Removal of Encroaching Conifers to Regenerate Degraded Aspen Stands in the Sierra Nevada

Aspen is considered a keystone species, and aspen communities are critical for maintaining biodiversity in western landscapes. Inventories of aspen stand health across the Eagle Lake Ranger District (ELRD), Lassen National Forest, California, U.S.A., indicate that 77% of stands are in decline and at risk of loss as defined by almost complete loss of mature aspen with little or no regeneration. This decline is due to competition from conifers establishing within aspen stands as a result of modification of natural fire regimes coupled with excessive browsing by livestock. Restoration treatments were implemented in four aspen stands in 1999 using mechanical equipment to remove competing conifers to enhance the growth environment for aspen. Recruitment and establishment of aspen stems were measured in treated stands (removal of competing conifers) and non‐treated stands (control) immediately prior to treatment and 2 and 4 years post‐treatment. There was a significant increase in total aspen stem density and in two of three aspen regeneration size classes for treated stands compared to controls. Pre‐treatment total aspen density was positively associated with total aspen density and density in all size classes of aspen (p < 0.001). The results demonstrate that mechanical removal of conifers is an effective treatment for restoring aspen.     

Saproxylic beetle diversity in a managed boreal forest: importance of stand characteristics and forestry conservation measures

Saproxylic beetles constitute a significant proportion of boreal forest biodiversity. However, the long history of timber production in Fennoscandia has significantly reduced the availability of dead wood and is considered a threat to the conservation of saproxylic beetle assemblages. Therefore, since the mid‐1990s dead wood retention in harvested stands has formed an integral part of silvicultural practices. However, the contribution of this biodiversity‐orientated management approach to conserving saproxylic beetle assemblages in boreal forest landscapes that include production forestry remains largely untested. We examined differences in resident saproxylic beetle assemblages among stands under different management in a boreal forest landscape in Central Sweden, and in particular stands managed according to new conservation‐orientated practices. We also investigated the relationship between beetle diversity and forest stand characteristics. Bark of coarse woody debris (CWD) was sieved for beetles in old managed stands, unmanaged nature reserves, and set‐aside areas, and clear‐cut stands harvested according to certification guidelines [new forestry (NF) clear‐cuts]. All stand types contributed significantly to the total diversity of beetles found. While stand size, position, and distance to nearest reserve were unimportant, both the quality and the quantity of CWD in stands contributed significantly to explaining beetle abundance and species richness. This extends the previous findings for red‐listed invertebrates, and shows that heterogeneous substrate quality and a range of management practices are necessary to maintain saproxylic beetle diversity in boreal forest landscapes that include production forestry. The unique abiotic conditions in combination with the abundant and varied CWD associated with NF clear‐cuts form an important component of forest stand heterogeneity for saproxylic beetles. It is thus essential that sufficient, diverse, CWD is retained in managed boreal landscapes to ensure the conservation of boreal saproxylic beetle assemblages.     

Forest floor properties across sharp compositional boundaries separating trembling aspen and jack pine stands in the southern boreal forest

Jack pine and trembling aspen are two early-seral boreal tree species with contrasting nutrient cycling strategies. Both species may form adjacent mono-specific stands separated by sharp compositional boundaries. We hypothesized that such boundaries result in wider functional ecotones. Spatial transitions in humus forms, forest floor chemistry and microbial communities were assessed across 32 m long transects set perpendicular to sharp compositional boundaries separating four jack pine and aspen stands. Split moving window analysis (SMWA) and moving window regression analysis (MWRA) were used to locate functional boundaries and ecotones. We found a gradual transition from moder (aspen) to mor (jack pine) humus spanning 16 m across the compositional boundary. An abrupt increase in forest floor water content at 3 m within jack pine stands was possibly due to aspen roots foraging for water beyond the boundary. The functional boundary and associated ecotone for forest floor pH, C:N ratio, Mg and ammonification were skewed toward jack pine stands, likely the result of aspen leaf dispersal. Low nitrification rates throughout jack pine stands and up to 11 m into aspen stands suggested that jack pine roots might extend far within aspen stands and produce metabolites that suppress nitrification. SMWA performed on the multivariate dataset of microbial fatty acids (FAs) revealed three distinct forest floor microbial communities that were skewed toward jack pine stands. Pine-type communities were associated to fungal FAs, pine-type and transition-type communities to non-fungal eukaryotic FAs, and aspen-type communities to bacterial FAs. Taken collectively, our data delimit a 24 m wide functional ecotone straddling sharp compositional boundaries separating trembling aspen and jack pine stands. We conclude that the functional diversity of boreal landscapes, where adjacent mono-specific stands are prevalent, is related to the patchiness of the landscape.     

