Competition among eucalyptus trees depends on genetic variation and resource supply

Genetic variation and environmental heterogeneity fundamentally shape the interactions between plants of the same species. According to the resource partitioning hypothesis, competition between neighbors intensifies as their similarity increases. Such competition may change in response to increasing supplies of limiting resources. We tested the resource partitioning hypothesis in stands of genetically identical (clone-origin) and genetically diverse (seed-origin) Eucalyptus trees with different water and nutrient supplies, using individual-based tree growth models. We found that genetic variation greatly reduced competitive interactions between neighboring trees, supporting the resource partitioning hypothesis. The importance of genetic variation for Eucalyptus growth patterns depended strongly on local stand structure and focal tree size. This suggests that spatial and temporal variation in the strength of species interactions leads to reversals in the growth rank of seed-origin and clone-origin trees. This study is one of the first to experimentally test the resource partitioning hypothesis for intergenotypic vs. intragenotypic interactions in trees. We provide evidence that variation at the level of genes, and not just species, is functionally important for driving individual and community-level processes in forested ecosystems.     

Tree species diversity and its relationship to stand parameters and geomorphology features in the eastern Black Sea region forests of Turkey

We studied the effects of stand parameters (crown closure, basal area, stand volume, age, mean stand diameter number of trees, and heterogeneity index) and geomorphology features (elevation, aspect and slope) on tree species diversity in an example of untreated natural mixed forest stands in the eastern Black Sea region of Turkey. Tree species diversity and basal area heterogeneity in forest ecosystems are quantified using the Shannon-Weaver and Simpson indices. The relationship between tree species diversity basal area heterogeneity stand parameters and geomorphology features are examined using regression analysis. Our work revealed that the relationship between tree species diversity and stand parameters is loose with a correlation coefficient between 0.02 and 0.70. The correlation of basal area heterogeneity with stand parameters fluctuated between 0.004 and 0.77 (R2). According to our results, stands with higher tree species diversity are characterised by higher mean stand diameter number of diameter classes, basal area and lower homogeneity index value. Considering the effect of geomorphology features on tree species or basal area heterogeneity we found that all investigated relationships are loose with R < or = 0.24. A significant correlation was detected only between tree species diversity and aspect. Future work is required to verify the detected trends in behaviour of tree species diversity if it is to estimate from the usual forest stand parameters and topography characteristics.     

Effects of Harvesting on Genetic Diversity in Old-Growth Eastern White Pine in Ontario, Canada

Genetic diversity measures at 54 isozyme loci coding for 16 enzymes in megagametophytes were compared between preharvest and postharvest gene pools of two adjacent virgin, old-growth (∼250 years) stands of eastern white pine ( Pinus strobus L.) in the Galloway Lake Old Pine Area of central Ontario. The concurrence of genetic diversity changes between the stands suggests that real and repeatable genetic erosion occurred in these gene pools as a result of harvesting. The total and mean number of alleles detected in each stand were reduced by approximately 25% after tree density reductions of 75%. The percentage of polymorphic loci dropped by about 33% from preharvest levels. About 40% of the low frequency (0.25> p ≥ 0.01) alleles and 80% of the rare ( p < 0.01) alleles were lost from each stand because of harvesting. Hypothetical multilocus gametic diversity was reduced by about 40% in each stand after harvesting. Latent genetic potential of each stand was reduced by about 50%, suggesting that the ability of these gene pools to adapt to changing environmental conditions may have been compromised. Heterozygosity estimates in the postharvest stands did not reflect reductions in allelic richness due to harvesting. Observed heterozygosity increased by 12% in one stand after harvesting, even though other genetic diversity measures decreased. Gene frequency changes due to harvesting imply that gene pools of naturally regenerated progeny stands may be quite different from the original parental stands. Silvicultural practices should ensure that the gene pools of remaining pristine old-growth stands have been reconstituted in the regenerating stands.     

Spatially explicit modeling of overstory manipulations in young forests: Effects on stand structure and light

Young forests can be manipulated in diverse ways to enhance their ecological values. We used stem maps from two dense, second-growth stands in western Washington and a spatially explicit light model (tRAYci) to simulate effects of five silvicultural manipulations on diameter distribution, species composition, spatial patterning, and light availability. Each treatment removed 30% of the basal area, but differed in how trees were selected for removal. Three primary treatments were thin from below (removing the smallest trees), random thin (removing trees randomly), and gap creation (removing all trees in circles ∼1 tree height in diameter). Two additional treatments combined elements of these approaches: random ecological thin (a mixture of thin from below and random thin) and structured ecological thin (a mixture of thin from below and gap creation).     

