Miles R. Silman

Benchmark map of forest carbon stocks in tropical regions across three continents

Developing countries are required to produce robust estimates of forest carbon stocks for successful implementation of climate change mitigation policies related to reducing emissions from deforestation and degradation (REDD). Here we present a “benchmark” map of biomass carbon stocks over 2.5 billion ha of forests on three continents, encompassing all tropical forests, for the early 2000s, which will be invaluable for REDD assessments at both project and national scales. We mapped the total carbon stock in live biomass (above- and belowground), using a combination of data from 4,079 in situ inventory plots and satellite light detection and ranging (Lidar) samples of forest structure to estimate carbon storage, plus optical and microwave imagery (1-km resolution) to extrapolate over the landscape. The total biomass carbon stock of forests in the study region is estimated to be 247 Gt C, with 193 Gt C stored aboveground and 54 Gt C stored belowground in roots. Forests in Latin America, sub-Saharan Africa, and Southeast Asia accounted for 49%, 25%, and 26% of the total stock, respectively. By analyzing the errors propagated through the estimation process, uncertainty at the pixel level (100 ha) ranged from ±6% to ±53%, but was constrained at the typical project (10,000 ha) and national (>1,000,000 ha) scales at ca. ±5% and ca. ±1%, respectively. The benchmark map illustrates regional patterns and provides methodologically comparable estimates of carbon stocks for 75 developing countries where previous assessments were either poor or incomplete.

SEED DISPERSAL NEAR AND FAR: PATTERNS ACROSS TEMPERATE AND TROPICAL FORESTS

Dispersal affects community dynamics and vegetation response to global change. Understanding these effects requires descriptions of dispersal at local and regional scales and statistical models that permit estimation. Classical models of dispersal describe local or long-distance dispersal, but not both. The lack of statistical methods means that models have rarely been fitted to seed dispersal in closed forests. We present a mixture model of dispersal that assumes a range of disperal patterns, both local and long distance. The bivariate Student’s t or “2Dt” follows from an assumption that the distance parameter in a Gaussian model varies randomly, thus having a density of its own. We use an inverse approach to “compete” our mixture model against classical alternatives, using seed rain databases from temperate broadleaf, temperate mixed-conifer, and tropical floodplain forests. For most species, the 2Dt model fits dispersal data better than do classical models. The superior fit results from the potential f...

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.

TREE SPECIES DISTRIBUTIONS IN AN UPPER AMAZONIAN FOREST

Not a single tree species distribution in the Amazon basin has been reliably mapped, though speculation regarding such distributions has been extensive. We present data from a network of 21 forest plots in Manu National Park, Peru, totaling >36 ha and sited over an area of -400 km2, to explore how tree species are distributed across upper Amazonia at a variety of spatial scales. For each of 825 tree species occurring in the plots we asked three questions: (1) Does the species have a large or small geographic range? (2) Is the species restricted to a single forest type, or is it found in several? (3) Is the species locally abundant anywhere or is it scarce everywhere? The answers served to classify a subset of species under Rabinowitzs classification scheme for rare species. Three main conclusions emerged. First, the great majority of tree species at Manu are geographically widespread. Every species identified to date occurs elsewhere in South America, outside the department of Madre de Dios; more than two-thirds of them have been collected 1500 km away in Amazonian Ecuador. Second, 15-26% of species appear to be restricted to a single forest type, when forest types are defined by historical river dynamics (i.e., terra firme forest, mature floodplain forest, swamp forest, and primary successional floodplain forest). The proportion of restricted species declined with increasing sampling effort, mak- ing 15% a more reliable figure. Third, while 88% of species occurred at densities of 1.5 individuals/ha. Extrapolating these results provides a first guess at how tree species are distributed across the western portion of the Amazon basin. We conclude with the suggestion that most tree species in the region are habitat generalists occurring over large areas of the Amazonian lowlands at low densities but large absolute population sizes.

Microbes do not follow the elevational diversity patterns of plants and animals

The elevational gradient in plant and animal diversity is one of the most widely documented patterns in ecology and, although no consensus explanation exists, many hypotheses have been proposed over the past century to explain these patterns. Historically, research on elevational diversity gradients has focused almost exclusively on plant and animal taxa. As a result, we do not know whether microbes exhibit elevational gradients in diversity that parallel those observed for macroscopic taxa. This represents a key knowledge gap in ecology, especially given the ubiquity, abundance, and functional importance of microbes. Here we show that, across a montane elevational gradient in eastern Peru, bacteria living in three distinct habitats (organic soil, mineral soil, and leaf surfaces) exhibit no significant elevational gradient in diversity (r2<0.17, P>0.1 in all cases), in direct contrast to the significant diversity changes observed for plant and animal taxa across the same montane gradient (r2>0.75, P<0.001 in all cases). This finding suggests that the biogeographical patterns exhibited by bacteria are fundamentally different from those of plants and animals, highlighting the need for the development of more inclusive concepts and theories in biogeography to explain these disparities.

