Primary lowland dipterocarp forest at Danum Valley, Sabah, Malaysia. Species composition and patterns in the understorey

The spatial distributions of species of tree 10 cm gbh were examined in two 4 ha plots and related to the local variation in topography and soil chemistry. The plots were similar in their species composition, particularly in terms of the densities of small trees, and they showed very similar edaphic characteristics. Size class distributions varied little within and between plots. Ordination of 0.25 ha subplots highlighted parallel gradients in the vegetation of both plots when the densities of trees 10 cm gbh were considered. Focusing on understorey trees in the 10-<50 cm gbh class at the 0.04 ha subplot scale showed a similar vegetation gradient in both plots closely associated with change from lower slope to ridge. No relationship with soil chemistry was found. On the ridges a special group of understorey species formed clumps and these species contributed importantly to the ordinations. Borneo has a regional history of occasionally severe droughts. It is suggested here that the observed patterns in the understorey are due to differential responses to low soil water supply, the ridges probably tending to dryness more than the lower slopes. Within the large and diverse family Euphorbiaceae, which dominates the understorey at Danum, there may be ecophysiological groupings of species. The long-term effects of disturbance interacting with local edaphic factors on forest structure and composition are discussed.     

Plurality of tree species responses to drought perturbation in Bornean tropical rain forest

Drought perturbation driven by the El Niño Southern Oscillation (ENSO) is a principal stochastic variable determining the dynamics of lowland rain forest in S.E. Asia. Mortality, recruitment and stem growth rates at Danum in Sabah (Malaysian Borneo) were recorded in two 4-ha plots (trees ≥ 10 cm gbh) for two periods, 1986–1996 and 1996–2001. Mortality and growth were also recorded in a sample of subplots for small trees (10 to <50 cm gbh) in two sub-periods, 1996–1999 and 1999–2001. Dynamics variables were employed to build indices of drought response for each of the 34 most abundant plot-level species (22 at the subplot level), these being interval-weighted percentage changes between periods and sub-periods. A significant yet complex effect of the strong 1997/1998 drought at the forest community level was shown by randomization procedures followed by multiple hypothesis testing. Despite a general resistance of the forest to drought, large and significant differences in short-term responses were apparent for several species. Using a diagrammatic form of stability analysis, different species showed immediate or lagged effects, high or low degrees of resilience or even oscillatory dynamics. In the context of the local topographic gradient, species’ responses define the newly termed perturbation response niche. The largest responses, particularly for recruitment and growth, were among the small trees, many of which are members of understorey taxa. The results bring with them a novel approach to understanding community dynamics: the kaleidoscopic complexity of idiosyncratic responses to stochastic perturbations suggests that plurality, rather than neutrality, of responses may be essential to understanding these tropical forests. The basis to the various responses lies with the mechanisms of tree-soil water relations which are physiologically predictable: the timing and intensity of the next drought, however, is not. To date, environmental stochasticity has been insufficiently incorporated into models of tropical forest dynamics, a step that might considerably improve the reality of theories about these globally important ecosystems.     

Neighbourhood abundance and small-tree survival in a lowland Bornean rainforest

Conspecific effects of neighbours on small-tree survival may have a role in tree population dynamics and community composition of tropical forests. This notion was tested with data from two 4-ha plots in lowland forest at Danum, Sabah (Borneo), for a 21-year interval (censuses at 1986, 1996, 2001, 2007). Species with ≥45 focal trees 10 to <100 cm stem girth per plot in 1986 were selected. Logistic regressions fitted mean focal tree size and mean inverse-distance-weighted basal area abundance of neighbours (within 20 m), for the periods over which each focus tree was alive. Coefficients of variation of neighbourhood basal area abundance, both spatially and temporally, quantified the changing environment of each focus tree. Fits were critically and individually evaluated, with corrections for spatial autocorrelation. Conspecific effects at Danum was generally very weak or non-existent: species’ mortality rates varied also across plots. The main reasons appear to be that (1) species were not dense enough to interact despite frequent although weak spatial aggregation, and their neighbourhoods were highly differing in species composition; and (2) these neighbourhoods were highly variable temporally, meaning that focus trees experienced stochastically fluctuating neighbourhood environments. Only one species, Dimorphocalyx muricatus, showed strong conspecific effects (varying between plots) which can be explained by its distinct ecology. This understorey species is highly aggregated on ridges and is drought-tolerant. That this functionally and habitat-specialized species, has implied intraspecific density-dependent feedback in its dynamics is a remarkable indication of the overall processes maintaining stability of the Danum forest.     