Competition and facilitation between tree species change with stand development

Processes governing tree interspecific interactions, such as facilitation and competition, may vary in strength over time. This study tried to unveil them by performing dendrometrical analyses on black spruce Picea mariana, trembling aspen Populus tremuloides and jack pine Pinus banksiana trees from pure and mixed mature boreal forest stands in the Clay Belt of northwestern Quebec and on the tills of northwestern Ontario. We cored 1430 trees and cut 120 for stem analysis across all stand composition types, tree species and study regions. Aspen annual growth rate was initially higher when mixed with conifers, but then progressively decreased over time compared to pure aspen stands, while jack pine growth rate did not differ with black spruce presence throughout all stages of stand development. When mixed with aspen, black spruce showed a contrary response to aspen, i.e. an initial loss in growth but a positive gain later. On the richer clay soil of the Quebec Clay Belt region, however, both aspen and spruce responses in mixed stands reversed between 37 and 54 years. Overall, our results demonstrate that interspecific interactions were present and tended to change with stand development and among species. Our results also suggest that the nature of interspecific interactions may differ with soil nutrient availability.     

Leaf litter and the small-scale distribution of carabid beetles (Coleoptera, Carabidae) in the boreal forest

Management practices favoring conifers at the expense of deciduous tree species, and the eradication of deciduous trees, especially aspen Populus tremula , from managed forests have resulted in population declines in several species in Fennoscandia. In addition to species depending on decaying wood of deciduous trees, earlier evidence suggests that leaf litter, especially that of aspen, is favored by many carabid species. We ran a four-year experiment in order to compare carabid assemblages of unchanged forest floor with artificially created leaf-litter plots in central Finland. A total of 18 plots (5 m in diameter) were established in three forest stands without aspen a few kilometers apart. Each stand had 3 litter plots (litter added) and 3 control plots, Pre-treatment samples were compared with those collected alter litter addition.
The litter addition affected the carabid-assemblage structure by increasing the catches of some species and decreasing the catch of one species. The number of carabid species was similar in control and litter plots. The litter effect was smaller than variation among forest stands and year-lo-year fluctuations. There was a strong temporal constancy among the plots: 'rich' plots remained 'rich' from year to year and similarly, 'poor' plots remained 'poor'.
The significant influence of leaf litter on carabid abundance can be attributable to both abiotic factors (microenvironmental conditions, especially humidity and temperature), and biotic ones (changes in niche structure, improved food supply). Leal litter seems to have an effect on carabid distribution patterns, and deciduous trees scattered among conifers are likely to be of importance on carabid fauna in boreal forests.     

Regeneration of planted conifers across climatic moisture gradients on the Canadian prairies: implications for distribution and climate change

The influence of dry climates on white spruce ( Picea glauca (Moench) Voss)) regeneration was examined by conducting surveys of seedlings and small trees that had regenerated naturally at 100 farm shelterbelts and plantations in southern Saskatchewan, Canada. The sites surveyed were located along a climate moisture gradient extending from the relatively moist boreal forest, across the aspen parkland, to the semi-arid prairie grasslands. Natural regeneration was greatest at sites in the boreal forest and northern aspen parkland, decreased in the southern aspen parkland, and was negligible in the grassland zone. Furthermore, the few seedlings found in the drier zones were usually in poor condition. Similar results were obtained for the introduced Colorado spruce ( Picea pungens Engelm.) and Scots pine ( Pinus sylvestris L.). It is concluded that the present climate of the southern parkland and grassland is too dry to permit natural regeneration of white spruce and other conifers. If increases in atmospheric CO₂ levels lead to a drier future climate in the southern boreal forest of western Canada, the ability of conifers to regenerate naturally may be significantly reduced.     

