Old-growth forest ecosystems comprise a mosaic of patches in different successional stages, with the fraction of the landscape in any particular state relatively constant over large temporal and spatial scales. The size distribution and return frequency of disturbance events, and subsequent recovery processes, determine to a large extent the spatial scale over which this old-growth steady state develops. Here, we characterize this mosaic for a Central Amazon forest by integrating field plot data, remote sensing disturbance probability distribution functions, and individual-based simulation modeling. Results demonstrate that a steady state of patches of varying successional age occurs over a relatively large spatial scale, with important implications for detecting temporal trends on plots that sample a small fraction of the landscape. Long highly significant stochastic runs averaging 1.0 Mg biomass·ha −1 ·y −1 were often punctuated by episodic disturbance events, resulting in a sawtooth time series of hectare-scale tree biomass. To maximize the detection of temporal trends for this Central Amazon site (e.g., driven by CO 2 fertilization), plots larger than 10 ha would provide the greatest sensitivity. A model-based analysis of fractional mortality across all gap sizes demonstrated that 9.1-16.9% of tree mortality was missing from plot-based approaches, underscoring the need to combine plot and remote-sensing methods for estimating net landscape carbon balance. Old-growth tropical forests can exhibit complex large-scale structure driven by disturbance and recovery cycles, with ecosystem and community attributes of hectare-scale plots exhibiting continuous dynamic departures from a steady-state condition.biodiversity | community composition | gap dynamics | NEP NEE NBP A common assumption in old-growth forest studies is that, in the absence of a directional forcing, ecosystem characteristics and tree species composition should exhibit some type of steady-state behavior (1). Thus, plot-based studies in old-growth tropical forests that observe changing tree species composition (2), increased liana abundance (3), faster turnover rates (4), and forest biomass accumulation (5, 6), are viewed as surprising departures from an expected steady-state condition. However, disturbance events can create a landscape with patches of varying successional age, and the extent to which forest plots representatively sample this mosaic remains an open question. An important issue is how to distinguish directional trends driven by a warming climate, or rising atmospheric CO 2 concentration, from smaller-scale stochastic patterns driven by disturbance and recovery cycles (7,8).Over long time periods, the disturbance regime of a forested region creates a shifting steady-state mosaic, represented by patches of different successional ages, with the fraction of the landscape in any particular state remaining relatively constant over time (9,10). In many tropical forests, gaps created by the windthrow of canopy trees is a major mode of disturb...

Canopy gaps created by wind-throw events, or blowdowns, create a complex mosaic of forest patches varying in disturbance intensity and recovery in the Central Amazon. Using field and remote sensing data, we investigated the short-term (four-year) effects of large (>2000 m2) blowdown gaps created during a single storm event in January 2005 near Manaus, Brazil, to study (i) how forest structure and composition vary with disturbance gradients and (ii) whether tree diversity is promoted by niche differentiation related to wind-throw events at the landscape scale. In the forest area affected by the blowdown, tree mortality ranged from 0 to 70%, and was highest on plateaus and slopes. Less impacted areas in the region affected by the blowdown had overlapping characteristics with a nearby unaffected forest in tree density (583±46 trees ha−1) (mean±99% Confidence Interval) and basal area (26.7±2.4 m2 ha−1). Highly impacted areas had tree density and basal area as low as 120 trees ha−1 and 14.9 m2 ha−1, respectively. In general, these structural measures correlated negatively with an index of tree mortality intensity derived from satellite imagery. Four years after the blowdown event, differences in size-distribution, fraction of resprouters, floristic composition and species diversity still correlated with disturbance measures such as tree mortality and gap size. Our results suggest that the gradients of wind disturbance intensity encompassed in large blowdown gaps (>2000 m2) promote tree diversity. Specialists for particular disturbance intensities existed along the entire gradient. The existence of species or genera taking an intermediate position between undisturbed and gap specialists led to a peak of rarefied richness and diversity at intermediate disturbance levels. A diverse set of species differing widely in requirements and recruitment strategies forms the initial post-disturbance cohort, thus lending a high resilience towards wind disturbances at the community level.