Structure and dynamics of a tropical dry forest in Ghana

Forest under low rainfall (averaging 745 mm yr^-1) on the Shai Hills in S.E. Ghana has redeveloped following cessation of farming in the 1890s. Forest stature is low, with a canopy at about 11 m, principally of three species, Diospyros abyssinica, D. mespiliformis and Millettia thonningii. Drypetes parvifolia and Vepris heterophylla are common understorey trees. Twelve species of woody liane were recorded. Species of thicket vegetation in the area were also present at low density. Most species are evergreen.Tree mortality averaged 2.3% yr^-1 and exceeded recruitment (1.5% yr^-1). Differences between species in mortality and recruitment were pronounced: canopy species showed a small decline in density; understorey species increased markedly and the thicket species declined. Seed production was very variable, but seedling establishment was very poor for all species. Seedling mortality was high (11% yr^-1) especially for small seedlings. These population trends probably represent the latter stages of succession of forest regrowth after farming about 100 years ago.Compared with tropical rain forest, Shai Hills forest has similar relative tree diameter growth (1–3.5% yr^-1), mortality and recruitment rates, and small-litter fall (5.52 t ha^-1 yr^-1).Shai Hills forest differs from rain forest by its short stature, relatively few (evergreen) tree species, poor regeneration from seed, high soil nutrient status and low rainfall. Similar forests have been reported in east Africa and in parts of New Guinea.Abbreviations dbh diameter at breast height (1.3 m) - gbh girth at breast height died May 1984     

The ecoclimatology of Danum, Sabah, in the context of the world's rainforest regions, with particular reference to dry periods and their impact

Climatic records for Danum for 1985-1998, elsewhere in Sabah since 1879, and long monthly rainfall series from other rainforest locations are used to place the climate, and particularly the dry period climatology, of Danum into a world rainforest context. The magnitude frequency and seasonality of dry periods are shown to vary greatly within the world's rainforest zone. The climate of Danum, which is aseasonal but subject, as in 1997-1998, to occasional drought, is intermediate between less drought-prone north-western Borneo and the more drought-prone east coast. Changes through time in drought magnitude frequency in Sabah and rainforest locations elsewhere in South-East Asia and in the Neotropics are compared. The 1997-1998 ENSO-related drought event in Sabah is placed into a historical context. The effects of drought on tree growth and mortality in the tropics are assessed and a model relating intensity and frequency of drought disturbance to forest structure and composition is discussed.     

Overstorey–Understorey Interactions Intensify After Drought-Induced Forest Die-Off: Long-Term Effects for Forest Structure and Composition

Severe drought events increasingly affect forests worldwide, but little is known about their long-term effects at the ecosystem level. Competition between trees and herbs (‘overstorey–understorey competition’) for soil water can reduce tree growth and regeneration success and may thereby alter forest structure and composition. However, these effects are typically ignored in modelling studies. To test the long-term impact of water competition by the herbaceous understorey on forest dynamics, we incorporated this process in the dynamic forest landscape model LandClim. Simulations were performed both with and without understorey under current and future climate scenarios (RCP4.5 and RCP8.5) in a drought-prone inner-Alpine valley in Switzerland. Under current climate, herbaceous understorey reduced tree regeneration biomass by up to 51%, particularly in drought-prone landscape positions (i.e., south-facing, low-elevation slopes), where it also caused a shift in forest composition towards drought-tolerant tree species (for example, Quercus pubescens). For adult trees, the understorey had a minor effect on growth. Under future climate change scenarios, increasing drought frequency and intensity resulted in large-scale mortality of canopy trees, which intensified the competitive interaction between the understorey and tree regeneration. At the driest landscape positions, a complete exclusion of tree regeneration and a shift towards an open, savannah-like vegetation occurred. Overall, our results demonstrate that water competition by the herbaceous understorey can cause long-lasting legacy effects on forest structure and composition across drought-prone landscapes, by affecting the vulnerable recruitment phase. Ignoring herbaceous vegetation may thus lead to a strong underestimation of future drought impacts on forests.     