Epiphytic macrolichens in managed and natural forest landscapes: a comparison at two spatial scales

To maintain biodiversity in managed forests we must understand how forestry affects various organisms across a wide range of spatial and temporal scales. We compared landscape structure, forest structure, and species richness and abundance of epiphytic macrolichens in three pairs of natural and managed boreal forest landscapes. Study landscapes (2500 ha) were located within and adjacent to three of the largest forest reserves in Sweden (Reivo, Muddus, Jelka). The structural heterogeneity within landscapes was higher in managed forests whereas within-stand structural heterogeneity was higher in natural landscapes. Species richness of macrolichens at the stand level (sample plot) was 23% higher in natural forests but there was no difference at the landscape level. Most (86%) of the common species were more frequent in natural landscapes. Lichen abundance (estimated by lichen litter) was two times higher in natural than in managed landscapes, 5.6 and 2.7 kg ha^-1 forest (pooled data), respectively. Both species richness and abundance were negatively related to cutting level (number and basal area of cut stumps) and positively related to stand variables (stand age, stem density and basal area). Lichen-rich forest stands were more numerous but covered a smaller area and were more isolated in managed landscapes. This may in turn have important consequences for dispersal of lichen propagules to second-growth forests. In conclusion, the results suggest that effects of forestry on epiphyte diversity and abundance are strongly related to the spatial scale (stand or landscape). To enhance biodiversity in managed forests we must increase structural heterogeneity at the whole range of spatial and temporal scales.     

Aspen succession and nitrogen loading: a case for epiphytic lichens as bioindicators in the Rocky Mountains,USA

Question: Can lichen communities be used to assess short‐ and long‐term factors affecting seral quaking aspen (Populus tremuloides) communities at the landscape scale? Location: Bear River Range, within the Rocky Mountains, in northern Utah and southern Idaho, USA. Method: Forty‐seven randomly selected mid‐elevation aspen stands were sampled for lichens and stand conditions. Plots were characterized according to tree species cover, basal area, stand age, bole scarring, tree damage, and presence of lichen species. We also recorded ammonia emissions with passive sensors at 25 urban and agricultural sites throughout an adjacent populated valley upwind of the forest stands. Nonmetric multidimensional scaling (NMS) ordination was used to evaluate an array of 20 variables suspected to influence lichen communities. Results: In NMS, forest succession explained most variance in lichen composition and abundance, although atmospheric nitrogen from local agricultural and urban sources also significantly influenced the lichen communities. Abundance of nitrophilous lichen species decreased with distance from peak ammonia sources and the urban center in all aspen succession classes. One lichen, Phaeophyscia nigricans, was found to be an effective bioindicator of nitrogen loading. Conclusions: Lichen communities in this landscape assessment of aspen forests showed clear responses to long‐term (stand succession) and short‐term (nitrogen deposition) influences. At the same time, several environmental factors (e.g. tree damage and scarring, distance to valley, topography, and stand age) had little influence on these same lichen communities. We recommend further use of epiphytic lichens as bioindicators of dynamic forest conditions.     

