David A. Neill

Drought sensitivity of the Amazon rainforest

Amazon forests are a key but poorly understood component of the global carbon cycle. If, as anticipated, they dry this century, they might accelerate climate change through carbon losses and changed surface energy balances. We used records from multiple long-term monitoring plots across Amazonia to assess forest responses to the intense 2005 drought, a possible analog of future events. Affected forest lost biomass, reversing a large long-term carbon sink, with the greatest impacts observed where the dry season was unusually intense. Relative to pre-2005 conditions, forest subjected to a 100-millimeter increase in water deficit lost 5.3 megagrams of aboveground biomass of carbon per hectare. The drought had a total biomass carbon impact of 1.2 to 1.6 petagrams (1.2 × 1015 to 1.6 × 1015 grams). Amazon forests therefore appear vulnerable to increasing moisture stress, with the potential for large carbon losses to exert feedback on climate change.

DOMINANCE AND DISTRIBUTION OF TREE SPECIES IN UPPER AMAZONIAN TERRA FIRME FORESTS

Amazonian forests are the largest and most diverse in the tropics, and much of the mystery surrounding their ecology can be traced to attempts to understand them through tiny local inventories. In this paper we bring together a large number of such inventories scattered across immense areas of western Amazonia in order to address simple questions about the distribution and abundance of tropical tree species in lowland terra firme forests there. The goal is to describe patterns of commonness and rarity at local (1 ha), landscape (∼104 km2), and regional (>106 km2) scales, and to fuse the results into a more complete picture of how tropical tree communities are structured. We present estimates of landscape-scale densities for ∼1400 taxa, based on data from tree plots scattered over large tracts of terra firme forest in eastern Ecuador and southeastern Peru. A database of morphological, ecological, and other traits of >1000 of these species compiled from the taxonomic literature is then used to explore how species that are common in the inventories differ from species that are rare. Although most species show landscape-scale densities of <1 individual/ha, most trees in both forests belong to a small set of ubiquitous common species. These common species combine high frequency with high local abundance, forming predictable oligarchies that dominate several thousand square kilometers of forest at each site. The common species comprising these oligarchies are a nonrandom subset of the two floras. At both sites a disproportionate number of common species are concentrated in the families Arecaceae, Moraceae, Myristicaceae, and Violaceae, and large-statured tree species are more likely to be common than small ones. Nearly a third of the 150 most common tree species in the Ecuadorean forest are also found among the 150 most common tree species in the Peruvian forest. For the 254 tree species shared by the two data sets, abundance in Ecuador is positively and significantly correlated with abundance ∼1400 km away in Peru. These findings challenge popular depictions of Amazonian vegetation as a small-scale mosaic of unpredictable composition and structure. Instead, they provide additional evidence that tropical tree communities are not qualitatively different from their temperate counterparts, where a few common species concentrated in a few higher taxa can dominate immense areas of forest. We hypothesize that most Amazonian forests are dominated at large scales by oligarchies similar in nature to the ones observed in Ecuador and Peru, and we argue that the patterns are more indicative of regulation of relative abundances by ecological factors than of nonequilibrium chance-based dynamics. The paper concludes with a discussion of the practical applications of predictable oligarchies over large areas of unexplored forest.

Increasing dominance of large lianas in Amazonian forests

Ecological orthodoxy suggests that old-growth forests should be close to dynamic equilibrium, but this view has been challenged by recent findings that neotropical forests are accumulating carbon and biomass, possibly in response to the increasing atmospheric concentrations of carbon dioxide. However, it is unclear whether the recent increase in tree biomass has been accompanied by a shift in community composition. Such changes could reduce or enhance the carbon storage potential of old-growth forests in the long term. Here we show that non-fragmented Amazon forests are experiencing a concerted increase in the density, basal area and mean size of woody climbing plants (lianas). Over the last two decades of the twentieth century the dominance of large lianas relative to trees has increased by 1.7–4.6% a year. Lianas enhance tree mortality and suppress tree growth, so their rapid increase implies that the tropical terrestrial carbon sink may shut down sooner than current models suggest. Predictions of future tropical carbon fluxes will need to account for the changing composition and dynamics of supposedly undisturbed forests.

