Increasing biomass in Amazonian forest plots

A previous study by Phillips et al. of changes in the biomass of permanent sample plots in Amazonian forests was used to infer the presence of a regional carbon sink. However, these results generated a vigorous debate about sampling and methodological issues. Therefore we present a new analysis of biomass change in old-growth Amazonian forest plots using updated inventory data. We find that across 59 sites, the above-ground dry biomass in trees that are more than 10 cm in diameter (AGB) has increased since plot establishment by 1.22 +/- 0.43 Mg per hectare per year (ha(-1) yr(-1), where 1 ha = 10(4) m2), or 0.98 +/- 0.38 Mg ha(-1) yr(-1) if individual plot values are weighted by the number of hectare years of monitoring. This significant increase is neither confounded by spatial or temporal variation in wood specific gravity, nor dependent on the allometric equation used to estimate AGB. The conclusion is also robust to uncertainty about diameter measurements for problematic trees: for 34 plots in western Amazon forests a significant increase in AGB is found even with a conservative assumption of zero growth for all trees where diameter measurements were made using optical methods and/or growth rates needed to be estimated following fieldwork. Overall, our results suggest a slightly greater rate of net stand-level change than was reported by Phillips et al. Considering the spatial and temporal scale of sampling and associated studies showing increases in forest growth and stem turnover, the results presented here suggest that the total biomass of these plots has on average increased and that there has been a regional-scale carbon sink in old-growth Amazonian forests during the previous two decades.     

Effects of individual size,local competition and canopy closure on the stem volume growth in a monoclonal Japanese cedar (<Emphasis Type="Italic">Cryptomeria japonica</Emphasis> D. Don) plantation

We studied how the dominant factor affecting stem volume growth changes during stand development in a monoclonal stand of Cryptomeria japonica D. Don. Stem analysis was used to compare growth history of trees in an unthinned plot (closed canopy) and a thinned plot (open canopy). In the unthinned plot, the dominant factor affecting stem volume growth was basal area (BA) before canopy closure, whereas neighborhood competition index (CI) was the dominant factor after canopy closure. In contrast, the dominant factor affecting stem volume in the thinned plot was BA throughout stand development. Spearman’s rank correlation coefficient between BA and CI continued to increase after canopy closure and size rank among individuals became increasingly fixed. Our results indicated that stem volume growth shifts from size-dependent to competition-dependent growth at canopy closure. The apparent correlation between tree size and growth rate observed in many previous studies may be the result of competition-mediated positive feedback between size and growth.     

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.     

Growth and mortality of trees in Amazonian tropical rain forest in Ecuador

Abstract. Structural changes are analysed in four samples representing 4 ha, two line transects and two hectare plots, of Amazonian tropical lowland rain forest in northern Ecuador. Only trees with a DBH ≥ 10 cm were included. A sample of floodplain forest in Añangu represents the largest turnover found in tropical forests (stand half-life = 23 yr). The line transect and hectare plot both of tierra firme forest in Añangu have the same turnover (37 yr) and were balanced for death and in-growth of both individuals and wood (basal area). The 1-ha tierra firme sample in Cuyabeno had a turnover of 67 yr and was in a growing phase. The floodplain line transect in Añangu was in a phase of structural breakdown. However, the floodplain line transect had the largest growth of basal area per tree (23.4 cm²/yr). The tierra firme samples had a growth of 9.6, 10.1, and 13.6 cm²/yr. Most of the dead trees fell with some uprooting in three of the four samples. However, no significant difference in the distribution of mode of death was found between the four samples. Death was independent of topography and the dead trees were randomly distributed. As the trees grow up they occupy more space and larger trees (DBH ≥ 15 cm) become more uniformly distributed, whereas smaller trees (DBH ≤ 15 cm) were randomly distributed. Our study confirms that plots of 1 ha are not sufficient to include representative samples of different stages of forest structure.     

Seed shadow, seedling recruitment, and spatial distribution of Buchenavia capitata (Combretaceae) in a fragment of the Brazilian Atlantic forest.

