Evaluating tree competition indices as predictors of basal area increment in western Montana forests

Fire hazard reduction treatments are commonly applied to mixed-species coniferous forests in western Montana, USA, to modify fuels structures and alter the competitive environments of individual trees. An improved understanding of how competition can be measured and how it conditions individual tree growth is needed for projecting the development of these forests, with and without treatment. Numerous studies have evaluated how competition affects tree growth and many indices have been developed to quantify the competition an individual tree experiences. These studies suggest that no single competition index or a single class of indices is universally superior; indices perform differently according to forest type and forest conditions. We chose several widely used distance-independent and distance-dependent competition indices, and also derived anisotropic distance-dependent indices from estimates of light interception by tree crowns. We evaluated the effectiveness of these competition measures for predicting basal area increment (BAI) of Pinus ponderosa, Pseudotsuga menziesii, and Larix occidentalis in western Montana. The best distance-dependent competition indices explained a larger proportion of growth variation than the best distance-independent indices (64% vs. 56%). This result indicates that competition is an important growth determinant in these forests and that competition varies locally, with variable tree densities and relatively complex stand structures creating heterogeneous neighborhood conditions. Competition indices derived from light interception models were only weakly correlated with other indices and performed poorly in terms of predicting tree growth. This result accords with previous observations that competition for light is not the primarily growth limitation for trees in the semi-arid conditions of western Montana. More sophisticated light availability models could be used to better assess variability in light interception and its marginal contribution to predictive accuracy of radial tree growth. Diameter and distance-dependent BAI models were developed for growth prediction at the species level and for all species combined.     

The effect of competition on individual tree basal area growth in mature stands of Pinus cooperi Blanco in Durango (Mexico)

In this paper, we evaluated how well-selected distance-dependent and distance-independent competition indices explain individual tree basal area growth of trees, growing in mature and even-aged stands of Pinus cooperi Blanco. A total of 18 competition measures were analyzed of which six do not need tree location (distance-independent) and 12 that utilize tree location (distance-dependent). The competition situation of a stand and/or an individual tree was studied using 11 different competitor selection methods. The mean square error reduction relative to no-competition was used to judge the performance of each competition index. It was found that the best distance-independent competition indices performed as well as the best distance-dependent competition indices. It was concluded that the BALMOD-index would be a good competition index to be incorporated into further individual tree basal area growth models for the study area.

Sugar maple (Acer saccharum Marsh.) growth is influenced by close conspecifics and skid trail proximity following selection harvest

In this study, we quantified the effects of local neighbourhood competition, light availability, and proximity to skid trails on the growth of sugar maple (Acer saccharum Marsh.) trees following selection harvest. We hypothesized that growth would increase with decreasing competition and increasing light availability, but that proximity to skid trails would negatively affect growth. A total of 300 sugar maples were sampled ∼10 years after selection harvesting in 18 stands in Témiscamingue (Québec, Canada). Detailed tree and skid trail maps were obtained in one 0.4 ha plot per stand. Square-root transformed radial growth data were fitted to a linear mixed model that included tree diameter, crown position, a neighbourhood competition index, light availability (estimated using the SORTIE light model), and distance to the nearest skid trail as explanatory variables. We considered various distance-dependent or -independent indices based on neighbourhood radii ranging from 6 to 12 m. The competition index that provided the best fit to the data was a distance-dependent index computed in a 6 m search radius, but a distance-independent version of the competition index provided an almost equivalent fit to data. Models corresponding to all combinations of main effects were fit to data using maximum likelihood, and weighted averages of parameter estimates were obtained using multimodel inference. All predictors had an influence on growth, with the exception of light. Radial growth decreased with increasing tree diameter, level of competition and proximity to skid trails, and varied among crown positions with trees in suppressed and intermediate positions having lower growth rates than codominants and dominants. Our results indicate that in selection managed stands, the radial growth of sugar maple trees depends on competition from close (≤6 m) conspecific neighbours, and is still affected by proximity to skid trails ∼10 years after harvesting. Such results underscore the importance of minimizing the extent of skid trail networks by careful pre-harvest planning of trail layout. We also conclude that the impact of heterogeneity among individual-tree neighbourhoods, such as those resulting from alternative spatial patterns of harvest, can usefully be integrated into models of post-harvest tree growth.     