Climate change is expected to increase the intensity of extreme precipitation events in Amazonia that in turn might produce more forest blowdowns associated with convective storms. Yet quantitative tree mortality associated with convective storms has never been reported across Amazonia, representing an important additional source of carbon to the atmosphere. Here we demonstrate that a single squall line (aligned cluster of convective storm cells) propagating across Amazonia in January, 2005, caused widespread forest tree mortality and may have contributed to the elevated mortality observed that year. Forest plot data demonstrated that the same year represented the second highest mortality rate over a 15‐year annual monitoring interval. Over the Manaus region, disturbed forest patches generated by the squall followed a power‐law distribution (scaling exponent α = 1.48) and produced a mortality of 0.3–0.5 million trees, equivalent to 30% of the observed annual deforestation reported in 2005 over the same area. Basin‐wide, potential tree mortality from this one event was estimated at 542 ± 121 million trees, equivalent to 23% of the mean annual biomass accumulation estimated for these forests. Our results highlight the vulnerability of Amazon trees to wind‐driven mortality associated with convective storms. Storm intensity is expected to increase with a warming climate, which would result in additional tree mortality and carbon release to the atmosphere, with the potential to further warm the climate system.

Widespread degradation of tropical forests is caused by a variety of disturbances that interact in ways that are not well understood. To explore potential synergies between edge effects, fire and windstorm damage as causes of Amazonian forest degradation, we quantified vegetation responses to a 30‐min, high‐intensity windstorm that in 2012, swept through a large‐scale fire experiment that borders an agricultural field. Our pre‐ and postwindstorm measurements include tree mortality rates and modes of death, above‐ground biomass, and airborne LiDAR‐based estimates of tree heights and canopy disturbance (i.e., number and size of gaps). The experimental area in the southeastern Amazonia includes three 50‐ha plots established in 2004 that were unburned (Control), burned annually (B1yr), or burned at 3‐year intervals (B3yr). The windstorm caused greater damage to trees (>10 cm DBH) in the burned plots (B1yr: 13 ± 9% of 785 trees; B3yr: 17 ± 13% of 433) than in the Control plot (8 ± 4% of 2,300; ± CI). It substantially reduced vegetation height by 14% in B1yr, 20% in B3yr and 12% in the Control plots, while it reduced above‐ground biomass by 18% of 77.7 Mg/ha (B1yr), 31% of 56.6 (B3yr), and 15% of 120 (Control). Tree damage was greatest near the agricultural field edge in all three plots, especially among large trees and in B3yr. Trunk snapping (70%) and uprooting (20%) were the most common modes of tree damage and mortality, with the height of trunk failure on the burned plots often corresponding with the height of historical fire scars. Of the windstorm‐damaged trees, 80% (B1yr), 90% (B3yr), and 57% (Control) were dead 4 years later. Trees that had crown damage experienced the least mortality (22%–60%), followed by those that were snapped (55%–94%) and uprooted (88%–94%). Synthesis. We demonstrate the synergistic effects of three kinds of disturbances on a tropical forest. Our results show that the effects of windstorms are exacerbated by prior degradation by fire and fragmentation. We highlight that understorey fires can produce long‐lasting effects on tropical forests not only by directly killing trees but also by increasing tree vulnerability to wind damage due to fire scars and a more open canopy.

