ALBERT

Description: Variable retention harvest systems are encouraged to promote complexity in managed forests, and aggregated retention has been suggested as a means of reducing moisture stress in residual trees. We studied the impacts of within-aggregate position on growth and foliar physiology to better understand the spatial dynamics of residual-tree responses to aggregated retention harvests in even-aged Pinus resinosa Ait. stands. Distance from edge and edge aspect influenced radial growth, volume increment, and growth efficiency, but only edge aspect affected foliar nitrogen content. Spatial variables had no significant relationships with foliar carbon isotope ratios (δ13C). Increases in radial growth, volume increment, and growth efficiency following harvesting were greatest near edges and in the northeastern quadrants of aggregates that received mechanical understory release treatments, and lowest in the southeastern quadrant of aggregates and near aggregate centers. Foliar nitrogen content was highest in the southwestern quadrants of aggregates that received understory release treatments, and lowest in the northwestern quadrants of aggregates. Our results suggest spatial relationships are important determinants of residual-tree responses to aggregated retention harvests.

Description: Emerald ash borer (EAB; Agrilus planipennis Fairmaire, 1888) has been a persistent disturbance for ash forests in the United States since 2002. Of particular concern is the impact that EAB will have on the ecosystem functioning of wetlands dominated by black ash (Fraxinus nigra Marsh.). In preparation, forest managers need reliable and complete maps of black ash dominated stands. Traditionally, forest survey data from the United States Forest Inventory and Analysis (FIA) Program have provided rigorous measures of tree species at large spatial extents but are limited when providing estimates for smaller management units (e.g., stands). Fortunately, geospatial data can extend forest survey information by generating predictions of forest attributes at scales finer than those of the FIA sampling grid. In this study, geospatial data were integrated with FIA data in a randomForest model to estimate and map black ash dominated stands in northern Minnesota in the United States. The model produced low error rates (overall error = 14.5%; area under the curve (AUC) = 0.92) and was strongly informed by predictors from soil saturation and phenology. These results improve upon FIA-based spatial estimates at national extents by providing forest managers with accurate, fine-scale maps (30 m spatial resolution) of black ash stand dominance that could ultimately support landscape-level EAB risk and vulnerability assessments.

Description: Characterizing the spatial distribution of tree mortality is critical to understanding forest dynamics, but empirical studies on these patterns under old-growth conditions are rare. This rarity is due in part to low mortality rates in old-growth forests, the study of which necessitates long observation periods, and the confounding influence of tree in-growth during such time spans. Here, we studied mortality of red pine ( Pinus resinosa Ait.) in five old-growth stands in Minnesota, USA, demonstrating the use of preexisting information of cohort age structures to account for in-growth after the most recent cohort establishment. Analyses of spatial point patterns, using both Ripley’s K-function and the pair correlation function, showed that tree mortality was essentially a random process, without evidence of contagious mortality patterns that are often expected for old-growth forests. Our analyses further demonstrated in practice that the distribution of dead trees may differ from that of the tree mortality events, which are constrained to occur within the initial distribution, and how mortality patterns can shape the spatial distribution of mature living trees, often attributed to aggregated regeneration patterns. These findings emphasize the need to disentangle the influence of the initial distribution of trees from that of actual tree mortality events.

Description: We studied longleaf pine (Pinuspalustris Mill.) ecosystems to determine causes and rates of overstory mortality, size of canopy disturbances, and the effects of disturbance on canopy structure. Further, we used redundancy analysis to relate variation in characteristics of mortality across a landscape to site and stand variables. We analyzed mortality that occurred from 1990 to 1994 in 70 second-growth plots that spanned a range of site conditions and stand structures, and in five large disturbances that occurred outside the random sample of plots. Half of pine mortality over 5 years in the 70 plots was from unknown causes. Lightning was the primary identifiable cause of mortality, followed by suppression and wind. Lightning mortality was most frequent on xeric sites, while windthrow was common on wet–mesic sites. Suppression mortality was frequent on wet–mesic sites and in higher density stands. Five-year mortality rates averaged 2.3 trees/ha, or 1.9% of original density. Most mortality consisted of single trees. Large disturbances (mostly from lightning) of 15–30 trees were rare, occurring once per 1000 ha in 5 years. Variation in amount of mortality and size of disturbance were unrelated to soil or stand structural characteristics. Low mortality rates from small-scale disturbances result in slow canopy turnover. These results indicate that large openings sufficient for natural regeneration of longleaf pine develop slowly in the absence of hurricanes. Silvicultural options for longleaf pine can be designed to mimic the canopy structure that results from natural canopy disturbances, which leave many live trees standing. Such options may be desirable if a goal of silviculture is to increase structural complexity in stands managed for timber.

Description: Understanding natural mortality patterns and processes of forest tree species is increasingly important given projected changes in mortality owing to global change. With this need in mind, the rate and spatial pattern of mortality was assessed over an 89-year period in a natural-origin Pinus resinosa (Aiton)-dominated system to assess these processes through advanced stages of stand development (stand age 120–209 years). Average annual mortality rates fluctuated through time, yet were within the range reported in other studies (0.60%–3.88% depending on species and sampling interval). Tree mortality was attributed to multiple agents, including the senescence of the short-lived Pinus banksiana Lamb., windthrow, root-rot fungi (Armillaria ostoyae (Romagn.) Herink), and perhaps infrequent droughts. Despite the often contagious nature of many disturbance agents, the overall spatial pattern of mortality events (the arrangement of dead trees within the fixed initial population of live trees) was random at all scales tested. Similarly, the current spatial pattern of dead trees was predominantly random, despite clustering at small scales (2–4 m). These findings underscore the importance of studying mortality rates, agents, and spatial patterns over long time periods to avoid misinterpreting stochastic mortality events, and their influence on longer term stand structure and development.

