Regional extent of an ecosystem engineer: earthworm invasion in northern hardwood forests.

The invasion of exotic earthworms into northern temperate and boreal forests previously devoid of earthworms is an important driver of ecosystem change. Earthworm invasion can cause significant changes in soil structure and communities, nutrient cycles, and the diversity and abundance of herbaceous plants. However, the regional extent and patterns of this invasion are poorly known. We conducted a regional survey in the Chippewa and Chequamegon National Forests, in Minnesota and Wisconsin, U.S.A., respectively, to measure the extent and patterns of earthworm invasion and their relationship to potential earthworm introduction sites. We sampled earthworms, soils, and vegetation in 20 mature, sugar maple-dominated forest stands in each national forest and analyzed the relationship between the presence of five earthworm taxonomic groups, habitat variables, and distance to the nearest potential introduction site. Earthworm invasion was extensive but incomplete in the two national forests. Four of the six earthworm taxonomic groups occurred in 55-95% of transects; however 20% of all transects were invaded by only one taxonomic group that has relatively minor ecological effects. Earthworm taxonomic groups exhibited a similar sequence of invasion found in other studies: Dendrobaena > Aporrectodea = Lumbricus juveniles > L. rubellus > L. terrestris. Distance to the nearest road was the best predictor of earthworm invasion in Wisconsin while distance to the nearest cabin was the best predictor in Minnesota. These data allow us to make preliminary assessments of landscape patterns of earthworm invasion. As an example, we estimate that 82% of upland mesic hardwood stands in the Wisconsin region are likely invaded by most taxonomic groups while only 3% are unlikely to be invaded at present. Distance to roads and cabins provides a coarse-scale predictor of earthworm invasion to focus stand-level assessments that will help forest managers better understand current and potential forest conditions and identify uninvaded areas that could serve as important refugia for plant species threatened by earthworm invasion.     

Changes in hardwood forest understory plant communities in response to European earthworm invasions

European earthworms are colonizing earthworm-free northern hardwood forests across North America. Leading edges of earthworm invasion provide an opportunity to investigate the response of understory plant communities to earthworm invasion and whether the species composition of the earthworm community influences that response. Four sugar maple-dominated forest sites with active earthworm invasions were identified in the Chippewa National Forest in north central Minnesota, USA. In each site, we established a 30 x 150 m sample grid that spanned a visible leading edge of earthworm invasion and sampled earthworm populations and understory vegetation over four years. Across leading edges of earthworm invasion, increasing total earthworm biomass was associated with decreasing diversity and abundance of herbaceous plants in two of four study sites, and the abundance and density of tree seedlings decreased in three of four study sites. Sample points with the most diverse earthworm species assemblage, independent of biomass, had the lowest plant diversity. Changes in understory plant community composition were most affected by increasing biomass of the earthworm species Lumbricus rubellus. Where L. rubellus was absent there was a diverse community of native herbaceous plants, but where L. rubellus biomass reached its maximum, the herbaceous-plant community was dominated by Carex pensylvanica and Arisaema triphyllum and, in some cases, was completely absent. Evidence from these forest sites suggests that earthworm invasion can lead to dramatic changes in the understory community and that the nature of these changes is influenced by the species composition of the invading earthworm community.     

Reduced Density and Nest Survival of Ground‐Nesting Songbirds Relative to Earthworm Invasions in Northern Hardwood Forests

Abstract: European earthworms (Lumbricus spp.) are spreading into previously earthworm‐free forests in the United States and Canada and causing substantial changes, including homogenization of soil structure, removal of the litter layer, and reduction in arthropod abundance and species richness of understory plants. Whether these changes affect songbirds that nest and forage on the forest floor is unknown. In stands with and without earthworms in the Chequamegon‐Nicolet National Forest, Wisconsin (U.S.A.), we surveyed for, monitored nests of, and measured attributes of habitat of Ovenbirds (Seiurus aurocapillus) and Hermit Thrushes (Catharus guttatus), both ground‐dwelling songbirds, and we sampled earthworms at survey points and nests. Bird surveys indicated significantly lower densities of Ovenbirds and Hermit Thrushes in relation to Lumbricus invasions at survey point and stand extents (3.1 and 15–20 ha, respectively). Modeling of Ovenbird nest survival (i.e., the probability that nestlings successfully fledge) indicated that lower survival probabilities were associated with increased sedge cover and decreased litter depth, factors that are related to Lumbricus invasions, possibly due to reduced nest concealment or arthropod abundance. Our findings provide compelling evidence that earthworm invasions may be associated with local declines of forest songbird populations.     