Wind loading of trees: influence of tree size and competition

Wind damage to forests is an important ecological disturbance factor. At the same time, it can have serious economic consequences due to a reduction in timber production. Current models for predicting the risk of wind damage are useful, but generally only focus on the “mean” tree within uniform stands. This paper presents measurements made of wind loading on trees of different sizes within four forest stands of different structure and management history, but all well-acclimated to current wind conditions. Each tree demonstrated a linear relationship between the maximum hourly turning moment and the square of the average hourly wind speed at the canopy top; we defined this ratio (the gradient of the line M _max vs. u ²) as the turning moment coefficient (T _C). T _C was correlated with tree size, in a relationship that differed little between the four forest sites despite the differences between the stands. The relationship between T _C and individual tree competition within each stand was investigated, using both distance-independent and distance-dependent competition indices. All sites showed decreasing T _C with increasing competition. However, the relationships differed between sites and would also be expected to change through time for a single site. The distance-dependent indices offered no improvement over the simpler, non-spatial indices that required only a diameter distribution. We suggest how, subject to further work, the results presented could be applied to calculate the risk of wind damage to trees of different sizes within a forest stand, and how the risk of wind damage to individual trees might change in response to thinning.     

Competition and tree crowns: A neighborhood analysis of three boreal tree species

Competition for canopy space is a fundamental structuring feature of forest ecosystems and remains an enduring focus of research attention. We used a spatial neighborhood approach to quantify the influence of local competition on the size of individual tree crowns in north-central British Columbia, where forests are dominated by subalpine fir (Abies lasiocarpa), lodgepole pine (Pinus contorta) and interior spruce (Picea glauca × engelmanii). Using maximum likelihood methods, we quantified crown radius and length as functions of tree size and competition, estimated by the species identity and spatial arrangement of neighboring trees. Tree crown size depended on tree bole size in all species. Given low levels of competition, pine displayed the widest, shortest tree crowns compared to the relatively long and narrow crowns found in spruce and fir. Sensitivity to crowding by neighbors declined with increasing tree height in all but the pine crown radius model. Five of the six selected best models included separate competition coefficients for each neighboring tree species, evidence that species generally differ in their competitive effects on neighboring tree crowns. The selected crown radius model for lodgepole pine, a shade-intolerant species, treated all neighbors as equivalent competitors. In all species, competition from neighbors exerted an important influence on crown size. Per-capita effects of competition across different sizes and species of neighbors and target trees varied, but subalpine fir generally displayed the strongest competitive effects on neighbors. Results from this study provide evidence that species differ both in their response to competition and in their competitive influence on neighbors, factors that may contribute to maintaining coexistence.     

Using a retrospective dynamic competition index to reconstruct forest succession

Understanding forest dynamics and stand structures is crucial for predicting forest succession. However, many forests have been altered due to century-long land-use practices, which complicates the reconstruction of past and current successional trajectories. For a better understanding of successional processes, we suggest studying the intra- and interspecific competition among single trees across time. We introduce a tree-ring based competition index to reconstruct the competitive dynamics of individual trees over time. This new retrospective dynamic competition index combines a temporal and a spatial component by calculating the yearly ratio between the basal area increments (bai) of the neighbouring trees and the subject tree. The new index is applied to mixed Scots pine (Pinus sylvestris L.) and pubescent oak (Quercus pubescens Willd.) stands in the inner-Alpine dry-valley Valais, for which a change in species composition is hypothesised. The aim is to analyse current stand structures in terms of recent changes in the competitive interactions at the single tree level and to relate these competitive dynamics to land-use change and increasing drought due to climate change. On five plots, the positions of 456 trees were recorded and increment cores were taken to derive bai data. The individual dynamic competition index curves were aggregated in clusters, which define typical patterns of competitive dynamics in both tree species. A large percentage of the trees (87% in oak, 70% in pine) were clustered into a group of trees with constant competition at a relatively low level. However, a smaller group of pines (20%) had recently faced increasing competition. In addition, stand structure analyses indicated a change towards a higher proportion of oak. This change in the competitive ability between oak and pine was found to be related to drought, in that oak had a competitive advantage in dry years. Furthermore, the high proportion of dead branches in pines with decreasing competitive abilities indicated increasing competition for light as a consequence of natural development towards a later successional stage that favours the more shade-tolerant oak. The new retrospective dynamic competition index proved to be promising in studying forest succession. The tree-ring based method allows us to identify changes in the competitive ability of single trees with a high temporal resolution and without repeated assessments.     