Hurricane Disturbance Alters Secondary Forest Recovery in Puerto Rico

Land-use history and large-scale disturbances interact to shape secondary forest structure and composition. How introduced species respond to disturbances such as hurricanes in post-agriculture forest recovery is of particular interest. To examine the effects of hurricane disturbance and previous land use on forest dynamics and composition, we revisited 37 secondary forest stands in former cattle pastures across Puerto Rico representing a range of exposure to the winds of Hurricane Georges in 1998. Stands ranged from 21 to>80 yr since agricultural abandonment and were measured 9 yr posthurricane. Stem density decreased as stands aged, while basal area and species richness tended to increase. Hurricane disturbance exerted contrasting effects on stand structure, contingent on stand age. In older stands, the basal area of large trees fell, shifting to a stand structure characteristic of younger stands, while the basal area of large trees tended to rise in younger stands with increasing hurricane disturbance. These results demonstrate that large-scale natural disturbances can alter the successional trajectory of secondary forest stands recovering from human land use, but stand age, precipitation and soil series were better predictors of changes in stand structure across all study sites. Species composition changed substantially between census intervals, but neither age nor hurricane disturbance consistently predicted species composition change. However, exposure to hurricane winds tended to decrease the abundance of the introduced tree Spathodea campanulata, particularly in smaller size classes. In all sites the abundance of the introduced tree Syzygium jambos showed a declining trend, again most strongly in smaller size classes, suggesting natural thinning through succession.     

Competition and regeneration in quaking aspen–white fir (Populus tremuloides–Abies concolor) forests in the Northern Sierra Nevada,USA

Question: How does competition between quaking aspen (Populus tremuloides) and white fir (Abies concolor) affect growth and spatial pattern of each species? Location: The northern Sierra Nevada, California, USA. Methods: In paired plots in mixed aspen‐ (n=3) or white fir‐dominated (n=2) stands, we mapped trees and saplings and recorded DBH, height, species, and condition and took increment cores. We tallied seedlings by species. Tree ring widths were used as a measure of basal area change over the last decade, and canopy openness was identified using hemispherical photographs. Linear mixed models were used to relate neighborhood indices of competition, stand, and tree‐level variables to diameter increment. Spatial patterns of stems were identified using the Neighborhood Density Function. Results: White fir radial growth was higher in aspen‐ than white fir‐dominated plots. Individual‐level variables were more important for white fir than for aspen growth, while variables representing competitive neighborhood were important only for aspen. The forest canopy was more open in aspen‐ than white fir‐dominated stands, but ample aspen seedlings were observed in all stands. Canopy stems of aspen and white fir were randomly distributed, but saplings and small trees were clumped. Aspen saplings were repelled by canopy aspen stems. Conclusions: Variation in canopy openness explained more stand–stand variation in white fir than aspen growth, but high light levels were correlated with recruitment of aspen seedlings to the sapling class. Radial growth of aspen was predicted by indices of neighborhood competition but not radial growth of white fir, indicating that spacing and stem arrangement was more important for aspen than white fir growth. Fire suppression has removed a major disturbance mechanism that promoted aspen persistence and reduced competition from encroaching conifers, and current forests favor species that regenerate best by advance regeneration (white fir).     

Spore deposition of wood-decaying fungi: importance of landscape composition

The abundance of many species of wood-decaying fungi has decreased dramatically in Swedish boreal forests over the last century. Therefore, we have investigated the relationship between the spore dispersal of wood-decaying fungi and two key features of landscape composition, namely the amount and age of old forest stands at different spatial scales. Spore deposition was monitored in two regions, using sampling methods based on recording the dikaryotisation of monokaryotic mycelia on nutrient agar and wood discs. The studied species, all mainly confined to Norway spruce in the boreal forest zone, were Fomitopsis pinicola, Fomitopsis rosea , Gloeoporus taxicola , Phlebia centrifuga and Trichaptum laricinum . The study of forestry intensity showed for both regions, that the spore deposition for F. pinicola , F. rosea and G. taxicola was higher in circular plots (radius 2-km) with a high proportion of old Norway spruce forest (>80 yr) than in plots with a lower proportion of old forest. Analysis of the variation in spore deposition of F. rosea and P. centrifuga in relation to the proportion of old spruce forest within 1, 2 and 3 km of the spore sampling point showed that the proportion of old forest within a 3-km radius explained more of the variation than the proportions within 1- and 2-km radii. In addition, the proportion of forest older than 140 yr explained more of the variation than the proportion of younger forests. Thus, the results show that the spore deposition of the studied species strongly depends on the landscape composition at both regional and local scales. Further, the spore deposition at the local scale was best explained by the proportion of >140 yr old spruce forest, which exceeds the common harvest rotation period.     