More asymmetric tree competition brings about more evapotranspiration and less runoff from the forest ecosystems: A simulation study

The present paper reports how stand size-structure dynamics due to competition between different-sized trees affect long-term forested water balance in Japanese cool-temperate planted stands (evergreen coniferous Cryptomeria japonica and deciduous coniferous Larix kaempferi stands) using a fully coupled multi-layered meteorological surface physics—terrestrial ecosystems model. The simulation captured the well-known annual variation in leaf area index (LAI) accurately with stand age in monocultured and even-aged stands; the occurrence of maximum LAI during the early growth stage and then a gradual decline followed by a steady state after the maximum LAI. The simulations also detected a high dependency of annual evapotranspiration (AETr) on LAI with stand age that is well known by prior observational researches. In the C. japonica (shade-tolerant late-successional species) stand, the relationship between annual net primary productivity of an individual tree (NPP_ind) and individual tree mass (w) changed from linear to a convex curve during self-thinning, indicating that the degree of asymmetric tree competition intensified with forest stand development. The higher degree of competitive asymmetry characterized by the convex-shaped NPPind-w relationship produced greater size inequality, i.e., the formation of trees stratified by height. Under such conditions, AETr and annual transpiration (ATr) were mainly regulated by larger trees. On the other hand, the NPPind-w relationships in the L. kaempferi (shade-intolerant early-successional species) stand were linear throughout the simulated period, indicating the lower degree of competitive asymmetry. Under such conditions, the growth of intermediate-sized trees was enhanced and these trees became a dominant source of AETr (and also ATr) during self-thinning. Furthermore, the sensitivity analysis of the effects of ecophysiological parameters such as foliage profile (i.e., vertical distribution of leaf area density) of an individual tree (distribution pattern is described by the parameter η), the maximum carboxylation velocity (V_cmax0) and biomass allocation pattern of individual plant growth (μ₁) on AETr, ATr and annual runoff (AR_off) showed that the temporal trends of AETr, ATr, AR_off and NPPind-w relationships were completely the same as those in the control simulations. However, the NPPind-w relationship during self-thinning indicated higher degrees of competitive asymmetry when η or V_cmax0 were greater than those in the control simulation and generated greater AETr and ATr and thus smaller AR_off. We found that more asymmetric tree competition brings about greater size inequality between different-sized trees and thus more evapotranspiration and less runoff in a forest stand. Overall, our simulation approach revealed that not only LAI dynamics but also plant competition, and thus size-structure dynamics, in a forest ecosystem are essential to long-term future projections of forested water balance.     

Tree mortality following partial harvests is determined by skidding proximity

Recently developed structural retention harvesting strategies aim to improve habitat and ecological services provided by managed forest stands by better emulating natural disturbances. The potential for elevated mortality of residual trees following such harvests remains a critical concern for forest managers, and may present a barrier to more widespread implementation of the approach. We used a harvest chronosequence combined with dendrochronological techniques and an individual-based neighborhood analysis to examine the rate and time course of residual-tree mortality in the first decade following operational partial "structural retention" harvests in the boreal forest of Ontario, Canada. In the first year after harvest, residual-tree mortality peaked at 12.6 times the preharvest rate. Subsequently, mortality declined rapidly and approached preharvest levels within 10 years. Proximity to skid trails was the most important predictor both of windthrow and standing death, which contributed roughly equally to total postharvest mortality. Local exposure further increased windthrow risk, while crowding enhanced the risk of standing mortality. Ten years after harvest, an average of 10.5% of residual trees had died as a result of elevated postharvest mortality. Predicted cumulative elevated mortality in the first decade after harvest ranged from 2.4% to 37% of residual trees across the observed gradient of skid trail proximity, indicating that postharvest mortality will remain at or below acceptable rates only if skidding impacts are minimized. These results represent an important step toward understanding how elevated mortality may influence stand dynamics and habitat supply following moderate-severity disturbances such as partial harvests, insect outbreaks, and windstorms.     

Thinning effects on production, root biomass and some soil properties in a young oriental beech stand in Artvin, Turkey

In this study, three thinning treatments were applied on a 0.54 ha young oriental beech (Fagus orientalis Lipsky) stand (25-30 years old) in Karadag, Artvin in the fall of 1999. The treatments were: (1) no thinning, (2) light thinning, and (3) heavy thinning. Prior to thinning, the stand averaged 15000 trees ha(-1) and 40.0 m2 ha(-1) of basal area, with an average mean diameter of 5 cm. Thinning reduced stand basal areas to about 31.1 and 24.9 m2 ha(-1) for the light and heavy thinning treatments, respectively. After 3-years, diameter increment was the highest in heavily thinned stand and the lowest in unthinned stand. The increment in stand basal area was about 10, 18 and 27% in the unthinned, lightly thinned, and heavily thinned stands, respectively. Fine root biomass significantly decreased with thinning and thinning had no significant effects on soil pH and soil organic matter content.     