On the detection of dynamic responses in a drought-perturbed tropical rainforest in Borneo

The dynamics of aseasonal lowland dipterocarp forest in Borneo is influenced by perturbation from droughts. These events might be increasing in frequency and intensity in the future. This paper describes drought-affected dynamics between 1986 and 2001 in Sabah, Malaysia, and considers how it is possible, reliably and accurately, to measure both coarse- and fine-scale responses of the forest. Some fundamental concerns about methodology and data analysis emerge. In two plots forming 8 ha, mortality, recruitment, and stem growth rates of trees ≥10 cm gbh (girth at breast height) were measured in a ‘pre-drought’ period (1986–1996), and in a period (1996–2001) including the 1997–1998 ENSO-drought. For 2.56 ha of subplots, mortality and growth rates of small trees (10–<50 cm gbh) were found also for two sub-periods (1996–1999, 1999–2001). A total of c. 19 K trees were recorded. Mortality rate increased by 25% while both recruitment and relative growth rates increased by 12% for all trees at the coarse scale. For small trees, at the fine scale, mortality increased by 6% and 9% from pre-drought to drought and on to ‘post-drought’ sub-periods. Relative growth rates correspondingly decreased by 38% and increased by 98%. Tree size and topography interacted in a complex manner with between-plot differences. The forest appears to have been sustained by off-setting elevated tree mortality by highly resilient stem growth. This last is seen as the key integrating tree variable which links the external driver (drought causing water stress) and population dynamics recorded as mortality and recruitment. Suitably sound measurements of stem girth, leading to valid growth rates, are needed to understand and model tree dynamic responses to perturbations. The proportion of sound data, however, is in part determined by the drought itself. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

El Niño droughts and their effects on tree species composition and diversity in tropical rain forests

In this study I investigated the effects of the extreme, 1997/98 El Niño related drought on tree mortality and understorey light conditions of logged and unlogged tropical rain forest in the Indonesian province of East Kalimantan (Borneo). My objectives were to test (1) whether drought had a significant effect on tree mortality and understorey light conditions, (2) whether this effect was greater in logged than in undisturbed forest, (3) if the expected change in tree mortality and light conditions had an effect on Macaranga pioneer seedling and sapling densities, and (4) which (a)biotic factors influenced tree mortality during the drought. The 1997/1998 drought led to an additional tree mortality of 11.2, 18.1, and 22.7% in undisturbed, old logged and recently logged forest, respectively. Mortality was highest in logged forests, due to extremely high mortality of pioneer Macaranga trees (65.4%). Canopy openness was significantly higher during the drought than during the non-drought year (6.0, 8.6 and 10.4 vs 3.7, 3.8 and 3.7 in undisturbed, old logged and recently logged forest, respectively) and was positively correlated with the number of dead standing trees. The increase in light in the understorey was accompanied by a 30 to 300-fold increase in pioneer Macaranga seedling densities. Factors affecting tree mortality during drought were (1) tree species successional status, (2) tree size, and (3) tree location with respect to soil moisture. Tree density and basal area per surface unit had no influence on tree mortality during drought. The results of this study show that extreme droughts, such as those associated with El Niño events, can affect the tree species composition and diversity of tropical forests in two ways: (1) by disproportionate mortality of certain tree species groups and tree size classes, and (2) by changing the light environment in the forest understorey, thereby affecting the recruitment and growth conditions of small and immature trees.     