Long-term decline of the Amazon carbon sink

Atmospheric carbon dioxide records indicate that the land surface has acted as a strong global carbon sink over recent decades, with a substantial fraction of this sink probably located in the tropics, particularly in the Amazon. Nevertheless, it is unclear how the terrestrial carbon sink will evolve as climate and atmospheric composition continue to change. Here we analyse the historical evolution of the biomass dynamics of the Amazon rainforest over three decades using a distributed network of 321 plots. While this analysis confirms that Amazon forests have acted as a long-term net biomass sink, we find a long-term decreasing trend of carbon accumulation. Rates of net increase in above-ground biomass declined by one-third during the past decade compared to the 1990s. This is a consequence of growth rate increases levelling off recently, while biomass mortality persistently increased throughout, leading to a shortening of carbon residence times. Potential drivers for the mortality increase include greater climate variability, and feedbacks of faster growth on mortality, resulting in shortened tree longevity. The observed decline of the Amazon sink diverges markedly from the recent increase in terrestrial carbon uptake at the global scale, and is contrary to expectations based on models.

An international network to monitor the structure, composition and dynamics of Amazonian forests (RAINFOR)

Abstract The Amazon basin is likely to be increasingly affected by environmental changes: higher temperatures, changes in precipitation, CO2 fertilization and habitat fragmentation. To examine the important ecological and biogeochemical consequences of these changes, we are developing an international network, RAINFOR, which aims to monitor forest biomass and dynamics across Amazonia in a co-ordinated fashion in order to understand their relationship to soil and climate. The network will focus on sample plots established by independent researchers, some providing data extending back several decades. We will also conduct rapid transect studies of poorly monitored regions. Field expeditions analysed local soil and plant properties in the first phase (2001–2002). Initial results suggest that the network has the potential to reveal much information on the continental-scale relations between forest and environment. The network will also serve as a forum for discussion between researchers, with the aim of standardising sampling techniques and methodologies that will enable Amazonian forests to be monitored in a coherent manner in the coming decades. Abbreviation: PSP = Permanent sample plot.

Markedly divergent estimates of Amazon forest carbon density from ground plots and satellites

Aim The accurate mapping of forest carbon stocks is essential for understanding the global carbon cycle, for assessing emissions from deforestation, and for rational land-use planning. Remote sensing (RS) is currently the key tool for this purpose, but RS does not estimate vegetation biomass directly, and thus may miss significant spatial variations in forest structure. We test the stated accuracy of pantropical carbon maps using a large independent field dataset. Location Tropical forests of the Amazon basin. The permanent archive of the field plot data can be accessed at: http://dx.doi.org/10.5521/FORESTPLOTS.NET/2014_1 Methods Two recent pantropical RS maps of vegetation carbon are compared to a unique ground-plot dataset, involving tree measurements in 413 large inventory plots located in nine countries. The RS maps were compared directly to field plots, and kriging of the field data was used to allow area-based comparisons. Results The two RS carbon maps fail to capture the main gradient in Amazon forest carbon detected using 413 ground plots, from the densely wooded tall forests of the north-east, to the light-wooded, shorter forests of the south-west. The differences between plots and RS maps far exceed the uncertainties given in these studies, with whole regions over- or under-estimated by > 25%, whereas regional uncertainties for the maps were reported to be < 5%. Main conclusions Pantropical biomass maps are widely used by governments and by projects aiming to reduce deforestation using carbon offsets, but may have significant regional biases. Carbon-mapping techniques must be revised to account for the known ecological variation in tree wood density and allometry to create maps suitable for carbon accounting. The use of single relationships between tree canopy height and above-ground biomass inevitably yields large, spatially correlated errors. This presents a significant challenge to both the forest conservation and remote sensing communities, because neither wood density nor species assemblages can be reliably mapped from space.

Amazon forest response to repeated droughts

The Amazon Basin has experienced more variable climate over the last decade, with a severe and widespread drought in 2005 causing large basin-wide losses of biomass. A drought of similar climatological magnitude occurred again in 2010; however, there has been no basin-wide ground-based evaluation of effects on vegetation. We examine to what extent the 2010 drought affected forest dynamics using ground-based observations of mortality and growth from an extensive forest plot network. We find that during the 2010 drought interval, forests did not gain biomass (net change: −0.43 Mg ha−1, confidence interval (CI): −1.11, 0.19, n = 97), regardless of whether forests experienced precipitation deficit anomalies. This contrasted with a long-term biomass sink during the baseline pre-2010 drought period (1998 to pre-2010) of 1.33 Mg ha−1 yr−1 (CI: 0.90, 1.74, p < 0.01). The resulting net impact of the 2010 drought (i.e., reversal of the baseline net sink) was −1.95 Mg ha−1 yr−1 (CI:−2.77, −1.18; p < 0.001). This net biomass impact was driven by an increase in biomass mortality (1.45 Mg ha−1 yr−1 CI: 0.66, 2.25, p < 0.001) and a decline in biomass productivity (−0.50 Mg ha−1 yr−1, CI:−0.78, −0.31; p < 0.001). Surprisingly, the magnitude of the losses through tree mortality was unrelated to estimated local precipitation anomalies and was independent of estimated local pre-2010 drought history. Thus, there was no evidence that pre-2010 droughts compounded the effects of the 2010 drought. We detected a systematic basin-wide impact of the 2010 drought on tree growth rates across Amazonia, which was related to the strength of the moisture deficit. This impact differed from the drought event in 2005 which did not affect productivity. Based on these ground data, live biomass in trees and corresponding estimates of live biomass in lianas and roots, we estimate that intact forests in Amazonia were carbon neutral in 2010 (−0.07 Pg C yr−1 CI:−0.42, 0.23), consistent with results from an independent analysis of airborne estimates of land-atmospheric fluxes during 2010. Relative to the long-term mean, the 2010 drought resulted in a reduction in biomass carbon uptake of 1.1 Pg C, compared to 1.6 Pg C for the 2005 event.

