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Can neutral theory predict the responses of Amazonian tree communities to forest fragmentation?

We use Hubbell's neutral theory to predict the impact of habitat fragmentation on Amazonian tree communities. For forest fragments isolated for about two decades, we generate neutral predictions for local species extinction, changes in species composition within fragments, and increases in the probability that any two trees within a fragment are conspecific. We tested these predictions using fragment and intact forest data from the Biological Dynamics of Forest Fragments Project in central Amazonia. To simulate complete demographic isolation, we excluded immigrants--species absent from a fragment or intact forest plot in the initial census but present in its last census--from our tests. The neutral theory correctly predicted the rate of species extinction from different plots as a function of the diversity and mortality rate of trees in each plot. However, the rate of change in species composition was much faster than predicted in fragments, indicating that different tree species respond differently to environmental changes. This violates the key assumption of neutral theory. When immigrants were included in our calculations, they increased the disparity between predicted and observed changes in fragments. Overall, neutral theory accurately predicted the pace of local extinctions in fragments but consistently underestimated changes in species composition.

Conservation of Natural Resources↗

Seasonal biochemical changes in coniferous forest canopies and their response to fertilization.

Seasonal changes in concentrations of total nitrogen, free amino acids, chlorophyll, starch and sugar were measured in foliage from fertilized and unfertilized conifer forests in New Mexico and Oregon. In the New Mexico Douglas-fir (Pseudotsuga menziesii var glauca (Beissn.) Franco) forest, fertilization resulted in elevated foliar nitrogen concentrations on all dates, from an average of 9 mg g(-1) in unfertilized trees to 14 mg g(-1) in fertilized trees. In the Oregon western hemlock (Tsuga heterophylla (Raf.) Sarg.) forest, fertilization increased total N by only 15%, from 13 mg g(-1) in unfertilized trees to 15 mg g(-1) in fertilized trees. Foliar nitrogen concentrations on a weight basis were lowest in winter and spring, but did not vary seasonally when expressed on a leaf area basis. Chlorophyll concentrations increased with fertilization and had greater seasonal variation than did total nitrogen concentrations. Chlorophyll concentrations were significantly higher during the growing season than in the winter and spring months. Fertilization did not result in major changes in the proportion of total nitrogen in chlorophyll at either the Oregon or the New Mexico site. Concentrations of free amino acids varied with date and fertilization treatment; in New Mexico, amino acids were highest in the winter sample, whereas in Oregon, they were lowest in winter and spring. At both sites, amino acid concentrations were significantly higher in fertilized trees than in control trees on most dates and the ratios of amino acid-N to total N were also significantly higher in fertilized trees. For both sites, starch concentrations were nearly zero for most of the year, but increased sharply just before bud break and initiation of new growth in the spring. Although fertilization resulted in increased nitrogen concentrations in foliage at both sites, the response in New Mexico was much greater than in Oregon. These results are in agreement with forest productivity data that suggest that growth in the New Mexico site is limited by nitrogen, whereas in the Oregon site it is not.

Journal Article↗

Transpiration of a boreal pine forest measured by branch bag, sap flow and micrometeorological methods.

Three independent methods were used to evaluate transpiration of a boreal forest: the branch bag, sap flow and eddy covariance methods. The branch bag method encloses several thousand needles and gives a continuous record of branch transpiration. The sap flow method provides a continuous record of sap velocity and an estimate of tree transpiration. The eddy covariance method typically measures evaporation rates between a forest and the atmosphere. We deployed an extra eddy covariance system below the forest to estimate canopy transpiration by difference. The three systems detected small water vapor fluxes despite a plentiful supply of energy to drive evaporation. We also observed that transpiration rates were low even when the soil was well supplied with water. Low rates of transpiration were attributed to the canopy's low leaf area index and the marked reduction in stomatal conductance as vapor pressure deficits increased. Water vapor fluxes, derived from the sap flow method, lagged behind those derived by the branch bag method by 1 to 2 h. The sap flow method also suffered from sampling errors caused by the non-uniformity of flow across the sapwood and the spatial variability of sapwood cross section throughout the forest. Despite technical difficulties associated with hourly measurements, daily totals of transpiration agreed well with values derived from micrometeorological systems.

