Effects of experimental lightgaps and topography on enrichment plantings in a central Amazonian secondary forest

Enrichment plantings into secondary forest are an important option in restoring species diversity and ecosystem services. However, little attention has been given to environmental requirements for species performance. This study evaluated the effects of lightgaps and topographic position on the growth and survival of four native tree species (Pouteria caimito, Garcinia macrophylla, Dipteryx odorata and Cynometra bauhiniaefolia) planted into a 26-year old secondary forest originating from abandoned pastures in the central Amazon Basin. Artificial lightgaps and control plots under closed canopy were uniformly distributed on plateaus and bottomlands near water bodies. Seedlings were planted randomly into the plots and monitored for 28 months. Seedling survival rate was high (93%) and did not differ among species. Overall, lightgaps produced a 38% increase in seedling height relative to the controls. Although the four species naturally occur in mature forest, two of the four grew significantly more in lightgaps than in closed canopy secondary forest. Overall, bottomlands facilitated greater seedling growth in height (38%) relative to plateaus, but only one species exhibited a significant increase. This study shows the importance of the environmental variability generated with canopy openings along the topographic gradient, suggesting that both the selection of species and microsite conditions of planting sites have to be considered important criteria in the recovery of degraded areas. (AU)

Outcrossing rate of Couratari multiflora (J. Smith) Eyma (Lecythidaceae), a low-density tropical tree species from a central Amazonian rainforest / Taxa de cruzamento de Couratari multiflora (J. Smith) Eyma (Lecythidaceae), uma espécie arbórea tropical de baixa densidade da Amazônia central

A multilocus mixed-mating model was used to evaluate the mating system of a population of Couratari multiflora, an emergent tree species found in low densities (1 individual/10 ha) in lowland forests of central Amazonia. We surveyed and observed phenologically 41 trees in an area of 400 ha. From these, only four mother trees were analyzed here because few of them set fruits, which also suffered high predation. No difference was observed between the population multilocus outcrossing rate (t mp = 0.953 ± 0.040) and the average single locus rate (t sp = 0.968 ± 0.132). The four mother trees were highly outcrossed (t m ~ 1). Two out of five loci showed departures from the Hardy-Weinberg Equilibrium (HWE) expectations, and the same results occurred with the mixed-mating model. Besides the low number of trees analyzed, the proportion of loci in HWE suggests random mating in the population. However, the pollen pool was heterogeneous among families, probably due to both the small sample number and the flowering of trees at different times of the flowering season. Reproductive phenology of the population and the results presented here suggest, at least for part of the population, a long-distance pollen movement, around 1,000 m.

Foi utilizado um modelo de cruzamento misto multilocos para analisar o sistema de cruzamento de uma população de Couratari multiflora, espécie arbórea emergente encontrada em baixas densidades (1 indivíduo/10 ha) nas florestas de "terra firme" da Amazônia central. Inventariamos e observamos fenologicamente 41 árvores em uma área de 400 ha. Dessas, somente quatro árvores-mãe foram analisadas, pois poucas árvores produziram frutos, os quais também sofreram alta predação. Não foi observada diferença entre a taxa de cruzamento multilocos (t mp = 0,953 ± 0,040) e a taxa média de loco único (t sp = 0,968 ± 0,132). As quatro árvores-mãe apresentaram alta taxa de cruzamento (t m ~ 1). Dois dos cinco locos analisados mostraram desvios das expectativas do Equilíbrio de Hardy-Weinberg (EHW), e os mesmos resultados ocorreram para o modelo misto de cruzamento. Apesar do baixo número de árvores analisadas, a alta proporção de locos em EHW sugere que a população tenha um sistema de cruzamento aleatório. Entretanto, o conjunto polínico foi heterogêneo entre famílias, provavelmente pelo pequeno número e florescimento das árvores analisadas em diferentes fases da estação de florescimento. A fenologia reprodutiva da população e os resultados apresentados neste estudo sugerem que ao menos parte da população cruza via movimento de pólen de longa distância, em torno de 1.000 m.

