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.     

Replicated throughfall exclusion experiment in an Indonesian perhumid rainforest: wood production,litter fall and fine root growth under simulated drought

Climate change scenarios predict increases in the frequency and duration of ENSO‐related droughts for parts of South‐East Asia until the end of this century exposing the remaining rainforests to increasing drought risk. A pan‐tropical review of recorded drought‐related tree mortalities in more than 100 monitoring plots before, during and after drought events suggested a higher drought‐vulnerability of trees in South‐East Asian than in Amazonian forests. Here, we present the results of a replicated (n = 3 plots) throughfall exclusion experiment in a perhumid tropical rainforest in Sulawesi, Indonesia. In this first large‐scale roof experiment outside semihumid eastern Amazonia, 60% of the throughfall was displaced during the first 8 months and 80% during the subsequent 17 months, exposing the forest to severe soil desiccation for about 17 months. In the experiment's second year, wood production decreased on average by 40% with largely different responses of the tree families (ranging from −100 to +100% change). Most sensitive were trees with high radial growth rates under moist conditions. In contrast, tree height was only a secondary factor and wood specific gravity had no influence on growth sensitivity. Fine root biomass was reduced by 35% after 25 months of soil desiccation while fine root necromass increased by 250% indicating elevated fine root mortality. Cumulative aboveground litter production was not significantly reduced in this period. The trees from this Indonesian perhumid rainforest revealed similar responses of wood and litter production and root dynamics as those in two semihumid Amazonian forests subjected to experimental drought. We conclude that trees from paleo‐ or neotropical forests growing in semihumid or perhumid climates may not differ systematically in their growth sensitivity and vitality under sublethal drought stress. Drought vulnerability may depend more on stem cambial activity in moist periods than on tree height or wood specific gravity.     

Development processes and growth pattern of Pinus densiflora stands in central eastern Korea

Stand growth and developmental processes were investigated in Pinus densiflora Siebold et Zucc. stands of different ages in the central eastern region of Korea. Stands were inventoried and five trees per stand were sampled for stem analysis, age estimation, and growth analysis. More than 80% of sampled trees in a stand were established within 3–5 years, and most stands had a single cohort structure. The initial growth of pine seedlings was slow, but the height growth accelerated beyond 2–3 m height, 5–10 years after establishment. Linear growth was maintained until 10–12 m height, at which suppressed trees fell behind and might die out. The young stand was composed of pure pines, while few pine seedlings and saplings were found in the understory of older stands. The peak of diameter growth rate occurred around 5–15 years after tree establishment, implying that competition begins during that period. The pine stand development follows four stages: (1) the young stage when the growth rate increases and peaks; (2) the height competition stage when trees focus on height growth for light while maintaining a narrow DBH and height distribution; (3) the differentiation stage when suppressed trees die out, and the DBH distribution becomes wider; and (4) the mature stage when stands have a multi-canopy structure with a wide DBH and height distribution, while the understory is dominated by other tree species. The changes in growth rates and stand structure through forest development would be implemented to predict alterations of above-ground carbon sequestration rates.     

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,...     

Dynamics and structural changes of an oak dominated Natural Forest Reserve in Austria

Natural forest reserves provide a rare opportunity to study forest dynamics after the cessation of human management. Inventories were carried out in 1996 and 2006 in an oak (Quercus spp.) dominated forest reserve formerly managed as coppice forest using the Bitterlich sampling method, an inventory method with a fixed angle of sight to select trees based on their stem diameter. The total living stand volume increased from 245.2 to 276.5 m³/ha (+12.8%) over the 10-year period. This net increase resulted from the growth of individual trees (+3.7%), the ingrowths of young trees (+17.7%) and tree mortality between 1996 and 2006 (−8.6%). Tree mortality included 14.8 m³/ha of standing deadwood and 6.2 m³/ha of fallen deadwood. Stand dynamics differed among tree species: the volume of oak (Quercus spp.) increased due to strong growth and low mortality, whereas hornbeam (Carpinus betulus) showed a decrease in stand volume due to high mortality and low growth. The findings suggest an increase in oak dominance at the expense of hornbeam although inventories repeated over longer time periods would be needed for confirmation. Our data indicate that the Bitterlich sampling method can be used for assessing tree species dynamics and structural changes in natural forest reserves, but some important processes (seedling recruitment, wood decomposition) would need to be investigated separately.