Journal Article↗

Net primary production of forests: a constant fraction of gross primary production?

Considerable progress has been made in our ability to model and measure annual gross primary production (GPP) by terrestrial vegetation. But challenges remain in estimating maintenance respiration (R(m)) and net primary production (NPP). To search for possible common relationships, we assembled annual carbon budgets from six evergreen and one deciduous forest in Oregon, USA, three pine plantations in New South Wales, Australia, a deciduous forest in Massachusetts, USA, and a Nothofagus forest on the South Island of New Zealand. At all 12 sites, a standard procedure was followed to estimate annual NPP of foliage, branches, stems, and roots, the carbon expended in synthesis of these organs (R(g)), their R(m), and that of previously produced foliage and sapwood in boles, branches, and large roots. In the survey, total NPP ranged from 120 to 1660 g C m(-2) year(-1), whereas the calculated fraction allocated to roots varied from 0.22 to 0.63. Comparative analysis indicated that the total NPP/GPP ratio was conservative (0.47 +/- 0.04 SD). This finding supports the possibility of greatly simplifying forest growth models. The constancy of the NPP/GPP ratio also provides an incentive to renew efforts to understand the environmental factors affecting partitioning of NPP above and belowground.

Journal Article↗

Use of middle infrared radiation to estimate the leaf area index of a boreal forest.

The leaf area index (LAI) of boreal forest can be estimated using reflected radiation recorded by satellite sensors. Measurements of visible and near infrared radiation are commonly used in the normalized difference vegetation index (NDVI) to estimate LAI. However, research, mainly in tropical forest, has demonstrated that LAI is related more closely to radiation of middle infrared wavelengths than of visible wavelengths. This paper derives a vegetation index, VI3, based on radiation from vegetation recorded at near and middle infrared wavelengths. For a boreal forest canopy, the relationship between VI3 and LAI was observed to be much stronger than that between NDVI and LAI. In addition, the LAI estimated using VI3 accounted for about 76% of the variation in field estimates of LAI, compared with about 46% when using the NDVI. We conclude that information provided by middle infrared radiation should be considered when estimating the leaf area index of boreal forest.

Journal Article↗

Photosynthetic utilization efficiency of absorbed photosynthetically active radiation by Scots pine and birch forest stands in the southern Taiga.

Absorption and utilization of photosynthetically active radiation (PAR) were investigated in Scots pine (Pinus sylvestris L.) and birch (Betula pendula Roth.) stands that were 41 years old at the end of the experimental period. Canopy depth of the Scots pine stand was about half that of the birch stand (6.5 versus 11.0 m), but absorption of PAR was similar in the two stands. The Scots pine forest canopy, with a leaf area index of 8.9, absorbed 90% of the incoming PAR (APAR), whereas the birch forest canopy, with a leaf area index of 5.9, absorbed 92% of APAR. During maximum foliage development, the upper Scots pine canopy absorbed more PAR than the upper birch canopy (75 versus 66%). The upper, middle and lower layers of the Scots pine canopy contained 37, 48 and 15% of the total needle surface area, respectively. The corresponding distribution of foliage surface area in the three layers of the birch canopy was 50, 30 and 20%, respectively. Measurements of photosynthetic rate were combined with estimates of leaf area index and stand phytomass to determine rates of primary production on a sunny day, a cloudy day, and on an annual basis. The energy equivalents of short- and long-term carbon gain were used with determinations of APAR to calculate photosynthetic utilization efficiency. Throughout the growing season, photosynthetic utilization efficiency of APAR in the upper canopy layer of the Scots pine forest was almost twice that in the lower canopy layer. In the birch forest, photosynthetic utilization efficiency was greater in the lower canopy layer than in the upper canopy layer. In all cases, utilization efficiency was higher in the birch stand than in the Scots pine stand (52 versus 29 J kJ(-1)). Taking account of respiration of the non-photosynthetic parts of each stand (night respiration of needles or leaves; respiration of branches, trunk and roots), estimated utilization efficiency of APAR for net primary production was 11 J kJ(-1) for Scots pine and 19 J kJ(-1) for birch. Solar conversion ratios, expressed as whole-plant net primary productivity per unit of APAR for the growing season, were 0.81 g MJ(-1) for Scots pine and 0.93 g MJ(-1) for birch.