Here we describe the seed shadow, seedling recruitment, ontogenetic structure and spatial distribution of Buchenavia capitata (an emergent canopy tree) in a 380-ha fragment of the Atlantic forest in northeast Brazil. In particular, we examine seed distribution around 10 parental trees and both seedling recruitment and mortality, during an 18 month period beneath and around parental trees. Moreover, we describe: (1) B. capitata occurrence within treefall gaps; (2) population structure in terms of ontogenetic stages for the whole site; and (3) spatial distribution of adults within an area of 51 hectares. 99% of seeds were found beneath parent crowns (n = 4,236) and seed density reached 14.6 +/- 29.9 seeds/m2 (0-140 seeds/m2). 49% of all seeds germinated but seedling mortality reached 100% after an 18 month period. In addition, saplings of B. capitata were not found in forest understory and within 30 treefall gaps (94-2,350 m2). The adults showed an average DBH of 69.3 +/- 22.1 cm, were 19.2 +/- 2.9 m tall and presented a clumped spatial distribution. B. capitata matched some of the features presented by shade intolerant trees or large-gap specialists, and we hypothesize that low rates or even lack of long distance seed dispersal events may be reducing the probability of B. capitata seeds reaching suitable habitats for successful seedling recruitment and growth. Because of that (1) seedlings face high levels of early mortality; (2) there is no sapling recruitment at the study site; and (3) local population faces senility and it is threatened by local extinction.     

Dynamics of the forest communities at Pasoh and Barro Colorado: comparing two 50-ha plots

Dynamics of the Pasoh forest in Peninsular Malaysia were assessed by drawing a comparison with a forest in Panama, Central America, whose dynamics have been thoroughly described. Census plots of 50 ha were established at both sites using standard methods. Tree mortality at Pasoh over an eight-year interval was 1.46% yr(-1) for all stems > or = 10 mm diameter at breast height (dbh), and 1.48% yr(-1) for stems > or = 100 mm dbh. Comparable figures at the Barro Colorado Island site in Panama (BCI) were 2.55% and 2.03%. Growth and recruitment rates were likewise considerably higher at BCI than at Pasoh. For example, in all trees 500-700 mm in dbh, mean BCI growth over the period 1985-1995 was 6 mm yr(-1), whereas mean Pasoh growth was about 3.5 mm yr(-1). Examining growth and mortality rates for individual species showed that the difference between the forests can be attributed to a few light-demanding pioneer species at BCI, which have very high growth and mortality; Pasoh is essentially lacking this guild. The bulk of the species in the two forests are shade-tolerant and have very similar mortality, growth and recruitment. The Pasoh forest is more stable than BCI's in another way as well: few of its tree populations changed much over the eight-year census interval. In contrast, at BCI, over 10% of the species had populations increasing or decreasing at a rate of >0.05 yr(-1) compared to just 2% of the species at Pasoh). The faster species turnover at BCI can probably be attributed to severe droughts that have plagued the forest periodically over the past 30 years; Pasoh has not suffered such extreme events recently. The dearth of pioneer species at Pasoh is associated with low-nutrient soil and slow litter breakdown, but the exact mechanisms behind this association remain poorly understood.     

Response of tree biomass and wood litter to disturbance in a Central Amazon forest

We developed an individual-based stochastic-empirical model to simulate the carbon dynamics of live and dead trees in a Central Amazon forest near Manaus, Brazil. The model is based on analyses of extensive field studies carried out on permanent forest inventory plots, and syntheses of published studies. New analyses included: (1) growth suppression of small trees, (2) maximum size (trunk base diameter) for 220 tree species, (3) the relationship between growth rate and wood density, and (4) the growth response of surviving trees to catastrophic mortality (from logging). The model simulates a forest inventory plot, and tracks recruitment, growth, and mortality of live trees, decomposition of dead trees (coarse litter), and how these processes vary with changing environmental conditions. Model predictions were tested against aggregated field data, and also compared with independent measurements including maximum tree age and coarse litter standing stocks. Spatial analyses demonstrated that a plot size of ~10 ha was required to accurately measure wood (live and dead) carbon balance. With the model accurately predicting relevant pools and fluxes, a number of model experiments were performed to predict forest carbon balance response to perturbations including: (1) increased productivity due to CO₂ fertilization, (2) a single semi-catastrophic (10%) mortality event, (3) increased recruitment and mortality (turnover) rates, and (4) the combined effects of increased turnover, increased tree growth rates, and decreased mean wood density of new recruits. Results demonstrated that carbon accumulation over the past few decades observed on tropical forest inventory plots (~0.5 Mg C ha^–1 year^–1) is not likely caused by CO₂ fertilization. A maximum 25% increase in woody tissue productivity with a doubling of atmospheric CO₂ only resulted in an accumulation rate of 0.05 Mg C ha^–1 year^–1 for the period 1980–2020 for a Central Amazon forest, or an order of magnitude less than observed on the inventory plots. In contrast, model parameterization based on extensive data from a logging experiment demonstrated a rapid increase in tree growth following disturbance, which could be misinterpreted as carbon sequestration if changes in coarse litter stocks were not considered. Combined results demonstrated that predictions of changes in forest carbon balance during the twenty-first century are highly dependent on assumptions of tree response to various perturbations, and underscores the importance of a close coupling of model and field investigations.     