Tree mortality is a key driver of forest community composition and carbon dynamics. Strong winds associated with severe convective storms are dominant natural drivers of tree mortality in the Amazon. Why forests vary with respect to their vulnerability to wind events and how the predicted increase in storm events might affect forest ecosystems within the Amazon are not well understood. We found that windthrows are common in the Amazon region extending from northwest (Peru, Colombia, Venezuela, and west Brazil) to central Brazil, with the highest occurrence of windthrows in the northwest Amazon. More frequent winds, produced by more frequent severe convective systems, in combination with well-known processes that limit the anchoring of trees in the soil, help to explain the higher vulnerability of the northwest Amazon forests to winds. Projected increases in the frequency and intensity of convective storms in the Amazon have the potential to increase wind-related tree mortality. A forest demographic model calibrated for the northwestern and the central Amazon showed that northwestern forests are more resilient to increased wind-related tree mortality than forests in the central Amazon. Our study emphasizes the importance of including wind-related tree mortality in model simulations for reliable predictions of the future of tropical forests and their effects on the Earth' system. A R 1995 Classification of multispectral images based on fractions of endmembers-application to land-cover change in the Brazilian amazon Remote Sens. Environ. 52 137-54 Aragao L E O C et al 2009 Above-and below-ground net primary productivity across ten Amazonian forests on contrasting soils Biogeosciences 6 2759-78 Baker T R et al 2004 Variation in wood density determines spatial patterns in Amazonian forest biomass Glob. Change Biol. 10 545-62 Boose E R, Serrano M I and Foster D R 2004 Landscape and regional impacts of hurricanes in Puerto Rico Ecol. Monogr. 74 335-52 Carlotto M J 1999 Reducing the effects of space-varying, wavelength-dependent scattering in multispectral imagery Int. J. Remote Sens. 20 3333-44 Chambers J Q, dos Santos J, Ribeiro R J and Higuchi N 2001 Tree damage, allometric relationships, and above-ground net primary production in central Amazon forest Forest Ecol.

Amazon forests account for ~25% of global land biomass and tropical tree species. In these forests, windthrows (i.e., snapped and uprooted trees) are a major natural disturbance, but the rates and mechanisms of recovery are not known. To provide a predictive framework for understanding the effects of windthrows on forest structure and functional composition (DBH ≥10 cm), we quantified biomass recovery as a function of windthrow severity (i.e., fraction of windthrow tree mortality on Landsat pixels, ranging from 0%–70%) and time since disturbance for terra‐firme forests in the Central Amazon. Forest monitoring allowed insights into the processes and mechanisms driving the net biomass change (i.e., increment minus loss) and shifts in functional composition. Windthrown areas recovering for between 4–27 years had biomass stocks as low as 65.2–91.7 Mg/ha or 23%–38% of those in nearby undisturbed forests (~255.6 Mg/ha, all sites). Even low windthrow severities (4%–20% tree mortality) caused decadal changes in biomass stocks and structure. While rates of biomass increment in recovering vegetation were nearly double (6.3 ± 1.4 Mg ha−1 year−1) those of undisturbed forests (~3.7 Mg ha−1 year−1), biomass loss due to post‐windthrow mortality was high (up to −7.5 ± 8.7 Mg ha−1 year−1, 8.5 years since disturbance) and unpredictable. Consequently, recovery to 90% of “pre‐disturbance” biomass takes up to 40 years. Resprouting trees contributed little to biomass recovery. Instead, light‐demanding, low‐density genera (e.g., Cecropia, Inga, Miconia, Pourouma, Tachigali, and Tapirira) were favored, resulting in substantial post‐windthrow species turnover. Shifts in functional composition demonstrate that windthrows affect the resilience of live tree biomass by favoring soft‐wooded species with shorter life spans that are more vulnerable to future disturbances. As the time required for forests to recover biomass is likely similar to the recurrence interval of windthrows triggering succession, windthrows have the potential to control landscape biomass/carbon dynamics and functional composition in Amazon forests.

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Lloyd et al. (2009) question the methods, concepts and conclusions of Fisher et al. (2008). We address these assertions, and place our work into a broader context. We demonstrate the veracity of Fisher et al., and further show that lack of data for intermediate-scale tree mortality disturbance events for old-growth tropical forests might prevent robust extrapolation of forest plot biomass accumulation data, and accurate estimates of distribution parameters such as power-law exponents (a)