Predation on artificial nests in a forest dominated landscape – the effects of nest type, patch size and edge structure

We studied the effects of forest patch size and forest edge structure on nest predation in a boreal coniferous forest landscape. The following predictions were tested. Nest predation should be higher in small than in large stands, in edges than in interior areas of forest stands, and in barren forest/clear–cut edges created by forestry than in natural forest/open marsh edges. Four types of artificial above ground nests (total of 261) were used; open cup nests with reindeer Rangifer t. tarandus hair, open cup nests with domestic hen Gallus domesticus feathers, and unlined open cup and nest–box nests. Nests were baited with one Japanese quail Coturnix coturnix japonica egg. Nest–boxes were depredated significantly less than open cup nests of all types. No edge- or stand size–related nest predation was found. The predation rate, regardless of the nest type, did not differ relative to the edge type and vegetation characteristics. However, better horizontal visibility of open cup nests due to more open vegetation structure increased predation risk in man–made edges compared to inherent edges. The results suggest that edge–related nest predation is absent or weak in forest dominated landscapes. This may be due to predator types present in the landscape and/or predators habitat use in forest dominated areas. Therefore, it might be that findings documented in other areas, such as in agricultural dominated landscapes, cannot be directly applied to managed forest landscapes.     

Landscape-Scale Disturbances Modified Bird Community Dynamics in Successional Forest Environment

Ecosystem-based forest management strives to develop silvicultural practices that best emulate natural disturbances such as wildfire to conserve biodiversity representative of natural forest ecosystems. Yet, current logging practices alter forest structure and reduce the proportion of old-growth forest and, consequently, can exert long-term effects on the dynamics of forest biota. The stand- and landscape-scale factors driving bird community dynamics in post-disturbance environment remain poorly understood. In this study, we examined bird community dynamics along successional gradients in boreal ecosystems originating from fire and logging in landscapes dominated by old-growth forest. We tested if bird species richness and community compositions in clear-cutting stands became comparable to those in natural stands after 70 years, and identified the relative contributions of stand- and landscape-scale forest attributes in bird community dynamics. Based on records of bird occurrences at 185 field sites in natural and clearcutting stands, we demonstrate that (1) both forest structures and bird communities underwent evident changes along successional gradients in post-clearcutting environment; (2) bird species richness and community composition in 60- to 70-years-old clearcutting stands still differed from those in 50- to 79-years-old natural stands, in spite of the fact that most forest attributes of clearcutting stands became comparable to those of natural stands after 40 years; and (3) landscape disturbances contributed more than stand characteristics in explaining the lack of convergence of mature forest species, residents, and short-distance migrants in post-clearcutting environment. Our study points out that more regards should be paid to improve the landscape configuration of the managed forests, and implies that old-growth forest retention within logged areas, combined with selection cutting and prolonged logging rotations, can better emulate fire and alleviate forest harvesting effects on bird community assemblages typical of natural boreal ecosystem.     