金苍林场蒙古栎天然中龄林竞争关系研究

以吉林省汪清林业局金仓林场2块面积为0.25 hm2的蒙古栎天然中龄林固定样地为对象,采用8个竞争指标,研究蒙古栎天然林种内、种间竞争状态。研究结果表明,该林分竞争激烈,蒙古栎与其他各树种存在种间竞争,但其主要竞争压力来自种内,种外竞争压力主要来自黑桦与白桦。其它树种的竞争主要为种间竞争。蒙古栎竞争能力处于中等,白桦与黑桦最强。各竞争指标结果存在差异性,但均能反映蒙古栎天然林竞争状态。蒙古栎竞争强度与胸径存在幂函数关系,竞争强度随个体大小的增大而减小。     

Five-year radial growth of red oaks in mixed bottomland hardwood stands

We studied the relationships among 5-year radial (diameter and basal area) growth of red oak (genus Quercus, subgenus Erythrobalanus) crop trees and predictor variables representing individual tree vigor, distance-dependant competition measures, and distance-independent competition measures. The red oaks we examined are representative of the commercially and ecologically important oak species of the bottomland hardwood forests of the southeastern US. The crown class score, a quantitative measure of crown class and tree vigor, performed best in accounting for the variability in tree diameter growth. Plot-level variables failed to account for a significant proportion of the variability in tree radial growth. The basal area of the first-order neighbors that were taller than the crop trees and located within 2.4 times the mean overstory crown radius had the highest negative correlation with crop tree 5-year radial growth. Red oaks were a major part of these competitors and likely exerted the greatest competitive pressure. However, crop tree radial growth was positively associated with the basal area of the red oaks which were indirect (second order) neighbors and which were taller than the crop trees. It is possible that indirect neighbors do not compete with the crop trees, but they likely compete with the direct competitors of the crop trees, thus having an indirect positive influence on crop tree growth. Such reasoning is consistent with previously observed spatial dependence up to four times the mean overstory crown radius. The findings may have implications for thinning hardwoods stands and crop tree management in that foresters need to take into account (1) oak intra-genus competition, (2) the negative competitive effect of direct neighbors, and (3) the potentially positive effect of the indirect neighbors, the competitors’ competitors.     

A model bridging distance-dependent and distance-independent tree models to simulate the growth of mixed forests

? It is widely believed that distance-independent tree models fail to take into account the complexity of mixed stands due to the fact that spatial structure often has a greater impact on growth and dynamics in mixed stands than in pure stands. On the other hand, distance-dependent tree models are difficult to use because they require a map of the stand, which is not only very costly but also impracticable in a routine management context.

? This paper reports the development of a model bridging distance-dependent and distanceindependent tree models, and that is designed to simulate the growth of a mixed forest. The model used distributions of the number of neighbours to reconstruct tree neighbourhoods and compute the competition indices needed as inputs to the growth model.

? Data were collected from a mixed forest of sessile oak and Scots pine in central France. The study showed that local competition indices explained a significant proportion of growth variability and that intraspecific competition was greater than interspecific competition. The model based on neighbourhood distributions gave consistent predictions compared to a distance-dependent model.

? This type of model could be used instead of distance-dependent models in management contexts.

