An analytical model of stand dynamics as a function of tree growth, mortality and recruitment: the shade tolerance-stand structure hypothesis revisited

Light competition and interspecific differences in shade tolerance are considered key determinants of forest stand structure and dynamics. Specifically two main stand diameter distribution types as a function of shade tolerance have been proposed based on empirical observations. All-aged stands of shade tolerant species tend to have steeply descending, monotonic diameter distributions (inverse J-shaped curves). Shade intolerant species in contrast typically exhibit normal (unimodal) tree diameter distributions due to high mortality rates of smaller suppressed trees. In this study we explore the generality of this hypothesis which implies a causal relationship between light competition or shade tolerance and stand structure. For this purpose we formulate a partial differential equation system of stand dynamics as a function of individual tree growth, recruitment and mortality which allows us to explore possible individual-based mechanisms--e.g. light competition-underlying observed patterns of stand structure--e.g. unimodal or inverse J-shaped equilibrium diameter curves. We find that contrary to expectations interspecific differences in growth patterns can result alone in any of the two diameter distributions types observed in the field. In particular, slow growing species can present unimodal equilibrium curves even in the absence of light competition. Moreover, light competition and shade intolerance evaluated both at the tree growth and mortality stages did not have a significant impact on stand structure that tended to converge systematically towards an inverse J-shaped curves for most tree growth scenarios. Realistic transient stand dynamics for even aged stands of shade intolerant species (unimodal curves) were only obtained when recruitment was completely suppressed, providing further evidence on the critical role played by juvenile stages of tree development (e.g. the sampling stage) on final forest structure and composition. The results also point out the relevance of partial differential equations systems as a tool for exploring the individual-level mechanisms underpinning forest structure, particularly in relation to more complex forest simulation models that are more difficult to analyze and to interpret from a biological point of view.     

Biotic and abiotic variables show little redundancy in explaining tree species distributions

Abiotic factors such as climate and soil determine the species fundamental niche, which is further constrained by biotic interactions such as interspecific competition. To parameterize this realized niche, species distribution models (SDMs) most often relate species occurrence data to abiotic variables, but few SDM studies include biotic predictors to help explain species distributions. Therefore, most predictions of species distributions under future climates assume implicitly that biotic interactions remain constant or exert only minor influence on large‐scale spatial distributions, which is also largely expected for species with high competitive ability. We examined the extent to which variance explained by SDMs can be attributed to abiotic or biotic predictors and how this depends on species traits. We fit generalized linear models for 11 common tree species in Switzerland using three different sets of predictor variables: biotic, abiotic, and the combination of both sets. We used variance partitioning to estimate the proportion of the variance explained by biotic and abiotic predictors, jointly and independently. Inclusion of biotic predictors improved the SDMs substantially. The joint contribution of biotic and abiotic predictors to explained deviance was relatively small (~9%) compared to the contribution of each predictor set individually (~20% each), indicating that the additional information on the realized niche brought by adding other species as predictors was largely independent of the abiotic (topo‐climatic) predictors. The influence of biotic predictors was relatively high for species preferably growing under low disturbance and low abiotic stress, species with long seed dispersal distances, species with high shade tolerance as juveniles and adults, and species that occur frequently and are dominant across the landscape. The influence of biotic variables on SDM performance indicates that community composition and other local biotic factors or abiotic processes not included in the abiotic predictors strongly influence prediction of species distributions. Improved prediction of species' potential distributions in future climates and communities may assist strategies for sustainable forest management.     