Habitat fragmentation, variable edge effects, and the landscape-divergence hypothesis.

Edge effects are major drivers of change in many fragmented landscapes, but are often highly variable in space and time. Here we assess variability in edge effects altering Amazon forest dynamics, plant community composition, invading species, and carbon storage, in the world's largest and longest-running experimental study of habitat fragmentation. Despite detailed knowledge of local landscape conditions, spatial variability in edge effects was only partially foreseeable: relatively predictable effects were caused by the differing proximity of plots to forest edge and varying matrix vegetation, but windstorms generated much random variability. Temporal variability in edge phenomena was also only partially predictable: forest dynamics varied somewhat with fragment age, but also fluctuated markedly over time, evidently because of sporadic droughts and windstorms. Given the acute sensitivity of habitat fragments to local landscape and weather dynamics, we predict that fragments within the same landscape will tend to converge in species composition, whereas those in different landscapes will diverge in composition. This 'landscape-divergence hypothesis', if generally valid, will have key implications for biodiversity-conservation strategies and for understanding the dynamics of fragmented ecosystems.

Rapid decay of tree-community composition in Amazonian forest fragments.

Forest fragmentation is considered a greater threat to vertebrates than to tree communities because individual trees are typically long-lived and require only small areas for survival. Here we show that forest fragmentation provokes surprisingly rapid and profound alterations in Amazonian tree-community composition. Results were derived from a 22-year study of exceptionally diverse tree communities in 40 1-ha plots in fragmented and intact forests, which were sampled repeatedly before and after fragment isolation. Within these plots, trajectories of change in abundance were assessed for 267 genera and 1,162 tree species. Abrupt shifts in floristic composition were driven by sharply accelerated tree mortality and recruitment within approximately 100 m of fragment margins, causing rapid species turnover and population declines or local extinctions of many large-seeded, slow-growing, and old-growth taxa; a striking increase in a smaller set of disturbance-adapted and abiotically dispersed species; and significant shifts in tree size distributions. Even among old-growth trees, species composition in fragments is being restructured substantially, with subcanopy species that rely on animal seed-dispersers and have obligate outbreeding being the most strongly disadvantaged. These diverse changes in tree communities are likely to have wide-ranging impacts on forest architecture, canopy-gap dynamics, plant-animal interactions, and forest carbon storage.

Can neutral theory predict the responses of Amazonian tree communities to forest fragmentation?

We use Hubbell's neutral theory to predict the impact of habitat fragmentation on Amazonian tree communities. For forest fragments isolated for about two decades, we generate neutral predictions for local species extinction, changes in species composition within fragments, and increases in the probability that any two trees within a fragment are conspecific. We tested these predictions using fragment and intact forest data from the Biological Dynamics of Forest Fragments Project in central Amazonia. To simulate complete demographic isolation, we excluded immigrants--species absent from a fragment or intact forest plot in the initial census but present in its last census--from our tests. The neutral theory correctly predicted the rate of species extinction from different plots as a function of the diversity and mortality rate of trees in each plot. However, the rate of change in species composition was much faster than predicted in fragments, indicating that different tree species respond differently to environmental changes. This violates the key assumption of neutral theory. When immigrants were included in our calculations, they increased the disparity between predicted and observed changes in fragments. Overall, neutral theory accurately predicted the pace of local extinctions in fragments but consistently underestimated changes in species composition.

Effects of the surrounding matrix on tree recruitment in Amazonian forest fragments.

