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The influence of the forest canopy on nutrient cycling.

Rates of key soil processes involved in recycling of nutrients in forests are governed by temperature and moisture conditions and by the chemical and physical nature of the litter. The forest canopy influences all of these factors and thus has a large influence on nutrient cycling. The increased availability of nutrients in soil in clearcuts illustrates how the canopy retains nutrients (especially N) on site, both by storing nutrients in foliage and through the steady input of available C in litter. The idea that faster decomposition is responsible for the flush of nitrate in clearcuts has not been supported by experimental evidence. Soil N availability increases in canopy gaps as small as 0.1 ha, so natural disturbances or partial harvesting practices that increase the complexity of the canopy by creating gaps will similarly increase the spatial variability in soil N cycling and availability within the forest. Canopy characteristics affect the amount and composition of leaf litter produced, which largely determines the amount of nutrients to be recycled and the resulting nutrient availability. Although effects of tree species on soil nutrient availability were thought to be brought about largely through differences in the decomposition rate of their foliar litter, recent studies indicate that the effect of tree species can be better predicted from the mass and nutrient content of litter produced, hence total nutrient return, than from litter decay rate. The greater canopy complexity in mixed species forests creates similar heterogeneity in nutritional characteristics of the forest floor. Site differences in slope position, parent material and soil texture lead to variation in species composition and productivity of forests, and thus in the nature and amount of litter produced. Through this positive feedback, the canopy accentuates inherent differences in site fertility.

Biomass↗

Trends in bole biomass accumulation, net primary production and tree mortality in Pseudotsuga menziesii forests of contrasting age.

Although it is generally accepted that the rate of accumulation of biomass declines as forests age, little is known about the relative contributions to this decline of changes in net primary production (NPP) and tree mortality. We used 10-15 years of observations of permanent plots in three small watersheds in and near the H.J. Andrews Experimental Forest, Oregon, to examine these issues. The three watersheds are of similar elevation and potential productivity and support young (29 years at last measurement), mature (approximately 100 years) and old (approximately 400 years) forest dominated by Pseudotsuga menziesii (Mirb.) Franco and Tsuga heterophylla (Raf.) Sarg. Accumulation of tree bole biomass was greatest in the young stand, reaching approximately 7 Mg ha-1 year-1 in the last measurement interval. Bole biomass accumulation was relatively constant (approximately 4-5 Mg ha-1 year-1) in the mature stand, and there was no net accumulation of bole biomass in the old-forest stand. The NPP of boles increased with time in the young stand, from approximately 3 to approximately 7 Mg ha-1 year-1, but was nearly constant in the mature and old-forest stands, at approximately 6 and 3-4 Mg ha-1 year-1, respectively. Mortality increased slowly in the young stand (from < 0.1 to 0.3 Mg ha-1 year-1), but fluctuated between 1-2 and 2-6 Mg ha-1 year-1 in the mature and old-forest stands, respectively. Thus, declining biomass accumulation with stand age reflects, in approximately equal amounts, both decreasing NPP and increasing mortality.

Biomass↗

Climate change and forests.

Factors governing long-term change in global temperature are reviewed. The magnitude and rate of change in global temperature resulting from current increases in the concentration of atmospheric greenhouse gases are considered in relation to their impact on forests. Movement in forest zone boundaries at a rate of 2.5 km year(-1) are possible, which is nearly ten times the rate forests have been known to move by natural reproduction. Climate models indicate that increased global temperature will affect rainfall distribution, lead to more frequent and more severe storms and increase climatic variability. Consequences for the world's forests include increased frequencies of fire and blow-down, and wide-spread decline. Increased atmospheric CO(2) concentrations may increase forest growth where the effect is not offset by reduced precipitation, but the overall effect of anticipated changes in global climate is likely to be widespread loss of forests.

Journal Article↗

Analytical model of carbon storage in the trees, soils, and wood products of managed forests.