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Zusammenfassung  In einem von Eiche (Quercus spp.) dominierten Naturwaldreservat wurde in den Jahren 1996 und 2006 auf einer Fl?che von 29 ha eine Inventur mittels der Winkelz?hlprobe nach Bitterlich durchgeführt. Der Gesamtvorrat erh?hte sich in der 10-j?hrigen Beobachtungsperiode von 245.2 m³/ha auf 276.5 m³/ha. Im Durchschnitt konnten 14.8 m³/ha stehendes Totholz und 6.2 m³/ha liegendes Totholz ermittelt werden. Die Mortalit?tsrate lag zwischen 1996 und 2006 bei 8.6%. Hainbuche (Carpinus betulus) zeigte aufgrund der hohen Mortalit?t und dem geringen Einwuchs im Vergleich zur Eiche und den anderen Baumarten eine Abnahme im Gesamtvorrat. Die Ergebnisse erlauben einen Einblick in die dynamischen Prozesse unter nahezu natürlichen Bedingungen. Die Eignung der Winkelz?hlprobe für die Beurteilung der Dynamik und die Analyse von strukturellen Ver?nderungen wird kritisch diskutiert.

     

A climate change‐induced threat to the ecological resilience of a subtropical monsoon evergreen broad‐leaved forest in Southern China

Recent studies have suggested that tropical forests may not be resilient against climate change in the long term, primarily owing to predicted reductions in rainfall and forest productivity, increased tree mortality, and declining forest biomass carbon sinks. These changes will be caused by drought‐induced water stress and ecosystem disturbances. Several recent studies have reported that climate change has increased tree mortality in temperate and boreal forests, or both mortality and recruitment rates in tropical forests. However, no study has yet examined these changes in the subtropical forests that account for the majority of China's forested land. In this study, we describe how the monsoon evergreen broad‐leaved forest has responded to global warming and drought stress using 32 years of data from forest observation plots. Due to an imbalance in mortality and recruitment, and changes in diameter growth rates between larger and smaller trees and among different functional groups, the average DBH of trees and forest biomass have decreased. Sap flow measurements also showed that larger trees were more stressed than smaller trees by the warming and drying environment. As a result, the monsoon evergreen broad‐leaved forest community is undergoing a transition from a forest dominated by a cohort of fewer and larger individuals to a forest dominated by a cohort of more and smaller individuals, with a different species composition, suggesting that subtropical forests are threatened by their lack of resilience against long‐term climate change.     

Structural dynamics of a natural mixed deciduous forest in western Thailand

Abstract. Structural dynamics of a natural tropical seasonal – mixed deciduous – forest were studied over a 4-yr period at Mae Klong Watershed Research Station, Kanchanaburi Province, western Thailand, with particular reference to the role of forest fires and undergrowth bamboos. All trees > 5 cm DBH in a permanent plot of 200 m × 200 m were censused every two years from 1992 to 1996. The forest was characterized by a low stem density and basal area and relatively high species diversity. Both the bamboo undergrowth and frequent forest fires could be dominant factors that prevent continuous regeneration. Recruitment, mortality, gain (growth of survival tree plus ingrowth) and loss in basal area (by tree death) during the four years were 6.70%/yr, 2.91%/yr, 1.22%/yr and 1.34%/yr, respectively. Mortality was size dependent; middle size trees (30–50 cm) had the lowest mortality, while the smallest (5–10 cm) had the highest mortality. Tree recruitment was observed particularly in the first two years, mostly in the area where die-back of undergrowth bamboo occurred. The bias of the spatial distribution of recruitment to the area of bamboo die-back was significant and stronger than that to the forest canopy gaps. Successful regeneration of trees which survive competition with other herbs and trees after dieback of bamboo could occur when repeated forest fires did not occur in subsequent years. It is suggested that both the fire disturbance regime and bamboo life-cycle greatly influence the structure and dynamics of this seasonal tropical forest.     