Testing indicators of sustainable forest management on understorey composition and diversity in southern Italy through variation partitioning

Tree species composition and stand structural complexity are valuable indicators of sustainable forest management. This article aims to investigate the relative influence of forest overstorey composition and structural attributes on understorey composition and diversity, taking into account also site characteristics and broad-scale environmental variables. We sampled vascular plant species composition and forest structure in 132 plots in the Cilento and Vallo di Diano National Park (southern Italy). Spearman’s non-parametric correlation coefficients were calculated between overstorey and understorey diversity indices, beech percentage, and altitude and environmental indices. A complete partitioning of the variation in understorey composition was then performed through canonical correspondence analysis considering four sets of variables: (1) overstorey composition, (2) structural attributes, (3) topography, and (4) landscape abiotic variables. Finally, we constructed a regression tree analysis of understorey species richness using the same explanatory variables. Understorey diversity indices were positively correlated with overstorey diversity indices and with environmental indices (i.e., light and soil heterogeneity). Overstorey and understorey diversity indices were negatively correlated with both altitude and the dominance of beech in the overstorey. Compositional variation was due primarily to overstorey composition and secondarily to structural attributes. Regression tree analysis revealed that altitude, overstorey species richness, and structural attributes play an important role in determining understorey species richness. According to our results, understorey composition and diversity are strongly related to overstorey composition and structural attributes. Indeed, the latter proved to be effective indicators of understorey characteristics in the study area.     

The effects of selective logging on the distribution of moths in a Bornean rainforest

The effects of selective logging on the diversity and species composition of moths were investigated by sampling from multiple sites in primary forest, both understorey and canopy, and logged forest at Danum Valley, Sabah, Malaysia. The diversity of individual sites was similar, although rarefied species richness of logged forest was 17% lower than for primary forest (understorey and canopy combined). There was significant heterogeneity in faunal composition and measures of similarity (NESS index) among primary forest understorey sites which may be as great as those between primary understorey and logged forest. The lowest similarity values were between primary forest understorey and canopy, indicating a distinct canopy fauna. A number of species encountered in the logged forest were confined to, or more abundant in, the canopy of primary forest. Approximately 10% of species were confined to primary forest across a range of species' abundances, suggesting this is a minimum estimate for the number of species lost following logging. The importance of accounting for heterogeneity within primary forest and sampling in the canopy when measuring the effects of disturbance on tropical forest communities are emphasized.     

Functional identity of overstorey tree height and understorey conservative traits drive aboveground biomass in a subtropical forest

The niche complementarity hypothesis has received empirical support but species differ in functional strategies for their contribution to ecosystem function, as predicted by the mass ratio hypothesis. Our understanding of how functional identity of conservative and acquisitive strategies of trees predicts aboveground biomass across forest strata (i.e. overstorey and understorey) remains unclear. Aboveground biomass, community-weighted mean (CWM − functional identity) of trait values (6 leaf and 2 stem traits), and soil physicochemical properties were estimated for 125 plots in a 5-ha subtropical forest in Eastern China. We used multiple linear regressions models to relate aboveground biomass to CWM indices at overstorey and understorey strata separately, and whole-community level. We finally employed the structural equation model to test for the effects of overstorey on understorey strata, in addition to the effects of soil physicochemical properties. Forest strata optimal models showed that overstorey strata had high aboveground biomass when they are dominated by functional identity of tree height, whereas high aboveground biomass in understorey strata was driven by functional identity of dense-wooded conservative strategy. Whole-community optimal model showed that communities dominated by functional identity of leaf dry matter content and mean leaf area had high aboveground biomass. Aboveground biomass was negatively related to soil nutrients across forest strata and whole-community level. The structural equation model showed that CWM of overstorey tree height did not affect understorey functional identity and aboveground biomass, when soil physicochemical properties were accounted. Soil nutrients had positive effect on functional identity of overstorey tree height whereas negative effect on functional identity of understorey dense-wooded strategy. This study highlights the fundamental roles of forest strata where overstorey and understorey strata contribute to their corresponding aboveground biomass with contrasting functional strategies across a range of soil nutrients. High aboveground biomass was potentially driven by functional identity of tree height through making use of plentiful soil nutrients at overstorey strata, whereas by conservative strategy at understorey strata through enduring nutrient-poor soils. To better understand the roles of functional identity of conservative and acquisitive strategies in driving ecosystem functions, it is worth to analyse forest strata separately.     