The carbon balance between managed forests and the atmosphere depends critically on the frequency and intensity of harvesting, and the lifetime of harvested products. To assess more quantitatively the nature of this dependence, a theoretical analysis, previously applied to carbon storage in trees and wood products only, is extended here to include the carbon in forest floor detritus and soil. A dimensionless combination of the parameters of the model, alpha, with critical value alpha(c), is identified such that for alpha < alpha(c), the conversion of old-growth forest to managed forest releases carbon to the atmosphere in the long term. Parameter alpha is given by the combination f(t)D/T(*), where f(t) is the fraction of old-growth forest carbon stored in trees, D is the residence time of harvested biomass (wood products and slash debris) within the system, and T(*) is the rotation period for maximum sustained yield (maximum mean annual increment). The critical value alpha(c), typically in the range 0.5-0.7, is derived for a variety of forest types. Parameter alpha determines the degree to which the carbon accumulated in harvested biomass offsets the loss of carbon in trees due to felling and in soils due to reduced litter input. When alpha > alpha(c), long-term carbon storage is optimized by harvesting for maximum sustained yield.

Journal Article↗

Effects of forest patch size on physiological stress and immunocompetence in an area-sensitive passerine, the Eurasian treecreeper (Certhia familiaris): an experiment.

We manipulated the primary brood size of Eurasian treecreepers (Certhia familiaris) breeding in different sized forest patches (0.5-12.8 ha) in moderately fragmented landscapes. We examined the effects of brood size manipulation (reduced, control, enlarged) and forest patch size on physiological stress (heterophil-lymphocyte ratios; H/L), body condition and cell-mediated immunocompetence (phytohaemagglutinin test). Nestlings' H/L ratios were negatively related to forest patch area in control and enlarged broods, whereas no effects were found in reduced broods. The effects of forest patch area were strongest in enlarged broods, which had, in general, twofold higher H/L ratios than control and reduced broods. The elevated H/L ratios were positively related to nestling mortality and negatively correlated with body-condition indices suggesting that the origin of stress in nestlings was mainly nutritional. Cell-mediated immunity of nestlings was not related to brood manipulation or to forest patch size. Also, the H/L ratios of adults were not related to brood manipulation or forest patch size. In addition, parental H/L ratios and body condition were not related to nestling H/L ratios. Our results suggest that during the breeding period the deleterious effects of habitat loss are seen explicitly in growing young.

Animals↗

Differences in the timing of reproduction between urban and forest European blackbirds (Turdus merula): result of phenotypic flexibility or genetic differences?

Species which have settled in urban environments are exposed to different conditions from their wild conspecifics. A previous comparative study of an urban and a forest-living European blackbird population had revealed a three weeks earlier onset of gonadal growth in urban individuals. These physiological adjustments are either the result of genetic differences that have evolved during the urbanization process, or of phenotypic flexibility resulting from the bird's exposure to the different environmental conditions of town or forest. To identify which of these two mechanisms causes the differences in reproductive timing, hand-reared birds originating from the urban and the forest populations were kept in identical conditions. The substantial differences in the timing of reproduction between urban and forest birds known from the field did not persist under laboratory conditions, indicating that temporal differences in reproductive timing between these two populations are mainly a result of phenotypic flexibility. Nevertheless, urban males initiated plasma luteinizing hormone (LH) secretion and testicular development earlier than forest males in their first reproductive season. Moreover, plasma LH concentration and follicle size declined earlier in urban females than in forest females, suggesting that genetic differences are also involved and might contribute to the variations in the timing of reproduction in the wild.

Animals↗

Coincident scales of forest feedback on climate and conservation in a diversity hot spot.