Growth and survivorship of dipterocarp seedlings: differences in shade persistence create a special case of dispersal limitation

A series of growth experiments and observations on natural populations have been carried out on dipterocarp species of contrasting ecology growing in artificial gaps and the forest understorey. These studies have demonstrated that although differences exist between species in photosynthetic and growth responses to the high-light environment, competition for light in canopy gaps is highly asymmetrical and tends to reinforce any pre-existing dominance hierarchy. We propose that differences in seedling persistence in forest canopy shade are highly influenced by species-specific biotic and abiotic interactions. Our experiments suggest that as seedlings, dipterocarp species trade off traits which enhance persistence and growth in shade against those that enhance their ability to exploit gaps. Less competitive species survive for progressively longer periods of time after a gregarious fruiting event. This leads to significant shifts with time in the number of species present in the seedling bank and hence in the importance of interspecific competition in determining which species dominates regrowth in gaps. We propose that this special case of dispersal limitation is more likely to account for coexistence of dipterocarp species than differences in growth responses to gaps of different size, with stochastic and environmental variables interacting to determine species distribution and abundance.     

Dispersal,establishment and survival of Ceriops tagal propagules in a north Australian mangrove forest

Studies of the ecology of mangroves show that a wide variety of factors, including salinity, desiccation, disturbance, competition and predation, may affect the distribution and abundance of species. Field studies were done to examine the relative importance of several of these factors in the establishment and early survival of Ceriops tagal, a species common in mid-to high-shore regions of mangrove forests in northern Australia. The fate of marked and tethered propagules was followed to estimate the range of dispersal and the intensity of predation. Propagules were artificially planted under different thicknesses of shade cloth (none, 30%, 80%) and in different habitats (clearing, forest, clearing-forest fringe) to examine the effects of light and soil conditions on survival and growth. Results suggested that dispersal was very limited: only 9% of marked propagules were ever found more than 3 m from the parent tree. Losses to predators were great, with 83% of tethered propagules being damaged or consumed within 3 months. On average, 56% of planted propagules survived for at least 6 weeks and 76% of these initiated growth. Survival in clearings was lower than in other habitats, with 29% fewer surviving six weeks and 48% fewer surviving 15 months. The growth of seedlings was correlated with soil temperature, but the effects of treatments were complex. Overall, results indicated that poor dispersal and establishment were the main factors likely to limit the colonisation and population growth of this species. Received: 12 September 1995/Accepted: 14 June 1996     

Multiple successional pathways of boreal forest stands in central Canada

Predicting forest composition change through time is a key challenge in forest management. While multiple successional pathways are theorized for boreal forests, empirical evidence is lacking, largely because succession has been inferred from chronosequence and dendrochronological methods. We tested the hypotheses that stands of compositionally similar overstory may follow multiple successional pathways depending on time since last stand‐replacing fire (TSF), edaphic conditions, and presence of intermediate disturbances. We used repeated measurements from combining sequential aerial photography and ground surveys for 361 boreal stands in central Canada. Stands were measured in 8–15 yr intervals over a ~ 60 yr period, covering a wide range of initial stand conditions. Multinomial logistic regression was used to analyze stand type transitions. With increasing TSF, stands dominated by shade‐intolerant Pinus banksiana, Populus sp., and Betula papyrifera demonstrated multiple pathways to stands dominated by shade‐tolerant Picea sp., Abies balsamea, and Thuja occidentalis. Their pathways seemed largely explained by neighborhood effects. Succession of stands dominated by shade‐tolerant species, with an exception of stands dominated by Picea sp., was not related to TSF, but rather dependent on edaphic conditions and presence of intermediate disturbances. Varying edaphic conditions caused divergent pathways with resource limited sites being dominated by nutrient‐poor tolerant species, and richer sites permitting invasion of early successional species and promoting species mixtures during succession. Intermediate disturbances promoted deciduous persistence and species diversity in A. balsamea and mixed‐conifer stands, but no evidence was detected to support “disturbance accelerated succession”. Our results demonstrate that in the prolonged absence of stand‐replacing disturbance boreal forest stands undergo multiple succession pathways. These pathways are regulated by neighborhood effects, resource availability, and presence of intermediate disturbance, but the relative importance of these regulators depends on initial stand type. The observed divergence of successional pathways supports the resource‐ratio hypothesis of plant succession.     