Age-related Decline in Forest Ecosystem Growth: An Individual-Tree, Stand-Structure Hypothesis

Forest growth is important both economically (yielding billions of dollars of annual revenues) and ecologically (with respect to ecosystem health and global carbon budgets). The growth of all forests follows a predictable general trend with age. In young forests, it accelerates as canopies develop; it then declines substantially soon after full canopy leaf area is reached. The classic explanation for the decline in growth invoked the increasing respiration costs required to sustain the larger masses of wood characteristic of older forests. Direct measurements of respiration have largely refuted this hypothesis, and recent work has focused on stand-level rates of resource supply, resource use, and growth. We developed and tested a hypothesis at the scale of individual trees (in relation to stand structure) to explain this declining stand-level rate of stem growth. According to our hypothesis, changes in stand structure allow dominant trees to sustain high rates of growth by increasing their acquisition of resources and using these resources efficiently (defined as stem growth per unit of resource used); smaller, nondominant trees grow more slowly as a result of their more limited acquisition of resources and a reduced rate of growth per unit of resource acquired. In combination, these two trends reduce overall stand growth. We tested this hypothesis by comparing growth, growth per unit of leaf area, and variation among trees within plots in two series of plantations of Eucalyptus in Brazil and by estimating individual-tree rates of growth and use of light, water, and nutrients in a plantation of Eucalyptus saligna in Hawaii. Our results supported the individual-tree hypothesis. We conclude that part of the universal age-related decline in forest growth derives from competition-related changes in stand structure and the resource-use efficiencies of individual trees. Received 19 February 2001; accepted 19 June 2001.     

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.     

Demography of the long‐lived conifer Agathis ovata in maquis and rainforest,New Caledonia

Abstract. The endemic New Caledonian conifer Agathis ovata occurs as an emergent tree in fire‐prone shrublands (maquis), and fire‐sensitive rainforest. Growth, survivorship and recruitment over 5 yr were compared for populations from forest and maquis on ultramafic substrates in New Caledonia to investigate whether demographic behaviour varied in response to the strongly contrasting forest and shrubland environments. Growth of seedlings and of small (30–100 cm height) and large (100 cm height; 5 cm DBH) saplings was slow, but varied significantly among stages, site types and years. The greatest difference in growth rates was among stages, seedlings growing 0.34 cm.yr^?1, small saplings 1.06 cm.yr^?1 and large saplings 2.13 cm.yr^?1. Tree DBH increased by only 0.05 cm.yr^?1 and, based on these rates, individuals with DBH of 30 cm are estimated to be more than 700 yr old. Few trees (3.5%) produced cones in any year and seedling recruitment was low, but some recruitment was recorded each year in both maquis and forest. Rates of recruitment per parent were highest in forest (1.28.yr^?1, cf 0.78.yr^?1), but the higher density of trees in maquis meant that overall recruitment was greater there (92 ha^?1.yr^?1, cf 56 ha^?1.yr^?1). Seedling mortality ranged from 0.9 to 2.9% among years with no significant difference between maquis and forest. No sapling mortality was recorded, but annual tree mortality ranged from 0 to 1.4%. Evidence from a recently burned site indicated that while trees may survive fire, seedlings and saplings do not. Post‐fire seedling recruitment per ha from surviving trees was four times lower than in unburned sites, but growth rates were four times higher. Similar demographic attributes, including high survivorship, low growth rate and low rates of recruitment over a long reproductive life, characterize Agathis ovata populations in both maquis and rainforest in New Caledonia and are indicative of a broad tolerance of light environments that is unusual among tree species. These demographic attributes help to explain the long‐term persistence of the species in these strongly contrasting habitats.     