     

The contribution of competition to tree mortality in old-growth coniferous forests

Competition is a well-documented contributor to tree mortality in temperate forests, with numerous studies documenting a relationship between tree death and the competitive environment. Models frequently rely on competition as the only non-random mechanism affecting tree mortality. However, for mature forests, competition may cease to be the primary driver of mortality.We use a large, long-term dataset to study the importance of competition in determining tree mortality in old-growth forests on the western slope of the Sierra Nevada of California, U.S.A. We make use of the comparative spatial configuration of dead and live trees, changes in tree spatial pattern through time, and field assessments of contributors to an individual tree's death to quantify competitive effects.Competition was apparently a significant contributor to tree mortality in these forests. Trees that died tended to be in more competitive environments than trees that survived, and suppression frequently appeared as a factor contributing to mortality. On the other hand, based on spatial pattern analyses, only three of 14 plots demonstrated compelling evidence that competition was dominating mortality. Most of the rest of the plots fell within the expectation for random mortality, and three fit neither the random nor the competition model. These results suggest that while competition is often playing a significant role in tree mortality processes in these forests it only infrequently governs those processes. In addition, the field assessments indicated a substantial presence of biotic mortality agents in trees that died.While competition is almost certainly important, demographics in these forests cannot accurately be characterized without a better grasp of other mortality processes. In particular, we likely need a better understanding of biotic agents and their interactions with one another and with competition.     

Do position and species identity of neighbours matter in 8–15-year-old post harvest mesic stands in the boreal mixedwood?

Neighbourhood competition indices (NCI), where position and species identity of neighbours are known, have been used to investigate growth and competitive interactions among adult trees. In this study, we used NCI in 8–15-year-old stands following clear-cutting in a boreal mixedwood forest of eastern Canada to improve our understanding of early successional forest dynamics. Trees of increasing diameter from the center (≥1 cm) to the edge (≥5 cm) were mapped in twenty-five circular 450 m² plots. Target trees (DBH ≥ 1 cm) were sampled in plot center to determine their annual radial stem growth. For each species, we compared a set of growth models using either a spatially explicit NCI or a non-spatial competition index. Both types of indices estimated a species-specific competition coefficient for each pair of competitor–target species. NCI were selected as the best competition model for all target species although differences in variance explained relative to the non-spatial index were small. This likely indicates that competition occurs at the local level but that the high density and the relative uniformity of these young stands creates similar neighbourhoods for most trees in a given stand. The effective neighbourhood radius for competitors varied among species and was smaller for shade tolerant species. Intraspecific neighbours were the strongest competitors for most species. Aspen (Populus tremuloides) was a weak competitor for all species as opposed to balsam fir (Abies balsamea) which was a strong competitor in all cases. These results are in contradiction with some widely used forest policies in North America (e.g. free-to-grow standards) that consider broadleaf species, such as aspen, as the strongest competitors. For these early successional forests, the decision regarding the use of spatial or non-spatial competition indices should rest on the intended use. For even-age management, spatial indices might not justify their use in high-density stands but they are needed for the simulation of novel harvest techniques creating complex stand structure.     

Spatiotemporally interactive growth dynamics in selected South African forests: Edaphoclimatic environment, crowding and climate effects

Stand-level tree diameter growth patterns were explored for evergreen moist forests in the southern Cape, South Africa. Results of standard multiple regression analyses, involving 934 permanent sample plots with data spanning a 10-year interval, revealed that stand-level increment of canopy species in the canopy layer (>30 cm dbh) was significantly determined by inherent species-specific growth capacities (species composition of the stand), water availability, forest matrix crowding and tree condition impairment (age-related manifestations of reduced vitality indicated by signs of crown die-back, damage and stem rot). In contrast, stand-level increment of trees of canopy species in the subcanopy layer (10-20 cm dbh) was prominently shaped by light availability, as mainly determined by the degree of canopy-level disturbance (mortality rate of trees >30 cm dbh), crowding (canopy-level overhead and forest matrix crowding) and proximity to conspecific adults (within 6-8 m). In addition to species-inherent and resource factors, considerable variation in stand-level growth resulted from site-climate interactions. For 507 of the permanent sample plots, increment data was available for two consecutive 10-year intervals; permitting the analysis of spatiotemporal interactions of growth patterns (repeated measures ANOVA). In the Knysna forests higher canopy-level increment rates were associated with the moister southerly facing slope sites in comparison with the drier northerly facing and ridge sites during the first increment period. During the second increment period, increment rates on the drier, but better illuminated sites had increased disproportionately. In contrast, in the Tsitsikamma forests, higher increment rates during the second increment period were encountered on moister flat bottomland sites (with extended periods of subsoil wetness) than on the comparatively drier southerly facing slope sites (increment period × site-based water availability × forests interaction). In both forests relatively higher growth performance of subcanopy-level trees during the second increment period was associated with stands experiencing conditions of enhanced light availability. Atmospheric temperatures were higher during the second increment period (mean periodic T_max: + 0.64 °C). The detected spatiotemporal interactions were interpreted as site × climate interactions where site-related conditions of favourable light or water availability resulted in enhanced temperature-linked growth responses during the second increment period. A metabolic performance trade-off model provided a framework for the interpretation of these complex site-climate interactions by placing the patterns of forest growth into an ecophysiological explanatory context.     