Selectivity by moose vs the spatial distribution of aspen: a natural experiment

Patch use theory predicts that herbivores perceive food as patches and spend more time in high quality patches, i.e. feeding sites providing a high net rate of intake of energy and/or limiting nutrients. The herbivores should accordingly not discriminate among food items in such high quality patches, and food items should thus be eaten in proportion to availability. In contrast, classical diet theory assumes food selection to take place at the level of individual plants, and predicts that the forager should concentrate on the most profitable item until availability drops below some critical threshold.
Here we address how the spatial distribution of European aspen Populus tremula , a highly preferred browse species, affects the selectivity by moose Alces alces at the patch and the tree level. The study was performed in a managed boreal forest landscape in coastal northern Sweden, where aspen occurs highly aggregated almost exclusively in discrete patches. We compared moose' selectivity for aspen and browsing intensity on aspen ramets and other browse species in aspen patches versus at randomly located sites.
Random sites and aspen stands were utilised equally by moose in terms of overall use of forage. There was no difference in total coverage of forage species and relative moose density. Selectivity for aspen was stronger at random sites than at aspen sites, and the browsing intensity on aspen was similar. We conclude that moose did not perceive aspen stands as discrete patches, and used aspen ramets more in accordance with diet theory. These findings agree with the idea that large generalist herbivores strive to maintain a mixed and balanced diet, causing rare species to be over-utilised (negative frequency-dependent food selection). By such selective feeding, moose may reinforce the aggregated distribution of aspen in the managed boreal forest landscape.     

Understory responses to restoration in aspen‐conifer forests around the Lake Tahoe Basin: residual stand attributes predict recovery

The removal of conifers from aspen (Populus tremuloides) stands is being undertaken throughout the western United States to restore aspen for local‐ and landscape‐level biodiversity. Current practices include mechanically removing conifers or hand thinning, piling, and burning cut conifers in and adjacent to aspen‐conifer stands. To evaluate the effectiveness of restoration treatments, we examined tree regeneration and herbaceous vegetation cover in thinned, thinned and pile burned, and non‐thinned control stands. Growth rates of small conifer saplings threatening to outcompete and replace aspen were also measured. Two to four years after pile burning, herbaceous vegetation cover within the footprint of burned piles (i.e. burn scars) was 35–73% of that in adjacent areas. Aspen was more likely to regenerate inside burn scars where fewer surrounding trees were true firs. Conifer seedlings were more likely to regenerate in burn scars where more of the surrounding trees were conifers (pine or fir) as opposed to aspen. Fir saplings had much slower growth than did aspen saplings. Overall, our findings show that restoration treatments are promoting desirable outcomes such as enhancing aspen regeneration but that follow‐up treatments will be needed to remove numerous conifer seedlings becoming established after restoration activities. Eliminating conifers, while they are small, growing slowly, and contributing little to fuel loads may be an economical way to prolong restoration treatment effectiveness.     

Stability in the patterns of long‐term development and growth of the Canadian spruce–moss forest

Aim The spruce–moss forest is the main forest ecosystem of the North American boreal forest. We used stand structure and fire data to examine the long‐term development and growth of the spruce–moss ecosystem. We evaluate the stability of the forest with time and the conditions needed for the continuing regeneration, growth and re‐establishment of black spruce (Picea mariana) trees. Location The study area occurs in Québec, Canada, and extends from 70°00′ to 72°00′ W and 47°30′ to 56°00′ N. Methods A spatial inventory of spruce–moss forest stands was performed along 34 transects. Nineteen spruce–moss forests were selected. A 500 m² quadrat at each site was used for radiocarbon and tree‐ring dating of time since last fire (TSLF). Size structure and tree regeneration in each stand were described based on diameter distribution of the dominant and co‐dominant tree species [black spruce and balsam fir (Abies balsamea)]. Results The TSLF of the studied forests ranges from 118 to 4870 cal. yr bp . Forests < 325 cal. yr bp are dominated by trees of the first post‐fire cohort and are not yet at equilibrium, whereas older forests show a reverse‐J diameter distribution typical of mature, old‐growth stands. The younger forests display faster height and radial growth‐rate patterns than the older forests, due to factors associated with long‐term forest development. Each of the stands examined established after severe fires that consumed all the soil organic material. Main conclusions Spruce–moss forests are able to self‐regenerate after fires that consume the organic layer, thus allowing seed regeneration at the soil surface. In the absence of fire the forests can remain in an equilibrium state. Once the forests mature, tree productivity eventually levels off and becomes stable. Further proof of the enduring stability of these forests, in between fire periods, lies in the ages of the stands. Stands with a TSLF of 325–4870 cal. yr bp all exhibited the same stand structure, tree growth rates and species characteristics. In the absence of fire, the spruce–moss forests are able to maintain themselves for thousands of years with no apparent degradation or change in forest type.     