Description: Longleaf pine - wiregrass (Pinus palustris Mill. - Aristida stricta Michx.) woodlands occupy sites ranging from deep, xeric sandhills to the edge of wetlands in the southeastern United States. Aboveground net primary productivity (ANPP) of the overstory and understory were determined for three replicate sites of three site types (xeric, intermediate, and wet-mesic) that span a wide environmental gradient. In addition, soil moisture (at 30 and 90 cm) and N mineralization (in situ buried bag incubations) were measured through an annual cycle. Longleaf pine - wiregrass ecosystems varied by nearly twofold in ANPP across complex gradients. Overstory and understory and total (overstory and understory) ANPP were positively correlated to soil moisture at 30 and 90 cm. The proportion of understory ANPP relative to the total ANPP did not increase across the environmental gradient as predicted by hypotheses that invoke niche differentiation in rooting habits of grasses and trees. Contrary to expectations, cumulative net N mineralization was negatively related to soil moisture. All ANPP estimates were significantly and negatively related to cumulative N-mineralization. Further work is needed to explore the mechanisms by which soil moisture regulates productivity across space, time, and for individual species. Additional experimentation through resource addition would allow for investigations into multiple resource limitations and how resource limitations vary depending on gradient position.

Description: Gradient studies of wetland forests have inferred that competition from upland tree species confines waterlogging-tolerant tree species to hydric environments. Little is known, however, about competition effects on individual-tree growth along stress gradients in wetland forests. We investigated tree growth and competition in mixed-species stands representing a waterlogging stress gradient in Fraxinus nigra Marsh. (black ash) forests in Minnesota, USA. Using competition indices, we examined how F. nigra basal area increment (BAI) responded to competition along the gradient and whether competition was size-asymmetric (as for light) or size-symmetric (as for soil resources). We modeled spatial distributions of F. nigra and associated tree species to assess how variation in species mixtures influenced competition. We found that although F. nigra BAI did not significantly differ with variations in site moisture, the importance of competition decreased as waterlogging stress increased. Competition across the gradient was primarily size-asymmetric (for light). Variation in species mixtures along the gradient was an important influence on competition. Some segregation of tree species occurred at all but the most upland site, where waterlogging stress was lowest and evidence of competition was greatest, confirming that competition from upland tree species confines F. nigra and potentially other waterlogging-tolerant species to hydric environments.

Description: Black ash wetlands are seriously threatened because of the invasive emerald ash borer (EAB). Wetland hydrology is likely to be modified following ash mortality, but the magnitude of hydrological impact following loss via EAB and alternative mitigation harvests is not clear. Our objective was to assess the water table response to simulated EAB and harvesting to determine if management actions will be needed to maintain ecosystem functions following EAB infestation. We applied four replicated treatments to 1.6 ha plots as follows: (1) control, (2) girdling of all black ash trees to simulate loss via EAB mortality, (3) group selection harvests (20% of stand in 0.04 ha gaps), and (4) clear-cut harvest. Water table (WT) elevations were monitored for 1 year pre-treatment and two years post-treatment. Clear-cutting delayed WT drawdown in both years of the study, and the WT was significantly higher than the control treatment, predominantly when WT depth was below 30 cm. The effect of the group selection treatment on WT response was muted compared to clear-cutting and also limited to periods when the WT depth was below 30 cm. These responses were attributed to establishment of shallow-rooted vegetation in cut areas, which would have limited influence on WT dynamics as depth increased. There was little effect of girdling on the WT in the first year post-treatment, but effects on the WT were very similar to clear-cutting in the second year and more pronounced when the WT was within 30 cm of the soil surface. These effects were attributed to reduced transpiration coupled with the presence of a partial canopy following girdling, which would have reduced vegetation establishment and evaporation compared to clear-cutting. Given the large influence of WT depth on vegetation dynamics and associated feedbacks to altered hydrology, these early results indicate a greater risk of ecosystem alteration following EAB mortality compared to clear-cut harvesting. Depending on local hydrologic regime, variation in precipitation patterns, and time for complete canopy loss, it may be necessary for managers to implement active mitigation strategies (e.g., group selection coupled with planting of alternative species) prior to EAB infestation to maintain ecosystem processes in these forested wetland systems.

Description: Growth dominance is a relatively new, simple, quantitative metric of within-stand individual tree growth patterns, and is defined as positive when larger trees in the stand display proportionally greater growth than smaller trees, and negative when smaller trees display proportionally greater growth than larger trees. We examined long-term silvicultural experiments in red pine ( Pinus resinosa Ait.) to characterize how stand age, thinning treatments (thinned from above, below, or both), and stocking levels (residual basal area) influence stand-level growth dominance through time. In stands thinned from below or from both above and below, growth dominance was not significantly different from zero at any age or stocking level. Growth dominance in stands thinned from above trended from negative at low stocking levels to positive at high stocking levels and was positive in young stands. Growth dominance in unthinned stands was positive and increased with age. These results suggest that growth dominance provides a useful tool for assessing the efficacy of thinning treatments designed to reduce competition between trees and promote high levels of productivity across a population, particularly among crop trees.