Above-ground forest biomass is not consistently related to wood density in tropical forests

Aim  It is increasingly accepted that the mean wood density of trees within a forest is tightly coupled to above-ground forest biomass. It is unknown, however, if a positive relationship between forest biomass and mean community wood density is a general phenomenon across forests. Understanding spatial variation in biomass as a function of wood density both within and among forests is important for predicting changes in stored carbon in response to global change, and here we evaluated the generality of a positive biomass–wood density relationship within and among six tropical forests.
Location  Costa Rica, Panama, Puerto Rico and Ecuador.
Methods  Individual stem data, including diameter at breast height and spatial position, for six forest dynamics plots were merged with an extensive wood density database. Individual stem biomass values were calculated from these data using published statistical models. Total above ground biomass, total basal area and mean community wood density were also quantified across a range of subcommunity plot sizes within each forest.
Results  Among forests, biomass did not vary with mean community wood density. The relationship between subcommunity biomass and mean wood density within a forest varied from negative to null to positive depending on the size of subcommunities and forest identity. The direction of correlation was determined by the associated total basal area–mean wood density correlation, the slope of which increased strongly with whole forest mean wood density.
Main conclusions  There is no general relationship between forest biomass and wood density, and in some forests, stored carbon is highest where wood density is lowest. Our results suggest that declining wood density, due to global change, will result in decreased or increased stored carbon in forests with high or low mean wood density, respectively.     

The above-ground coarse wood productivity of 104 Neotropical forest plots

The net primary production of tropical forests and its partitioning between long‐lived carbon pools (wood) and shorter‐lived pools (leaves, fine roots) are of considerable importance in the global carbon cycle. However, these terms have only been studied at a handful of field sites, and with no consistent calculation methodology. Here we calculate above‐ground coarse wood carbon productivity for 104 forest plots in lowland New World humid tropical forests, using a consistent calculation methodology that incorporates corrections for spatial variations in tree‐size distributions and wood density, and for census interval length. Mean wood density is found to be lower in more productive forests. We estimate that above‐ground coarse wood productivity varies by more than a factor of three (between 1.5 and 5.5 Mg C ha^?1 a^?1) across the Neotropical plots, with a mean value of 3.1 Mg C ha^?1 a^?1. There appear to be no obvious relationships between wood productivity and rainfall, dry season length or sunshine, but there is some hint of increased productivity at lower temperatures. There is, however, also strong evidence for a positive relationship between wood productivity and soil fertility. Fertile soils tend to become more common towards the Andes and at slightly higher than average elevations, so the apparent temperature/productivity relationship is probably not a direct one. Coarse wood productivity accounts for only a fraction of overall tropical forest net primary productivity, but the available data indicate that it is approximately proportional to total above‐ground productivity. We speculate that the large variation in wood productivity is unlikely to directly imply an equivalent variation in gross primary production. Instead a shifting balance in carbon allocation between respiration, wood carbon and fine root production seems the more likely explanation.     

The spatial distribution of forest biomass in the Brazilian Amazon: a comparison of estimates

The amount of carbon released to the atmosphere as a result of deforestation is determined, in part, by the amount of carbon held in the biomass of the forests converted to other uses. Uncertainty in forest biomass is responsible for much of the uncertainty in current estimates of the flux of carbon from land‐use change. In the present contribution several estimates of forest biomass are compared for the Brazilian Amazon, based on spatial interpolations of direct measurements, relationships to climatic variables, and remote sensing data. Three questions were posed: First, do the methods yield similar estimates? Second, do they yield similar spatial patterns of distribution of biomass? And, third, what factors need most attention if we are to predict more accurately the distribution of forest biomass over large areas? The answer to the first two questions is that estimates of biomass for Brazil's Amazonian forests (including dead and belowground biomass) vary by more than a factor of two, from a low of 39 PgC to a high of 93 PgC. Furthermore, the estimates disagree as to the regions of high and low biomass. The lack of agreement among estimates confirms the need for reliable determination of aboveground biomass over large areas. Potential methods include direct measurement of biomass through forest inventories with improved allometric regression equations, dynamic modelling of forest recovery following observed stand‐replacing disturbances, and estimation of aboveground biomass from airborne or satellite‐based instruments sensitive to the vertical structure plant canopies.     