Low dynamics, high longevity and persistence of sessile structural species dwelling on Mediterranean coralligenous outcrops

There is still limited understanding of the processes underlying benthic species dynamics in marine coastal habitats, which are of disproportionate importance in terms of productivity and biodiversity. The life-history traits of long-lived benthic species in these habitats are particularly poorly documented. In this study, we assessed decadal patterns of population dynamics for ten sponge and anthozoan species that play key structural roles in coralligenous outcrops (～25 m depth) in two areas of the NW Mediterranean Sea. This study was based on examination of a unique long-term photographic series, which allowed analysis of population dynamics over extensive spatial and time spans for the very first time. Specifically, 671 individuals were censused annually over periods of 25-, 15-, and 5-years. This long-term study quantitatively revealed a common life-history pattern among the ten studied species, despite the fact they present different growth forms. Low mortality rates (3.4% yr(-1) for all species combined) and infrequent recruitment events (mean value of 3.1±0.5 SE recruits yr(-1)) provided only a very small fraction of the new colonies required to maintain population sizes. Overall, annual mortality and recruitment rates did not differ significantly among years; however, some species displayed important mortality events and recruitment pulses, indicating variability among species. Based on the growth rates of these 10 species, we projected their longevity and, obtained a mean estimated age of 25-200 years. Finally, the low to moderate turnover rates (mean value 0.80% yr(-1)) observed among the coralligenous species were in agreement with their low dynamics and persistence. These results offer solid baseline data and reveal that these habitats are among the most vulnerable to the current increases of anthropogenic disturbances.     

Forest regeneration following ungulate removal in a montane Hawaiian wet forest

Most Hawaiian forests lack resiliency following disturbance due to the presence of non‐native and invasive plant and animal species. The montane wet forest within Hakalau Forest National Wildlife Refuge on Hawai'i island has a long history of ungulate disturbance but portions of the refuge were fenced and most ungulates excluded by the early 1990s. We examined patterns of regeneration within two 100 ha study sites in this forest following the removal of ungulates and in the absence of invasive woody tree species to determine, in part, if passive restoration techniques can be successful under these conditions. We characterized growth, mortality, and basal area (BA) changes for approximately 7,100 marked individuals of all native tree species present in two surveys over a 17–18‐year period within two hundred 30 m diameter forest plots. Considerable recruitment within plots of new trees of all species significantly changed size class distributions and erased deficits in small‐sized trees observed during the first survey, particularly for the codominant canopy tree, koa (Acacia koa). Overall, growth of established dominant 'ōhi'a trees (Metrosideros polymorpha) and recruitment of mid‐canopy trees contributed to increases in BA while high levels of mortality for large A. koa trees contributed to decreased BA. This resulted in a slight increase in BA between the two surveys (+1.9%). This study demonstrates that fencing and ungulate removal may have rescued the A. koa population by facilitating the first real pulse in recruitment in over a century, and that passive restoration can be a successful management strategy in this forest.     

Pattern and process in Amazon tree turnover, 1976-2001

Previous work has shown that tree turnover, tree biomass and large liana densities have increased in mature tropical forest plots in the late twentieth century. These results point to a concerted shift in forest ecological processes that may already be having significant impacts on terrestrial carbon stocks, fluxes and biodiversity. However, the findings have proved controversial, partly because a rather limited number of permanent plots have been monitored for rather short periods. The aim of this paper is to characterize regional-scale patterns of 'tree turnover' (the rate with which trees die and recruit into a population) by using improved datasets now available for Amazonia that span the past 25 years. Specifically, we assess whether concerted changes in turnover are occurring, and if so whether they are general throughout the Amazon or restricted to one region or environmental zone. In addition, we ask whether they are driven by changes in recruitment, mortality or both. We find that: (i) trees 10 cm or more in diameter recruit and die twice as fast on the richer soils of southern and western Amazonia than on the poorer soils of eastern and central Amazonia; (ii) turnover rates have increased throughout Amazonia over the past two decades; (iii) mortality and recruitment rates have both increased significantly in every region and environmental zone, with the exception of mortality in eastern Amazonia; (iv) recruitment rates have consistently exceeded mortality rates; (v) absolute increases in recruitment and mortality rates are greatest in western Amazonian sites; and (vi) mortality appears to be lagging recruitment at regional scales. These spatial patterns and temporal trends are not caused by obvious artefacts in the data or the analyses. The trends cannot be directly driven by a mortality driver (such as increased drought or fragmentation-related death) because the biomass in these forests has simultaneously increased. Our findings therefore indicate that long-acting and widespread environmental changes are stimulating the growth and productivity of Amazon forests.     

