large-scale influx of European settlers. Compared to the tree species composition described in the General Land Office records, the six contemporary management alternatives considered all would lead to a lower abundance of pine species, a higher abundance of red/black oak species, and a slightly higher abundance of white oak species after 200 years. Impacts of management on tree species composition varied with physiographic settings. The projected relative abundance of pine differed significantly across the five physiographic classes over the first 40 years of the simulation. In the medium term (simulation years 41-100) the projected relative pine abundance differed significantly among only four physiographic classes. In the long term (simulation years 100-200) the projected relative pine abundance differed for only one physiographic class. In contrast, differences among physiographic classes in the relative abundance of black oaks and white oaks increased over time. In general, the expected long-term differences in relative tree species abundance among six proposed alternative management plans are small compared to shifts in tree species composition that have occurred from 1850 to the present.

Untangling positive and negative biotic interactions: views from above and below ground in a forest ecosystem.

In ecological communities, the outcome of plant-plant interactions represents the net effect of positive and negative interactions occurring above and below ground. Untangling these complex relationships can provide a better understanding of mechanisms that underlie plant-plant interactions and enhance our ability to predict population, community, and ecosystem effects of biotic interactions. In forested ecosystems, tree seedlings interact with established vegetation, but the mechanisms and outcomes of these interactions are not well understood. To explore such mechanisms, we manipulated above- and belowground interactions among tree seedlings, shrubs, and trees and monitored seedling survival and growth of six species (Pinus banksiana, Betula papyrifera, P. resinosa, Quercus rubra, P. strobus, and Acer rubrum) in mature pine-dominated forest in northern Minnesota, USA. The forest had a moderately open canopy and sandy soils. Understory manipulations were implemented in the forest interior and in large gaps and included removal of shrubs (no interactions), tieback of shrubs (belowground), removal of shrubs with addition of shade (aboveground), and unmanipulated shrubs (both below- and aboveground). We found that shrubs either suppressed or facilitated seedling survival and growth depending on the seedling species, source of interaction (e.g., above- or belowground), and ecological context (e.g., gap or forest interior). In general, shrubs strongly influenced survival and growth in gaps, with more modest effects in the forest interior. In gaps, the presence of shrub roots markedly decreased seedling growth and survival, supporting the idea that belowground competition may be more important in dry, nutrient-poor sites. Shrub shade effects were neutral for three species and facilitative for the other three. Facilitation was more likely for shade-tolerant species. In the forest interior, shrub shade negatively affected seedling survival for the most shade-intolerant species. For several species the net effect of shrubs masked the existence of both positive and negative interactions above and below ground. Our results highlight the complexity of plant-plant interactions, demonstrate that outcomes of these interactions vary with the nature of resource limitation and the ecophysiology of the species involved, and suggest that ecological theory that rests on particular notions of plant-plant interactions (e.g., competition) should consider simultaneous positive and negative interactions occurring above and below ground.

Wood {delta}13C, {delta}18O and radial growth responses of residual red pine to variable retention harvesting.

Variable retention harvests are used to enhance the development of structural complexity in managed forests by retaining living trees and other structural legacies from the pre-harvest ecosystem. While harvesting should increase resource availability to residual trees, greater crown exposure may also increase environmental stress, which makes it difficult to predict growth in different structural environments. We used stable carbon isotope ratios (delta(13)C) of annual rings from red pine trees (Pinus resinosa Ait.) as an index of intrinsic water use efficiency (iWUE), the ratio of photosynthetic carbon assimilation (A) to stomatal conductance (g(s)), to better understand how differences in physiological performance relate to growth responses following harvests that left residuals dispersed, aggregated between small (0.1 ha) gaps or aggregated between large (0.3 ha) gaps. Stable oxygen isotope ratios (delta(18)O) were used as an index of g(s) to investigate the drivers behind changes in iWUE. Retention harvesting did not appear to affect delta(13)C or delta(18)O at the stand scale when compared to unharvested control stands, but there was a significant, negative correlation between residual tree delta(13)C and plot basal area in the second and third years after harvesting that suggests declining iWUE as overstory competition increases. Residual tree delta(18)O was similar across treatments and basal areas. Trees in variable retention harvests showed small but positive increases in radial growth from the pre-treatment to post-treatment measurement periods, while radial growth declined in unharvested control stands. There were no significant differences in radial growth among retention treatments. Our results suggest residual red pine in relatively open environments benefit from greater A but do not show evidence of changes in g(s) that would indicate altered water relations.

Integration of satellite imagery and forest inventory in mapping dominant and associated species at a regional scale.

To achieve the overall objective of restoring natural environment and sustainable resource usability, each forest management practice effect needs to be predicted using a simulation model. Previous simulation efforts were typically confined to public land. Comprehensive forest management practices entail incorporating interactions between public and private land. To make inclusion of private land into management planning feasible at the regional scale, this study uses a new method of combining Forest Inventory and Analysis (FIA) data with remotely sensed forest group data to retrieve detailed species composition and age information for the Missouri Ozark Highlands. Remote sensed forest group and land form data inferred from topography were integrated to produce distinct combinations (ecotypes). Forest types and size classes were assigned to ecotypes based on their proportions in the FIA data. Then tree species and tree age determined from FIA subplots stratified by forest type and size class were assigned to pixels for the entire study area. The resulting species composition map can improve simulation model performance in that it has spatially explicit and continuous information of dominant and associated species, and tree ages that are unavailable from either satellite imagery or forest inventory data. In addition, the resulting species map revealed that public land and private land in Ozark Highlands differ in species composition and stand size. Shortleaf pine is a co-dominant species in public land, whereas it becomes a minor species in private land. Public forest is older than private forest. Both public and private forests have deviated from historical forest condition in terms of species composition. Based on possible reasons causing the deviation discussed in this study, corresponding management avenues that can assist in restoring natural environment were recommended.

delta13C and delta18O trends across overstory environments in whole foliage and cellulose of three Pinus species.

Stable isotope ratios of carbon (delta(13)C) and oxygen (delta(18)O) are increasingly used to investigate environmental influences on plant physiology. Cellulose is often isolated for isotopic studies, but some authors have questioned the value of this process. We studied trends in delta(13)C and delta(18)O of whole foliage and holocellulose from seedlings of three Pinus species across three overstory environments to evaluate the benefits of holocellulose extraction in the context of a traditional ecological experiment. Both tissue types showed increasing delta(13)C from closed-canopy controls to thinned plots to 0.3 ha canopy gaps, and no change in delta(18)O between overstory environments. delta(13)C of P. resinosa and P. strobus was greater than delta(13)C of P. banksiana in whole foliage and holocellulose samples, and there were no differences in delta(18)O associated with species in either tissue type. Our results suggest whole foliage and holocellulose provide similar information about isotopic trends across broad environmental gradients and between species, but holocellulose may be better suited for studying differences in stable isotope composition between multiple species across several treatments.