Alexandre A. Oliveira

Pervasive alteration of tree communities in undisturbed Amazonian forests

Amazonian rainforests are some of the most species-rich tree communities on earth. Here we show that, over the past two decades, forests in a central Amazonian landscape have experienced highly nonrandom changes in dynamics and composition. Our analyses are based on a network of 18 permanent plots unaffected by any detectable disturbance. Within these plots, rates of tree mortality, recruitment and growth have increased over time. Of 115 relatively abundant tree genera, 27 changed significantly in population density or basal area—a value nearly 14 times greater than that expected by chance. An independent, eight-year study in nearby forests corroborates these shifts in composition. Contrary to recent predictions, we observed no increase in pioneer trees. However, genera of faster-growing trees, including many canopy and emergent species, are increasing in dominance or density, whereas genera of slower-growing trees, including many subcanopy species, are declining. Rising atmospheric CO2 concentrations may explain these changes, although the effects of this and other large-scale environmental alterations remain uncertain. These compositional changes could have important impacts on the carbon storage, dynamics and biota of Amazonian forests.

A central Amazonian terra firme forest. I. High tree species richness on poor soils

Tree size, density, and species richness were established for three one-hectare plots of terra firme forest in central Amazonian Brazil. In the three hectares, 1916 individual trees with DBH 10 ≥ cm were sampled. A total of 58 families, 181 genera, and 513 species were determined. Hectare A had 285 species, 138 genera, and 47 families; hectare B 280 species, 123 genera, and 48 families; and hectare C 280 species, 125 genera, and 44 families. Comparably high species richness in Amazonia has heretofore only been reported from western Amazonia. This dispels the idea that high species richness can only develop in areas with rich soils and relatively high rainfall. It is suggested that such high species richness is the result of a combination of habitat heterogeneity and geological history. These high diversity forests, because they occur on nutrient poor soils, can be protected with little or no impact on development in the region because the soils are essentially useless for agriculture and for supporting long-term cattle pasture.

Biomass resilience of Neotropical secondary forests

Land-use change occurs nowhere more rapidly than in the tropics, where the imbalance between deforestation and forest regrowth has large consequences for the global carbon cycle. However, considerable uncertainty remains about the rate of biomass recovery in secondary forests, and how these rates are influenced by climate, landscape, and prior land use. Here we analyse aboveground biomass recovery during secondary succession in 45 forest sites and about 1,500 forest plots covering the major environmental gradients in the Neotropics. The studied secondary forests are highly productive and resilient. Aboveground biomass recovery after 20 years was on average 122 megagrams per hectare (Mg ha−1), corresponding to a net carbon uptake of 3.05 Mg C ha−1 yr−1, 11 times the uptake rate of old-growth forests. Aboveground biomass stocks took a median time of 66 years to recover to 90% of old-growth values. Aboveground biomass recovery after 20 years varied 11.3-fold (from 20 to 225 Mg ha−1) across sites, and this recovery increased with water availability (higher local rainfall and lower climatic water deficit). We present a biomass recovery map of Latin America, which illustrates geographical and climatic variation in carbon sequestration potential during forest regrowth. The map will support policies to minimize forest loss in areas where biomass resilience is naturally low (such as seasonally dry forest regions) and promote forest regeneration and restoration in humid tropical lowland areas with high biomass resilience.

Geographic distribution of tree species occurring in the region of Manaus, Brazil: implications for regional diversity and conservation

The alpha-diversity of trees found in the region of Manaus, Brazil is among the highest recorded for one-hectare plots in Amazonia or any tropical forest. Based on a survey of the distributions of 2541 Neotropical tree species, we analyzed the geographic distribution of 364 species of terra firme forest trees that occur in the region and that are not edaphic specialists. Fifteen distinct distribution patterns were recognized for trees occurring in Manaus. The great majority of species (84.9%) have continuous, somewhat restricted distributions, 35 (9.6%) show broad distributions and 20 species (5.5%) show disjunction between Amazonia and Eastern Brazil. A remarkable 150 (41.2%) of these species showed the region of Manaus as one of their distribution limits. Using the same pool of 2541 species distributions, the percentage of species with a distribution limit in Manaus was compared with that for other localities known to be centers of botanical collection. The null hypothesis that the difference in proportion of species with distribution limits among these localities and Manaus is insignificant was rejected. We conclude that the results are not an artifact of collecting density, that Manaus is indeed a crossroads of distinct phytogeographic regions, and that this explains part of the high species diversity of trees in the region of Manaus. A number of scenarios proposed for the Pleistocene in Amazonia postulate some degree of fragmentation of Amazonian forests or at least populations. As much as these theories may conflict with each other in some respects, they are compatible with the concept of Manaus as a region of re-convergence of isolated or disrupted floras and faunas. The significance of the vicinity of Manaus in the history of the Amazon flora and its current status as a repository for surprisingly high tree diversity highlights the need to make this region a conservation priority.