Neighborhood analyses of canopy tree competition along environmental gradients in New England forests.

We use permanent-plot data from the USDA Forest Service's Forest Inventory and Analysis (FIA) program for an analysis of the effects of competition on tree growth along environmental gradients for the 14 most abundant tree species in forests of northern New England, USA. Our analysis estimates actual growth for each individual tree of a given species as a function of average potential diameter growth modified by three sets of scalars that quantify the effects on growth of (1) initial target tree size (dbh), (2) local environmental conditions, and (3) crowding by neighboring trees. Potential growth of seven of the 14 species varied along at least one of the two environmental axes identified by an ordination of relative abundance of species in plots. The relative abundances of a number of species were significantly displaced from sites where they showed maximum potential growth. In all of these cases, abundance was displaced to the more resource-poor end of the environmental gradient (either low fertility or low moisture). The pattern was most pronounced among early successional species, whereas late-successional species reached their greatest abundance on sites where they also showed the highest growth in the absence of competition. The analysis also provides empirical estimates of the strength of intraspecific and interspecific competitive effects of neighbors. For all but one of the species, our results led us to reject the hypothesis that all species of competitors have equivalent effects on a target species. Most of the individual pairwise interactions were strongly asymmetric. There was a clear competitive hierarchy among the four most shade-tolerant species, and a separate competitive hierarchy among the shade-intolerant species. Our results suggest that timber yield following selective logging will vary dramatically depending on the configuration of the residual canopy, because of interspecific variation in the magnitude of both the competitive effects of different species of neighbors and the competitive responses of different species of target trees to neighbors. The matrix of competition coefficients suggests that there may be clear benefits in managing for specific mixtures of species within local neighborhoods within stands.     

A diameter increment model for Crimean pine (Pinus nigra Arnold) and Calabrian pine (Pinus brutia Ten.) stands in Isparta region, Turkey

The purpose of this research was to test the precision of a diameter increment model for the estimation of future periodic diameter increment. Individual trees of Crimean pine (Pinus nigra Arnold) and Calabrian pine (Pinus brutia Ten.) located in both natural and plantation stands were selected. For that reason, normal closed canopy, pure, even-aged and undisturbed stands were examined. In 2002, plots were sampled in three natural and three plantation stands in Isparta region of Turkey. The number of sampling points in sample plots ranged from 19 to 55. In each sampling point, a subject tree and six competitors were selected. In each sampling point, subject tree and competitor trees were stem mapped (x and y coordinate system), and diameter (dbh), total height, age, and 10-yrs radial increment recorded. The predictors of a distance dependent diameter increment model were chosen that included tree level (diameter (d), competition index (CI), and age (t)) and stand level (basal area (G), and site index (SI)) characteristics as well as their transformations. The best fit index of the regression model was pursued in trials with variable combinations. The models explained 65%, 60%, 68% and 50% of the variation in individual tree diameter increment of Crimean pine and Calabrian pine for both natural and plantations stands, respectively. These models can be estimated diameter increment of individual trees at highly significant level (p<0.001).     

Forest structure and light regimes following moderate wind storms: implications for multi-cohort management.

Moderate-severity disturbances appear to be common throughout much of North America, but they have received relatively little detailed study compared to catastrophic disturbances and small gap dynamics. In this study, we examined the immediate impact of moderate-intensity wind storms on stand structure, opening sizes, and light regimes in three hemlock-hardwood forests of northeastern Wisconsin. These were compared to three stands managed by single-tree and group selection, the predominant forest management system for northern hardwoods in the region. Wind storms removed an average of 41% of the stand basal area, compared to 27% removed by uneven-aged harvests, but both disturbances removed trees from a wide range of size classes. The removal of nearly half of the large trees by wind in two old-growth stands caused partial retrogression to mature forest structure, which has been hypothesized to be a major disturbance pathway in the region. Wind storms resulted in residual stand conditions that were much more heterogeneous than in managed stands. Gap sizes ranged from less than 10 m2 up to 5000 m2 in wind-disturbed stands, whereas the largest opening observed in managed stands was only 200 m2. Wind-disturbed stands had, on average, double the available solar radiation at the forest floor compared to managed stands. Solar radiation levels were also more heterogeneous in wind-disturbed stands, with six times more variability at small scales (0.1225 ha) and 15 times more variability at the whole-stand level. Modification of uneven-aged management regimes to include occasional harvests of variable intensity and spatial pattern may help avoid the decline in species diversity that tends to occur after many decades of conventional uneven-aged management. At the same time, a multi-cohort system with these properties would retain a high degree of average crown cover, promote structural heterogeneity typical of old-growth forests, and maintain dominance by late-successional species.     