Little is known about how the surrounding modified matrix affects tree recruitment in fragmented forests. We contrasted effects of two different matrix types, Vismia- and Cecropia-dominated regrowth, on recruitment of pioneer tree species in forest fragments in central Amazonia. Our analyses were based on 22, 1-ha plots in seven experimental forest fragments ranging in size from 1 to 100 ha. By 13 to 17 years after fragmentation, the population density of pioneer trees was significantly higher in plots surrounded by Vismia regrowth than in plots surrounded by Cecropia regrowth, and the species composition and dominance of pioneers differed markedly between the two matrix types. Cecropia sciadophylla was the most abundant pioneer in fragments surrounded by Cecropia regrowth (constituting nearly 50% of all pioneer trees), whereas densities of species in Vismia-surrounded fragments were distributed more evenly. Thus the surrounding matrix had a strong influence on patterns of tree recruitment in Amazonian forest fragments.

Rain forest fragmentation and the proliferation of successional trees.

The effects of habitat fragmentation on diverse tropical tree communities are poorly understood. Over a 20-year period we monitored the density of 52 tree species in nine predominantly successional genera (Annona, Bellucia, Cecropia, Croton, Goupia, Jacaranda, Miconia, Pourouma, Vismia) in fragmented and continuous Amazonian forests. We also evaluated the relative importance of soil, topographic, forest dynamic, and landscape variables in explaining the abundance and species composition of successional trees. Data were collected within 66 permanent 1-ha plots within a large (approximately 1000 km2) experimental landscape, with forest fragments ranging from 1 to 100 ha in area. Prior to forest fragmentation, successional trees were uncommon, typically comprising 2-3% of all trees (> or =10 cm diameter at breast height [1.3 m above the ground surface]) in each plot. Following fragmentation, the density and basal area of successional trees increased rapidly. By 13-17 years after fragmentation, successional trees had tripled in abundance in fragment and edge plots and constituted more than a quarter of all trees in some plots. Fragment age had strong, positive effects on the density and basal area of successional trees, with no indication of a plateau in these variables, suggesting that successional species could become even more abundant in fragments over time. Nonetheless, the 52 species differed greatly in their responses to fragmentation and forest edges. Some disturbance-favoring pioneers (e.g., Cecropia sciadophylla, Vismia guianensis, V. amazonica, V. bemerguii, Miconia cf. crassinervia) increased by >1000% in density on edge plots, whereas over a third (19 of 52) of all species remained constant or declined in numbers. Species responses to fragmentation were effectively predicted by their median growth rate in nearby intact forest, suggesting that faster-growing species have a strong advantage in forest fragments. An ordination analysis revealed three main gradients in successional-species composition across our study area. Species gradients were most strongly influenced by the standlevel rate of tree mortality on each plot and by the number of nearby forest edges. Species-composition also varied significantly among different cattle ranches, which differed in their surrounding matrices and disturbance histories. These same variables were also the best predictors of total successional-tree abundance and species richness. Successional-tree assemblages in fragment interior plots (>150 m from edge), which are subjected to fragment area effects but not edge effects, did not differ significantly from those in intact forest, indicating that area effects per se had little influence on successional trees. Soils and topography also had little discernable effect on these species. Collectively, our results indicate that successional-tree species proliferate rapidly in fragmented Amazonian forests, largely as a result of chronically elevated tree mortality near forest edges and possibly an increased seed rain from successional plants growing in nearby degraded habitats. The proliferation of fast-growing successional trees and correlated decline of old-growth trees will have important effects on species composition, forest dynamics, carbon storage, and nutrient cycling in fragmented forests.

Efeitos de área e de borda sobre a estrutura florestal em fragmentos de floresta de terra-firme após 13-17 anos de isolamento / Area and edge effects on forest structure in Amazonian forest fragments after 13-17 years of isolation

