Effect of inventory plot patterns in the floristic analysis of tropical woodland and dense forest

This study was set up to examine the effect of plot patterns on the accuracy of phytosociological characterization of tropical vegetation. Fifteen and twenty square plots of 1 ha were demarcated, respectively, in woodland and dense forest in Bénin. Each 1 ha plot was divided into 100 quadrats of one 100 m². Species of trees in each quadrat were identified and recorded. The cost in terms of time required to record tree species in each 1 ha plot and five random quadrats in a 1 ha plot were also recorded to compute the mean inventory effort for a team of three foresters. From the 100 quadrats in a 1 ha plot, fourteen independent subplots of square and rectangular plots with different sizes were considered by grouping together adjacent quadrats of 100 m². Eigenanalysis was carried out to compare the subplots. Moreover, the relationship between the relative loss of accuracy (RLA) and the size of subplots was modelled. Plot size highly influenced the RLA (P < 0.05). Findings indicated that the square plots of 1500 and 1000 m² with an inventory effort of 0.35 and 0.20 man‐days per subplot, respectively in tropical dense forests and woodlands appeared to be the most efficient in the phytosociological characterization of woody vegetation.     

Soil is the main predictor of secondary rain forest estimated aboveground biomass across a Neotropical landscape

We studied the relative effects of landscape configuration, environmental variables, forest age, and spatial variables on estimated aboveground biomass (AGB) in Costa Rican secondary rain forests patches. We measured trees ≥5 cm dbh in 24, 0.25 ha plots and estimated AGB for trees 5–24.9 cm dbh and for trees >25 cm dbh using two allometric equations based on multispecies models using tree dbh and wood‐specific gravity. AGB averaged 87.3 Mg/ha for the 24 plots (not including remnant trees) and 123.4 Mg/ha including remnant trees (20 plots). There was no effect of forest age on AGB. Variation partitioning analysis showed that soils, climate, landscape configuration, and space together explained 61% of tree AGB variance. When controlling for the effects of the other three variables, only soils remained significant. Soil properties, specifically K and Cu, had the strongest independent effect on AGB (variation partitioning, R² = 0.17, p = 0.0310), indicating that in this landscape, AGB variation in secondary forest patches is influenced by soil chemical properties. Elucidating the relative influence of soils in AGB variation is critical for understanding changes associated with land cover modification across Neotropical landscapes, as it could have important consequences for land use planning since secondary forests are considered carbon sinks. Abstract in Spanish is available with online material.     

Above-ground biomass and vegetation attributes in the forest-savannah mosaic of Togo,West Africa

Despite the increasing interest in the role of African savannah and woodlands on the global carbon cycle, little is known about the above-ground biomass (AGB) and the factors affecting it in these ecosystems in West Africa. We estimated AGB in different vegetation types of a forest–savannah mosaic in Togo, and we investigated the relationship between AGB, structural and diversity attributes. We also assessed the effects of using the ≥5 or ≥10 cm diameter threshold on AGB estimates. We sampled tree diameter, height and species of all trees ≥5 cm diameter following standardised protocols in 160 plots of 50 × 20 m (50 × 10 m for riparian). Above-ground biomass (AGB) (all trees ≥5 cm diameter) ranged from 6.2 Mg/ha in shrub savannah to 292 Mg/ha in riparian forest and showed significant differences between vegetation types. Differences in AGB were related to structural attributes, with little influence of diversity attributes. The effects of minimum tree diameter size (5 or 10 cm) on AGB estimates were negligible. At a landscape level, closed-canopy and open forests stored important quantities of carbon. We highlight the importance of the forest–savannah mosaic as a large carbon pool, which could be released if converted to another land cover type.     

Estimation of aboveground biomass and net biomass increment in a cool temperate forest on a landscape scale

I clarified aboveground biomass (AGB), net biomass increment (NBI) and its spatial heterogeneity in a cool temperate forest on a landscape scale (>2,200 ha). The relationships among AGB, NBI, and the size frequency distribution of trees of each stand were examined by combining an analysis of vegetation using aerial photographs, and data from 146 inventory plots (28.8 ha in total). This area included natural broad-leaved stands, harvested broad-leaved stands, and artificial conifer plantations. A –3/2 power distribution density function was applied to the individual mass frequency distribution of each plot. Estimated AGB in carbon (C) equivalent was 480–5,615 g C m^–2 (3,130 g C m^–2 on average), and NBI was –98 to 436 g C m^–2 year^–1 (83.0 g C m^–2 year^–1 on average). NBI had a single significant relationship with the reciprocal of theoretical maximum individual mass, while NBI was not significantly related to AGB. My results showed that, on a landscape scale, AGB and NBI strongly depend on the size structure of forest stands.     

