Understanding the processes that determine above-ground biomass (AGB) in Amazonian forests is important for predicting the sensitivity of these ecosystems to environmental change and for designing and evaluating dynamic global vegetation models (DGVMs). AGB is determined by inputs from woody productivity [woody net primary productivity (NPP)] and the rate at which carbon is lost through tree mortality. Here, we test whether two direct metrics of tree mortality (the absolute rate of woody biomass loss and the rate of stem mortality) and/or woody NPP, control variation in AGB among 167 plots in intact forest across Amazonia. We then compare these relationships and the observed variation in AGB and woody NPP with the predictions of four DGVMs. The observations show that stem mortality rates, rather than absolute rates of woody biomass loss, are the most important predictor of AGB, which is consistent with the importance of stand size structure for determining spatial variation in AGB. The relationship between stem mortality rates and AGB varies among different regions of Amazonia, indicating that variation in wood density and height/diameter relationships also influences AGB. In contrast to previous findings, we find that woody NPP is not correlated with stem mortality rates and is weakly positively correlated with AGB. Across the four models, basinwide average AGB is similar to the mean of the observations. However, the models consistently overestimate woody NPP and poorly represent the spatial patterns of both AGB and woody NPP estimated using plot data. In marked contrast to the observations, DGVMs typically show strong positive relationships between woody NPP and AGB. Resolving these differences will require incorporating forest size structure, mechanistic models of stem mortality and variation in functional composition in DGVMs.
Tropical tree height-diameter (<i>H:D</i>) relationships may vary by forest type and region making large-scale estimates of above-ground biomass subject to bias if they ignore these differences in stem allometry. We have therefore developed a new global tropical forest database consisting of 39 955 concurrent <i>H</i> and <i>D</i> measurements encompassing 283 sites in 22 tropical countries. Utilising this database, our objectives were: <br><br> 1. to determine if <i>H:D</i> relationships differ by geographic region and forest type (wet to dry forests, including zones of tension where forest and savanna overlap).<br><br> 2. to ascertain if the <i>H:D</i> relationship is modulated by climate and/or forest structural characteristics (e.g. stand-level basal area, <i>A</i>).<br><br> 3. to develop <i>H:D</i> allometric equations and evaluate biases to reduce error in future local-to-global estimates of tropical forest biomass. <br><br> Annual precipitation coefficient of variation (<i>P</i><sub>V</sub>), dry season length (<i>S</i><sub>D</sub>), and mean annual air temperature (<i>T</i><sub>A</sub>) emerged as key drivers of variation in <i>H:D</i> relationships at the pantropical and region scales. Vegetation structure also played a role with trees in forests of a high <i>A</i> being, on average, taller at any given <i>D</i>. After the effects of environment and forest structure are taken into account, two main regional groups can be identified. Forests in Asia, Africa and the Guyana Shield all have, on average, similar <i>H:D</i> relationships, but with trees in the forests of much of the Amazon Basin and tropical Australia typically being shorter at any given <i>D</i> than their counterparts elsewhere. <br><br> The region-environment-structure model with the lowest Akaike's information criterion and lowest deviation estimated stand-level <i>H</i> across all plots to within a median –2.7 to 0.9% of the true value. Some of the plot-to-plot variability in <i>H:D</i> relationships not accounted for by this model could be attributed to variations in soil physical conditions. Other things being equal, trees tend to be more slender in the absence of soil physical constraints, especially at smaller <i>D</i>. Pantropical and continental-level models provided only poor estimates of <i>H</i>, especially when the roles of climate and stand structure in modulating <i>H:D</i> allometry were not simultaneously taken into account
Abstract. Through interpretations of remote-sensing data and/or theoretical propositions, the idea that forest and savanna represent "alternative stable states" is gaining increasing acceptance. Filling an observational gap, we present detailed stratified floristic and structural analyses for forest and savanna stands located mostly within zones of transition (where both vegetation types occur in close proximity) in Africa, South America and Australia. Woody plant leaf area index variation was related to tree canopy cover in a similar way for both savanna and forest with substantial overlap between the two vegetation types. As total woody plant canopy cover increased, so did the relative contribution of middle and lower strata of woody vegetation. Herbaceous layer cover declined as woody cover increased. This pattern of understorey grasses and herbs progressively replaced by shrubs as the canopy closes over was found for both savanna and forests and on all continents. Thus, once subordinate woody canopy layers are taken into account, a less marked transition in woody plant cover across the savanna–forest-species discontinuum is observed compared to that inferred when trees of a basal diameter > 0.1 m are considered in isolation. This is especially the case for shrub-dominated savannas and in taller savannas approaching canopy closure. An increased contribution of forest species to the total subordinate cover is also observed as savanna stand canopy closure occurs. Despite similarities in canopy-cover characteristics, woody vegetation in Africa and Australia attained greater heights and stored a greater amount of above-ground biomass than in South America. Up to three times as much above-ground biomass is stored in forests compared to savannas under equivalent climatic conditions. Savanna–forest transition zones were also found to typically occur at higher precipitation regimes for South America than for Africa. Nevertheless, consistent across all three continents coexistence was found to be confined to a well-defined edaphic–climate envelope with soil and climate the key determinants of the relative location of forest and savanna stands. Moreover, when considered in conjunction with the appropriate water availability metrics, it emerges that soil exchangeable cations exert considerable control on woody canopy-cover extent as measured in our pan-continental (forest + savanna) data set. Taken together these observations do not lend support to the notion of alternate stable states mediated through fire feedbacks as the prime force shaping the distribution of the two dominant vegetation types of the tropical lands.