濒危植物泰山柳种内与种间的竞争关系

以整合的基径代替胸径,采用Hegyi单木竞争指数模型对灌木泰山柳进行种内与种间的竞争关系分析。结果表明:泰山柳的种内、种间竞争强度分别占总竞争强度(86.11)的35.01%、64.99%,说明泰山柳的竞争压力主要来自种间。随着泰山柳基径的增加,种内、种间竞争强度均逐渐降低。对泰山柳产生竞争影响的树木共9种,其竞争强度为泰山柳>花楸树>紫椴>油松>连翘>湖北海棠>天目琼花>金花忍冬>巧玲花。整个林分和伴生树种竞争指数都与对象木基径近似地服从幂函数关系(I C=AB-K),根据函数模型可以看出,当泰山柳基径达到10 cm后,竞争强度趋于平稳,而基径小于10 cm的泰山柳竞争强度激烈,可以清除其周围部分竞争木,给予足够的生存空间,从而达到保护的目的。     

Structural diversity of English yew (<Emphasis Type="Italic">Taxus baccata</Emphasis> L.) populations

In Europe, the English yew species (Taxus baccata L.) is endangered. Intensive human land-use, including forest management, has caused a decrease of the yew populations all over Europe. In Austria, gene conservation forests are used for the in situ conservation of populations of this rare tree species by silvicultural treatments. In order to improve the conservation management in these gene conservation forests, this study addresses the relation between competition and viability of yew populations through the use of structural diversity indices. The structural indices, which include mingling, tree–tree distance, diameter, and tree height differentiation, were determined for a structural group of four trees as well as the neighbouring trees of the male and female yews at the monitoring plots on a regular grid in three gene conservation forests. Although the three study sites provided quite different environmental conditions for English yew, the vitality of each individual yew was influenced by the inter-specific competition of the neighbouring tree species at all sites. Low vitality was associated with a small mean distance to neighbours and large tree height differentiation. In conclusion, we suggest that a combination of different structural indicators is needed for an integrative assessment of conservation status in the gene conservation forests. This would help improve the evaluation of the impact management has on yew population viability.     

Belowground competition in a broad-leaved temperate mixed forest: pattern analysis and experiments in a four-species stand

We investigated fine root biomass and distribution patterns in a species-rich temperate Carpinus–Quercus–Fagus–Tilia forest and searched for experimental evidence of symmetry or asymmetry in belowground competition. We conducted extensive root coring and applied the recently introduced in situ-root growth chamber technique for quantifying fine root growth under experimentally altered intra- and interspecific root neighbourhoods in the intact stand. In 75% of all soil cores, fine roots of more than two tree species were present indicating a broad overlap of the root systems of neighbouring trees. Quercus trees had more than ten times less fine root biomass in relation to aboveground biomass or productivity (stem growth) and a much higher leaf area index/root area index ratio than Carpinus, Fagus and Tilia trees. The root growth chamber experiments indicated a high belowground competitive ability of Fagus in interspecific interactions, but a low one of Quercus. We conclude that (1) interspecific root competition is ubiquitous in this mixed stand, (2) root competition between trees can be clearly asymmetric, and (3) tree species may be ranked according to their belowground competitive ability. Fagus was found to be the most successful species in belowground competition which matches with its superiority in aboveground competition in this forest community.     