吉林蛟河针阔混交林林层结构对生产力的影响

探讨不同林层物种多样性和林分结构与森林生产力之间的关系,为森林可持续经营管理策略优化提供理论支持。以吉林蛟河30 hm²针阔混交林固定监测样地中的38432株木本植物的野外调查数据为研究对象,利用结构方程模型探讨了不同林层物种多样性和林分结构多样性对森林生产力的作用机制。结果表明：（1）林冠层林分结构多样性和生产力之间呈负相关关系,路径系数为-0.138;林下层林分结构多样性和生产力之间呈正相关关系,路径系数为0.083。（2）林冠层物种多样性和生产力之间呈正相关关系,路径系数为0.099;林下层物种多样性和生产力之间关系不显著,且物种多样性在林冠层和林下层呈正相关关系,路径系数为0.147。物种多样性和林分结构多样性与森林生产力之间不存在唯一的关系,这种关系的大小、方向和潜在机制与林层有关。为此,在森林经营过程中要充分考虑不同林层物种多样性和林分结构多样性与森林生产力之间的关系。     

Changes in forest land use and management in Sabah, Malaysian Borneo, 1990-2010, with a focus on the Danum Valley region

In an earlier special issue of this journal, Marsh & Greer summarized forest land use in Sabah at that time and gave an introduction to the Danum Valley Conservation Area. Since that assessment, during the period 1990-2010, the forests of Sabah and particularly those of the ca 10 000 km(2) concession managed on behalf of the State by Yayasan Sabah (the Sabah Foundation) have been subject to continual, industrial harvesting, including the premature re-logging of extensive tracts of previously only once-logged forest and large-scale conversion of natural forests to agricultural plantations. Over the same period, however, significant areas of previously unprotected pristine forest have been formally gazetted as conservation areas, while much of the forest to the north, the south and the east of the Danum Valley Conservation Area (the Ulu Segama and Malua Forest Reserves) has been given added protection and new forest restoration initiatives have been launched. This paper analyses these forest-management and land-use changes in Sabah during the period 1990-2010, with a focus on the Yayasan Sabah Forest Management Area. Important new conservation and forest restoration and rehabilitation initiatives within its borders are given particular emphasis.     

Spatial pattern and process in forest stands within the Virginia piedmont

Abstract. Question: Underlying ecological processes have often been inferred from the analysis of spatial patterns in ecosystems. Using an individual‐based model, we evaluate whether basic assumptions of species’life‐history, drought‐susceptibility, and shade tolerance generate dynamics that replicate patterns between and within forest stands. Location: Virginia piedmont, USA. Method: Model verification examines the transition in forest composition and stand structure between mesic, intermediate and xeric sites. At each site, tree location, diameter, and status were recorded in square plots ranging from 0.25 to 1.0 ha. Model validation examines the simulated spatial pattern of individual trees at scales of 1–25 m within each forest site using a univariate Ripley's K function. Results: 7512 live and dead trees were surveyed across all sites. All sites exhibit a consistent, significant shift in pattern for live trees by size, progressing from a clumped understorey (trees ± 0.1 m in diameter) to a uniform overstorey (trees > 0.25 m). Simulation results reflect not only the general shift in pattern of trees at appropriate scales within sites, but also the general transition in species composition and stand structure between sites. Conclusions: This shift has been observed in other forest ecosystems and interpreted as a result of competition; however, this hypothesis has seldom been evaluated using simulation models. These results support the hypothesis that forest pattern in the Virginia piedmont results from competition involving species’life‐history attributes driven by soil moisture availability between sites and light availability within sites.     