Above-ground tropical tree biomass and carbon storage estimates commonly ignore tree height. We estimate the effect of incorporating height (<i>H</i>) on forest biomass estimates using 37 625 concomitant <i>H</i> and diameter measurements (<i>n</i> = 327 plots) and 1816 harvested trees (<i>n</i> = 21 plots) tropics-wide to answer the following questions: <br><br> 1. For trees of known biomass (from destructive harvests) which <i>H</i>-model form and geographic scale (plot, region, and continent) most reduces biomass estimate uncertainty? <br><br> 2. How much does including <i>H</i> relationship estimates derived in (1) reduce uncertainty in biomass estimates across 327 plots spanning four continents? <br><br> 3. What effect does the inclusion of <i>H</i> in biomass estimates have on plot- and continental-scale forest biomass estimates? <br><br> The mean relative error in biomass estimates of the destructively harvested trees was half (mean 0.06) when including <i>H</i>, compared to excluding <i>H</i> (mean 0.13). The power- and Weibull-<i>H</i> asymptotic model provided the greatest reduction in uncertainty, with the regional Weibull-<i>H</i> model preferred because it reduces uncertainty in smaller-diameter classes that contain the bulk of biomass per hectare in most forests. Propagating the relationships from destructively harvested tree biomass to each of the 327 plots from across the tropics shows errors are reduced from 41.8 Mg ha<sup>−1</sup> (range 6.6 to 112.4) to 8.0 Mg ha<sup>−1</sup> (−2.5 to 23.0) when including $H$. For all plots, above-ground live biomass was 52.2±17.3 Mg ha<sup>−1</sup> lower when including <i>H</i> estimates (13%), with the greatest reductions in estimated biomass in Brazilian Shield forests and relatively no change in the Guyana Shield, central Africa and southeast Asia. We show fundamentally different stand structure across the four forested tropical continents, which affects biomass reductions due to $H$. African forests store a greater portion of total biomass in large-diameter trees and trees are on average larger in diameter. This contrasts to forests on all other continents where smaller-diameter trees contain the greatest fractions of total biomass. After accounting for variation in $H$, total biomass per hectare is greatest in Australia, the Guyana Shield, and Asia and lowest in W. Africa, W. Amazonia, and the Brazilian Shield (descending order). Thus, if closed canopy tropical forests span 1668 million km<sup>2</sup> and store 285 Pg C, then the overestimate is 35 Pg C if <i>H</i> is ignored, and the sampled plots are an unbiased statistical representation of all tropical forest in terms of biomass and height factors. Our results show that tree $H$ is an important allometric factor that needs to be included in future forest...