The regional variation of aboveground live biomass in old-growth Amazonian forests

The biomass of tropical forests plays an important role in the global carbon cycle, both as a dynamic reservoir of carbon, and as a source of carbon dioxide to the atmosphere in areas undergoing deforestation. However, the absolute magnitude and environmental determinants of tropical forest biomass are still poorly understood. Here, we present a new synthesis and interpolation of the basal area and aboveground live biomass of old‐growth lowland tropical forests across South America, based on data from 227 forest plots, many previously unpublished. Forest biomass was analyzed in terms of two uncorrelated factors: basal area and mean wood density. Basal area is strongly affected by local landscape factors, but is relatively invariant at regional scale in moist tropical forests, and declines significantly at the dry periphery of the forest zone. Mean wood density is inversely correlated with forest dynamics, being lower in the dynamic forests of western Amazonia and high in the slow‐growing forests of eastern Amazonia. The combination of these two factors results in biomass being highest in the moderately seasonal, slow growing forests of central Amazonia and the Guyanas (up to 350 Mg dry weight ha^?1) and declining to 200–250 Mg dry weight ha^?1 at the western, southern and eastern margins. Overall, we estimate the total aboveground live biomass of intact Amazonian rainforests (area 5.76 × 10⁶ km² in 2000) to be 93±23 Pg C, taking into account lianas and small trees. Including dead biomass and belowground biomass would increase this value by approximately 10% and 21%, respectively, but the spatial variation of these additional terms still needs to be quantified.     

Variation in above‐ground forest biomass across broad climatic gradients

Aim An understanding of the relationship between forest biomass and climate is needed to predict the impacts of climate change on carbon stores. Biomass patterns have been characterized at geographically or climatically restricted scales, making it unclear if biomass is limited by climate in any general way at continental to global scales. Using a dataset spanning multiple climatic regions we evaluate the generality of published biomass–climate correlations. We also combine metabolic theory and hydraulic limits to plant growth to first derive and then test predictions for how forest biomass should vary with maximum individual tree biomass and the ecosystem water deficit. Location Temperate forests and dry, moist and wet tropical forests across North, Central and South America. Methods A forest biomass model was derived from allometric functions and power‐law size distributions. Biomass and climate were correlated using extensive forest plot (276 0.1‐ha plots), wood density and climate datasets. Climate variables included mean annual temperature, annual precipitation, their ratio, precipitation of the driest quarter, potential and actual evapotranspiration, and the ecosystem water deficit. The water deficit uniquely summarizes water balance by integrating water inputs from precipitation with water losses due to solar energy. Results Climate generally explained little variation in forest biomass, and mixed support was found for published biomass–climate relationships. Our theory indicated that maximum individual biomass governs forest biomass and is constrained by water deficit. Indeed, forest biomass was tightly coupled to maximum individual biomass and the upper bound of maximum individual biomass declined steeply with water deficit. Water deficit similarly constrained the upper bound of forest biomass, with most forests below the constraint. Main conclusions The results suggest that: (1) biomass–climate models developed at restricted geographic/climatic scales may not hold at broader scales; (2) maximum individual biomass is strongly related to forest biomass, suggesting that process‐based models should focus on maximum individual biomass; (3) the ecosystem water deficit constrains biomass, but realized biomass often falls below the constraint; such that (4) biomass is not strongly limited by climate in most forests so that forest biomass may not predictably respond to changes in mean climate.     

The relationship between tree biodiversity and biomass dynamics changes with tropical forest succession

Theory predicts shifts in the magnitude and direction of biodiversity effects on ecosystem function (BEF) over succession, but this theory remains largely untested. We studied the relationship between aboveground tree biomass dynamics (Δbiomass) and multiple dimensions of biodiversity over 8–16 years in eight successional rainforests. We tested whether successional changes in diversity–Δbiomass correlations reflect predictions of niche theories. Diversity–Δbiomass correlations were positive early but weak later in succession, suggesting saturation of niche space with increasing diversity. Early in succession, phylogenetic diversity and functional diversity in two leaf traits exhibited the strongest positive correlations with Δbiomass, indicating complementarity or positive selection effects. In mid‐successional stands, high biodiversity was associated with greater mortality‐driven biomass loss, i.e. negative selection effects, suggesting successional niche trade‐offs and loss of fast‐growing pioneer species. Our results demonstrate that BEF relationships are dynamic across succession, thus successional context is essential to understanding BEF in a given system.     