Susceptibility of a Northern Hardwood Forest to Exotic Earthworm Invasion

Abstract:  Numerous exotic earthworm species are colonizing northern hardwood forests of North America, where no native earthworms exist. Upon invasion, earthworms have been shown to alter the surface soil environment and plant populations and communities. We sought to identify land-use factors in the Ottawa National Forest (ONF), Michigan (U.S.A.), that contribute to earthworm invasion in forest dominated by sugar maple ( Acer saccharum Marsh.) so that the susceptibility to additional colonization could be evaluated. We sampled earthworm communities in Sylvania Wilderness Area, a unique old-growth hardwood forest, and nonwilderness sites influenced by recreational fishing, recent timber harvesting, or roads. All the nonwilderness sites contained one to five species of exotic earthworms. In contrast, only 50% of wilderness sites contained exotic earthworms, all of a single species. Nonwilderness sites also had thinner litter and duff layers, higher soil C and N content, and higher nitrogen mineralization potentials than Sylvania sites. Two central differences between Sylvania and nonwilderness sites were that all nonwilderness sites were in close contact with roads and had a history of timber harvest, whereas these factors were not present in Sylvania Wilderness Area. Using average rates of colonization, we constructed two geographic information system models to estimate the percentage of sugar maple on the ONF falling within a theoretical 100-year invasion distance of roads and of second-growth sugar maple as relative indices of susceptibility to invasion. Both models indicated high susceptibility to invasion, with 91.7% and 98.9% of sugar maple habitat falling within a theoretical 100-year invasion distance of roads or historical harvests, respectively.     

Patterns of litter disappearance in a northern hardwood forest invaded by exotic earthworms.

A field study was conducted to evaluate the effects of exotic earthworm invasions on the rates of leaf litter disappearance in a northern hardwood forest in southcentral New York, USA. Specifically, we assessed whether differences in litter quality and the species composition of exotic earthworm communities affected leaf litter disappearance rates. Two forest sites with contrasting communities of exotic earthworms were selected, and disappearance rates of sugar maple and red oak litter were estimated in litter boxes in adjacent earthworm-free, transition, and earthworm-invaded plots within each site. After 540 days in the field, 1.7-3 times more litter remained in the reference plots than in the earthworm-invaded plots. In the earthworm-invaded plots, rates of disappearance of sugar maple litter were higher than for oak litter during the first year, but by the end of the experiment, the amount of sugar maple and oak litter remaining in the earthworm-invaded plots was identical within each site. The composition of the earthworm communities significantly affected the patterns of litter disappearance. In the site dominated by the anecic earthworm Lumbricus terrestris and the endogeic Aporrectodea tuberculata, the percentage of litter remaining after 540 days (approximately 17%) was significantly less than at the site dominated by L. rubellus and Octolasion tyrtaeum (approximately 27%). This difference may be attributed to the differences in feeding behavior of the two litter-feeding species: L. terrestris buries entire leaves in vertical burrows, whereas L. rubellus usually feeds on litter at the soil surface, leaving behind leaf petioles and veins. Our results showed that earthworms not only accelerate litter disappearance rates, but also may reduce the differences in decomposition rates that result from different litter qualities at later stages of decay. Similarly, our results indicate that earthworm effects on decomposition vary with earthworm community composition. Furthermore, because earthworm invasion can involve a predictable shift in community structure along invasion fronts or through time, the community dynamics of invasion are important in predicting the spatial and temporal effects of earthworm invasion on litter decomposition, especially at later stages of decay.     