Variations in resistance to canopy disturbances and their interactions with the spatial structure of major species in a cool‐temperate forest

Question: How does typhoon‐related disturbance (more specifically, disturbance in the understorey due to tree‐fall and branch‐fall) affect different species mortality rates in a vertically well‐structured forest community? Location: Cool‐temperate, old‐growth forest in the Daisen Forest Reserve, Japan. Methods: We investigated the canopy dynamics and mortality rate trends of trees ≥5 cm diameter at breast height in a 4‐ha study plot, and analysed the effects of tree diameter and spatial structure on the mortality risks for major tree species in the understorey. Results: Significant differences were found in the mortality rates and proportions of injured dead stems between census periods, which were more pronounced in the understorey than in the canopy. Acer micranthum, which showed increased mortality during typhoon disturbance periods, had a clumped distribution. In contrast, Acer japonicum and Viburnum furcatum, which showed similar mortality rates between census periods, had a loosely clumped spatial distribution and a negative association with canopy trees, respectively. In the understorey stems of Acanthopanax sciadophylloides and Fagus crenata, whose spatial distribution patterns depended on canopy gaps, significant increases in mortality rates were observed only during severe typhoon‐related disturbance periods. Conclusions: The sensitivity of trees to typhoon‐related canopy disturbance is more pronounced in the lower layers of vertically structured forest communities. Differences in mortality patterns generated through the combined effects of spatial variation in disturbance regime and species‐specific spatial distribution patterns (spatial aggregation, association with canopy trees, and canopy gap dependency) contribute to the co‐existence of understorey species in forest communities that are subject to typhoon‐related disturbance.     

特大干旱对树木死亡的影响——以美国德克萨斯州东部森林为例

美国德克萨斯州在2011年经历了史上最严重的干旱,这一事件造成约3亿多株树木死亡。在大时空尺度上(面积约9×10~6 hm~2,时间跨度近20年),基于近1800个森林样地,4次周期性调查中的约209663株树木,使用主成份分析(PCA)和广义线性混合效应模型(GLM)回归,对树木死亡的时空差异及其干旱强度与长度对树木死亡造成的中长期复杂影响进行了研究。采用树木密度、树木基面积、林地年龄、样地调查时间间隔作为树木间的竞争指标,分析了造成大旱前后周期水平和年度水平上的样地树木死亡差异的原因。综合分析了不同地理区域、树木种组、胸径大小和林地起源的4个划分标准下树木死亡对死亡率的相对贡献。结果表明：松属树木的死亡率最低(7.92%);高度低、胸径小的树木的死亡率较大,分别为29.79%和26.00%。人工林的树木死亡率(10.26%)低于天然林(13.47%);西海湾平原生态区树木的死亡率在干旱后达到最大(22.27%);西南区的树木死亡率在干旱后也达到最大(13.78%);海拔和纬度对树木死亡率影响不明显。德州东部森林整体死亡格局形成原因较为复杂,各地理区域、林地起源、树木大小和不同树种,...     

Stochasticity of species assemblage of canopy trees and understorey plants in a temperate secondary forest created by major disturbances

Detrended canonical correspondence analysis (DCCA) was applied to explore the species assemblage of plants in a temperate secondary forest that was created by major disturbances. The DCCA showed vague relationships between species dominance and environmental factors for canopy tree species even when rare species were excluded from the analysis. For the highest dominant species of the understorey, the scores of the first axis, which correlated with the species richness of overstorey trees, decreased. This fact showed that the assemblage of canopy trees affects, through the differences in leaf phenology or leaf characteristics of canopy trees, the life history of dominant understorey plants. The studys results suggest that competition colonization might be more important for canopy trees during the developing stage of the forest if the disturbance occurs on a large scale, and that colonization from the local species pool determines species assemblage on a landscape scale.     