Linking responses of native and invasive plants to hurricane disturbances: implications for coastal plant community structure

Hurricane disturbances produce significant changes in forest microclimates, creating opportunities for seedling regeneration of native and invasive plant species alike. However, there is limited information on how changes in microclimates and pre-existing forest conditions affect native and invasive plants responses to hurricane disturbances. In this manipulative study, we examined the responses of three common shrub/small stature tree species, two of which are native to the coastal region of the southeastern USA (Baccharis halimifolia and Morella cerifera) and one that is invasive (Triadica sebifera), to two key components of hurricane disturbance (canopy damage and saline storm surge). In a greenhouse, we grew seedlings of these species under a range of shade levels that mimicked pre-and post-hurricane canopy conditions for wet pine forest and mixed hardwood forest, two forest communities common in coastal areas of the southeastern USA. Seedlings were subjected to saline storm surges equivalent to full strength sea water for 3 days. Seedling responses (mortality and growth) to the treatments were monitored for 16 months. All species benefitted from higher canopy openness. Storm surge effects were short-lived and seedlings readily recovered under high light conditions. The storm surge had stronger negative effects on survival and growth of all species when coupled with high shade, suggesting storm surge has greater negative impacts on seedlings where hurricane winds cause minimal or no canopy damage. The invasive T. sebifera was by far more shade tolerant than the natives. Survival of T. sebifera seedlings under highly shaded conditions may provide it a competitive edge over native species during community reassembly following tropical storms. Differential responses of native and invasive species to hurricane disturbances will have profound consequences on community structure across coastal forest stands, and may be regulated by legacies of prior disturbances, community structure, extent of canopy damage, and species’ tolerance to specific microclimates.

     

Structure and dynamics of subalpine forests in the Wang Lang Natural Reserve, Sichuan, China

Population structure (size, age, spatial patterns) and radial growth patterns are used to analyze regeneration patterns of Abies faxoniana, Betula albosinensis, Betula utilis, Larix potaninii, Picea purpurea, and Sabina saltuaria and reconstruct disturbance history in 8 subalpine forest stands in Wang Lang Natural Reserve, Sichuan, China. In old-growth stands tree regeneration occurs in tree-fall gaps whereby A. faxoniana, Betula sp., P. purpurea, and S. saltuaria persist at stand scales by gap-phase regeneration. Clump sizes of young populations are similar to canopy gap sizes but clumps sizes vary among species. Young Betula patches are larger than those of A. faxoniana suggesting that gap-partitioning by size contributes to species coexistence in mixed stands. Picea purpurea and S. saltuaria are longer lived than A. faxoniana which may compensate for lower recruitment and prevent their replacement by A. faxoniana. Tree regeneration and community structure are also influenced by the understory bamboo Fargesia denudata. Seedlings, saplings, and shrub density all decline with an increase in bamboo cover. Species that regenerate in old-growth forest also regenerate after flooding as do species that establish only on bare substrates (i.e. Larix potaninii, Prunus sp.). Structural and compositional patterns in Wang Lang forests are a reflection of disturbance history, canopy species life history attributes such as dispersal ability, shade tolerance, growth rates, and longevity, and competition of trees and shrubs with understory bamboos.     

Tree Size Inequality Reduces Forest Productivity: An Analysis Combining Inventory Data for Ten European Species and a Light Competition Model