Relative effect of litter quality, forest type and their interaction on leaf decomposition in south-east Brazilian forests

Decomposition was studied in a reciprocal litter transplant experiment to examine the effects of forest type, litter quality and their interaction on leaf decomposition in four tropical forests in south-east Brazil. Litterbags were used to measure decomposition of leaves of one tree species from each forest type:Calophyllum brasiliensefrom restinga forest; Guapira opposita from Atlantic forest; Esenbeckia leiocarpa from semi-deciduous forest; and Copaifera langsdorffii fromcerrad˜ ao.Decompositionratesinrainforests(Atlanticandrestinga)weretwiceasfastasthoseinseasonalforests (semi-deciduous and cerrad˜ ao), suggesting that intensity and distribution of precipitation are important predictors of decomposition rates at regional scales. Decomposition rates varied by species, in the following order: E. leiocarpa > C. langsdorffii > G. opposita > C. brasiliense. However, there was no correlation between decomposition rates and chemical litter quality parameters: C:N, C:P, lignin concentration and lignin:N. The interaction between forest type andlitterqualitywaspositivemainlybecauseC.langsdorffiidecomposedfasterthanexpectedinitsnativeforest.Thisis apotentialindicationofadecomposersadaptationtospecificsubstratesinatropicalforest.Thesefindingssuggestthat besides climate, interactions between decomposers and plants might play an essential role in decomposition processes and it must be better understood.

Litterfall and leaf decomposition in forest fragments under different successional phases on the Atlantic Plateau of the state of Sao Paulo, Brazil

Litterfall and litter decomposition are vital processes in tropical forests because they regulate nutrient cycling. Nutrient cycling can be altered by forest fragmentation. The Atlantic Forest is one of the most threatened biomes in the world due to human occupation over the last 500 years. This scenario has resulted in fragments of different size, age and regeneration phase. To investigate differences in litterfall and leaf decomposition between forest successional phases, we compared six forest fragments at three different successional phases and an area of mature forest on the Atlantic Plateau of Sao Paulo, Brazil. We sampled litter monthly from November 2008 to October 2009. We used litterbags to calculate leaf decomposition rate of an exotic species, Tipuana tipu (Fabaceae), over the same period litter sampling was performed. Litterfall was higher in the earliest successional area. This pattern may be related to the structural properties of the forest fragments, especially the higher abundance of pioneer species, which have higher productivity and are typical of early successional areas. However, we have not found significant differences in the decomposition rates between the studied areas, which may be caused by rapid stabilization of the decomposition environment (combined effect of microclimatic conditions and the decomposers activities). This result indicates that the leaf decomposition process have already been restored to levels observed in mature forests after a few decades of regeneration, although litterfall has not been entirely restored. This study emphasizes the importance of secondary forests for restoration of ecosystem processes on a regional scale.

Structure, diversity, and spatial patterns in a permanent plot of a high Restinga forest in Southeastern Brazil

(Structure, composition, and diversity of a high Restinga forest in the Southern Atlantic Forest) We assessed the structure, diversity and distribution of tree species in a 10.24-ha permanent plot of high Restinga forest (HRF) in southeastern Brazil. We sampled 15,040 individuals belonging to 45 families, 87 genera and 116 species (density= 1,468 trees ha -1 , and basal area= 28.0 m 2 ha -1 ). Mean richness was lower than other types of tropical forests and other HRFs that have been less intensively sampled, which is probably due to the younger age and hydromorphy of the soils in the study site. Tree density, basal area, species composition, and diversity all varied considerably depending on the size class considered. The studied plot had a density similar to other tropical forests but with a smaller basal area (fewer individuals > 60 cm in diameter), and higher density of smaller trees. Multi-stemmed individuals were not frequent (11% of the trees), though they were widespread among species (74% of the species). Spatial distribution was predominantly clumped, a pattern common to other types of tropical forests. Besides growing over white-sand soils, the PEIC plot and other white-sand forests of the world also had very similar patterns of community structure, richness and diversity. Therefore, we consider the HRF as a recognizable subtype of white-sand forest.