Journal Article↗

Process-based models for forest ecosystem management: current state of the art and challenges for practical implementation.

Recent progress toward the application of process-based models in forestmanagement includes the development of evaluation and parameter estimation methods suitable for models with causal structure, and the accumulation of data that can be used in model evaluation. The current state of the art of process modeling is discussed in the context of forest ecosystem management. We argue that the carbon balance approach is readily applicable for projecting forest yield and productivity, and review several carbon balance models for estimating stand productivity and individual tree growth and competition. We propose that to develop operational models, it is necessary to accept that all models may have both empirical and causal components at the system level. We present examples of hybrid carbon balance models and consider issues that currently require incorporation of empirical information at the system level. We review model calibration and validation methods that take account of the hybrid character of models. The operational implementation of process-based models to practical forest management is discussed. Methods of decision-making in forest management are gradually moving toward a more general, analytical approach, and it seems likely that models that include some process-oriented components will soon be used in forestry enterprises. This development is likely to run parallel with the further development of ecophysiologically based models.

Journal Article↗

Forest growth and species distribution in a changing climate.

Climate change has many potential effects on plants, some detrimental to growth, others beneficial. Increasing CO(2) concentration can increase photosynthetic rates, with the greatest increases likely to be in C(3) plants growing in warm dry conditions. Increasing temperature directly affects plant growth through effects on photosynthetic and respiration rates. However, plants have a considerable ability to adapt to changing conditions and can tolerate extremely high temperatures, provided that adequate water is available. Increasing temperature may increase vapor pressure deficits of the air, and thereby increase transpiration rates from most plant canopies. Effects are likely to vary among plant communities, with forests generally experiencing greater increases in transpiration rates than grasslands. These increases in transpiration are likely to be reduced by stomatal closure in response to increasing CO(2) concentration. In many areas, precipitation will probably increase with global warming; however, these increases may be insufficient to meet the increased transpirational demand by plant canopies. Increasing temperature is likely to increase soil organic matter decomposition rates so that nutrients may be more readily mineralized and made available to plants. In highly fertile systems, this could lead to nutrient losses through leaching. For different combinations of increases in temperature and CO(2) concentration, and for systems primarily affected by water or nutrient limitations, different overall effects on plant productivity can be expected. Responses will be negative in some circumstances and positive in others, but on the whole, catastrophic changes to forest growth seem unlikely under most conditions. In contrast,ecological consequences of climate change are potentially more serious. The distribution of many species tends to be limited to a narrow range of environmental conditions. Climate conditions over much of a species' current natural range may therefore become unsuitable, leading to significant decline of forests or of particular species within forests.

Journal Article↗

Leaf optical properties in Venezuelan cloud forest trees.