The study was carried out on the gallery forest of the Bacaba stream situated in the Municipal Ecological Reserve ‘Mário Viana’ (14°43′S, 52°21′W) in Nova Xavantina, Eastern Mato Grosso, Brazil. Three sections of the gallery (upper, middle and lower) running downstream and differing in slope were surveyed by stratified sampling. Fortyseven nested 10m × 10m plots were analysed in each section, giving a total sampling area of 1.41ha overall. All trees or lianas ≥ 15cm girth at breast height were recorded and a total of 129 species belonging to 105 genera and 47 families were found. Diversity was high, with the Shannon index ranging from 3.84 nats/individual in the lower section to 4.08 in the middle section. The most important families (IVI) were Caesalpiniaceae (upper and middle sections) and Arecaceae (lower section), and the most important species were Diospyros obovata (upper section), Hymenaea courbaril var. stilbocarpa (middle section) and Mauritia flexuosa (lower section). Morisita and Sørensen indices of similarity were calculated. The floristic composition was complex and included species in common with a number of Brazilian forest types and with cerrado (savanna), as well as many widespread species, but stronger links with Amazonian forests could be detected. This is to be expected since the area lies in the ecotonal zone of the cerrado and Amazonian forest biomes and the Bacaba stream itself is a tributary of the Mortes–Araguaia–Amazon river system.
Abstract. Sampling along a precipitation gradient in tropical America extending from ca. 0.8 to 2.0 m a−1, savanna soils had consistently lower exchangeable cation concentrations and higher C/N ratios than nearby forest plots. These soil differences were also reflected in canopy averaged leaf traits with savanna trees typically having higher leaf mass per unit area but lower mass-based nitrogen (Nm) and potassium (Km). Both Nm and Km also increased with declining mean annual precipitation (PA), but most area-based leaf traits such as leaf photosynthetic capacity showed no systematic variation with PA or vegetation type. Despite this invariance, when taken in conjunction with other measures such mean canopy height, area-based soil exchangeable potassium content, [K]sa, proved to be an excellent predictor of several photosynthetic properties (including 13C isotope discrimination). Moreover, when considered in a multivariate context with PA and soil plant available water storage capacity (θP) as covariates, [K]sa also proved to be an excellent predictor of stand-level canopy area, providing drastically improved fits as compared to models considering just PA and/or θP. Neither calcium, magnesium nor soil pH could substitute for potassium when tested as alternative model predictors (ΔAIC > 10). Nor for any model could simple soil texture metrics such as sand or clay content substitute for either [K]sa or θP. Taken in conjunction with recent work in Africa and the forests of the Amazon Basin this suggests – in combination with some newly conceptualised interacting effects of PA and θP also presented here – a critical role for potassium as a modulator of tropical vegetation structure and function.
The occurrence of cerrado (as tree and shrub savanna is called in Brazil) and forest formations side by side is common at the southern margin of the Brazilian Amazonian Forest, and previous studies have demonstrated the advance of forests over cerrado areas. The aim of the present study is to provide an accurate documentation of the transition process between the two major biomes. Tree data (≥ 5 cm diameter at 0.3 m above soil level) from three plots of cerrado sensu stricto lying near three of cerradão (the taller, denser form of cerrado) were inventoried starting in 2002 in an area of 1.5 ha made up of 150 subplots of 10 × 10 m (50 in each area). This showed that the most important species of the cerradão were invading areas previously occupied by smaller, lower forms of cerrado (although it is sometimes difficult to define which are ‘forest’ and which ‘cerrado’ species as many are flexible in size – for instance Emmotum nitens can often be intermediate, establishing in cerrado that develops into cerradão and on to forest). Some typical species such as Eriotheca gracilipes and Emmotum nitens, established since the first inventories, have increased their populations (between 27 and 210%). Tachigali vulgaris, a typical, weedy, adventive species of the Cerrado–Amazonian Forest transition, showed the largest increase in abundance in areas of cerrado sensu stricto (between 100 and 1200%), and is probably the most important pioneer species in the initial advance of the forest into cerrado at the Southern Amazonian border.
In the Cerrado-Amazon ecotone in central Brazil, recent studies suggest some encroachment of forest into savanna, but how, where, and why this might be occurring is unclear. To better understand this phenomenon, we assessed changes in the structure and dynamics of tree species in three vegetation types at the Cerrado-Amazon ecotone that are potentially susceptible to encroachment: open cerrado (OC), typical cerrado (TC) and dense woodland (DW). We estimated changes in density, basal area and aboveground biomass of trees with diameter >10 cm over four inventories carried out between 2008 and 2015 and classified the species according to their preferred habitat (savanna, generalist, or forest). There was an increase in all structural parameters assessed in all vegetation types, with recruitment and gains in basal area and biomass greater than mortality and losses. Thus, there were net gains between the first and final inventories in density (OC: 3.
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