Population dynamics of four understorey shrub species during a 7‐yr period in a primary beech forest

Abstract. Shrubs persist in the understorey layer of forests throughout their lives, while tall trees remain there only during the juvenile stage and then grow into the canopy layer. Thus demographic parameters (recruitment‐, mortality‐, and growth‐rates) of shrub species are expected to differ from those of tall tree species. We investigated aspects of the demography of four dominant deciduous‐shrub species (Viburnum furcatum, Lindera umbellata var. membranacea, Magnolia salicifolia, and Hydrangea paniculata) in Fagus crenata forests at the beginning and at the end of a 7‐yr period in a 1‐ha permanent plot. For each species, the number of stems changed little (within ± 10%) during the study period, while total basal area increased markedly (11.7–33.8%), because (1) new stems continuously recruited by vegetative growth replaced the substantial number of dead stems, and (2) vegetative stems grew vigorously, probably due to nutrient support from parents. The results indicate that these four understorey shrub species have high ability to maintain their population size in the shaded forest understorey. While in each species more than half of the dead stems were standing dead, a substantial proportion of the dead stems (9.0–38.5%) showed signs of mechanical damage, such as stem breakage and suppression by fallen branches or trees. Snow pressure that resulted in decumbent stems was also an important mortality agent for V. furcatum (20.7%) and L. umbellata var. membranacea (5.6%). Probability of damage was constant across all DBH‐classes for all study species. In each species, newly recruited stems and dead stems were spatially aggregated, largely due to habits of vegetative growth and mechanical damage, respectively. This study revealed that several demographic traits, resulting from recruitment by vegetative growth and death by mechanical damage, were shrub‐species specific and markedly different from those of tall tree species.     

Interesting times on Krakatau: stand dynamics in the 1990s

The Krakatau Islands, Indonesia, have provided an opportunity for ecologists to track primary succession from the 'clean slate' of 1883, through forest closure in the 1920s, to the contemporary period, in which successional changes take the form of alterations in composition and stature of forest stands rather than gross changes in ecosystem type. This paper reports on permanent forest plots established on the islands in 1989, and fully surveyed again in both 1992 and 1997. Since 1989, the plots have been subject to natural disturbance phenomena in the form of varying combinations of, for example, deposition of volcanic ejecta, landslides, lightning strikes, storm damage and drought. These effects have been concentrated between 1992 and 1997, during which the volcano Anak Krakatau has deposited ash on the islands of Sertung and Panjang, but not on Rakata. Data on stand responses are presented for growth rates (dbh (diameter at breast height, 1.3 m) increment), stem recruitment and mortality, biomass changes partitioned into mortality, ingrowth and growth of established trees), and compositional shifts. The discussion focuses on evaluation of questions and successional models framed earlier in the programme. One general finding is that the stand dominants as of 1989 have tended to decline in number within the plots, generally through low levels of recruitment failing to balance rates of mortality. The effects of disturbance to the plots appear to be evident in terms of mortality and recruitment, dbh increment, and changes in biomass. The patterns of change in the eight plots are quite varied, such that relatively few generalizations are possible. The difficulties of establishing meaningful baseline rates for tree growth and stand biomass are discussed.     