Leaf optical properties and related leaf characteristics were compared for thirteen cloud forest tree species differing in successional status. Sun leaves were sampled for the eight pioneer species and sun and shade leaves were sampled for the five climax species. Sun leaves had a slightly higher absorptance than shade leaves, although differences were small. Sun leaves had a higher leaf mass per unit area (LMA) and a lower chlorophyll concentration per unit leaf mass, resulting in similar chlorophyll concentrations per unit leaf area and hence similar light harvesting capacities as shade leaves. However, shade leaves realized a higher efficiency of absorptance per unit leaf biomass than sun leaves. There were few differences in leaf characteristics of sun leaves between the climax and pioneer species. Absorptance values of cloud forest species were comparable with values reported for rain forest and more seasonal forest species. Intraspecific variation in leaf absorptance was largely the result of variation in LMA, whereas interspecific variation in leaf absorptance was largely a result of variation in chlorophyll concentration per unit leaf area.

Journal Article↗

Thinning effects on carbon allocation to fine roots in a Quercus ilex forest.

The dynamics of the fine root system are relevant to calculations of the carbon balance of the ecosystem, and there is also a need to quantify changes in this component caused by disturbances. Mediterranean forest systems have historically been coppiced to obtain charcoal. As a result of the resprouting capacity of holm oaks (Quercus ilex L.), these forests present more than 50% of their biomass below ground (stump + roots > 1 cm in diameter), but the effects of thinning on the fine root system are unknown. Fine root biomass, production, mortality and longevity were studied in a control and a thinned Mediterranean holm oak forest by minirhizotron methodology. Observations of fine roots started 2 years after thinning and continued for almost 3 years. Extraction of 80% of the former basal area of the forest greatly affected carbon allocation patterns. Biomass increased by more than 100%, production increased by 76%, mortality increased by 32% and longevity decreased by around 2 weeks. The greatest differences between treatments were associated with differences in growth during autumn months in the top 40 cm of soil, particularly between 10 and 20 cm depth.

Biomass↗

Above- and belowground biomass and net primary production in a 73-year-old Scots pine forest.

We estimated above- and belowground biomass and net primary production (NPP) of a 73-year-old Scots pine (Pinus sylvestris L.) forest stand in the Belgian Campine region. Total biomass for the stand was 176 Mg ha(-1), of which 74.4% was found in stems. The root system contained 12.6% of total biomass, most of it in coarse roots (> 5 mm). Fine roots (< 5 mm) comprised only about 1.7% of total biomass, and more than 50% of fine root biomass was retrieved in the litter layer and the upper 15 cm of the mineral soil. The ratio of belowground biomass to aboveground biomass was 0.14, which is lower than that of other Scots pine forests and other coniferous forests. Between 1995 and 2001, mean annual NPP was 11.2 Mg ha(-1) year(-1), of which 68.7% was allocated to aboveground compartments. Stems, needles and cones made relatively high contributions to total NPP compared with branches. However, branch NPP was possibly underestimated because litterfall of big branches was neglected. The proportion of total NPP in belowground components was 31.3%. Coarse root NPP (2% of total) was low compared with its biomass. Fine root NPP was 3.3 Mg ha(-1) year(-1), representing about 29.5% of total NPP; however, the estimate of fine root NPP is much more uncertain than NPP of aboveground compartments. The ratio NPP/GPP (gross primary production) was 0.32, which was low compared with other coniferous forests.

Belgium↗

Enhancement of understory productivity by asynchronous phenology with overstory competitors in a temperate deciduous forest.

Some saplings and shrubs growing in the understory of temperate deciduous forests extend their periods of leaf display beyond that of the overstory, resulting in periods when understory radiation, and hence productivity, are not limited by the overstory canopy. To assess the importance of the duration of leaf display on the productivity of understory and overstory trees of deciduous forests in the north eastern United States, we applied the simulation model, BIOME-BGC with climate data for Hubbard Brook Experimental Forest, New Hampshire, USA and mean ecophysiological data for species of deciduous, temperate forests. Extension of the overstory leaf display period increased overstory leaf area index (LAI) by only 3 to 4% and productivity by only 2 to 4%. In contrast, extending the growing season of the understory relative to the overstory by one week in both spring and fall, increased understory LAI by 35% and productivity by 32%. A 2-week extension of the growing period in both spring and fall increased understory LAI by 53% and productivity by 55%.