Fire Interval Effects on Successional Trajectory in Boreal Forests of Northwest Canada

Although succession may follow multiple pathways in a given environment, the causes of such variation are often elusive. This paper describes how changes in fire interval mediate successional trajectory in conifer-dominated boreal forests of northwestern Canada. Tree densities were measured 5 and 19 years after fire in permanent plots and related to pre-fire vegetation, site and fire characteristics. In stands that were greater than 75 years of age when they burned, recruitment density of conifers was significantly correlated with pre-fire species basal area, supporting the expectation of stand self-replacement as the most common successional pathway in these forests. In contrast, stands that were under 25 years of age at the time of burning had significantly reduced conifer recruitment, but showed no change in recruitment of trembling aspen (Populus tremuloides). As a result, young-burned stands had a much higher probability of regenerating to deciduous dominance than mature-burned stands, despite the dominance of both groups by spruce (Picea mariana and Picea glauca) and pine (Pinus contorta) before the fire. Once initiated, deciduous-dominated stands may be maintained across subsequent fire cycles through mechanisms such as low on-site availability of conifer seed, competition with the aspen canopy, and rapid asexual regeneration of aspen after fire. We suggest that climate-related increases in fire frequency could trigger more frequent shifts from conifer to deciduous-dominated successional trajectories in the future, with consequent effects on multiple ecosystem processes.     

Contrasting effects of habitat quantity and quality on moth communities in fragmented landscapes

Habitat loss is commonly identified as a major threat to the loss of global biodiversity. In this study, we expand on our previous work by addressing the question of how lepidopteran species richness and composition vary among remnants of North American eastern deciduous forest located within agricultural or pastoral landscapes. Specifically, we tested the relative roles of habitat quantity (measured as stand area and percent forest in the greater landscape) and habitat quality (measured as tree species diversity) as determinants of moth species richness. We sampled >19 000 individuals comprising 493 moth species from 21 forest sites in two forested ecoregions. In the unglaciated Western Allegheny Plateau, the species richness of moths with woody host plants diminished as forest stand size and percent forest in the landscape decreased, but the total species richness and abundance of moths were unaffected by stand size, percent forest in the landscape, or tree species diversity. In contrast, the overall species richness and abundance of moths in the glaciated North Central Tillplain were affected primarily by tree species diversity and secondarily by forest size. Higher tree species diversity may reduce species loss from smaller forest stands, suggesting that small, diverse forests can support comparable numbers of species to those in less diverse, large stands. Smaller forests, however, contained a disproportionate number of moth species that possess larvae known to feed on herbaceous vegetation. Thus, although woody plant feeding moths are lost from forests with changes in stand area, new species appear capable of recolonizing smaller fragments from the surrounding habitat matrix. Our study further suggests that when species replacement occurs, local patch size and habitat quality may be more important than landscape context in determining the community structure of forest Lepidoptera.     

Tree species dynamics and disturbance in three Swedish boreal forest stands during the last two thousand years

Abstract. Data from three forest stands for the past 2000 yr show how the shade-intolerant species Pinus sylvestris and Betula pubescens maintain significant populations in the Swedish boreal landscape. Age structure data from a northern stand close to the range limits of Picea abies and Pinus complement a local pollen diagram, and reveal cyclic population fluctuations which can be related to periods of climatic stress and fire. Pollen data from two southern stands show that high fire frequencies in the past prevented the expansion of Picea populations. Reduction of the fire frequency during the last 200 yr has favoured Picea. A long time perspective reveals the population dynamics of long-lived species and indicates the controlling factors. Such knowledge permits assessment of the current status and likely future of forest stands.