Effects of residual trees on tree height of 18- and 19-year-old Cryptomeria japonica planted in group selection openings

This study examined the effects of residual trees on the height of 18- and 19-year-old Cryptomeria japonica planted in group selection openings of about 0.1 ha at Yufuin in Oita Prefecture in southern Japan. We first developed a general model expressing variation in the height for all of the planted trees within the openings from indices of both between-cohort competition (the effects of residual trees) and within-cohort competition (among the even-aged planted trees). The between-cohort competition index was calculated both with and without taking into account the direction that residual trees were positioned from subject trees. The two models suggested that residual trees located near the northern edge of the opening were not effective competitors for light for the planted trees within the opening. In addition, our results indicated that both between-cohort competition and within-cohort competition are important for explaining tree height in closed stands within group selection openings. Using the general model, we then estimated the potential loss of height growth for locally dominant trees within the openings that could be attributed only to the influence of residual trees. These results indicated that in addition to the effect of opening size, both the degree of crowding and the direction of residual trees from the planted trees are important factors affecting the height of individual trees planted in group selection openings.     

川西南山地云南松过熟林上层枯立木和活立木竞争强度研究

邻体干扰是种群自疏的原因。但在云南松过熟林中,邻体干扰是否影响到枯立木的死亡呢?本文选用Weiner指数、Hegyi指数和张跃西指数计算竞争强度,研究了云南松过熟林径级相当的枯立木和活立木的邻体竞争强度。将8 m范围内邻体分为三类:云南松、壳斗科类和其它类,分别占42.83%、20.65%、36.52%(枯立木)和44.29%、22.49%、33.22%(活立木)。分别受枯立木和活立木周围邻体的分布格局差异的影响,邻体对云南松枯立木的种内竞争和种间竞争相当,对云南松活立木的竞争以种内竞争占优势。邻体对云南松枯立木的竞争强度小于活立木的,但不存在统计学上的显著差异。由于枯立木死亡后大小不变,三个指数所得枯立木竞争强度均大于枯立木死亡时的竞争强度。因此,活立木的竞争强度比目前枯立木死亡时受到的竞争强度还要大,这表明枯立木死亡受邻体竞争影响的可能性较低。由于Weiner指数、Hegyi指数和张跃西指数的构建参数不同,同样数据所得竞争强度各不相同,随距离的变化趋势也各异,但在计算云南松枯立木和活立木竞争强度时获得了一致结果。     

近天然落叶松-云冷杉林树种竞争关系的变化

以4块落叶松-云冷杉间伐实验样地为对象,采用6个与距离无关的竞争指标,研究林木竞争状态在20年间(1986～2006年)的变化及间伐的影响。结果表明,落叶松为主要树种,竞争能力最强,其竞争压力主要来自种内,红松和其他阔叶树的竞争能力有逐渐增强的趋势,因此,随着林分生长,落叶松的竞争主体地位受到了来自其他针叶树云冷杉、红松及阔叶树种的威胁。该林分竞争关系近期相对稳定,但有从落叶松-云冷杉林向阔叶树及红松林发展的趋势。间伐影响种间竞争关系和林分平均竞争水平,影响程度与间伐对象的选择与间伐强度有关。林分竞争水平随间伐强度的增加而降低,但会逐渐恢复。     

Structural diversity of forest communities on Baihuashan Mountain, Beijing

The structural diversity of forests on Baihuashan Mountain, Beijing, was surveyed by a plotless method combined with branch and leaf coverage estimation in the different layers. New structural indices were constructed, calculated and compared among different communities. On the basis of previous work, structural diversity of forest communities at the stand level was described by a vertical complexity index and a horizontal heterogeneity index. From a correlational analysis among the new indices and other commonly used biodiversity indices, we concluded that the new indices are closely related to a tree height inequality index and the Shannon-Wiener index of the tree and shrub layer, which indicated that the new indices were good at indicating structural diversity in the different forests on Baihuashan Mountain. The results show that, in natural forests, structural diversity of pioneer communities is much lower than in late successive communities. In plantations, structural diversity is determined by the stage of development and tree species. Tending would increase horizontal heterogeneity and decrease vertical complexity. __________ Translated from Journal of Beijing Forestry University, 2007, 29(1): 7–13 [译自: 北京林业大学学报]