Effects of population density on forest structure and species richness and diversity of trees in Kampong Thom Province, Cambodia

This study examined differences in stand structure, tree species richness, and tree species diversity in relation to population density in Kampong Thom Province, Cambodia. Tree data were obtained from a 1997 forest inventory involving 60 clusters (540 plots) systematically distributed over 30% of the provincial forest area. Spatially referenced population data were obtained from the 1998 national population census. The average number of trees per cluster was 356/ha, the average basal area, 23 m²/ha, the average stand volume, 217 m³/ha, and the average aboveground biomass, 273 Mg/ha for all trees of DBH 10 cm and larger. The average species richness per cluster was 37 species, while average species diversity was measured as 0.916 using Simpson’s index and 2.98 by Shannon’s index. Significant negative correlations were generally found between population density surrounding clusters and tree density, basal area, stand volume, aboveground biomass, and species richness and diversity for three examined diameter classes (DBH of 10–30, ≥30, and ≥10 cm). As the distance from clusters for calculating population density increased, the correlation levels increased up to 5 or 7 km, depending on the variables and diameter class, and then stayed relatively constant for stand structure variables and decreased for species richness and diversity. The results indicate that evidence of disturbance was more pronounced at higher population density up to around 5 to 7 km. We suggest that introduction of greater controls on human disturbance should be a high priority for resource management and conservation in Kampong Thom Province and, presumably, Cambodia as a whole.     

Seasonally flooded,and terra firme in northern Congo: Insights on their structure,diversity and biomass

In the Congo basin, considerable uncertainty remains about the amount and spatial variation of carbon stocks. We studied two types of seasonally flooded forests (dominated by Guibourtia demeusei and Lophira alata) and nearby terra firme forests in northern Congo. We sampled 1.25 ha per forest type and a total of 1,400 trees ≥5 cm diameter. AGB ranged from 207–343 Mg/ha, with no significant differences between forest types. Few significant differences were observed in vegetation structure or tree diversity between forest types. Species richness and stem density of small trees were lower, and dominance was higher in Guibourtia plots, which are subject to greater flooding than Lophira plots. Guibourtia was absent from smaller diameter class in Guibourtia forests; and Uapaca spp. were more abundant in terra firme than in seasonally flooded plots. We show that both types of seasonally flooded forests store important quantities of AGB and should also be considered in forest conservation programmes. We recommend more research on seasonally flooded forests, on larger geographical extent, which assesses flood depth and duration, and measures tree height in the field, as we took a conservative approach to AGB estimates, and AGB could be even greater than we report here.     

Error propagation and scaling for tropical forest biomass estimates

The above-ground biomass (AGB) of tropical forests is a crucial variable for ecologists, biogeochemists, foresters and policymakers. Tree inventories are an efficient way of assessing forest carbon stocks and emissions to the atmosphere during deforestation. To make correct inferences about long-term changes in biomass stocks, it is essential to know the uncertainty associated with AGB estimates, yet this uncertainty is rarely evaluated carefully. Here, we quantify four types of uncertainty that could lead to statistical error in AGB estimates: (i) error due to tree measurement; (ii) error due to the choice of an allometric model relating AGB to other tree dimensions; (iii) sampling uncertainty, related to the size of the study plot; (iv) representativeness of a network of small plots across a vast forest landscape. In previous studies, these sources of error were reported but rarely integrated into a consistent framework. We estimate all four terms in a 50 hectare (ha, where 1 ha = 10(4) m2) plot on Barro Colorado Island, Panama, and in a network of 1 ha plots scattered across central Panama. We find that the most important source of error is currently related to the choice of the allometric model. More work should be devoted to improving the predictive power of allometric models for biomass.     