Earthworm Invasion as the Driving Force Behind Plant Invasion and Community Change in Northeastern North American Forests

Abstract:  Identification of factors that drive changes in plant community structure and contribute to decline and endangerment of native plant species is essential to the development of appropriate management strategies. Introduced species are assumed to be driving causes of shifts in native plant communities, but unequivocal evidence supporting this view is frequently lacking. We measured native vegetation, non-native earthworm biomass, and leaf-litter volume in 15 forests in the presence and absence of 3 non-native plant species ( Microstegium vimineum, Alliaria petiolata, Berberis thunbergii ) to assess the general impact of non-native plant and earthworm invasions on native plant communities in northeastern United States. Non-native plant cover was positively correlated with total native plant cover and non-native earthworm biomass. Earthworm biomass was negatively associated with cover of native woody and most herbaceous plants and with litter volume. Graminoid cover was positively associated with non-native earthworm biomass and non-native plant cover. These earthworm-associated responses were detected at all sites despite differences in earthworm species and abundance, composition of the native plant community, identity of invasive plant species, and geographic region. These patterns suggest earthworm invasion, rather than non-native plant invasion, is the driving force behind changes in forest plant communities in northeastern North America, including declines in native plant species, and earthworm invasions appear to facilitate plant invasions in these forests. Thus, a focus on management of invasive plant species may be insufficient to protect northeastern forest understory species.     

Declines in Woodland Salamander Abundance Associated with Non-Native Earthworm and Plant Invasions

Abstract:  Factors that negatively affect the quality of wildlife habitat are a major concern for conservation. Non-native species invasions, in particular, are perceived as a global threat to the quality of wildlife habitat. Recent evidence indicates that some changes to understory plant communities in northern temperate forests of North America, including invasions by 3 non-native plant species, are facilitated by non-native earthworm invasion. Furthermore, non-native earthworm invasions cause a reduction in leaf litter on the forest floor, and the loss of forest leaf litter is commonly associated with declines in forest fauna, including amphibians. We conducted a mark-recapture study of woodland salamander abundance across plant invasion fronts at 10 sites to determine whether earthworm or plant invasions were associated with reduced salamander abundance. Salamander abundance declined exponentially with decreasing leaf litter volume. There was no significant relationship between invasive plant cover and salamander abundance, independent of the effects of leaf litter loss due to earthworm invasion. An analysis of selected salamander prey abundance (excluding earthworms) at 4 sites showed that prey abundance declined with declining leaf litter. The loss of leaf litter layers due to non-native earthworm invasions appears to be negatively affecting woodland salamander abundance, in part, because of declines in the abundance of small arthropods that are a stable resource for salamanders. Our results demonstrate that earthworm invasions pose a significant threat to woodland amphibian fauna in the northeastern United States, and that plant invasions are symptomatic of degraded amphibian habitat but are not necessarily drivers of habitat degradation.     

Indirect Effects of Pandemic Deer Overabundance Inferred from Caterpillar‐Host Relations

Externally feeding phytophagous insect larvae (i.e., caterpillars, here, larval Lepidoptera and sawflies, Hymenoptera: Symphyta) are important canopy herbivores and prey resources in temperate deciduous forests. However, composition of forest trees has changed dramatically in the eastern United States since 1900. In particular, browsing by high densities of white‐tailed deer (Odocoileus virginianus) has resulted in forests dominated by browse‐tolerant species, such as black cherry (Prunus serotina), and greatly reduced relative abundance of other tree species, notably pin cherry (Prunus pensylvanica) and birches (Betula spp.). To quantify effects of these changes on caterpillars, we sampled caterpillars from 960 branch tips of the 8 tree species that comprise 95% of trees in Allegheny hardwood forests: red maple (Acer rubrum), striped maple (Acer pensylvanicum), sugar maple (Acer saccharum), sweet birch (Betula lenta), yellow birch (Betula allegheniensis), American beech (Fagus grandifolia), black cherry, and pin cherry. We collected 547 caterpillar specimens that belonged to 66 Lepidoptera and 10 Hymenoptera species. Caterpillar density, species richness, and community composition differed significantly among tree species sampled. Pin cherry, nearly eliminated at high deer density, had the highest density and diversity of caterpillars. Pin cherry shared a common caterpillar community with black cherry, which was distinct from those of other tree hosts. As high deer density continues to replace diverse forests of cherries, maples, birches, and beech with monodominant stands of black cherry, up to 66% of caterpillar species may be eliminated. Hence, deer‐induced changes in forest vegetation are likely to ricochet back up forest food webs and therefore negatively affect species that depend on caterpillars and moths for food and pollination. Efectos Indirectos de la Sobreabundancia de Venados Pandémicos Inferida de Relaciones Orugas‐Huéspedes     