Plant structural diversity is usually considered as beneficial for ecosystem functioning. For instance, numerous studies have reported positive species diversity-productivity relationships in plant communities. However, other aspects of structural diversity such as individual size inequality have been far less investigated. In forests, tree size inequality impacts directly tree growth and asymmetric competition, but consequences on forest productivity are still indeterminate. In addition, the effect of tree size inequality on productivity is likely to vary with species shade-tolerance, a key ecological characteristic controlling asymmetric competition and light resource acquisition. Using plot data from the French National Geographic Agency, we studied the response of stand productivity to size inequality for ten forest species differing in shade tolerance. We fitted a basal area stand production model that included abiotic factors, stand density, stand development stage and a tree size inequality index. Then, using a forest dynamics model we explored whether mechanisms of light interception and light use efficiency could explain the tree size inequality effect observed for three of the ten species studied. Size inequality negatively affected basal area increment for seven out of the ten species investigated. However, this effect was not related to the shade tolerance of these species. According to the model simulations, the negative tree size inequality effect could result both from reduced total stand light interception and reduced light use efficiency. Our results demonstrate that negative relationships between size inequality and productivity may be the rule in tree populations. The lack of effect of shade tolerance indicates compensatory mechanisms between effect on light availability and response to light availability. Such a pattern deserves further investigations for mixed forests where complementarity effects between species are involved. When studying the effect of structural diversity on ecosystem productivity, tree size inequality is a major facet that should be taken into account.     

Strategy Space and the Disturbance Spectrum: A Life-History Model for Tree Species Coexistence

The disturbance spectrum consists of disturbance patterns differing in type, size, intensity, and frequency. It is proposed that tree life-history traits are adaptations to particular disturbance regimes. Four independent axes are proposed to define the dominant dimensions of tree strategy space: shade tolerance, tree height, capacity for vegetative reproduction, and seed dispersal distance. A fitness model was developed to elucidate interactions between the proposed life-history traits. The model shows how alternate life-history sets can coexist when disturbance patterns fluctuate in space and time. Variable disturbance regimes were shown, based on data and simulation results, to enhance species coexistence, as predicted. The strategy space model accurately predicts the number of common tree species for the eastern United States, boreal Canada, and southwestern pi?on-juniper woodlands. The model also provides an explanation for latitudinal gradients in tree species richness in North America and Europe. The proposed model predicts a relationship between disturbance characteristics and the species composition of a forest that allows for the coexistence of large numbers of species. The life-history traits of size, growth rate, life span, shade tolerance, age of reproduction, seed dispersal distance, and vegetative reproduction are all incorporated into the model.     

Multiple Environmental Controls on Cockroach Assemblage Structure in a Tropical Rain Forest

Arthropod abundance and diversity are remarkable in tropical forests, but are also spatially patchy. This has been attributed either to resources, predators, abiotic conditions or disturbances, but whether such factors may simultaneously shape arthropod assemblage structure is little known. We used cockroaches to test for multiple environmental controls on assemblage structure in 25 km² of Amazonian forest. We performed nocturnal, direct searches for cockroaches in 30 plots (250 m × 2 m) during two seasons, and gathered data on biotic and abiotic factors from previous studies. Cockroach abundance increased with dry litter mass, a measure of resource amount, while species richness increased with litter phosphorus content, a measure of resource availability. Cockroach abundance and species richness decreased with ant relative abundance. Cockroach species composition changed along the gradient of: (1) soil clay content, which correlates with a broad differentiation between flood‐prone and non‐flooded forest; (2) soil relative moisture, consistent with known interspecific variation in desiccation tolerance; and (3) according to the abundance of ants, a potential predator. Turnover in species composition was correlated with abiotic conditions—sorting species according to physiological requirements and to disturbance‐related life history traits—and to ants' selective pressure. Cockroach abundance, diversity, and composition seem to be controlled by distinct sets of environmental factors, but predators which were represented by ants, emerged as a common factor underlying cockroach distribution. Such patterns of community structure may have been previously overlooked by undue focus on single or a few factors, and may be common to tropical forest arthropods.     