Intraspecific leaf trait variability along a boreal-to-tropical community diversity gradient

Disentangling the mechanisms that shape community assembly across diversity gradients is a central matter in ecology. While many studies have explored community assembly through species average trait values, there is a growing understanding that intraspecific trait variation (ITV) can also play a critical role in species coexistence. Classic biodiversity theory hypothesizes that higher diversity at species-rich sites can arise from narrower niches relative to species-poor sites, which would be reflected in reduced ITV as species richness increases. To explore how ITV in woody plant communities changes with species richness, we compiled leaf trait data (leaf size and specific leaf area) in a total of 521 woody plant species from 21 forest communities that differed dramatically in species richness, ranging from boreal to tropical rainforests. At each forest, we assessed ITV as an estimate of species niche breadth and we quantified the degree of trait overlap among co-occurring species as a measure of species functional similarity. We found ITV was relatively invariant across the species richness gradient. In addition, we found that species functional similarity increased with diversity. Contrary to the expectation from classic biodiversity theory, our results rather suggest that neutral processes or equalizing mechanisms can be acting as potential drivers shaping community assembly in hyperdiverse forests.

The importance of mesofauna and decomposition environment on leaf decomposition in three forests in southeastern Brazil

We examined the effects of soil mesofauna and the litter decomposition environment (above and belowground) on leaf decomposition rates in three forest types in southeastern Brazil. To estimate decomposition experimentally, we used litterbags with a standard substrate in a full-factorial experimental design. We used model selection to compare three decomposition models and also to infer the importance of forest type, decomposition environment, mesofauna, and their interactions on the decomposition process. Rather than the frequently used simple and double-exponential models, the best model to describe our dataset was the exponential deceleration model, which assumed a single organic compartment with an exponential decrease of the decomposition rate. Decomposition was higher in the wet than in the seasonal forest, and the differences between forest types were stronger aboveground. Regarding litter decomposition environment, decomposition was predominantly higher below than aboveground, but the magnitude of this effect was higher in the seasonal than in wet forests. Mesofauna exclusion treatments had slower decomposition, except aboveground into the Semi-deciduous Forest, where the mesofauna presence did not affect decomposition. Furthermore, the effect of mesofauna was stronger in the wet forests and belowground. Overall, our results suggest that, in a regional scale, both decomposers activity and the positive effect of soil mesofauna in decomposition are constrained by abiotic factors, such as moisture conditions.

Does extreme environmental severity promote plant facilitation? An experimental field test in a subtropical coastal dune

The stress gradient hypothesis (SGH) postulates how the balance between plant competition and facilitation shifts along environmental gradients. Early formulations of the SGH predicted that facilitation should increase monotonically with stress. However, a recent theoretical refinement of the SGH postulates stronger facilitation under moderate stress, followed by a decreasing role of facilitation in the most severe environments. We conducted field experiments along the most severe part of a coastal dune gradient in southeast Brazil to test the effect of stress on the intensity and importance of the net interactions between two tree species. First, we compared the performance of distinct life stages of Ternstroemia brasiliensis in the presence and absence of Guapira opposita adults along a beach-to-inland gradient, a gradient of environmental severity. To test the effect of one stress factor in particular, we also manipulated water availability, a limiting resource due to the sandy soils. At the most severe part of the coastal gradient (i.e. closest to the seashore), both intensity and importance of the interaction between G. opposita and T. brasiliensis were negatively related to stress, with a pattern consistent across distinct life stages of the target species. However, the sign of the net interaction depended on the life stage of the target species. Our results provide empirical evidence that the role of facilitation tends to wane, leading to neutral or even negative net interactions between species as stress reaches its maximum, as predicted by the recent refinements of the SGH.