Diversity in specific gravity and water content of wood among Bornean tropical rainforest trees

Wood properties were measured for trees in lowland dipterocarp forests in West Kalimantan. In 1993 and 1994, 353 samples of 286 species were collected from trunk base of trees of approximately 5 cm in diameter, and the specific gravities (SG: oven dry weight/fresh volume) and water contents of wood including bark were measured. The SG of each species ranged from 0.21 to 0.84, and the mean ± SD was 0.53 ± 0.13. The wide range of SG suggests that the forest had a high diversity in wood properties. The most dominant and diversified genus in this area was Shorea, and the SG of 15 species varied from 0.21 to 0.71. The range covered SG of pioneer (six Macaranga, 0.29–0.43) and small trees in primary forests (nine Eugenia and 10 Xanthophyllum, 0.55–0.77). The SG average for tree species of secondary forests of 2–6 years old was 0.31. It was significantly smaller than that of primary forests (0.58). In a primary dipterocarp forest plot, light-wood species grew faster in diameter than heavy-wood species. Water content ranged from 0.26 to 0.76. Heavy wood had low water content. Among light-wood species, some (Shorea, Artocarpus) had low water contents and others (Ficus) had high water contents. Some riverine trees also had high water contents. These wood properties appear strongly related to the life history of trees and successional stage.     

Effect of wood density and water permeability on wood decomposition rates of 32 Bornean rainforest trees

Aims A better understanding of wood litter decomposition is essential for predicting responses of forest ecosystems to global climate change. Recent studies suggest that chemical properties of wood litters, rather than physical ones such as wood density, are more important for interspecific differences in wood decomposition rates. However, empirical data are still limited, especially for tropical trees. In addition, decomposition rate of wood litter often varies with time, which makes interspecific comparison difficult. We studied the wood decomposition of 32 rainforest trees to elucidate (i) the degree of interspecific variation in wood decomposition rate of a given size and configuration and (ii) if initial wood density and water permeability are consistent predictors of the overall decomposition rate and its pattern over time.Methods A common garden decomposition experiment was conducted in a tropical rainforest in Malaysian Borneo for 32 native tree species. Small wood sticks were set on the forest floor and the weight loss was monitored monthly for 2.7 years.Important findings We found large variation in the wood decomposition rate (a 49-fold range), suggesting that we need to consider this variation when calculating community-level carbon dynamics of tropical rain forests. The physical traits of wood, i.e. wood density and water permeability, were related to wood decomposition rate and its pattern over time. Decomposition half-time related positively and negatively to initial wood density and water permeability, respectively. The time-dependent-rate model fitted better for 18 species (56% of the study species) that had higher water permeabilities than the others, suggesting that micelle porosity in wood relates to temporal changes in decomposition rate.     

Testing for changes in biomass dynamics in large‐scale forest datasets

Tropical forest responses to climate and atmospheric change are critical to the future of the global carbon budget. Recent studies have reported increases in estimated above‐ground biomass (EAGB) stocks, productivity, and mortality in old‐growth tropical forests. These increases could reflect a shift in forest functioning due to global change and/or long‐lasting recovery from past disturbance. We introduce a novel approach to disentangle the relative contributions of these mechanisms by decomposing changes in whole‐plot biomass fluxes into contributions from changes in the distribution of gap‐successional stages and changes in fluxes for a given stage. Using 30 years of forest dynamic data at Barro Colorado Island, Panama, we investigated temporal variation in EAGB fluxes as a function of initial EAGB (EAGB_i) in 10 × 10 m quadrats. Productivity and mortality fluxes both increased strongly with initial quadrat EAGB. The distribution of EAGB (and thus EAGB_i) across quadrats hardly varied over 30 years (and seven censuses). EAGB fluxes as a function of EAGB_i varied largely and significantly among census intervals, with notably higher productivity in 1985–1990 associated with recovery from the 1982–1983 El Niño event. Variation in whole‐plot fluxes among census intervals was explained overwhelmingly by variation in fluxes as a function of EAGB_i, with essentially no contribution from changes in EAGB_i distributions. The high observed temporal variation in productivity and mortality suggests that this forest is very sensitive to climate variability. There was no consistent long‐term trend in productivity, mortality, or biomass in this forest over 30 years, although the temporal variability in productivity and mortality was so strong that it could well mask a substantial trend. Accurate prediction of future tropical forest carbon budgets will require accounting for disturbance‐recovery dynamics and understanding temporal variability in productivity and mortality.     