Effects of landscape design of forest reserves on Saproxylic beetle diversity

Increasing the density of natural reserves in the forest landscape may provide conservation benefits for biodiversity within and beyond reserve borders. We used 2 French data sets on saproxylic beetles and landscape cover of forest reserves (LCFR) to test this hypothesis: national standardized data derived from 252 assessment plots in managed and reserve stands in 9 lowland and 5 highland forests and data from the lowland Rambouillet forest, a forested landscape where a pioneer conservation policy led to creation of a dense network of reserves. Abundance of rare and common saproxylic species and total saproxylic species richness were higher in forest reserves than in adjacent managed stands only in highland forests. In the lowland regional case study, as LCFR increased total species richness and common species abundance in reserves increased. In this case study, when there were two or more reserve patches, rare species abundance inside reserves was higher and common species richness in managed stands was higher than when there was a single large reserve. Spillover and habitat amount affected ecological processes underlying these landscape reserve effects. When LCFR positively affected species richness and abundance in reserves or managed stands, >12‐20% reserve cover led to the highest species diversity and abundance. This result is consistent with the target of 17% forested land area in reserves set at the Nagoya biodiversity summit in 2010. Therefore, to preserve biodiversity we recommend at least doubling the current proportion of forest reserves in European forested landscapes.     

Effects of Timber Harvesting on Southern Appalachian Salamanders

We compared the species richness and abundance of salamanders on six recent clearcuts (< 10 years old) with that of salamanders on 34 mature forest stands (>50 years old) in southern Appalachian forests in western North Carolina, U.S.A. Catches of salamanders from plots in mature forest stands were about five times higher than those on recent clearcuts. Almost all species and major taxonomic groups of salamanders were adversely affected by timber removal. Mean number of species collected per plot was about twice as great in mature forest stands as in clearcuts. Analyses of stand age versus salamander catch for 47 plots indicate that 50–70 years are required for populations to return to predisturbance levels following cutting. We conservatively estimate that clearcutting in U.S. national forests in western North Carolina results in a loss of nearly 14 millton salamanders annually. It also is chronically reducing regional populations by more than a quarter of a billion salamanders (9%) below that which could be sustained if mature forests were not cut.     

Recovery of Faunal Communities During Tropical Forest Regeneration

Abstract:  As mature tropical forests are cleared, secondary forests may play an important role in the conservation of animal species, depending on how fast animal communities recover during forest regeneration. I reviewed published studies on the recovery of animal species richness and composition during tropical forest regeneration. In 38 of the 39 data sets I examined, conversion of forest to agriculture or pasture substantially reduced species richness. Given suitable conditions for forest recovery, the species richness of the animal taxa considered can be predicted to resemble that of mature forests roughly 20–40 years after land abandonment. At least for ants and birds, however, recovery of species composition appears to take substantially longer than recovery of species richness. Because species richness for many taxa appears to recover relatively rapidly in secondary forests, conservation of secondary forests may be an effective investment in future diversity. The slower recovery of species composition indicates, however, that some species will require stands of mature forest to persist.     