Ecosystem↗

Parameter sensitivity and uncertainty of the forest carbon flux model FORUG: a Monte Carlo analysis.

The Monte Carlo technique can be used to propagate input variable uncertainty and parameter uncertainty through a model to determine output uncertainty. However, to carry out Monte Carlo simulations, the uncertainty distributions or the probability density functions (PDFs) of the model parameters and input variables must be known. This remains one of the bottlenecks in current uncertainty research in forest carbon flux modeling. Because forest carbon flux models involve many parameters, we questioned whether it is necessary to take into account all parameters in the uncertainty analysis. A sensitivity analysis can determine the parameters contributing most to the overall model output uncertainty. This paper illustrates the usefulness of the Monte Carlo simulation technique for ranking parameters for sensitivity and uncertainty in process-based forest flux models. The uncertainty of the output (net ecosystem exchange, NEE) of the FORUG model was estimated for the Hesse beech forest (1997). Based on the arbitrary uncertainty of ten key parameters, a standard deviation of 0.88 Mg C ha(-1) year(-1) NEE was found which is equal to 24% of the mean value of NEE. Sensitivity analysis showed that the overall output uncertainty of the FORUG model can largely be determined by accounting for the uncertainty of only a few key parameters. The results led to the identification of the key FORUG parameters and to the recommendation for a process-based description of the soil respiration process in the FORUG model.

Carbon↗

Physiological and phenological responses of oak seedlings to oak forest soil in the absence of trees.

Established trees influence the growth and physiology of seedlings by altering above- and belowground conditions; however, tree influences on seedling physiology via belowground interactions are not well understood. We used soil transfers to an open field to examine the belowground influences of a Quercus ellipsoidalis E.J. Hill dominated forest on Q. ellipsoidalis seedling mycorrhizal infection, nutrient uptake, growth and photosynthesis over three years. After two years, seedlings planted with large quantities of forest soil (HF treatment) had greater leaf mass and foliar N concentrations than seedlings receiving smaller quantities of forest soil (LF) and control treatments. Mycorrhizal infection was greater in the HF treatment after one year compared with the LF and control treatments, with a positive correlation of foliar N and mycorrhizal infection in Year 2. There were marked effects of treatments on seedling spring phenology with HF seedlings breaking bud up to 17 days earlier than seedlings in the other treatments. The HF seedlings also had more rapid leaf expansion and larger leaves, and an increase in net photosynthetic rates. These results highlight complex linkages between above- and belowground physiology: forest soil had substantial effects on seedling physiology, including traits such as phenology that have previously been considered to be under aboveground control. Belowground influences of trees on conspecific seedlings may play a critical role in early seedling establishment.

Biomass↗

The concept of closure in calculating carbon balance of forests: accounting for differences in spatial and temporal scales of component processes.

The concept of closure, which has been used extensively to calculate the heat and water budgets of forests, has led to advances in the understanding of how heat and water balances are controlled. The successful construction of closed heat and water budgets of forests can be attributed to the large scale of input and output fluxes relative to changes in storage, the comparative ease with which such fluxes can be measured, and the regularity with which forests cycle through discharge and recharge on diurnal and annual cycles. The applicability of simple, closed budget analysis for carbon balance of forests is less certain because changes in internal storage, which are of major significance, occur at different time scales for growth and decomposition. We conclude that carbon balance might best be modeled as an open system. For long-term predictions, the most important processes to consider are acclimation and adaptation, which occur at different spatial and temporal scales. Construction of a modeling procedure to accommodate this variation is discussed.

Journal Article↗

Amplicon-based analyses of single-nucleotide polymorphisms reveal the genetic structure of a forest insect baculovirus.