Disappearance of overstorey and understorey litter in an open eucalypt forest

Litter accumulation dynamics and first year rates of disappearance were investigated for leaves of overstorey and understorey species in the mixed eucalypt forest in the Griffith University study area, Queensland, Australia. The average biomass of the litter layer, was 10.2 t/ha. The wood and overstorey leaf litter formed 62.7% of the accumulated litter biomass, and were spread continuously across the ground. The distribution of shrub litter was discontinuous, being concentrated in localized patches beneath individual plants. The litter bag and tethered leaf techniques were used to measure the rate of disappearance of overstorey and shrub leaves. The ‘pairedquadrat’ technique was used to measure the weight loss of the grasses. The small leaves of the dominant shrub, Pultenaea villosa Willd. disappeared most rapidly, followed by the overstorey leaves, grasses and Acacia leaves. Fragmentation by physical factors and litter fauna appeared to be the major factors responsible for the disappearance of the overstorey leaves during the first year of exposure. The data suggest that removal of leaf constituents by leaching and microbial decomposition were more important for the shrub litter than the overstorey leaves. Fractional disappearance rates (loss constants) obtained from the short term weight loss measurements, and calculated using the steady-state model of litter accumulation (k=L/X), overestimated the rate of litter disappearance and litter decomposition. For the overstorey leaves in particular, the loss constants also overestimated the rate of loss of material from the litter layer since the fragmented and consumed tissues accumulated in the fraction of comminuted fragments before moving into the humus/soil subsystem. A compartment model of the components of the litter layer in the mixed eucalypt forest is presented. It incorporates overstorey and understorey litter accession, accumulation and disappearance data. The adoption of a two dimensional decomposition/accumulation matrix is suggested as an appropriate framework within which to simulate the dynamics of the litter subsystem in mixed eucalypt forest ecosystems.     

Sapling dynamics in a southeastern Texas floodplain forest

Abstract. How much temporal variation in recruitment, mortality and change in size class occurs in the sapling layer of mature temperate forests in the absence of large‐scale exogenous disturbance? Using 15 years of data from a flood‐plain forest in Big Thicket National Preserve, we found that year‐to‐year variation in demographic parameters was greater than we originally expected. Death rates were generally more variable than recruitment rates, and were much more variable for large saplings than for small ones. Small saplings of the 10 most common species had at least one year when they experienced two to eight times their long‐term mean recruitment and death rates. Large saplings had at least one year when they experienced three to 10 times their long‐term mean death rates and at least one year with two to seven times the long‐term mean recruitment rate. Temporal patterns in sapling death rates were related to flooding patterns, while temporal patterns in recruitment were related to the Palmer Drought Severity Index, an indicator of drought severity and soil moisture availability. We also identified apparently synchronous patterns of demographic response among less flood‐tolerant species which differed from the responses of more flood‐tolerant species. We demonstrated the effects of both climatic variation and light variation in affecting stand‐wide sapling demographics in a forest where canopy gaps are important for regeneration, and where chronic understorey disturbance favours growth over survivorship as a sapling strategy.     

Species richness and canopy productivity of Australian plant communities

The species richness (number of vascular plants per hectare) of Australian plant communities (containing a mosaic of gap, regeneration, maturation and senescent phases) is correlated with the annual biomass productivity of the overstorey canopy.The annual production of leaves and stem in the canopy of the plant community is shown to be limited by the requirements of photosynthesis (particularly light and the availability of water) and the length of the growing season.The species richness of Australian plant communities is the product of the blance between the dominance of the overstorey and the response of the understorey to the shading of the overstorey. For all climatic regions and zones the species richness of the overstorey of the plant community is shown to be exponentially related to the annual shoot growth of the overstorey canopy, until the latitudinal or altitudinal tree line is reached. With latitudinal increase outside the tropics, overstorey canopies of forest communities absorb increasingly more of the incident solar radiation. markedly reducing the species richness of the understorey strata. In contrast, in these latitudes the overstorey of plant communities with widely spaced trees or tall shrubs will absorb far less solar radiation, thus enabling the species richness of the understorey to be maintained.