Tree allometry and improved estimation of carbon stocks and balance in tropical forests

Tropical forests hold large stores of carbon, yet uncertainty remains regarding their quantitative contribution to the global carbon cycle. One approach to quantifying carbon biomass stores consists in inferring changes from long-term forest inventory plots. Regression models are used to convert inventory data into an estimate of aboveground biomass (AGB). We provide a critical reassessment of the quality and the robustness of these models across tropical forest types, using a large dataset of 2,410 trees ≥ 5 cm diameter, directly harvested in 27 study sites across the tropics. Proportional relationships between aboveground biomass and the product of wood density, trunk cross-sectional area, and total height are constructed. We also develop a regression model involving wood density and stem diameter only. Our models were tested for secondary and old-growth forests, for dry, moist and wet forests, for lowland and montane forests, and for mangrove forests. The most important predictors of AGB of a tree were, in decreasing order of importance, its trunk diameter, wood specific gravity, total height, and forest type (dry, moist, or wet). Overestimates prevailed, giving a bias of 0.5–6.5% when errors were averaged across all stands. Our regression models can be used reliably to predict aboveground tree biomass across a broad range of tropical forests. Because they are based on an unprecedented dataset, these models should improve the quality of tropical biomass estimates, and bring consensus about the contribution of the tropical forest biome and tropical deforestation to the global carbon cycle. Electronic Supplementary Material Supplementary material is available for this article at     

Investigating above-ground biomass in old-growth and secondary montane forests of the Cameroon Highlands

Tropical montane forests can store and sequester substantial amounts of carbon in above-ground biomass (AGB), but variations in this storage related to location or degradation have not been quantified in the Cameroon Highlands. We established 25 permanent plots (20 m × 40 m) and sampled all trees ≥10 cm diameter following standard RAINFOR protocols. We estimated AGB and investigated variations related to taxonomic and structural forest attributes, including the height–diameter allometry in five forest types (four old-growth dominated by different species and one secondary forest). Secondary forests had significantly lower AGB than old-growth forests (49.4 ± 2.5 vs. >476.3 ± 168.7 Mg/ha, respectively), mostly related to lower basal area and tree height. Significant differences in species composition but not in forest structure or AGB were found between the four types of old-growth forests studied, located at different altitudes and mountains. We discuss the importance of these montane forests for carbon storage and, considering their high diversity and current threats, their potential for carbon finance mechanisms related to both avoided deforestation and forest restoration.     

Accounting for tree spatial distribution in a comparison of plot sizes and shapes in dense forest and woodland in Benin (West Africa)

The study examined simultaneously, the effect of tree spatial distribution, inventory plot size and shape on the estimation error of basal area in two contrasting environments. Twenty and fifteen square plots of 1 ha each (divided into 100 quadrats of 0.01 ha) were randomly set in dense forest and woodland, respectively. Thirteen subplots of various shapes and sizes were obtained from the association of adjacent quadrats. Estimation error was calculated using residual mean square of one‐way ANOVA, based on replications of subplot within 1 ha plots. Tree spatial distribution was measured using Green index. Weighted linear regression and mixed effect models were applied to Box & Cox transformed data. In general, the estimation error of basal area decreased with increase in subplot size. However, the effects of tree spatial distribution and plot shape varied with the vegetation type. Where trees tended to be aggregated, estimation error increased with degree of aggregation, and rectangular plots of 0.24 ha produced an acceptable precision. It was concluded that 0.24 ha rectangular plots can be used in tropical environments where the target parameters vary constantly according to one direction, while square plots of the same size are optimal for reliable analysis in case of randomness.     

Mass‐ratio and complementarity effects simultaneously drive aboveground biomass in temperate Quercus forests through stand structure