Amplicon-based next-generation sequencing (aNGS) is a powerful tool in diagnostics and genetic studies. We developed an aNGS approach to study the population structure of the Lymantria dispar multiple nucleopolyhedrovirus (LdMNPV), a specific pathogen of the spongy moth Lymantria dispar, a devastating lepidopteran pest in European, Asian, and American deciduous forests. Naturally occurring pathogens, such as LdMNPV, are frequently reported to cause epizootics and a rapid decline of insect pest populations. DNA samples of pooled LdMNPV-infected larvae from forest regions in Northern Bavaria (Germany) were subjected to whole genome sequencing (WGS) and aNGS optimization. Then, five marker regions were identified in the genome of LdMNPV for PCR amplification, covering 21 highly specific single-nucleotide polymorphism (SNP) positions that enabled comprehensive analysis at the intra- and intersample levels. These markers were used in aNGS analyses of 70 single larvae collected in 12 forest sites, followed by SNP-based hierarchical clustering on principal components (HCPC). This approach identified three LdMNPV population clusters consisting of homogenous (pure) and heterogeneous (mixed) LdMNPV samples. To explain the genetic variability within each sample, a model based on linear optimization was developed and validated by comparing the predictions from aNGS and WGS data. The analyses showed that LdMNPV from Bavarian forests carried genetic variants highly similar to those present in the commercial product Gypchek&#xae;, developed for biocontrol. The distribution of genetic characteristics showed some trends of geographic and temporal prevalence, which are indicative of short-distance and long-distance transmission. The aNGS approach offers a fast, cost-effective, and comprehensive insight into the natural population structure of LdMNPV.

insects↗

Changes in the use and management of forests for abating carbon emissions: issues and challenges under the Kyoto Protocol.

The global carbon cycle is significantly influenced by changes in the use and management of forests and agriculture. Humans have the potential through changes in land use and management to alter the magnitude of forest-carbon stocks and the direction of forest-carbon fluxes. However, controversy over the use of biological means to absorb or reduce emissions of CO(2) (often referred to as carbon 'sinks') has arisen in the context of the Kyoto Protocol. The controversy is based primarily on two arguments: sinks may allow developed nations to delay or avoid actions to reduce fossil fuel emissions, and the technical and operational difficulties are too threatening to the successful implementation of land use and forestry projects for providing carbon offsets. Here we discuss the importance of including carbon sinks in efforts to address global warming and the consequent additional social, environmental and economic benefits to host countries. Activities in tropical forest lands provide the lowest cost methods both of reducing emissions and reducing atmospheric concentrations of greenhouse gases. We conclude that the various objections raised as to the inclusion of carbon sinks to ameliorate climate change can be addressed by existing techniques and technology. Carbon sinks provide a practical available method of achieving meaningful reductions in atmospheric concentrations of carbon dioxide while at the same time contribute to national sustainable development goals.

Air Pollutants↗

Comparative aspects of diet in Amazonian forest-dwellers.

Recent research shows that lowland forests of the Amazon Basin differ in numerous ways including features of climate and soils, faunal composition and forest structure, composition and phenology. Such differences strongly suggest that single-factor models used to explain features of human ecology in Amazonia may be too limited. A comparative study of the dietary ecology of four forest-living indigenous groups in Brazil (Arara, Parakana, Arawete, Mayoruna) revealed a number of differences. Primary crops, as well as animal types most utilized as prey, were found to differ markedly between groups. Although some differences can be accounted for by general environmental factors, no compelling single environmental factor can explain why any one group could not behave dietarily in ways more similar to another. Many of these intergroup dietary differences appear to represent a type of cultural character displacement that aids in distinguishing the members of one group from another. As all human groups, through the medium of culture, are actual or potential occupants of the same dietary niche, each group may distance itself from potential dietary rivals through cultural conventions. This behaviour may be justified, as the lack of overlap between forest-living groups in combination with generally intense intergroup hostility suggests that the biomass and distribution patterns of critical dietary resources in this environment may set limits to viable population size for particular areas.

Animals↗