Effects of Widespread Drought‐Induced Aspen Mortality on Understory Plants

Forest die‐off around the world is expected to increase in coming decades as temperature increases due to climate change. Forest die‐off will likely affect understory plant communities, which have substantial influence on regional biological diversity, ecosystem function, and land–atmosphere interactions, but how die‐off alters these plant communities is largely unknown. We examined changes in understory plant communities following a widespread, drought‐induced die‐off of trembling aspen (Populus tremuloides) in the western United States. We assessed shrub and herbaceous cover and volume in quadrats in 55 plots located across a wide range of levels of aspen mortality. We measured species richness and composition of herbaceous plant communities by recording species presence and absence in 12 sets of paired (1 healthy, 1 dying) aspen plots. Although understory composition in healthy and dying stands was heterogeneous across the landscape, shrub abundance, cover, and volume were higher and abundance of herbaceous species, cover, and volume were lower in dying aspen stands. Shrub cover and volume increased from 2009 to 2011 in dying stands, which suggests that shrub growth and expansion is ongoing. Species richness of herbs declined by 23% in dying stands. Composition of herbs differed significantly between dying and healthy stands. Richness of non‐native species did not differ between stand types. The understory community in dying aspen stands was not similar to other shrub‐dominated plant communities in the region and may constitute a novel community. Our results suggest that changes in understory plant communities as forests die off could be a significant indirect effect of climate change on biological diversity and forest communities. Efectos de la Mortalidad Extensiva de Álamos Inducida por Sequía sobre Plantas del Sotobosque     

Vegetation Responses along Edge-to-Interior Gradients in the Mixed Hardwood Forests of the Roanoke River Basin, North Carolina

Compared with forest interiors, forest edges typically have a different plant species composition and community structure, a phenomenon known as "edge effect." Edge effects make the functional interior area of a forest smaller than its actual area. The objective of this study was to estimate how far the effects of agriculturally maintained edges penetrate the mixed hardwood forests of the Roanoke River Basin, North Carolina. I determined percentage cover for all vascular plant species in 10-by-100-meter belt transects on north-facing or south-facing edges of four relatively undisturbed forests. Changes in the percentage cover of individual species, the relative cover of exotic species, and species richness all indicated that edge effects penetrate deeper on south-facing edges (to 60 meters) than on north-facing edges (to 20 meters). Analyses of species responses to the edge showed a number of species to be edge oriented, but no species was found to be interior oriented. The results of multivariate analyses (ordination and cluster analysis) suggested that edge effects could be detected to 50 meters on south-facing edges and 10–30 meters on north-facing edges. These results allow us to better understand the difference between a forest's actual area and its functional interior area.     

Thinning and burning result in low-level invasion by nonnative plants but neutral effects on natives.

Many historically fire-adapted forests are now highly susceptible to damage from insects, pathogens, and stand-replacing fires. As a result, managers are employing treatments to reduce fuel loadings and to restore the structure, species, and processes that characterized these forests prior to widespread fire suppression, logging, and grazing. However, the consequences of these activities for understory plant communities are not well understood. We examined the effects of thinning and prescribed fire on plant composition and diversity in Pinus ponderosa forests of eastern Washington (USA). Data on abundance and richness of native and nonnative plants were collected in 70 stands in the Colville, Okanogan, and Wenatchee National Forests. Stands represented one of four treatments: thinning, burning, thinning followed by burning, or control; treatments had been conducted 3-19 years before sampling. Multi-response permutation procedures revealed no significant effect of thinning or burning on understory plant composition. Similarly, there were no significant differences among treatments in cover or richness of native plants. In contrast, nonnative plants showed small, but highly significant, increases in cover and richness in response to both thinning and burning. In the combined treatment, cover of nonnative plants averaged 2% (5% of total plant cover) but did not exceed 7% (16% of total cover) at any site. Cover and richness of nonnative herbs showed small increases with intensity of disturbance and time since treatment. Nonnative plants were significantly less abundant in treated stands than on adjacent roadsides or skid trails, and cover within these potential source areas explained little of the variation in abundance within treated stands. Although thinning and burning may promote invasion of nonnative plants in these forests, our data suggest that their abundance is limited and relatively stable on most sites.     

Biotic Impoverishment and Homogenization in Unfragmented Forest Understory Communities