Forests play a key role in regulating the global carbon cycle, a substantial portion of which is stored in aboveground biomass (AGB). It is well understood that biodiversity can increase the biomass through complementarity and mass‐ratio effects, and the contribution of environmental factors and stand structure attributes to AGB was also observed. However, the relative influence of these factors in determining the AGB of Quercus forests remains poorly understood. Using a large dataset retrieved from 523 permanent forest inventory plots across Northeast China, we examined the effects of integrated multiple tree species diversity components (i.e., species richness, functional, and phylogenetic diversity), functional traits composition, environmental factors (climate and soil), stand age, and structure attributes (stand density, tree size diversity) on AGB based on structural equation models. We found that species richness and phylogenetic diversity both were not correlated with AGB. However, functional diversity positively affected AGB via an indirect effect in line with the complementarity effect. Moreover, the community‐weighted mean of specific leaf area and height increased AGB directly and indirectly, respectively; demonstrating the mass‐ratio effect. Furthermore, stand age, density, and tree size diversity were more important modulators of AGB than biodiversity. Our study highlights that biodiversity–AGB interaction is dependent on the regulation of stand structure that can be even more important for maintaining high biomass than biodiversity in temperate Quercus forests.     

Insights into regional patterns of Amazonian forest structure,diversity, and dominance from three large terra-firme forest dynamics plots

We analyze forest structure, diversity, and dominance in three large-scale Amazonian forest dynamics plots located in Northwestern (Yasuni and Amacayacu) and central (Manaus) Amazonia, to evaluate their consistency with prevailing wisdom regarding geographic variation and the shape of species abundance distributions, and to assess the robustness of among-site patterns to plot area, minimum tree size, and treatment of morphospecies. We utilized data for 441,088 trees (DBH ≥1 cm) in three 25-ha forest dynamics plots. Manaus had significantly higher biomass and mean wood density than Yasuni and Amacayacu. At the 1-ha scale, species richness averaged 649 for trees ≥1 cm DBH, and was lower in Amacayacu than in Manaus or Yasuni; however, at the 25-ha scale the rankings shifted, with Yasuni < Amacayacu < Manaus. Within each site, Fisher’s alpha initially increased with plot area to 1–10 ha, and then showed divergent patterns at larger areas depending on the site and minimum size. Abundance distributions were better fit by lognormal than by logseries distributions. Results were robust to the treatment of morphospecies. Overall, regional patterns in Amazonian tree species diversity vary with the spatial scale of analysis and the minimum tree size. The minimum area to capture local diversity is 2 ha for trees ≥1 cm DBH, or 10 ha for trees ≥10 cm DBH. The underlying species abundance distribution for Amazonian tree communities is lognormal, consistent with the idea that the rarest species have not yet been sampled. Enhanced sampling intensity is needed to fill the still large voids we have in plant diversity in Amazon forests.     

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.     

Woody plant communities of isolated Afromontane cloud forests in Taita Hills,Kenya

In the Taita Hills in southern Kenya, remnants of the original Afromontane forest vegetation are restricted to isolated mountain peaks. To assess the level of degradation and the need for forest restoration, we examined how forest plant communities and their indicator species vary between and within remnant patches of cloud forest. We used ordinal abundance data to compare plant communities in eight forest fragments. We also analyzed data on the diversity and abundance of trees in 57 0.1 ha plots to compare tree communities within and between the largest two of these fragments, Ngangao (120 ha) and Mbololo (220 ha). The extant vegetation of the Taita Hills at landscape scale consists of secondary moist montane to intermediate montane forest. There was a high species dissimilarity between fragments (69%). Variation in species composition coincided with an abiotic gradient related to elevation. At plot level, secondary successional species and species of forest edges were most abundant and most frequent. Inferred clusters of plots almost entirely coincided with the two forest fragments. Indicator species associated with forest margins and gaps were more frequent in the smaller of the two forest fragments, while indicators for the larger fragment were more typical for less disturbed moist forest. Abiotic site variability but also different levels of disturbance determine site-specific variants of the montane forest. Conservation efforts should not only focus on maintaining forest quantity (size), but also on forest quality (species composition). Late-successional rainforest species are underrepresented in the woody plant communities of the Taita Hills and assisting restoration of viable populations of cloud forest climax tree species is urgently needed.     