Long-term growth dynamics of natural forests in Hokkaido,northern Japan

Abstract. The long-term growth dynamics of natural forest stands on the island of Hokkaido were described on the basis of an analysis of data from 38 permanent plots spanning 15–22 yr. Stand structure was characterized by basal area, stem density and tree size variability. To detect trends in stand structure, regression models for recruitment rate (per ha per yr), mortality rate and the rate of change in stem density and tree size variability were developed by a stepwise method using initial basal area, stem density, tree size variability, species composition summarized by LNMDS ordination, altitude, annual mean temperature, annual precipitation, type of understorey vegetation, topography and slope aspect as candidates for predictor variables. The same analyses were conducted for basal area increment (net growth) and its components: survivor growth = basal area gain by growth of surviving individuals and mortality = basal area loss by death of individuals. Stem density remained generally unchanged; recruitment was relatively low even in very sparse stands. Stand basal area generally increased as survivor growth was approximately double the mortality. Recruitment rate was strongly affected by the presence of dwarf bamboo (Sasa spp.) vegetation on the forest floor which inhibited tree regeneration. Mortality rate was density-dependent; dense stands had higher mortality than sparse stands. Density change rate (recruitment rate - mortality rate) was, therefore, determined by both the type of understorey vegetation and stem density. Survivor growth was high in stands with high stem density and basal area. Mortality was dependent on basal area and altitude. Net basal area increment (net growth) was dependent only on stem density with other factors that influenced survivor growth and mortality omitted. Tree size variability decreased in stands with high tree size variability whereas it increased in stands with low size variability. Based on the obtained models for density change rate and net basal area increment, trajectories of stands were illustrated on a log-log diagram of stem density and basal area. The predicted differences in trajectories as affected by the understorey vegetation type indicated the importance of dwarf bamboo vegetation for forest dynamics on Hokkaido.     

Frequent,low‐amplitude disturbances drive high tree turnover rates on a remote,cyclone‐prone Polynesian island

Aim How important are frequent, low‐intensity disturbances to tree community dynamics of a cyclone‐prone forest? We tested the following hypotheses concerning the ‘inter‐cataclysm’ period on a remote Polynesian island: (1) tree turnover would be high and recruitment rates would be significantly higher than mortality; (2) low‐intensity disturbance would result in a marginal increase in tree mortality in the short term; (3) turnover would vary among species and would be associated with plant traits linked to differences in life history; and (4) mortality and recruitment events would be spatially non‐random. Location Tutuila, a volcanic island in the Samoan Archipelago, Polynesia. Methods We censused the tree (stem diameter ≥ 10 cm) community in 3.9 ha of tropical forest three times over a 10‐year period, 1998–2008. We calculated annual mortality, recruitment and turnover rates for 36 tree species. We tested for non‐random spatial patterns and predictors of mortality, and non‐random spatial patterns of tree recruitment. A 2004 cyclone passing within 400 km allowed us to measure the effects of a non‐cataclysmic disturbance on vital rates. Results Annual turnover was 2.8% and annual recruitment was 3.6%; these are some of the highest rates in the tropics, and likely to be a response to a cyclone that passed < 50 km from Tutuila in 1991. Species turnover rates over 10 years were negatively correlated with wood specific gravity, and positively correlated with annual stem diameter increment. Mortality was spatially aggregated, and a function of site, species and an individual’s growth rate. Recruitment was highest on ground with low slope. The low‐magnitude cyclone disturbance in 2004 defoliated 29% of all trees, but killed only 1.8% of trees immediately and increased annual mortality over 5 years by 0.7%. Main conclusions The inter‐cataclysm period on Tutuila is characterized by frequent, low‐amplitude disturbances that promote high rates of tree recruitment and create a dynamic, non‐equilibrium or disturbed island disequilibrium tree community. Species with low wood density and fast growth rates have enhanced opportunities for recruitment between cataclysms, but also higher probabilities of dying. Our results suggest that increases in the frequency of cyclone activity could shift relative abundances towards disturbance‐specialist species and new forest turnover rates.     

Species Dynamics During Early Secondary Forest Succession: Recruitment, Mortality and Species Turnover

The "Initial Floristic Composition" hypothesis is applied to secondary tropical rain forest succession in abandoned agricultural fields with light previous land-use and close to seed sources. This hypothesis predicts that both pioneer and shade-tolerant species colonize a site directly after abandonment, and as the canopy closes, the recruitment of pioneers sharply declines, while recruitment of shade-tolerant species continues. It also predicts higher mortality among pioneers. Consequently, recruited and dead trees are expected to differ in species composition, with highest species richness for the recruits. During 18 mo, we monitored recruitment and mortality of trees with height ≥ 1.5 m in eight plots in abandoned cornfields with initial fallow age of 1–5 yr, in SE Mexico. Shade-tolerant species established in the first years of succession, albeit in low numbers. As predicted, recruited and dead trees differed in species richness and composition, and in shade-tolerant frequency. In contrast to our expectations, over 50 percent of recruits were from pioneer species, as high stand-level mortality opened new opportunities for continued pioneer colonization. Species turnover starts very early in succession but is not always a gradual and continuous process, complicating prevailing succession models. The strong spatial and temporal variability of succession emphasizes the need to monitor these dynamics in permanent plots across a range of initial stand ages, with multiple plots in a given age class.     