西双版纳热带季节雨林粗木质物残体储量及其空间分布

粗木质物残体(CWD)在森林生态系统的物质循环和碳动态中发挥着重要作用,但目前国内对热带地区CWD储量与其空间分布关系上的研究还处于起步阶段。对西双版纳1hm2热带季节雨林样地调查发现,≥10cm的粗木质物残体占森林总凋落物量的68.02%,其中45.72%的CWD处于腐解中后期。由于多数大径级的CWD分布于15°～25°的坡地上,且其饱和持水力随腐解等级的增加而上升,因此,CWD将有助于改善坡地生境下的水肥条件。CWD的分布受风向和本地湿热环境的影响,但掉落指向则更倾向于随机过程。CWD产生和分布上的普遍性有可能对森林生境异质性和生物多样性产生重大影响。     

Roots,nutrients and their relationship to spatial patterns

Ecosystem sustainability and resilience after a disturbance may be regulated by processes occurring at smaller spatial scales. The matrix of different spatial environments are created by (1) individual plants that accumulate higher concentrations of specific nutrients, trace elements or defensive plant secondary chemicals and thereby modify the chemistry of their ecological space and/or rates of processes, (2) the presence of structures (e.g., coarse woody debris) that may buffer some micro-environments from disturbances by functioning as a hospitable environment or as a reservoir for mycorrhizal fungi to sustain them into the next phase of stand development, and (3) chemical changes in soils during soil development which may result in distinct soil chemical environments. The response of the plants or change in the sustainability of carbon and nutrient cycles may be expressed more strongly at this smaller ecological space of an individual plant and furthermore must be frequently examined separately by the above- and belowground space of that individual.This paper will present three case studies from temperate and tropical forest ecosystems which suggest the importance of studying plant growth and nutrient and trace element cycling by stratifying sampling to encompass the mosaic patterns of existing spatial variability within the ecosystem. The examples show how individual plant species are able to create ecologically distinct spatial environments because of their distribution patterns within the landscape, how nutrient transfers in roots respond to the chemical variations in the soil, and how roots and mycorrhizal fungi are able to maintain themselves in the mosaic of coarse woody debris remaining on a site after the elimination of aboveground tree biomass.     

Environmental correlates of tree biomass, basal area, wood specific gravity and stem density gradients in Borneo's tropical forests

Aim Tropical forests have been recognized as important global carbon sinks and sources. However, many uncertainties about the spatial distribution of live tree above‐ground biomass (AGB) remain, mostly due to limited availability of AGB field data. Recent studies in the Amazon have already shown the importance of large sample size for accurate AGB gradient analysis. Here we use a large stem density, basal area, community wood density and AGB dataset to study and explain their spatial patterns in an Asian tropical forest. Location Borneo, Southeast Asia. Methods We combined stem density, basal area, community wood density and AGB data from 83 locations in Borneo with an environmental database containing elevation, climate and soil variables. The Akaike information criterion was used to select models and environmental variables that best explained the observed values of stem density, basal area, community wood density and AGB. These models were used to extrapolate these parameters across Borneo. Results We found that wood density, stem density, basal area and AGB respond significantly, but differentially, to the environment. AGB was only correlated with basal area, but not with stem density and community wood specific gravity. Main conclusions Unlike results from Amazonian forests, soil fertility was an important positive correlate for AGB in Borneo while community wood density, which is a main driver of AGB in the Neotropics, did not correlate with AGB in Borneo. Also, Borneo's average AGB of 457.1 Mg ha^?1 was c. 60% higher than the Amazonian average of 288.6 Mg ha^?1. We find evidence that this difference might be partly explained by the high density of large wind‐dispersed Dipterocarpaceae in Borneo, which need to be tall and emergent to disperse their seeds. Our results emphasize the importance of Bornean forests as carbon sinks and sources due to their high carbon storage capacity.     

Species richness and wood production: a positive association in Mediterranean forests

A major debate in the study of biodiversity concerns its influence on ecosystem functioning. We compared whether wood production in forests was associated with tree functional group identity (i.e. deciduous, conifer or sclerophylous), tree species richness (1–≥ 5) and tree functional group richness (1–3) by comparing more than 5000 permanent plots distributed across Catalonia (NE Spain). Deciduous forests were more productive than coniferous and sclerophylous forests. Wood production increased with tree species richness. However, functional group richness increased wood production only in sclerophylous forests. When other forest structure, environmental variables and management practices were included in the analysis, tree functional group identity and species richness still remained significant, while functional species richness did not. Our survey indicates that across a regional scale, and across a broad range of environmental conditions, a significant positive association exists between local tree species richness and wood production at least in typical early successional Mediterranean-type forests.     