Ontogenetic shifts in plant–plant interactions in a rare cycad within angiosperm communities

Gymnosperms and angiosperms can co-occur within the same habitats but key plant traits are thought to give angiosperms an evolutionary competitive advantage in many ecological settings. We studied ontogenetic changes in competitive and facilitative interactions between a rare gymnosperm (Dioon sonorense, our target species) and different plant and abiotic neighbours (conspecific-cycads, heterospecific-angiosperms, or abiotic-rocks) from 2007 to 2010 in an arid environment of northwestern Mexico. We monitored survival and growth of seedlings, juveniles, and adults of the cycad Dioon sonorense to evaluate how cycad survival and relative height growth rate (RHGR) responded to intra- and interspecific competition, canopy openness, and nearest neighbour. We tested spatial associations among D. sonorense life stages and angiosperm species and measured ontogenetic shifts in cycad shade tolerance. Canopy openness decreased cycad survival while intraspecific competition decreased survival and RHGR during early ontogeny. Seedling survival was higher in association with rocks and heterospecific neighbours where intraspecific competition was lower. Shade tolerance decreased with cycad ontogeny reflecting the spatial association of advanced stages with more open canopies. Interspecific facilitation during early ontogeny of our target species may promote its persistence in spite of increasing interspecific competition in later stages. We provide empirical support to the long-standing assumption that marginal rocky habitats serve as refugia from angiosperm competition for slow-growing gymnosperms such as cycads. The lack of knowledge of plant–plant interactions in rare or endangered species may hinder developing efficient conservation strategies (e.g. managing for sustained canopy cover), especially under the ongoing land use and climatic changes.     

干扰对动物传播森林植物种子有效性影响的研究进展

干扰在森林生态系统中普遍存在,并影响森林的更新和演替.动物传播种子是种子更新的必经阶段,其对森林干扰的响应在一定程度上能够预测未来的森林群落组成和结构变化,对于明确森林演替方向具有重要意义.本文论述了森林干扰对动物传播种子有效性(包括动物传播种子的数量和质量)影响研究的生态学意义,全面揭示了自然干扰(火干扰、林窗干扰等)和人为干扰(生境破碎化、狩猎、采伐等)对动物传播种子数量、传播距离以及传播后幼苗更新影响的研究进展,指出干扰通过影响动物种群动态,进而造成动物传播种子数量发生了改变,动物传播种子的距离对干扰的响应基本表现出轻微负相关;干扰对传播后幼苗更新的影响结果因干扰类型的不同而复杂多变,干扰迹地环境因子的变化也影响着传播后的种子萌发和幼苗更新.干扰对动物传播种子有效性影响研究中存在的问题,主要表现为火干扰迹地恢复过程、增益性的干扰(如抚育、间伐、林窗)等对种子传播有效性影响研究的匮乏,以及忽略了温带森林内的干扰对动物传播种子的影响等.今后,应开展干扰对种子传播有效性的长期研究;对于干扰多发地带的森林,应高度重视增益性干扰影响动物传播植物种子的研究.     

Water level fluctuations and dynamics of amphibious plants at Lake Constance: Long-term study and simulation

Inundations of lakeshores are classical examples of how disturbance can influence community diversity and composition. As the occurrence and intensity of flooding are predicted to change dramatically as a result of climate change, predicting the consequences of such changes has become a major task for community ecology. Here we present abundance data of five species that comprise a species-poor community of high conservation value at lakeshores of Lake Constance over 17 years, during which one of the longest flood periods and the lowest water levels since 1890 occurred. We used simple regression models and increasingly sophisticated Markov chain models plus non-linear parameter estimation to put down abundance changes to direct effects of flooding on population-dynamic parameters and to indirect effects of flooding through modification of interspecific competition. We found a negative effect of flood duration on abundance changes for the non-specialist species Agrostis stolonifera and Phalaris arundinacea, but no effect on Carex acuta. The specialist species, Ranunculus reptans but not Littorella uniflora showed a positive effect of flooding. Data analysis revealed an unambiguous competitive hierarchy with the two graminoid species (C. acuta, P. arundinacea) being superior, and the habitat specialists being most sensitive to interspecific competition. We used estimated parameters to project the community dynamics under different flooding regimes. Long-term projection showed that the original community is threatened by two non-specialist species (C. acuta and P. arundinacea). Even if this forecast was influenced by various model limitations, it may indicate irreversible changes in soil fertility during the phase of high eutrophication between 1950 and 1980. Our study demonstrated that long-term abundance relevés combined with Markov modelling and predictive simulations are an important counterpart to detailed short-term studies. The combination of empirical and theoretical methods elucidates the interaction of biotic and abiotic factors in community change.     