Combining tree-ring analyses on stems and coarse roots to study the growth dynamics of forest trees: a case study on Norway spruce (Picea abies [L.] H. Karst)

We show the potential of a new method combining tree-ring analyses on stems and on coarse roots of individual trees in order to advance the understanding of growth dynamics in forest trees. To this end, we studied the root–shoot allometry of trees and its dependence on site conditions. Along a gradient in water supply in Southern Germany from dry to moist sites we selected 43 Norway spruce trees (Picea abies [L.] H. Karst.) aged 65–100 years. Increment cores were taken from stem and main roots revealing aboveground and belowground growth course over the last 34 years. Annual growth rates in roots and stems and their allometric relationships were applied as surrogate variables for tree resource allocation to aboveground and belowground organs. The mean sensitivities of both stem and root chronologies were found to be site-specific, and increased from the moist through the dry sites. No temporal offset between aboveground and belowground growth reactions to climate conditions was found in Norway spruce at any of the sites. These results suggest that the root–shoot allometry depends on the specific site conditions only at the driest site, following the optimal biomass partitioning theory (the more restricted the water supply, the more organic matter allocation into the belowground organs).     

Long-Term Growth and Succession in Restored and Natural Mangrove Forests in Southwestern Florida

We compared colonization, growth and succession from 1989 to 2000 in a restored mangrove site and in gap and closed canopy sites in a natural mangrove forest. The restored site was created in 1982 and planted with Rhizophora mangle (≈2 m⁻²) propagules. By 1989, Laguncularia racemosa, with densities up to 12.9 tree m⁻², was a dominant in all plots, although densities were greater at edge plots relative to inner plots, and near open water (west plots) relative to further inland (east plots), and in tall mangrove plots relative to scrub plots. Rhizophora mangle (1989 tree densities about 2 m⁻²) was a codominant in inner and scrub plots, while Avicennia germinans had the lowest densities (<1 tree m⁻²) in all plots. From 1989 to 2000 L. racemosa experienced reduced recruitment and apparent density-dependent mortality of canopy individuals in plots with high initial densities. Scrub plots experienced high rates of colonization by R. mangle and L. racemosa, rapid growth in height of all species (1989–1996), followed by a dieoff of L. racemosa in later years (1997–2000) as the canopy came to resemble that of tall mangrove plots. Colonization and growth rates were lower in gap and closed canopy regions of the natural forest relative to rates in the restored site. After 11 years, densities of L. racemosa were 10–20× lower and R. mangle slightly less in the gap relative to densities in tall mangrove plots in the restored site at the same age. Although the restored stand had converged with the natural forest by 2000 in terms of some factors such as species richness, vegetation cover, litterfall, and light penetration, trees were still much smaller and stem densities much higher. Full development of mature structure and ecological function will likely require decades more development.     

Modeling decay rates of dead wood in a neotropical forest

Variation of dead wood decay rates among tropical trees remains one source of uncertainty in global models of the carbon cycle. Taking advantage of a broad forest plot network surveyed for tree mortality over a 23-year period, we measured the remaining fraction of boles from 367 dead trees from 26 neotropical species widely varying in wood density (0.23–1.24 g cm⁻³) and tree circumference at death time (31.5–272.0 cm). We modeled decay rates within a Bayesian framework assuming a first order differential equation to model the decomposition process and tested for the effects of forest management (selective logging vs. unexploited), of mode of death (standing vs. downed) and of topographical levels (bottomlands vs. hillsides vs. hilltops) on wood decay rates. The general decay model predicts the observed remaining fraction of dead wood (R ² = 60%) with only two biological predictors: tree circumference at death time and wood specific density. Neither selective logging nor local topography had a differential effect on wood decay rates. Including the mode of death into the model revealed that standing dead trees decomposed faster than downed dead trees, but the gain of model accuracy remains rather marginal. Overall, these results suggest that the release of carbon from tropical dead trees to the atmosphere can be simply estimated using tree circumference at death time and wood density.     