A seasonal evergreen forest in Belize: unusually high tree species richness for northern Central America

Most of the undisturbed forests of northern Central America remain undescribed. Some studies predict that tree species richness in lowland forests of northern Central America should be much lower than in similar forest types in southern Central America. This paper describes the physical and biological structure of two permanent, one-hectare plots on a valley floor in the Bladen Nature Reserve, Belize and compares these plots with other forests in the Neotropics. The plots have 91 and 89 species of trees ≥ 10 cm diameter (327 and 358 individuals, respectively) and comparable or higher measures of alpha-diversity than other such forests in Central America. The recorded tree species richness from other forests in northern Central America may be lower than those in southern Belize. The plots are dominated by Mortoniella pittieri Woodson (Apocynaceae), a disjunct from Costa Rica and Nicaragua. More than 50% of the tree species have distributions ranging into Bolivia or Peru, and >75% are wide-ranging species. The plots have a lower tree density and a higher proportion of large trees (>70 cm dbh) than other wet Neotropical forests. Like other Neotropical forests, tree species in the plots are generally aggregated and have a high proportion of vertebrate-dispersed fruits. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 138 , 275–296.     

Tree Mortality and Collecting Botanical Vouchers in Tropical Forests1

There is growing concern about the potential impact of reseatchers on tropical forest ecology, but few data. The aim of this paper is to examine the effects of collecting botanical specimens from tropical forest trees on their subsequent survivorship, using mortality data from plots in Amazonian Peru that were established in 1989 and reinventoried in 1994. In total, 2017 trees were originally tagged and collections were made from 948 trees. Making voucher collections always involved using unsterilized telescopic plant collecting poles to cut representative small branches, and sometimes also involved using iron-spiked tree-climbing gear to gain access to the canopy. Annual mortality in the four plots averaged 1.99 percent. Among the whole population of dicotyledenous trees, there was no detectable difference between the mortality rate of collected trees (1.96%) and noncollected trees (2.29%). We conclude that in spite of the physical damage caused to collected trees, collecting voucher specimens from tropical moist forest trees may not affect their survivorship, at least in the short-term. Further studies are needed to fully evaluate the potential impacts of research activities on petmanent forest plots in the tropics.     

Disentangling stand and environmental correlates of aboveground biomass in Amazonian forests

Tropical forests contain an important proportion of the carbon stored in terrestrial vegetation, but estimated aboveground biomass (AGB) in tropical forests varies two‐fold, with little consensus on the relative importance of climate, soil and forest structure in explaining spatial patterns. Here, we present analyses from a plot network designed to examine differences among contrasting forest habitats (terra firme, seasonally flooded, and white‐sand forests) that span the gradient of climate and soil conditions of the Amazon basin. We installed 0.5‐ha plots in 74 sites representing the three lowland forest habitats in both Loreto, Peru and French Guiana, and we integrated data describing climate, soil physical and chemical characteristics and stand variables, including local measures of wood specific gravity (WSG). We use a hierarchical model to separate the contributions of stand variables from climate and soil variables in explaining spatial variation in AGB. AGB differed among both habitats and regions, varying from 78 Mg ha^?1 in white‐sand forest in Peru to 605 Mg ha^?1 in terra firme clay forest of French Guiana. Stand variables including tree size and basal area, and to a lesser extent WSG, were strong predictors of spatial variation in AGB. In contrast, soil and climate variables explained little overall variation in AGB, though they did co‐vary to a limited extent with stand parameters that explained AGB. Our results suggest that positive feedbacks in forest structure and turnover control AGB in Amazonian forests, with richer soils (Peruvian terra firme and all seasonally flooded habitats) supporting smaller trees with lower wood density and moderate soils (French Guianan terra firme) supporting many larger trees with high wood density. The weak direct relationships we observed between soil and climate variables and AGB suggest that the most appropriate approaches to landscape scale modeling of AGB in the Amazon would be based on remote sensing methods to map stand structure.     

Tree Height Reduction After Selective Logging in a Tropical Forest

By harvesting scattered large trees, selective logging increases light availability and thereby stimulates growth and crown expansion at early‐life stage among remnant trees. We assessed the effects of logging on total and merchantable bole (i.e., lowest branch at crown base) heights on 952 tropical canopy trees in French Guiana. We observed reductions in both total (mean, ?2.3 m) and bole (mean, ?2.0 m) heights more than a decade after selective logging. Depending on local logging intensity, height reductions resulted in 2–13 percent decreases in aboveground tree biomass and 3–17 percent decreases in bole volume. These results highlight the adverse effects of logging at both tree and stand levels. This decrease in height is a further threat to future provision of key environmental services, such as timber production and carbon sequestration.