Tropical forest wood production: a cross-continental comparison

Summary: Tropical forest above-ground wood production (AGWP) varies substantially along environmental gradients. Some evidence suggests that AGWP may vary between regions and specifically that Asian forests have particularly high AGWP. However, comparisons across biogeographic regions using standardized methods are lacking, limiting our assessment of pan-tropical variation in AGWP and potential causes. We sampled AGWP in NW Amazon (17 long-term forest plots) and N Borneo (11 plots), both with abundant year-round precipitation. Within each region, forests growing on a broad range of edaphic conditions were sampled using standardized soil and forest measurement techniques. Plot-level AGWP was 49% greater in Borneo than in Amazonia (9.73 ± 0.56 vs. 6.53 ± 0.34 Mg dry mass ha -1 a -1 , respectively; regional mean ± 1 SE). AGWP was positively associated with soil fertility (PCA axes, sum of bases and total P). After controlling for the edaphic environment, AGWP remained significantly higher in Bornean plots. Differences in AGWP were largely attributable to differing height-diameter allometry in the two regions and the abundance of large trees in Borneo. This may be explained, in part, by the greater solar radiation in Borneo compared with NW Amazonia. Trees belonging to the dominant SE Asian family, Dipterocarpaceae, gained woody biomass faster than otherwise equivalent, neighbouring non-dipterocarps, implying that the exceptional production of Bornean forests may be driven by floristic elements. This dominant SE Asian family may partition biomass differently or be more efficient at harvesting resources and in converting them to woody biomass. Synthesis. N Bornean forests have much greater AGWP rates than those in NW Amazon when soil conditions and rainfall are controlled for. Greater resource availability and the highly productive dipterocarps may, in combination, explain why Asian forests produce wood half as fast again as comparable forests in the Amazon. Our results also suggest that taxonomic groups differ in their fundamental ability to capture carbon and that different tropical regions may therefore have different carbon uptake capacities due to biogeographic history. North Bornean forests have much greater AGWP rates than those in north-western Amazon when soil conditions and rainfall are controlled for. Greater resource availability and the highly productive dipterocarps may, in combination, explain why these Asian forests produce wood half as fast again as comparable forests in the Amazon. Our results also suggest that taxonomic groups differ in their fundamental ability to capture carbon and that different tropical regions may therefore have different carbon uptake capacities due to biogeographic history.

Spatial patterns of above-ground structure, biomass and composition in a network of six Andean elevation transects

Background: The Amazon to Andes transition zone provides large expanses of relatively pristine forest wilderness across environmental gradients. Such elevational gradients are an excellent natural laboratory for establishing long-term interactions between forest ecosystems and environmental parameters, which is valuable for understanding ecosystem responses to environmental change. Aims: This study presents data on elevational trends of forest structure (biomass, basal area, height, stem density), species richness, and composition from six elevational transects in the Andes. Methods: We analysed the spatial patterns of forest structure, above-ground biomass and composition from 76 permanent plots, ranging from lowland Amazonian rain forest to high-elevation cloud forests in Ecuador, Peru, and Bolivia. Results: Forest above-ground woody biomass stocks ranged from 247 Mg ha−1 (Peru, 210 m) to 86 Mg ha−1 (Peru, 3450 m), with significantly decreasing trends of tree height and biomass and an increasing trend of stem density with increasing elevation. We observed an increase in forest richness at three taxonomic levels at mid-elevation, followed by a decrease in richness within the cloud immersion zone. Conclusions: The transects show an increase in stem density, a decline in tree height and above-ground coarse wood biomass and a hump-shaped trend in species richness with increasing elevation. These results suggest that environmental change could lead to significant shifts in the properties of these ecosystems over time.

Variation in stem mortality rates determines patterns of above-ground biomass in Amazonian forests: implications for dynamic global vegetation models

Abstract 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, basin‐wide 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.