The effect of altitude,patch size and disturbance on species richness and density of lianas in montane forest patches

The species richness and density of lianas (woody vines) in tropical forests is determined by various abiotic and biotic factors. Factors such as altitude, forest patch size and the degree of forest disturbance are known to exert strong influences on liana species richness and density. We investigated how liana species richness and density were concurrently influenced by altitude (1700–2360 m), forest patch size, forest patch location (edge or interior) and disturbance intensity in the tropical montane evergreen forests, of the Nilgiri and Palni hills, Western Ghats, southern India. All woody lianas (≥1 cm dbh) were enumerated in plots of 30 × 30 m in small, medium and large forest patches, which were located along an altitudinal gradient ranging from 1700 to 2360 m. A total of 1980 individual lianas were recorded, belonging to 45 species, 32 genera and 21 families, from a total sampling area of 13.86 ha (across 154 plots). Liana species richness and density decreased significantly with increasing altitude and increased with increasing forest patch size. Within forest patches, the proportion of forest edge or interior habitat influenced liana distribution and succession especially when compared across the patch size categories. Liana species richness and density also varied along the altitudinal gradient when examined using eco-physiological guilds (i.e. shade tolerance, dispersal mode and climbing mechanism). The species richness and density of lianas within these ecological guilds responded negatively to increasing altitude and positively to increasing patch size and additionally displayed differing sensitivities to forest disturbance. Importantly, the degree of forest disturbance significantly altered the relationship between liana species richness and density to increasing altitude and patches size, and as such is likely the primary influence on liana response to montane forest succession. Our findings suggest that managing forest disturbance in the examined montane forests would assist in conserving local liana diversity across the examined altitudinal range.     

The Effects of Seed Dispersal on the Simulation of Long-Term Forest Landscape Change

The study of forest landscape change requires an understanding of the complex interactions of both spatial and temporal factors. Traditionally, forest gap models have been used to simulate change on small and independent plots. While gap models are useful in examining forest ecological dynamics across temporal scales, large, spatial processes, such as seed dispersal, cannot be realistically simulated across large landscapes. To simulate seed dispersal, spatially explicit landscape models that track individual species distribution are needed. We used such a model, LANDIS, to illustrate the implications of seed dispersal for simulating forest landscape change. On an artificial open landscape with a uniform environment, circular-shaped tree species establishment patterns resulted from the simulations, with areas near seed sources more densely covered than areas further from seed sources. Because LANDIS simulates at 10-y time steps, this pattern reflects an integration of various possible dispersal shapes and establishment that are caused by the annual variations in climate and other environmental variables. On real landscapes, these patterns driven only by species dispersal radii are obscured by other factors, such as species competition, disturbance, and landscape structure. To further demonstrate the effects of seed dispersal, we chose a fairly disturbed and fragmented forest landscape (approximately 500,000 ha) in northern Wisconsin. We compared the simulation results of a map with tree species (seed source locations) realistically parameterized (the real scenario) against a randomly parameterized species map (the random scenario). Differences in the initial seed source distribution lead to different simulation results of species abundance with species abundance starting at identical levels under the two scenarios. This is particularly true for the first half of the model run (0–250 y). Under the random scenario, infrequently occurring and shade tolerant species tend to be overestimated, while midabundant and midshade tolerant species tend to be underestimated. The over- and underestimation of species abundance diminish when examining long-term (500 y) landscape dynamics, because stochastic factors, such as fire, tend to make the landscapes under both scenarios converge. However, differences in spatial patterns, and especially species age-cohort distributions, can persist under the two scenarios for several hundred years. Received 24 November 1998; accepted 17 March 1999.     