POPULATION REGULATION OF A DOMINANT RAIN FOREST TREE BY A MAJOR SEED PREDATOR

We take advantage of a fortuitous local extinction and recolonization of white-lipped peccaries (WLPs) at the Cocha Cashu Biological Station in southeastern Peru to assess the impact of this high-biomass seed predator on the recruitment of a dominant member of the tree community, the palm Astrocaryum murumuru. WLPs were common at Cocha Cashu in the mid-1970s. In 1978, the species vanished from the entire region and did not reappear until 1990. To assess the impacts of the presence-absence of WLPs on Astrocaryum recruitment, we conducted transect counts of both the number and spatial distribution of palm seedlings in 1978 (when WLPs were present), in 1990 (after a 12-yr absence of WLPs), and in 1999 (after 10 yr of recovery). Other factors affecting recruitment, such as climate variability, differences in tree fecundity, and fluctuations of vertebrate and invertebrate seed predators were also examined as alternative hypotheses for any changes in seedling spatial distribution and abundance. In the absence of WLPs, the density of Astrocaryum seedlings increased 1.7-fold, and the spatial distribution of seedlings with respect to safe sites was significantly altered. After the return of WLPs to the study area, seedling density dropped to its former level, and the spatial distribution of seedlings with respect to safe sites returned to the same pattern found 21 yr earlier in 1978. None of the other factors investigated varied systematically with the census intervals. These results demonstrate that the absence of a single, albeit important, member of a diverse seed predator guild can have a major impact on the demography of a common tree species. We infer that individual seed predators can play central roles in regulating the demography of tropical trees, as envisioned many years ago by Janzen and Connell, and that their extirpation or extinction can have cascading effects in tropical ecosystems.

Sparse Pre-Columbian Human Habitation in Western Amazonia

Population Limits Extensive pre-Columbian populations inhabited the central and eastern Amazon basin, as evidenced by the clearing and modification of forests. McMichael et al. (p. 1429) examined how far inland such activities may have extended by sampling soils across western Amazonia, including river bluffs, which were heavily occupied downstream. Little evidence of human disturbance across a wide region was found by looking for charcoal layers (which would suggest use of fire) and phytoliths, which trace local plants and would indicate the presence of crops. Furthermore, no ceramics or tools were found. Thus, pre-Columbian human populations seem to have been sparse in western Amazonia. Analysis of soils in western Amazonia finds little evidence for pre-Columbian human occupations there. Locally extensive pre-Columbian human occupation and modification occurred in the forests of the central and eastern Amazon Basin, but whether comparable impacts extend westward and into the vast terra firme (interfluvial) zones, remains unclear. We analyzed soils from 55 sites across central and western Amazonia to assess the history of human occupation. Sparse occurrences of charcoal and the lack of phytoliths from agricultural and disturbance species in the soils during pre-Columbian times indicated that human impacts on interfluvial forests were small, infrequent, and highly localized. No human artifacts or modified soils were found at any site surveyed. Riverine bluff areas also appeared less heavily occupied and disturbed than similar settings elsewhere. Our data indicate that human impacts on Amazonian forests were heterogeneous across this vast landscape.

Fire, climate change and biodiversity in Amazonia: a Late-Holocene perspective

Fire is an important and arguably unnatural component of many wet Amazonian and Andean forest systems. Soil charcoal has been used to infer widespread human use of landscapes prior to European Conquest. An analysis of Amazonian soil carbon records reveals that the records have distinct spatial and temporal patterns, suggesting that either fires were only set in moderately seasonal areas of Amazonia or that strongly seasonal and aseasonal areas are undersampled. Synthesizing data from 300 charcoal records, an age–frequency diagram reveals peaks of fire apparently coinciding with some periods of very strong El Niño activity. However, the El Niño record does not always provide an accurate prediction of fire timing, and a better match is found in the record of insolation minima. After the time of European contact, fires became much scarcer within Amazonia. In both the Amazonia and the Andes, modern fire pattern is strongly allied to human activity. On the flank of the Andes, forests that have never burned are being eroded by fire spreading downslope from grasslands. Species of these same forests are being forced to migrate upslope due to warming and will encounter a firm artificial fire boundary of human activity.

Holocene fire and occupation in Amazonia: records from two lake districts

While large-scale pre-Columbian human occupation and ecological disturbance have been demonstrated close to major Amazonian waterways, less is known of sites in terra firme settings. Palaeoecological analyses of two lake districts in central and western Amazonia reveal long histories of occupation and land use. At both locations, human activity was centred on one of the lakes, while the others were either lightly used or unused. These analyses indicate that the scale of human impacts in these terra firme settings is localized and probably strongly influenced by the presence of a permanent open-water body. Evidence is found of forest clearance and cultivation of maize and manioc. These data are directly relevant to the resilience of Amazonian conservation, as they do not support the contention that all of Amazonia is a ‘built landscape’ and therefore a product of past human land use.

Amazonian exploitation revisited: ecological asymmetry and the policy pendulum

The influence of pre-Columbian human populations on Amazonian ecosystems is being actively debated. The longstanding view that Amazonia was only minimally impacted by human actions has been challenged, and a new paradigm of Amazonia as a “manufactured landscape” is emerging. If such disturbance was the norm until just 500 years ago, Amazonian ecosystems could be far more ecologically resilient to disturbance than previously supposed. Alternatively, if the “manufactured landscape” label is an overstatement, then policy that assumes such resilience may cause substantial and long-lasting ecological damage. We present paleoecological data suggesting a middle path, in which some areas were heavily modified, but most of Amazonia was minimally impacted. Bluffs adjacent to main river channels and highly seasonal areas appear to have been the most extensively settled locations. Away from areas where humans lived, their influence on ecosystems was very local. Consequently, we see no evidence suggesting that large a...