As estimativas de densidade e biomassa de árvores vivas com DAP > 10 cm e arvoretas 1-9,9 cm de DAP, liteira lenhosa grossa caída (LCG diâmetro > 10 cm), árvores mortas em pé (> 10 de DAP) e liteira lenhosa fina caída (LCF - 2,5 9,9 cm de diâmetro) foram quantificadas em 56 parcelas permanentes de 1 ha, distribuídas em quatro categorias de tamanho de fragmento - fragmentos de 1 ha (4 parcelas), fragmentos de 10 ha (12 parcelas) e fragmentos de 100 ha (14 parcelas) e floresta contínua (19 parcelas) e em duas classes de distância da borda - < 300 m de distância da borda (29 parcelas) e > 300 m (21 parcelas). A densidade e a biomassa de árvores e arvoretas de espécies de estágios sucessionais mais avançados não diferiram significativamente entre as diferentes categorias de tamanho e entre as duas distâncias da borda. Por outro lado, fragmentos florestais e locais < 300 m de distâcia da borda tiveram maior biomassa e densidade de árvores e arvoretas de espécies pioneiras do que floresta contínua e locais > 300 m da borda, respectivamente. Fragmentos florestais apresentaram maior quantidade de LCG e LCF do que a floresta contínua. Houve também diferenças significativas entre ambas as distâncias da borda para a quantidade de LCG e LCF e necromassa total. Uma análise de covariância mostrou que não houve efeito de tamanho do fragmento, mas a distância da borda teve um efeito significativo sobre a quantidade de LCG e LCF. A quantidade de LCG e LCF foi correlacionada negativamente com a distância da borda - locais mais próximos à borda tiveram cerca de 40 por cento e 60 por cento mais LCG do que locais mais distantes.

Density and biomass of live trees >10 cm DBH and saplings 1-9.9 cm DBH, coarse woody debris (LCG diameter > 10 cm), fine woody debris (LCF diameter 2.5-9.9 cm), and standing dead trees (> 10 cm DBH) were quantified in 56 permanent, 1-ha sample plots. These plots are located in four 1- (4 plots), three 10- (12 plots) and two 100- (14 plots) forest fragments in size and nearby continuous forests (19 plots) as well as in two classes of distance from the edges - < 300 m (29 plots) and > 300 m (21 plots). Density and biomass of primary species did not differ significantly among the four size categories and the two edge distance classes. However, forest fragments and distance < 300 m from the edges had more biomass and density of pioneer trees and saplings than did continuous forest and distance > 300 m from the edge, respectively. There were no significant differences among the size categories for standing dead trees. Forest fragments, however, had more quantity of LCG and LCF than did continuous forests. Moreover, distances < 300 m from the edges had higher quantity of LCG and LCF and total necromass than did distances > 300 m. We performed an ANCOVA to assess whether differences in LCG and LCF in fragments were due to proximity of forest borders. An ANCOVA showed that there was no significant effect of fragment size on necromass, but a significant effect of edge distance on both LCG and LCF. The quantity of LCG and LCF was correlated negatively with edge distance sites close to the edge presented over 40-60 percent more LCG than sites far from the edges in both forest fragments and continuous forests.

Biomass change in an Atlantic tropical moist forest: the ENSO effect in permanent sample plots over a 22-year period.

There are a number of controversies surrounding both biomass estimation and carbon balance in tropical forests. Here we use long-term (from 1978 through 2000) data from five 0.5-ha permanent sample plots (PSPs) within a large tract of relatively undisturbed Atlantic moist forest in southeastern Brazil to quantify the biomass increment (DeltaM(I)), and change in total stand biomass (DeltaM(stand)), from mortality, recruitment, and growth data for trees >/=10 cm diameter at breast height (DBH). Despite receiving an average of only 1,200 mm annual precipitation, total forests biomass (334.5+/-11.3 Mg ha(-1)) was comparable to moist tropical forests with much greater precipitation. Over this relatively long-term study, forest biomass experienced rapid declines associated with El Niño events, followed by gradual biomass accumulation. Over short time intervals that overlook extreme events, these dynamics can be misinterpreted as net biomass accumulation. However for the 22 years of this study, there was a small reduction in forest biomass, averaging -1.2 Mg ha(-1) year(-1) (+/-3.1). Strong climatic disturbances can severely reduce forest biomass, and if the frequency and intensity of these events increases beyond historical averages, these changing disturbance regimes have the capacity to significantly reduce forest biomass, resulting in a net source of carbon to the atmosphere.

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.