Abstract:  Ecological change is often hard to document because of a lack of reliable baseline data. Several recent then-versus-now surveys of temperate forest and grassland communities demonstrate losses of local plant species, but most are based on data from a single site. We resurveyed understory communities in 62 upland forest stands in northern Wisconsin (U.S.A.) for which quantitative baseline data exist from 50 years ago. These stands are within a largely unfragmented region but vary in species composition and successional stage. We collected data on changes in (1) total and native species richness, (2) the ratio of exotic to native species, (3) the relative abundance of habitat generalists, and (4) community similarity among sites. We also compared how these rates of change varied over time. Over the past 50 years, native species density declined an average of 18.5% at the 20-m² scale, whereas the ratio of exotic species to native species increased at 80% of all sites. Habitat generalists increased, and habitat specialists declined, accounting in part for an 8.7% rise in average similarity in species composition among sites. Most of these changes cannot be related to succession, habitat loss, or invasion by exotic species. Areas without deer hunting showed the greatest declines in native species density, with parks and research natural areas faring no better than unprotected stands. Animal-pollinated and animal-dispersed species also declined, particularly at unhunted sites. These results demonstrate the power of quantitative multistand data for assessing ecological change and identify overabundant deer as a key driver of community change. Because maintaining forest habitats alone fails to preserve plant diversity at local scales, local biotic simplification seems likely to continue in the region unless active efforts are taken to protect diversity.     

Invasibility and Effects of Amur Honeysuckle in Southwestern Ohio Forests

The Asian exotic Amur honeysuckle ( Lonicera maackii [Rupr.] Herder) has become the dominant shrub in many forests in southwestern Ohio and in some other locations in the eastern United States. Our research focused on the invasibility of forest communities and relationships of L. maackii to the abundance of tree seedlings and herbs. We surveyed 93 forest stands near Oxford, Ohio (USA) to determine L. maackii cover, time since invasion, tree canopy cover, tree basal area, and a shade tolerance index. Stepwise multiple regression indicated that greater than one-half of the variation (r ² = 0.550) in Lonicera cover was correlated to five variables (in descending order of importance): tree canopy cover, distance from Oxford, shade tolerance index, tree basal area, and time since invasion. The results suggest that high light levels and proximity to an abundant seed source increase forest invasibility. Tree seedling density, species richness of seedlings, and herb cover were all inversely related to L. maackii cover. When Lonicera becomes abundant, future structure and composition of forests could be affected and local populations of herbs threatened.     

Rapid assessment of postfire plant invasions in coniferous forests of the western United States.

Fire is a natural part of most forest ecosystems in the western United States, but its effects on nonnative plant invasion have only recently been studied. Also, forest managers are engaging in fuel reduction projects to lessen fire severity, often without considering potential negative ecological consequences such as nonnative plant species introductions. Increased availability of light, nutrients, and bare ground have all been associated with high-severity fires and fuel treatments and are known to aid in the establishment of nonnative plant species. We use vegetation and environmental data collected after wildfires at seven sites in coniferous forests in the western United States to study responses of nonnative plants to wildfire. We compared burned vs. unburned plots and plots treated with mechanical thinning and/or prescribed burning vs. untreated plots for nonnative plant species richness and cover and used correlation analyses to infer the effect of abiotic site conditions on invasibility. Wildfire was responsible for significant increases in nonnative species richness and cover, and a significant decrease in native cover. Mechanical thinning and prescribed fire fuel treatments were associated with significant changes in plant species composition at some sites. Treatment effects across sites were minimal and inconclusive due to significant site and site x treatment interaction effects caused by variation between sites including differences in treatment and fire severities and initial conditions (e.g., nonnative species sources). We used canonical correspondence analysis (CCA) to determine what combinations of environmental variables best explained patterns of nonnative plant species richness and cover. Variables related to fire severity, soil nutrients, and elevation explained most of the variation in species composition. Nonnative species were generally associated with sites with higher fire severity, elevation, percentage of bare ground, and lower soil nutrient levels and lower canopy cover. Early assessments of postfire stand conditions can guide rapid responses to nonnative plant invasions.     

Demography of northern flying squirrels informs ecosystem management of western interior forests.