The dynamic relationship between vegetation and climate is now widely acknowledged. Climate influences the distribution of vegetation; and through a number of feedback mechanisms vegetation affects climate. This implies that land-use changes such as deforestation will have climatic consequences. However, the spatial scales at which such feedbacks occur remain largely unknown. Here, we use a large database of precipitation and tree cover records for an area of the biodiversity-rich Atlantic forest region in south eastern Brazil to investigate the forest-rainfall feedback at a range of spatial scales from ca 10(1)-10(4) km2. We show that the strength of the feedback increases up to scales of at least 10(3) km2, with the climate at a particular locality influenced by the pattern of landcover extending over a large area. Thus, smaller forest fragments, even if well protected, may suffer degradation due to the climate responding to land-use change in the surrounding area. Atlantic forest vertebrate taxa also require large areas of forest to support viable populations. Areas of forest of ca 10(3) km2 would be large enough to support such populations at the same time as minimizing the risk of climatic feedbacks resulting from deforestation.

Animals↗

Dynamics of the forest communities at Pasoh and Barro Colorado: comparing two 50-ha plots.

Dynamics of the Pasoh forest in Peninsular Malaysia were assessed by drawing a comparison with a forest in Panama, Central America, whose dynamics have been thoroughly described. Census plots of 50 ha were established at both sites using standard methods. Tree mortality at Pasoh over an eight-year interval was 1.46% yr(-1) for all stems > or = 10 mm diameter at breast height (dbh), and 1.48% yr(-1) for stems > or = 100 mm dbh. Comparable figures at the Barro Colorado Island site in Panama (BCI) were 2.55% and 2.03%. Growth and recruitment rates were likewise considerably higher at BCI than at Pasoh. For example, in all trees 500-700 mm in dbh, mean BCI growth over the period 1985-1995 was 6 mm yr(-1), whereas mean Pasoh growth was about 3.5 mm yr(-1). Examining growth and mortality rates for individual species showed that the difference between the forests can be attributed to a few light-demanding pioneer species at BCI, which have very high growth and mortality; Pasoh is essentially lacking this guild. The bulk of the species in the two forests are shade-tolerant and have very similar mortality, growth and recruitment. The Pasoh forest is more stable than BCI's in another way as well: few of its tree populations changed much over the eight-year census interval. In contrast, at BCI, over 10% of the species had populations increasing or decreasing at a rate of >0.05 yr(-1) compared to just 2% of the species at Pasoh). The faster species turnover at BCI can probably be attributed to severe droughts that have plagued the forest periodically over the past 30 years; Pasoh has not suffered such extreme events recently. The dearth of pioneer species at Pasoh is associated with low-nutrient soil and slow litter breakdown, but the exact mechanisms behind this association remain poorly understood.

Ecosystem↗

Contrasting simulated past and future responses of the Amazonian forest to atmospheric change.

Modelling simulations of palaeoclimate and past vegetation form and function can contribute to global change research by constraining predictions of potential earth system responses to future warming, and by providing useful insights into the ecophysiological tolerances and threshold responses of plants to varying degrees of atmospheric change. We contrasted HadCM3LC simulations of Amazonian forest at the last glacial maximum (LGM; 21 kyr ago) and a Younger Dryas-like period (13-12 kyr ago) with predicted responses of future warming to provide estimates of the climatic limits under which the Amazon forest remains relatively stable. Our simulations indicate that despite lower atmospheric CO2 concentrations and increased aridity during the LGM, Amazonia remains mostly forested, and that the cooling climate of the Younger Dryas-like period in fact causes a trend toward increased above-ground carbon balance relative to today. The vegetation feedbacks responsible for maintaining forest integrity in past climates (i.e. decreased evapotranspiration and reduced plant respiration) cannot be maintained into the future. Although elevated atmospheric CO2 contributes to a positive enhancement of plant carbon and water balance, decreased stomatal conductance and increased plant and soil respiration cause a positive feedback that amplifies localized drying and climate warming. We speculate that the Amazonian forest is currently near its critical resiliency threshold, and that even minor climate warming may be sufficient to promote deleterious feedbacks on forest integrity.

Atmosphere↗

Tropical forests and global atmospheric change: a synthesis.