Tree turnover in a premontane neotropical forest (1998–2009) in Costa Rica

Tropical premontane forests between 700 and 1,400 m.a.s.l. represent a particular component of the gamma diversity of neotropical ecosystems; however, the extent of information about their dynamics lags behind the more studied lowland rain forests. Data from three 1-ha permanent plots in a premontane forest in Costa Rica collected during an 11-year period (1998–2009) suggested a high tree turnover rate for this ecosystem (high mortality rate, λ = 2.4% and annual recruitment, μ = 2.6%). The floristic composition did not significantly change during the study period, but its high dynamism (2.4%) exceeded that of several reported values from highly diverse neotropical lowland rain forests. The documented decrease in abundance (8.6%) and basal area (14.3%) of trees ≥10 cm in DBH differs from the general trend of increase described for several lowland tropical rain forests in recent decades. We detected a significant population reduction (>15% of individuals from 1998 to 2009) in several relatively abundant tree species, whereas the populations of the three most dominant species remained nearly constant. The high tree turnover recorded for this premontane forest might not have affected tree diversity; but it might be promoting recruitment and growth of some tree species that may eventually become over-dominant in this ecosystem.     

Distribution and population structure of four Central Amazonian high-várzea timber species

Amazonian white-water (várzea) floodplains harbor many commercially important timber species which in Brazil are harvested following regulations of the Federal Environmental Agency (IBAMA). Although it is well-known that tree physiology, growth, and species distribution of Amazonian floodplain trees is linked to the heights and durations of the periodical inundations, information about timber stocks and population dynamics is lacking for most tree species. We investigated timber stocks and the population structure of four intensely logged tree species in a western Brazilian várzea forest on an area totaling 7.5 ha. Spatial distribution was investigated in all trees as a function of inundation height and duration and the distance to the river channel, and additionally for saplings (trees <10 cm diameter at breast height––DBH) as a function of the relative photosynthetically active radiation (rPAR). The diameter-class distribution in Hura crepitans and Ocotea cymbarum indicated that populations are subject to density variations that possibly are traced to small-scale flood variability. In all species, saplings concentrated at higher topographic elevations than the mature tree populations, which suggest that the physical ‘escape’ from a flooded environment is an important acclimation to flooding. While Ocotea cymbarum and Guarea guidonia were high-density wood species characterized by random dispersion and a pronounced shade-tolerance, Hura crepitans and Sterculia apetala presented lower wood density, aggregated dispersion, and were more light-demanding. All species presented exploitable stems according to the current harvest regulations, with elevated abundances in comparison to other Amazonian forest types. However, stem densities are below the harvest rates indicating that the harvest regulations are not sustainable. We recommend that the forest management in várzea forests should include specific establishment rates of timber species in dependence of the peculiar site conditions to achieve sustainability.     

Effects of flooding regime and woody bamboos on tree community dynamics in a section of tropical semideciduous forest in South-Eastern Brazil

The influence of a population of the understorey woody bamboo Merostachys riedeliana and different flooding regimes on tree community dynamics in a section of tropical semideciduous forest in South-Eastern Brazil was examined. A forest section with an area of 1.6 ha composed of 71 adjacent plots was located on a slope ending at the river margin. The section was divided into five topographical sectors according to the mean duration of river floods. In 1991 and 1998 all trees with a diameter at the base of the trunk ≥ 5 cm were measured, identified and tagged, and all live bamboo culms were counted. Annualised estimates of the rates of tree mortality and recruitment, gain and loss of tree basal area, and change in bamboo density were calculated for each of the 71 plots and five topographical sectors as well as for diameter classes and tree species. To segregate patterns arising from spatially autocorrelated events, geostatistical analyses were used prior to statistical comparisons and correlations. In general, mortality rates were not compensated by recruitment rates but there was a net increase in basal area in all sectors, suggesting that the tree community as a whole was in a building phase. Tree community dynamics of the point bar forest (Depression and Levée sectors) differed from that of the upland forest (Ridgetop, Middle Slope and Lower Slope sectors) in the extremely high rates of gain in basal area. The predominant and specialised species, Inga vera and Salix humboldtiana, are probably favoured by relaxed competition in an environment stressed by long-lasting floods. In the upland forest, mortality rates were highest at the Middle Slope, particularly for smaller trees, while recruitment rates were lowest. As bamboo clumps were concentrated in this sector, the locally higher instability in the tree community probably resulted from the direct interference of bamboos. The density of bamboo culms in the upland forest was negatively correlated with the rates of tree recruitment and gain in basal area, and positively correlated with tree mortality rates. Bamboos therefore seemed to restrict the recruitment, growth and survival of trees. This revised version was published online in June 2006 with corrections to the Cover Date.     