Species richness and stand stability in conifer forests of the Sierra Nevada

Theoretical and empirical studies have long suggested that stability and complexity are intimately related, but evidence from long-lived systems at large scales is lacking. Stability can either be driven by complex species interactions, or it can be driven by the presence/absence and abundance of a species best able to perform a specific ecosystem function. We use 64 years of stand productivity measures in forest systems composed of four dominant conifer tree species to contrast the effect of species richness and abundance on three stability measures. To perform this contrast, we measured the annual growth increments of > 900 trees in mixed and pure forest stands to test three hypotheses: increased species richness will (1) decrease stand variance, (2) increase stand resistance to drought events, and (3) increase stand resilience to drought events. In each case, the alternate hypothesis was that species richness had no effect, but that species composition and abundance within a stand drove variance, resistance, and resilience. In pure stands, the four species demonstrated significant differences in productivity, and in their resistance and resilience to drought events. The two pine species were the most drought resistant and resilient, whereas mountain hemlock was the least resistant and resilient, and red fir was intermediate. For community measures we found a moderately significant (P = 0.08) increase in the community coefficient of variation and a significant (P = 0.03) increase in resilience with increased species richness, but no significant relationship between species richness and community resistance, though the variance in community resistance to drought decreased with species richness. Community resistance to drought was significantly (P = 0.001) correlated to the relative abundance of lodgepole pine, the most resistant species. We propose that resistance is driven by competition for a single limiting resource, with negative diversity effects. In contrast resilience measures the capacity of communities to partition resources in the absence of a single limiting resource, demonstrating positive diversity effects.     

Do evergreen and deciduous trees have different effects on net N mineralization in soil?

Evergreen and deciduous plants are widely expected to have different impacts on soil nitrogen (N) availability because of differences in leaf litter chemistry and ensuing effects on net N mineralization (N(min)). We evaluated this hypothesis by compiling published data on net N(min) rates beneath co-occurring stands of evergreen and deciduous trees. The compiled data included 35 sets of co-occurring stands in temperate and boreal forests. Evergreen and deciduous stands did not have consistently divergent effects on net N(min) rates; net N(min) beneath deciduous trees was higher when comparing natural stands (19 contrasts), but equivalent to evergreens in plantations (16 contrasts). We also compared net N(min) rates beneath pairs of co-occurring genera. Most pairs of genera did not differ consistently, i.e., tree species from one genus had higher net N(min) at some sites and lower net N(min) at other sites. Moreover, several common deciduous genera (Acer, Betula, Populus) and deciduous Quercus spp. did not typically have higher net N(min) rates than common evergreen genera (Pinus, Picea). There are several reasons why tree effects on net N(min) are poorly predicted by leaf habit and phylogeny. For example, the amount of N mineralized from decomposing leaves might be less than the amount of N mineralized from organic matter pools that are less affected by leaf litter traits, such as dead roots and soil organic matter. Also, effects of plant traits and plant groups on net N(min) probably depend on site-specific factors such as stand age and soil type.     

Large ontogenetic declines in intra-crown leaf area index in two temperate deciduous tree species

The widespread occurrence of age-related changes in leaf morphology and allocation suggests that the leaf area index of individual trees (intra-crown LAI) may decline late in ontogeny. We used direct, within-canopy measurements to quantify the LAI of canopy trees with exposed crowns of two temperate deciduous species. Intra-crown LAI declined from approximately 7 to 4 in Acer saccharum, and from approximately 9.5 to 6.5 in Betula alleghaniensis, as tree size increased (from 15 to 72 cm diameter at breast height [dbh]). For A. saccharum, age (which varied from 30 to 160 years) was a significantly better predictor of LAI decline than dbh. We also modeled the effect of ontogenetic declines in LAI on understory light availability and found that light transmission increases significantly as canopy trees grow and mature. Our results thus suggest that gradual declines in LAI with tree age may play an important and overlooked role in contributing to the heterogeneity of sub-canopy light regimes in mature forests.     

Tree growth inference and prediction from diameter censuses and ring widths.