We present a personal perspective on the highlights of the Theme Issue 'Tropical forests and global atmospheric change'. We highlight the key findings on the contemporary rate of climatic change in the tropics, the evidence-gained from field studies-of large-scale and rapid change in the dynamics and biomass of old-growth forests, and evidence of how climate change and fragmentation can interact to increase the vulnerability of plants and animals to fires. A range of opinions exists concerning the possible cause of these observed changes, but examination of the spatial 'fingerprint' of observed change may help to identify the driving mechanism(s). Studies of changes in tropical forest regions since the last glacial maximum show the sensitivity of species composition and ecology to atmospheric changes. Model studies of change in forest vegetation highlight the potential importance of temperature or drought thresholds that could lead to substantial forest decline in the near future. During the coming century, the Earth's remaining tropical forests face the combined pressures of direct human impacts and a climatic and atmospheric situation not experienced for at least 20 million years. Understanding and monitoring of their response to this atmospheric change are essential if we are to maximize their conservation options.

Atmosphere↗

A case study of nitrogen saturation in western U.S. forests.

Virtually complete nitrification of the available ammonium in soil and nitrification activity in the forest floor are important factors predisposing forests in the San Bernardino Mountains of southern California to nitrogen (N) saturation. As a result, inorganic N in the soil solution is dominated by nitrate. High nitrification rates also generate elevated nitric oxide (NO) emissions from soil. High-base cation saturation of these soils means that soil calcium depletion or effects associated with soil acidification are not an immediate risk for forest health as has been postulated for mesic forests in the eastern U.S. Physiological disturbance (e.g., altered carbon [C] cycling, reduced fine root biomass, premature needle abscission) of ozone-sensitive ponderosa pine trees exposed to high N deposition and high ozone levels appear to be the greater threat to forest sustainability. However, N deposition appears to offset the aboveground growth depression effects of ozone exposure. High nitrification activity reported for many western ecosystems suggests that with chronic N inputs these systems are prone to N saturation and hydrologic and gaseous losses of N. High runoff during the winter wet season in California forests under a Mediterranean climate may further predispose these watersheds to high nitrate leachate losses. After 4 years of N fertilization at a severely N saturated site in the San Bernardino Mountains, bole growth unexpectedly increased. Reduced C allocation below- ground at this site, presumably in response to ozone or N or both pollutants, may enhance the bole growth response to added N.

California↗

Rotation forest: A new classifier ensemble method.

We propose a method for generating classifier ensembles based on feature extraction. To create the training data for a base classifier, the feature set is randomly split into K subsets (K is a parameter of the algorithm) and Principal Component Analysis (PCA) is applied to each subset. All principal components are retained in order to preserve the variability information in the data. Thus, K axis rotations take place to form the new features for a base classifier. The idea of the rotation approach is to encourage simultaneously individual accuracy and diversity within the ensemble. Diversity is promoted through the feature extraction for each base classifier. Decision trees were chosen here because they are sensitive to rotation of the feature axes, hence the name "forest." Accuracy is sought by keeping all principal components and also using the whole data set to train each base classifier. Using WEKA, we examined the Rotation Forest ensemble on a random selection of 33 benchmark data sets from the UCI repository and compared it with Bagging, AdaBoost, and Random Forest. The results were favorable to Rotation Forest and prompted an investigation into diversity-accuracy landscape of the ensemble models. Diversity-error diagrams revealed that Rotation Forest ensembles construct individual classifiers which are more accurate than these in AdaBoost and Random Forest, and more diverse than these in Bagging, sometimes more accurate as well.

Algorithms↗

Rain forest promotes trophic interactions and diversity of trap-nesting Hymenoptera in adjacent agroforestry.