Factors affecting fall down rates of dead aspen (Populus tremuloides) biomass following severe drought in west‐central Canada

Increases in mortality of trembling aspen (Populus tremuloides Michx.) have been recorded across large areas of western North America following recent periods of exceptionally severe drought. The resultant increase in standing, dead tree biomass represents a significant potential source of carbon emissions to the atmosphere, but the timing of emissions is partially driven by dead‐wood dynamics which include the fall down and breakage of dead aspen stems. The rate at which dead trees fall to the ground also strongly influences the period over which forest dieback episodes can be detected by aerial surveys or satellite remote sensing observations. Over a 12‐year period (2000–2012), we monitored the annual status of 1010 aspen trees that died during and following a severe regional drought within 25 study areas across west‐central Canada. Observations of stem fall down and breakage (snapping) were used to estimate woody biomass transfer from standing to downed dead wood as a function of years since tree death. For the region as a whole, we estimated that >80% of standing dead aspen biomass had fallen after 10 years. Overall, the rate of fall down was minimal during the year following stem death, but thereafter fall rates followed a negative exponential equation with k = 0.20 per year. However, there was high between‐site variation in the rate of fall down (k = 0.08–0.37 per year). The analysis showed that fall down rates were positively correlated with stand age, site windiness, and the incidence of decay fungi (Phellinus tremulae (Bond.) Bond. and Boris.) and wood‐boring insects. These factors are thus likely to influence the rate of carbon emissions from dead trees following periods of climate‐related forest die‐off episodes.     

Neighborhood interactions influencing tree population dynamics in nonpyrogenous boreal forest in northern Finland

Open-canopy moss-rich woodlands dominated by Picea abies and Betula pubescens in northern Finland may undergo cyclic development with reciprocal replacement of the tree species due to the positive feedbacks on soil conditions. Although relations to the abiotic environment are well understood, intra- and interspecific interactions during development of sparse boreal forests have received less attention. We studied tree regeneration, growth and survival with respect to size and density of neighboring trees in four stands representing roughly four stages of the Picea–Betula forest cycle. We conducted spatial analysis (Ripley’s K-function) of mapped locations of live and dead stems to reconstruct the distribution of stems prior to mortality, and to infer possible causes of tree population decline. The prevalence of standing dead stems enabled us to test if mortality was associated with density and size of neighboring trees. Logistic regression was used to test for relationships between tree survival and local crowding indices. We also examined spatial autocorrelations of individual size characteristics to determine the mode and spatial extent of tree interactions. Picea abies had reduced recruitment in open areas occupied by mosses and dwarf-shrubs, and preferentially regenerated near B. pubescens, whereas B. pubescens formed small clumps (and occasionally these consisted entirely of stems from a single tree) that showed local repulsion from large P. abies trees. Size of neighboring trees was the primary determinant of individual growth and survival, whereas neighborhood density per se had only a weak effect. Picea abies had negatively correlated sizes among close neighbors (0–4 m radius) indicating that dominant trees suppress their smaller neighbors. Negative autocorrelations prevailed at the transition stages where the patches of smaller trees were concentrated around evenly spaced large trees. Tree sizes became spatially independent at the mature phase. We hypothesize that both low light and soil nutrient availability causes the P. abies population decline. Dominant trees in this high latitude forest have large light depletion zones and shallow root system to promote strong above- and below-ground competition with younger trees. Higher mortality rates within canopy patches were not compensated for by recruitment in gaps, causing P. abies population to decline steadily.     