We studied northern flying squirrel (Glaucomys sabrinus) demography in the eastern Washington Cascade Range to test hypotheses about regional and local abundance patterns and to inform managers of the possible effects of fire and fuels management on flying squirrels. We quantified habitat characteristics and squirrel density, population trends, and demography in three typical forest cover types over a four-year period. We had 2034 captures of flying squirrels over 41 000 trap nights from 1997 through 2000 and marked 879 squirrels for mark-recapture population analysis. Ponderosa pine (Pinus ponderosa) forest appeared to be poorer habitat for flying squirrels than young or mature mixed-conifer forest. About 35% fewer individuals were captured in open pine forest than in dry mixed-conifer Douglas-fir (Pseudotsuga menziesii) and grand fir (Abies grandis) forests. Home ranges were 85% larger in pine forest (4.6 ha) than in mixed-conifer forests (2.5 ha). Similarly, population density (Huggins estimator) in ponderosa pine forest was half (1.1 squirrels/ha) that of mixed-conifer forest (2.2 squirrels/ha). Tree canopy cover was the single best correlate of squirrel density (r = 0.77), with an apparent threshold of 55% canopy cover separating stands with low- from high-density populations. Pradel estimates of annual recruitment were lower in open pine (0.28) than in young (0.35) and mature (0.37) forest. High recruitment was most strongly associated with high understory plant species richness and truffle biomass. Annual survival rates ranged from 45% to 59% and did not vary among cover types. Survival was most strongly associated with understory species richness and forage lichen biomass. Maximum snow depth had a strong negative effect on survival. Rate of per capita increase showed a density-dependent response. Thinning and prescribed burning in ponderosa pine and dry mixed conifer forests to restore stable fire regimes and forest structure might reduce flying squirrel densities at stand levels by reducing forest canopy, woody debris, and the diversity or biomass of understory plants, truffles, and lichens. Those impacts might be ameliorated by patchy harvesting and the retention of large trees, woody debris, and mistletoe brooms. Negative stand-level impacts would be traded for increased resistance and resilience of dry-forest landscapes to now-common, large-scale stand replacement fires.     

Forests Too Deer: Edge Effects in Northern Wisconsin

Abstract: Browsing by white-tailed deer (Odocoileus virginianus) can profoundly affect the abundance and population structure of several woody and herbaceous plant species. Enclosure studies and population surveys reveal that past and current deer densities as low as 4 deer/km² may prevent regeneration of the once common woody species, Canada yew (Taxus canadensis), eastern hemlock (Tsuja canadensis), and white cedar Puja occidentalis), as well as several herbaceous species. Prior to European settlement, forests in northern Wisconsin contained relatively sparse deer populations (<4/km²), but extensive timber cutting in the late nineteenth century boosted deer populations. Continued habitat fragmentation resulting from scattered timber harvests and the creation of "wildlife openings" to improve deer forage maintain these high densities throughout much of the Northeast.
Because deer wander widely, the effects of high deer densities penetrate deeply into remaining stands of old and mature forest, greatly modifying their composition Thus, abundant early successional and "edge" habitat, and the high deer densities they engender, represent significant external threats to these plant communities. We hypothesize that establishing large (200–400 km²) continuous areas of maturing forest, especially in conjunction with increased hunting, could reduce local deer densities and so provide a simple and inexpensive method for retaining species sensitive to the deleterious effects of browsing.     

Slope aspect modifies community responses to clear-cutting in boreal forests

Slope aspect modifies microclimate and influences ecological processes and spatial distribution of species across forest landscapes, but the impact of slope aspect on community responses to disturbance is poorly understood. Such insight is necessary to understand landscape community dynamics and resilience. We compared bryophyte (liverworts and mosses) communities in matched 0.02-ha plots of four boreal stand types in central Sweden: recently clear-felled and mature stands dominated by Norway spruce in south-facing and north-facing slopes. Differences between forests and clear-cuts were interpreted as effects of clear-cutting, and differences between south- and north-facing slopes as effects of aspect. In response to clear-cutting, bryophyte cover and composition changed more in south-facing slopes. Only one out of ten significantly declining species in south-facing slopes also declined significantly in north-facing slopes. North-facing slopes lost fewer bryophyte species, and among those, fewer forest species and fewer species associated with wood and bark. In north-facing slopes, the average proportions of mosses and liverworts shared between the forest and the clear-cut plot were 88% and 74%, respectively. Corresponding numbers for south-facing slopes were 79% and 33%. In addition, more bryophyte species were added in north- than south-facing slopes after clear-cutting, somewhat reducing the difference in compositional change between aspects. South- and north-facing mature forests differed in species composition, mostly due to higher richness of mosses in south-facing slopes. The smaller changes in bryophyte communities on north-facing slopes in response to clear-cutting have implications for ecosystem dynamics and management as high local survival may enhance landscape-level resilience.