1. Human alteration of natural ecosystems to agroecosystems continues to accelerate in tropical countries. The resulting world-wide decline of rain forest causes a mosaic landscape, comprising simple and complex agroecosystems and patchily distributed rain forest fragments of different quality. Landscape context and agricultural management can be expected to affect both species diversity and ecosystem services by trophic interactions. 2. In Central Sulawesi, Indonesia, 24 agroforestry systems, differing in the distance to the nearest natural forest (0-1415 m), light intensity (37.5-899.6 W/m(-2)) and number of vascular plant species (7-40 species) were studied. Ten standardized trap nests for bees and wasps, made from reed and knotweed internodes, were exposed in each study site. Occupied nests were collected every month, over a period totalling 15 months. 3. A total of 13,617 brood cells were reared to produce adults of 14 trap-nesting species and 25 natural enemy species, which were mostly parasitoids. The total number of species was affected negatively by increasing distance from forest and increased with light intensity of agroforestry systems. The parasitoids in particular appeared to benefit from nearby forests. Over a 500-m distance, the number of parasitoid species decreased from eight to five, and parasitism rates from 12% to 4%. 4. The results show that diversity and parasitism, as a higher trophic interaction and ecosystem service, are enhanced by (i) improved connectivity of agroecosystems with natural habitats such as agroforestry adjacent to rain forest and (ii) management practices to increase light availability in agroforestry, which also enhances richness of flowering plants in the understorey.

Animals↗

When log-dwellers meet loggers: impacts of forest fragmentation on two endemic log-dwelling beetles in southeastern Australia.

Anthropogenic activities continue to cause massive fragmentation and reduction of forest area worldwide. With fragmentation and reduction of habitat recognized as the greatest threats to biodiversity, the implementation of improved, informed and conservation-based forestry practices is essential, and requires a greater understanding of the responses of different organisms to forest fragmentation. While genetic techniques can add invaluable insights to fragmentation studies they have rarely been employed, particularly for multiple species. In the present study, we combined genetic information, obtained from allozyme loci and anonymous single copy nuclear DNA markers, with ecological data to investigate the impacts of forest fragmentation on two log-dwelling beetles with different life histories, in an 'islands of bush in a sea of pine' model, at Tumut in New South Wales, Australia. Both the relatively mobile (i.e. has high dispersal ability and/or broad habitat range) Adelium calosomoides and the less mobile Apasis puncticeps showed reduced mobility and gene flow in fragmented compared to continuous forest: there was significantly greater isolation by distance and stronger local structure revealed by spatial autocorrelation in fragmented forest. Analysis of patch and species characteristics revealed that genetic and demographic structure may be influenced by log degradation class for both species, and number of potential dispersal barriers, distance from continuous forest and desiccation intolerance/moisture preference for Ap. puncticeps. Thus the pine plantation matrix poses a barrier or filter for gene flow and mobility in both beetle species.

Animals↗

Forest fragmentation effects on patch occupancy and population viability of herbaceous plant species.

Habitat fragmentation is one of the major threats to species diversity. In this review, we discuss how the genetic and demographic structure of fragmented populations of herbaceous forest plant species is affected by increased genetic drift and inbreeding, reduced mate availability, altered interactions with pollinators, and changed environmental conditions through edge effects. Reported changes in population genetic and demographic structure of fragmented plant populations have, however, not resulted in large-scale extinction of forest plants. The main reason for this is very likely the long-term persistence of small and isolated forest plant populations due to prolonged clonal growth and long generation times. Consequently, the persistence of small forest plant populations in a changing landscape may have resulted in an extinction debt, that is, in a distribution of forest plant species reflecting the historical landscape configuration rather than the present one. In some cases, fragmentation appears to affect ecosystem integrity rather than short-term population viability due to the opposition of different fragmentation-induced ecological effects. We finally discuss extinction and colonization dynamics of forest plant species at the regional scale and suggest that the use of the metapopulation concept, both because of its heuristic power and conservation applications, may be fruitful.

Biodiversity↗

The Northwest Forest Plan: origins, components, implementation experience, and suggestions for change.