Tree population changes in a tropical dry evergreen forest of south India over a decade (1992–2002)

Changes in species composition and density of trees >10 cm gbh in a tropical dry evergreen forest in Puthupet, south India are interpreted for the period between 1992 and 2002. A 1-ha plot was inventoried in 1992 and was recensused in 2002. During the 10-year interval tree taxa diversity as well as stand density increased, but the basal area value decreased. Tree species richness increased by 21% (from 24 to 29 species) by an addition of eight species and local extinction of three species. The tree density increased just by eight individuals (from 1330 stems ha⁻¹ in 1992 to 1338 ha⁻¹ in 2002), but the basal area decreased by 8% (from 37.5 to 34.5 m² ha⁻¹). Many species (11 numbers) have increased in abundance rather than decreased. Many surviving species seem to have considerable stability in abundance at the local scale. The density of smaller stems (10 29 cm gbh) increased by 15.3%, while that of the larger trees decreased drastically (81.6%). Ninety percent of the missing stems were from the middlestorey of the forest. Tree density changes among the three ecological guilds revealed a decrease in stem density and an increase in basal area in the lowerstorey; while the middlestorey exhibited a reverse trend. Family-wise, tree density changes revealed that the majority of families (67%) showed an increase in stem density. Long-term studies on tree population changes are essential to estimate tree mortality and recruitment rates, which will provide a greater insight in tropical forest dynamics.     

Primary forest dynamics in lowland dipterocarp forest at Danum Valley, Sabah, Malaysia, and the role of the understorey

Changes in species composition in two 4-ha plots of lowland dipterocarp rainforest at Danum, Sabah, were measured over ten years (1986-1996) for trees > or = 10 cm girth at breast height (gbh). Each included a lower-slope to ridge gradient. The period lay between two drought events of moderate intensity but the forest showed no large lasting responses, suggesting that its species were well adapted to this regime. Mortality and recruitment rates were not unusual in global or regional comparisons. The forest continued to aggrade from its relatively (for Sabah) low basal area in 1986 and, together with the very open upper canopy structure and an abundance of lianas, this suggests a forest in a late stage of recovery from a major disturbance, yet one continually affected by smaller recent setbacks. Mortality and recruitment rates were not related to population size in 1986, but across subplots recruitment was positively correlated with the density and basal area of small trees (10-< 50cm gbh) forming the dense understorey. Neither rate was related to topography. While species with larger mean gbh had greater relative growth rates (rgr) than smaller ones, subplot mean recruitment rates were correlated with rgr among small trees. Separating understorey species (typically the Euphorbiaceae) from the overstorey (Dipterocarpaceae) showed marked differences in change in mortality with increasing gbh: in the former it increased, in the latter it decreased. Forest processes are centred on this understorey quasi-stratum. The two replicate plots showed a high correspondence in the mortality, recruitment, population changes and growth rates of small trees for the 49 most abundant species in common to both. Overstorey species had higher rgrs than understorey ones, but both showed considerable ranges in mortality and recruitment rates. The supposed trade-off in traits, viz slower rgr, shade tolerance and lower population turnover in the understorey group versus faster potential growth rate, high light responsiveness and high turnover in the overstorey group, was only partly met, as some understorey species were also very dynamic. The forest at Danum, under such a disturbance-recovery regime, can be viewed as having a dynamic equilibrium in functional and structural terms. A second trade-off in shade-tolerance versus drought-tolerance is suggested for among the understorey species. A two-storey (or vertical component) model is proposed where the understorcy-overstorey species' ratio of small stems (currently 2:1) is maintained by a major feedback process. The understorey appears to be an important part of this forest, giving resilience against drought and protecting the overstorey saplings in the long term. This view could be valuable for understanding forest responses to climate change where drought frequency in Borneo is predicted to intensify in the coming decades.