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.     

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.     

Rainforest fragmentation and the demography of the economically important palm <Emphasis Type="Italic">Oenocarpus bacaba</Emphasis> in central Amazonia

We summarize a long-term study of the effects of edge creation on establishment of the economically important arboreal palm Oenocarpus bacaba in an experimentally fragmented landscape in central Amazonia. Recruitment and mortality of large individuals (≥10 cm diameter-at-breast-height) were recorded within 21 1-ha plots in fragmented and intact forests for periods of up to 22 years. In addition, 12 small (0.7 × 14 m) sub-plots within each 1-ha plot were used to enumerate the abundance of seedlings and saplings (5–400 cm tall). On average, the recruitment of large trees was over two times faster near forest edges, leading to a sharp (90%) increase in the mean population density of large individuals near forest edges, whereas the density of larger trees remained constant in the forest interior. Overall seedling and sapling density was significantly lower in edge than interior plots, but edge plots had a much higher proportion of larger (>100 cm tall) saplings. Our findings demonstrate that forest edges can have complex effects on tree demography and that one must consider all tree life stages in order to effectively assess their effects on plant populations.     

Liana cutting for restoring tropical forests: a rare palaeotropical trial

Liana growth following forest disturbance is threatening the tropical carbon sink by delaying or preventing recovery. Tree growth can be stimulated by liana cutting; however, its applicability for conservation management remains uncertain, particularly in Africa (the least‐studied continent for ecological restoration) and against pervasive barriers such as wildfires. We conducted a small‐scale trial to investigate tree sapling regeneration following liana cutting in a lowland African forest prone to low intensity wildfires. We employed a BACI design comprising eighteen 25 m² plots of sapling trees in liana‐infested areas. After 5 years of liana cutting, we saw greater recruitment, stem growth and net biomass. Wildfires caused 51% mortality and probably masked liana cutting influences on species and survival, but may have encouraged stem recruitment through interaction with liana cutting. Incorporating our data into a first quantitative review of previous studies, we found that tree growth, recruitment and net growth rates were all consistently higher where lianas were either absent or removed (respectively: 80%, 215%, 633%; n = 14, 3, 4). Tree growth impacts were approximately equivalent across size‐classes and continents. We give recommendations for improved plot and sample sizes, but conclude that liana cutting is a promising restoration method for lowland tropical forests, including Africa.     

Effect of liana cutting on water potential and growth of adult<Emphasis Type="Italic"> Senna multijuga</Emphasis> (Caesalpinioideae) trees in a Bolivian tropical forest

Lianas, or woody climbing plants, are a major constituent of seasonally dry tropical forests, and are thought to impact negatively their host trees. In this study we evaluated whether liana presence was associated with reduced leaf water potentials and growth in adult Senna multijuga trees during the dry season in a lowland Bolivian forest. We used leaf water potentials in trees as a first approach to assess trees’ water status, under the assumption that leaf water potentials become more negative when water losses (via transpiration) exceed gains (by uptake). We measured relative growth in girth at 1.5 m height (gbh) to quantify tree growth. At the beginning of the 1996 dry season (early June), we selected 20 S. multijuga trees 10–20 cm dbh, and measured their gbh. We also recorded pre-dawn and mid-day leaf water potentials in these trees. In ten experimental trees all lianas were then cut, while the remaining trees were used as controls. Pre-dawn and mid-day water potentials were re-measured 1 day after liana-cutting, and then every week in all trees for 1 month and then at 3 and 5 months, until the beginning of the next rainy season (November); gbh was measured again in July 1997 to estimate relative growth rate. Liana removal was associated with less negative pre-dawn (–0.3 vs –0.4 MPa) and mid-day (–0.5 vs –0.7 MPa) water potentials in trees during the dry season. This difference appeared as early as 1 day after cutting, and disappeared once the rainy season began. Liana-cut trees grew more (0.4 mm/mm year) than liana-uncut trees (0.2 mm/mm year). These findings suggest that lianas may interfere with water availability to these trees during the dry season, and may also hinder tree growth. Received: 16 November 1999 / Accepted: 23 March 2000