基于TM影像、森林资源清查数据和人工神经网络的森林碳空间分布模拟

森林是陆地生态系统中最大的碳库,在全球碳平衡和减缓全球气候变化方面发挥着不可替代的作用。当前主要利用森林资源清查数据和优势树种材积源-生物量的关系进行碳储量估算,在此基础上有效结合遥感影像数据将会更好的满足相关部门对国家和区域森林碳储量计算的需求。利用临安市2004年森林资源清查的930个样地数据和同年度Landsat TM影像数据,提取6个波段灰度值以及与碳储量相关性相对较大的3个波段组合,结合人工神经网络对研究区森林碳储量及其分布进行有效模拟。结果显示,用误差反向传播算法训练神经网络较好的重建了森林碳密度空间分布和变化,森林碳地上部分模拟结果与样地实测值之间的一致性好,全区域模拟结果森林碳平均值为0.98Mg(10.89Mg/hm²),总体森林碳密度模拟结果低于样地平均值约13%,进一步验证了人工神经网络在对大范围森林碳估算与模拟上具有较好的效果,为区域森林碳储量的估测研究提供有效的方法支持。     

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.     

Forest biomass stocks and dynamics across the subtropical Andes

Forest biomass plays an important role in the global carbon cycle. Therefore, understanding the factors that control forest biomass stocks and dynamics is a key challenge in the context of global change. We analyzed data from 60 forest plots in the subtropical Andes (22–27.5° S and 300–2300 m asl) to describe patterns and identify drivers of aboveground biomass (AGB) stocks and dynamics. We found that AGB stocks remained roughly constant with elevation due to compensating changes in basal area (which increased with elevation) and plot‐mean wood specific gravity (which decreased with elevation). AGB gain and loss rates both decreased with elevation and were explained mainly by temperature and rainfall (positive effects on both AGB gains and losses). AGB gain was also correlated with forest‐use history and weakly correlated with forest structure. Mean annual temperature and rainfall showed minor effects on AGB stocks and AGB change (gains minus losses) over recent decades. Although AGB change was only weakly correlated with climate variables, increases in AGB gains and losses with increasing rainfall—together with observed increases in rainfall in the subtropical Andes—suggest that these forests may become increasingly dynamic in the future. Abstract in Spanish is available with online material     

Field observed relationships between biodiversity and ecosystem functioning during secondary succession in a tropical lowland rainforest

The relationship between biodiversity and ecosystem functioning (BEF) is one of the most concerned topics in ecology. However, most of the studies have been conducted in controlled experiments in grasslands, few observational field studies have been carried out in forests. In this paper, we report variations of species diversity, functional diversity and aboveground biomass (AGB) for woody plants (trees and shrubs) along a chronosequence of four successional stages (18-year-old fallow, 30-year-old fallow, 60-year-old fallow, and old-growth forest) in a tropical lowland rainforest recovered after shifting cultivation on Hainan Island, China. Fifty randomly selected sample plots of 20 m × 20 m were investigated in each of the four successional stages. Four functional traits (specific leaf area, wood density, maximum species height and leaf dry matter content) were measured for each woody plants species and the relationships between species/functional diversity and AGB during secondary succession were explored. The results showed that both plant diversity and AGB recovered gradually with the secondary succession. AGB was positively correlated with both species and functional diversity in each stage of succession. Consistent with many controlled experimental results in grasslands, our observational field study confirms that ecosystem functioning is closely related to biodiversity during secondary succession in species rich tropical forests.     

How drought events during the last century have impacted biomass carbon in Amazonian rainforests

During the last two decades, inventory data show that droughts have reduced biomass carbon sink of the Amazon forest by causing mortality to exceed growth. However, process-based models have struggled to include drought-induced responses of growth and mortality and have not been evaluated against plot data. A process-based model, ORCHIDEE-CAN-NHA, including forest demography with tree cohorts, plant hydraulic architecture and drought-induced tree mortality, was applied over Amazonia rainforests forced by gridded climate fields and rising CO₂ from 1901 to 2019. The model reproduced the decelerating signal of net carbon sink and drought sensitivity of aboveground biomass (AGB) growth and mortality observed at forest plots across selected Amazon intact forests for 2005 and 2010. We predicted a larger mortality rate and a more negative sensitivity of the net carbon sink during the 2015/16 El Niño compared with the former droughts. 2015/16 was indeed the most severe drought since 1901 regarding both AGB loss and area experiencing a severe carbon loss. We found that even if climate change did increase mortality, elevated CO₂ contributed to balance the biomass mortality, since CO₂-induced stomatal closure reduces transpiration, thus, offsets increased transpiration from CO₂-induced higher foliage area.