Estimation of tree growth is based on sparse observations of tree diameter, ring widths, or increments read from a dendrometer. From annual measurements on a few trees (e.g., increment cores) or sporadic measurements from many trees (e.g., diameter censuses on mapped plots), relationships with resources, tree size, and climate are extrapolated to whole stands. There has been no way to formally integrate different types of data and problems of estimation that result from (1) multiple sources of observation error, which frequently result in impossible estimates of negative growth, (2) the fact that data are typically sparse (a few trees or a few years), whereas inference is needed broadly (many trees over many years), (3) the fact that some unknown fraction of the variance is shared across the population, and (4) the fact that growth rates of trees within competing stands are not independent. We develop a hierarchical Bayes state space model for tree growth that addresses all of these challenges, allowing for formal inference that is consistent with the available data and the assumption that growth is nonnegative. Prediction follows directly, incorporating the full uncertainty from inference with scenarios for "filling the gaps" for past growth rates and for future conditions affecting growth. An example involving multiple species and multiple stands with tree-ring data and up to 14 years of tree census data illustrates how different levels of information at the tree and stand level contribute to inference and prediction.     

Importance of conserving large and old trees to continuity of tree-related microhabitats

Protecting structural features, such as tree-related microhabitats (TreMs), is a cost-effective tool crucial for biodiversity conservation applicable to large forested landscapes. Although the development of TreMs is influenced by tree diameter, species, and vitality, the relationships between tree age and TreM profile remain poorly understood. Using a tree-ring-based approach and a large data set of 8038 trees, we modeled the effects of tree age, diameter, and site characteristics on TreM richness and occurrence across some of the most intact primary temperate forests in Europe, including mixed beech and spruce forests. We observed an overall increase in TreM richness on old and large trees in both forest types. The occurrence of specific TreM groups was variably related to tree age and diameter, but some TreM groups (e.g., epiphytes) had a stronger positive relationship with tree species and elevation. Although many TreM groups were positively associated with tree age and diameter, only two TreM groups in spruce stands reacted exclusively to tree age (insect galleries and exposed sapwood) without responding to diameter. Thus, the retention of trees for conservation purposes based on tree diameter appears to be a generally feasible approach with a rather low risk of underrepresentation of TreMs. Because greater tree age and diameter positively affected TreM development, placing a greater emphasis on conserving large trees and allowing them to reach older ages, for example, through the establishment of conservation reserves, would better maintain the continuity of TreM resource and associated biodiversity. However, this approach may be difficult due to the widespread intensification of forest management and global climate change.     

Modelling succession of Eastern Canadian mixedwood forest

The succession of a mixed species stand of the Acadian Forest was simulated by adopting an approach taken by Botkin et al. (1972) (JABOWA) and later modified by Shugart and West (1977) (FORET). The model simulates the dynamic aspects of successional behaviour of the stand by projecting the current stage of vegetation composition (size of each individual of each species present on a 1/12 ha plot) with an element of randomness. The growth of each tree is modeled as a function of its size (represented by its diameter at breast height (dbh), height and leaf area index), the climate, the tree's tolerance to shade and the competition for available resources and space. Stand aboveground biomass (metric ton/ha), leaf area index, number of trees/ha and species composition of the stand were simulated under seven different conditions: (a) normal treatment, (b) increasing biomass maximum 30%, (c) decreasing biomass maximum 30%, (d) increasing degree-days 30%, (e) decreasing degree-days 30%, (f) increasing light extinction coefficient 30% and (g) decreasing light extinction coefficient 30%. Under the first set of conditions (normal treatment), the aboveground biomass reached a maximum value during the first hundred years, decreased during the second, and remained stable for the rest of the 500-year simulation period. The leaf area showed a similar pattern. The total number of trees/ha decreased sharply during the first fifty years and reached a stable value by the end of the first hundred years. Successional patterns and species competitive abilities were interpreted from the accumulated biomass of each species over the simulation period under the different treatments.Species composition of the stand under the normal treatment showed dominance of deciduous species at early stages of succession, with conifer species being increased and becoming dominant at the end of the simulation period. When the climate was raised 30% (warmer) over the average, the deciduous species continued their dominance over the course of the simulation period. Other simulated species dynamics also appeared to follow what is known about their successional behaviour in the field.     

细叶桉造林密度试验初报

在细叶桉造林试验中，设计了7种密度，结果表明，在造林的当年，高密度林分能较早形成林分环境，有利于早期生长，而后来其个体间较早开始进行营养空间的竞争，部分个体生长受到抑制。随着时间的推移，高密度林分不利于冠径的发展。相应地低密度林分呈现出径生长较快的趋势，并且这种趋势随着树木长大而愈来愈明显。单株材积随密度的变化趋势与径的变化规律相一致。林分的蓄积量更大程度地受株数的作用，密度越大，蓄积量越大。建议细叶桉小径材和纸浆材的造林密度为2500株／hm^2。