Characterizing Tropical Tree Species Growth Strategies: Learning from Inter-Individual Variability and Scale Invariance

Understanding how tropical tree species differ in their growth strategies is critical to predict forest dynamics and assess species coexistence. Although tree growth is highly variable in tropical forests, species maximum growth is often considered as a major axis synthesizing species strategies, with fast-growing pioneer and slow-growing shade tolerant species as emblematic representatives. We used a hierarchical linear mixed model and 21-years long tree diameter increment series in a monsoon forest of the Western Ghats, India, to characterize species growth strategies and question whether maximum growth summarizes these strategies. We quantified both species responses to biotic and abiotic factors and individual tree effects unexplained by these factors. Growth responses to competition and tree size appeared highly variable among species which led to reversals in performance ranking along those two gradients. However, species-specific responses largely overlapped due to large unexplained variability resulting mostly from inter-individual growth differences consistent over time. On average one-third of the variability captured by our model was explained by covariates. This emphasizes the high dimensionality of the tree growth process, i.e. the fact that trees differ in many dimensions (genetics, life history) influencing their growth response to environmental gradients, some being unmeasured or unmeasurable. In addition, intraspecific variability increased as a power function of species maximum growth partly as a result of higher absolute responses of fast-growing species to competition and tree size. However, covariates explained on average the same proportion of intraspecific variability for slow- and fast-growing species, which showed the same range of relative responses to competition and tree size. These results reflect a scale invariance of the growth process, underlining that slow- and fast-growing species exhibit the same range of growth strategies.     

大兴安岭主要森林类型林分空间结构优化模拟

以大兴安岭地区4种主要林型[天然落叶松纯林(LF)、天然白桦纯林(BF)、针阔混交林(CBMF)和针叶混交林(CMF)]的1 hm²固定样地调查数据为基础,选择常用林分空间[混交度(M)、大小比数(U)、角尺度(W)、竞争指数(CI)]和非空间结构参数[林分生长活力(DC)、林木稳定性(DH)],并结合熵值-AHP赋权法,构建单木综合抚育采伐指数(T),运用Excel VBA编制模拟抚育采伐过程;系统比较各林型不同抚育采伐强度(分别为10%、20%和30%)下T值,并确定各林型的最优抚育采伐强度.结果表明:初始状态下,各林型平均W均为0.57,属于典型聚集分布;各林分平均U在0.50～0.51,林木整体生长的优势程度处于典型的中庸状态;各林分平均M整体较低,其中,混交林明显大于纯林;林分内平均CI大于2.0,林木生长竞争压力较大;LF整体稳定性、活力均显著高于其他林分;整体来说,BF的经营迫切性程度显著高于其他林分.从同一林型内相邻抚育采伐强度间的T值增长率来看,LF的最佳抚育采伐强度为30%,其余3种林型的最佳抚育采伐强度均为10%,T值增长率分别为9.7%、7.9%、6.6%和3.9%;而从不同林型下T值大小和郁闭度来看,BF的最佳抚育采伐强度为20%,其余林型的最优抚育采伐强度均为30%,此时T值分别提高了28.9%、16.4%、17.5%和9.2%.抚育采伐后林分整体结构得到不同程度的改善,其中,林分M显著增大,林木水平空间分布格局趋向随机分布,优势树种优势程度增加,样地内林木竞争压力明显减小,DC略有降低,DH得到提升.     