In the 1990s the federal forests in the Pacific Northwest underwent the largest shift in management focus since their creation, from providing a sustained yield of timber to conserving biodiversity, with an emphasis on endangered species. Triggered by a legal challenge to the federal protection strategy for the Northern Spotted Owl (Strix occidentalis caurina), this shift was facilitated by a sequence of science assessments that culminated in the development of the Northwest Forest Plan. The plan, adopted in 1994, called for an extensive system of late-successional and riparian reserves along with some timber harvest on the intervening lands under a set of controls and safeguards. It has proven more successful in stopping actions harmful to conservation of old-growth forests and aquatic systems than in achieving restoration goals and economic and social goals. We make three suggestions that will allow the plan to achieve its goals: (1) recognize that the Northwest Forest Plan has evolved into an integrative conservation strategy, (2) conserve old-growth trees and forests wherever they occur and (3) manage federal forests as dynamic ecosystems.

Conservation of Natural Resources↗

Conserving old-growth forest diversity in disturbance-prone landscapes.

A decade after its creation, the Northwest Forest Plan is contributing to the conservation of old-growth forests on federal land. However the success and outlook for the plan are questionable in the dry provinces, where losses of old growth to wildfire have been relatively high and risks of further loss remain. We summarize the state of knowledge of old-growth forests in the plan area, identify challenges to conserve them, and suggest some conservation approaches that might better meet the goals of the plan. Historically, old-growth forests in these provinces ranged from open, patchy stands, maintained by frequent low-severity fire, to a mosaic of dense and open stands maintained by mixed-severity fires. Old-growth structure and composition were spatially heterogeneous, varied strongly with topography and elevation, and were shaped by a complex disturbance regime of fire, insects, and disease. With fire suppression and cutting of large pines (Pinus spp.) and Douglas-firs (Pseudotsuga menziesii [Mirbel] Franco), old-growth diversity has declined and dense understories have developed across large areas. Challenges to conserving these forests include a lack of definitions needed for planning of fire-dependent old-growth stands and landscapes, and conflicts in conservation goals that can be resolved only at the landscape level. Fire suppression has increased the area of the dense, older forest favored by Northern Spotted Owls (Strix occidentalis caurina) but increased the probability of high-severity fire. The plan allows for fuel reduction in late-successional reserves; fuel treatments, however apparently have not happened at a high enough rate or been applied in a landscape-level approach. Landscape-level strategies are needed that prioritize fuel treatments by vegetation zones, develop shaded fuel breaks in strategic positions, and thin and apply prescribed fire to reduce ladder fuels around remaining old trees. Evaluations of the current and alternative strategies are needed to determine whether the current reserve-matrix approach is the best strategy to meet plan goals in these dynamic landscapes.

Biodiversity↗

Preemptive and salvage harvesting of New England forests: when doing nothing is a viable alternative.

One unexpected consequence of natural disturbances in forested areas is that managers often initiate activities that may impose greater ecosystem impacts than the disturbances themselves. By salvage logging areas affected by windstorms or other impacts, by harvesting host trees in advance of insect infestation or disease, or by preemptively harvesting forests in an attempt to improve their resilience to future disturbances and stresses, managers initiate substantial changes in the ecosystem structure and function. Much of this activity is undertaken in the absence of information on the qualitative and quantitative differences between disturbance impacts and harvesting. To provide insight for such decisions we evaluated the ecosystem consequences of two major disturbance processes in New England (US.A.)--intense windstorms and invasive pests and pathogens-and contrasted them with impacts from preemptive and salvage harvesting. Despite dramatic physical changes in forest structure resulting from hurricane impacts and insect infestation, little disruption of biogeochemical processes or other ecosystem functions typically follows these disturbances. Indeed, the physical and organic structures produced by these disturbances are important natural features providing habitat and landscape heterogeneity that are often missing due to centuries of land use. From an ecosystem perspective there are strong arguments against preemptive and salvage logging or the attempt through silvicultural means to improve the resistance or resilience of forests to disturbance and stress. There are often valid motivations for salvage or preemptive logging including financial considerations, human safety, and a desire to shape the long-term composition and resource-production characteristics of forests. Nonetheless, there are many ecological benefits derived from leaving forests alone when they are affected or threatened by disturbances and pest and pathogen outbreaks.

Animals↗