Matthew D. Powers

Ungulate Carcasses Perforate Ecological Filters and Create Biogeochemical Hotspots in Forest Herbaceous Layers Allowing Trees a Competitive Advantage

Ecological filters and nutrient heterogeneity are important in the function of ecosystems. Herbaceous layers alter forest ecosystems by filtering tree species during early stages of tree reproduction and influencing nutrient cycling. Important aspects about how tree species successfully establish below and extend above this ecological filter are unanswered in forest ecology. We experimentally tested the effects of large ungulate carcasses on the filtering function of herbaceous layers. Even well-utilized carcasses created unexpected disturbances that reduced herbaceous cover, which effectively perforated the herbaceous layer filter that can differentially influence tree reproduction. Carcasses also created lasting biogeochemical “hotspots” in forest soils that may help maintain plant biodiversity by creating resource heterogeneity and shifting competitive relationships. Because the spatial distribution of carcasses is influenced by predators, these data establish an unrecognized link between large carnivores, prey carcasses, and ecosystem processes. This link supports a novel understanding of disturbance by large herbivores in forest ecosystems by demonstrating an important interaction between predator–prey functional traits and tree seedling dynamics on either side of a major ecological filter.

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.

Wood δ13C, δ18O 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.

Spatial dynamics of radial growth and growth efficiency in residual Pinus resinosa following aggregated retention harvesting

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 southw...

δ13C and δ18O trends across overstory environments in whole foliage and cellulose of three Pinus species

Stable isotope ratios of carbon (δ13C) and oxygen (δ18O) 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 δ13C and δ18O 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 δ13C from closed-canopy controls to thinned plots to 0.3 ha canopy gaps, and no change in δ18O between overstory environments. δ13C of P. resinosa and P. strobus was greater than δ13C of P. banksiana in whole foliage and holocellulose samples, and there were no differences in δ18O 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.

Disturbance dynamics influence Carex pensylvanica abundance in a northern hardwood forest

Abstract We examined relationships involving the abundance of Carex pensylvanica Lam., a native sedge that can form dense mats in northern hardwood understories, and three types of disturbances: forest management, deer herbivory, and exotic earthworm activity. Stands managed using even-aged silvicultural systems in areas with high deer densities (> 20 deer km−2) had greater C. pensylvanica cover than stands managed using even-aged silvicultural systems in areas with lower deer densities (10–20 deer km−2), or unharvested second-growth communities. Stands managed using uneven-aged silvicultural systems were not significantly different from any other treatment group, regardless of deer density. There was a significant management × deer × earthworm interaction, and earthworm density had a positive relationship with C. pensylvanica cover in most management history × deer density combinations. High levels of disturbance associated with at least two of the three factors considered in this study were associated with high C. pensylvanica cover, but patterns across any single variable were inconsistent. These results elucidate the intricate nature of invasive plant dynamics, and provide a baseline for more detailed studies of the causal mechanisms underlying sedge mat formation in northern hardwood forests.