Determinants of tree survival at local scale in a sub-tropical forest

Tree survival is a critical driver of stand dynamics, influencing forest structure and composition. Many local-scale drivers (tree size, abiotic and biotic factors) have been proposed as being important in explaining patterns of tree survival, but their contributions are still unknown. We examined the relative importance of these local drivers on tree survival using generalized linear mixed models in an old-growth sub-tropical forest in south China at three levels (community, guild, and species). Among the variables tested, tree size was typically the most important driver of tree survival, followed by abiotic and then biotic variables. Tree size has a strongly positive effect on tree survival for small trees (10–30 cm dbh) and shade-tolerant tree species. Of the abiotic factors tested, elevation tended to be more important in affecting tree survival than other topographic variables. Abiotic factors generally influenced survival of species with relatively high abundances, for individuals in smaller size classes and for mid-tolerant species. Among biotic factors, we found that the mortality of tree species was not driven by density- and frequency-dependent effects in this sub-tropical forest, as indicated by the results of both total basal area of neighbors and the proportion of conspecific neighbors in our study. We conclude that the relative importance of variables driving patterns of tree survival varied greatly among tree size classes, species guilds, shade tolerance, density, and abundance classes in this sub-tropical forest. These results also provide critical information for future studies of forest dynamics and offer insight into forest management in this region.     

Aggregated spatial distributions of species in a subtropical karst forest,southwestern China

Aims Spatial distribution patterns of species reflect not only the ecological processes but also the habitat features that are related to species distribution. In karst topography, species distribution patterns provide more specific information about their environments. The objectives of this study are as follows: (i) to analyse and explain the spatial distribution patterns of conspecific trees in an old-growth subtropical karst forest; (ii) to investigate pattern changes at different spatial scales; (iii) to test the spatial pattern similarity (or dissimilarity) between trees at different abundances, diameter at breast height classes, canopy layers and different functional groups (shade tolerance and seed dispersal mode); (iv) to examine whether habitat heterogeneity has an important effect on the species spatial distribution.Methods The spatial distributions of woody species with ≥20 individuals in a 1-ha subtropical karst forest plot at Maolan in southwestern China were quantified using the relative neighbourhood density Ω based on the average density of conspecific species in a circular neighbourhood around each species.Important findings Aggregated distribution is the dominant pattern in the karst forest, but the ratio of aggregated species in total species number decreases with an increase in spatial scale. Less abundant species are more aggregated than most abundant species. Aggregation is weaker in larger diameter classes, which is consistent with the prediction of self-thinning. Seed dispersal mode influences spatial patterns, with species dispersed by animals being less aggregated than those dispersed by wind and gravity. Other species functional traits (e.g. shade tolerance) also influence the species spatial distributions. Moreover, differences among species habitat associations, e.g. with rocky outcrops, play a significant role in species spatial distributions. These results indicate that habitat heterogeneity, seed dispersal limitation and self-thinning primarily contribute to the species spatial distributions in this subtropical karst forest.     

中亚热带濒危植物毛红椿和南方红豆杉种内与种间竞争差异

濒危植物是全球生物多样性的重要组成部分,为了弄清中亚热带地区不同生长型濒危乔木树种的竞争模式及其机制,研究了江西九岭山国家级自然保护区毛红椿和南方红豆杉的种内与种间竞争差异。结果表明: 阳性落叶树种毛红椿种群的竞争压力主要来自于同种个体,种内竞争指数占总竞争指数的66.4%,耐阴常绿树种南方红豆杉种群主要受种间竞争的影响,种间竞争指数占总竞争指数的68.7%。2种濒危植物受到的种内、种间以及总竞争强度均随着林木径级的增大而逐渐减小,表明竞争压力由幼林向成熟林逐渐降低。在生长过程中,毛红椿以种内竞争引起的自疏作用为主导,南方红豆杉则由种间竞争引起的他疏作用为主导。毛红椿和南方红豆杉种群在更新过程中,小径级个体都要经历强度较大的种内或种间竞争选择才能发育为成树。由于主要竞争压力的来源差异,对于生活在中亚热带同一区域内不同生长型的濒危植物应该采取不同的生物多样性保育措施。