Giants of the Amazon: How does environmental variation drive the diversity patterns of large trees?

For more than three decades, major efforts in sampling and analyzing tree diversity in South America have focused almost exclusively on trees with stems of at least 10 and 2.5 cm diameter, showing highest species diversity in the wetter western and northern Amazon forests. By contrast, little attention has been paid to patterns and drivers of diversity in the largest canopy and emergent trees, which is surprising given these have dominant ecological functions. Here, we use a machine learning approach to quantify the importance of environmental factors and apply it to generate spatial predictions of the species diversity of all trees (dbh ≥ 10 cm) and for very large trees (dbh ≥ 70 cm) using data from 243 forest plots (108,450 trees and 2832 species) distributed across different forest types and biogeographic regions of the Brazilian Amazon. The diversity of large trees and of all trees was significantly associated with three environmental factors, but in contrasting ways across regions and forest types. Environmental variables associated with disturbances, for example, the lightning flash rate and wind speed, as well as the fraction of photosynthetically active radiation, tend to govern the diversity of large trees. Upland rainforests in the Guiana Shield and Roraima regions had a high diversity of large trees. By contrast, variables associated with resources tend to govern tree diversity in general. Places such as the province of Imeri and the northern portion of the province of Madeira stand out for their high diversity of species in general. Climatic and topographic stability and functional adaptation mechanisms promote ideal conditions for species diversity. Finally, we mapped general patterns of tree species diversity in the Brazilian Amazon, which differ substantially depending on size class.

Evolutionary heritage influences Amazon tree ecology.

Lineages tend to retain ecological characteristics of their ancestors through time. However, for some traits, selection during evolutionary history may have also played a role in determining trait values. To address the relative importance of these processes requires large-scale quantification of traits and evolutionary relationships among species. The Amazonian tree flora comprises a high diversity of angiosperm lineages and species with widely differing life-history characteristics, providing an excellent system to investigate the combined influences of evolutionary heritage and selection in determining trait variation. We used trait data related to the major axes of life-history variation among tropical trees (e.g. growth and mortality rates) from 577 inventory plots in closed-canopy forest, mapped onto a phylogenetic hypothesis spanning more than 300 genera including all major angiosperm clades to test for evolutionary constraints on traits. We found significant phylogenetic signal (PS) for all traits, consistent with evolutionarily related genera having more similar characteristics than expected by chance. Although there is also evidence for repeated evolution of pioneer and shade tolerant life-history strategies within independent lineages, the existence of significant PS allows clearer predictions of the links between evolutionary diversity, ecosystem function and the response of tropical forests to global change.

Tropical forest lianas have greater non-structural carbohydrate concentrations in the stem xylem than trees.

Lianas (woody vines) are important components of tropical forests and are known to compete with host trees for resources, decrease tree growth and increase tree mortality. Given the observed increases in liana abundance in some forests and their impacts on forest function, an integrated understanding of carbon dynamics of lianas and liana-infested host trees is critical for improved prediction of tropical forest responses to climate change. Non-structural carbohydrates (NSC) are the main substrate for plant metabolism (e.g., growth, respiration), and have been implicated in enabling tree survival under environmental stress, but little is known of how they vary among life-forms or of how liana infestation impacts host tree NSC. We quantified stem total NSC (NSC) concentrations and its fractions (starch and soluble sugars) in trees without liana infestation, trees with more than 50% of the canopy covered by lianas, and the lianas infesting those trees. We hypothesized that i) liana infestation depletes NSC storage in host trees by reducing carbon assimilation due to competition for resources; ii) trees and lianas, which greatly differ in functional traits related to water transport and carbon uptake, would also have large differences in NSC storage, and that As water availability has a significant role in NSC dynamics of Amazonian tree species, we tested these hypotheses within a moist site in western Amazonia and a drier forest site in southern Amazonia. We did not find any difference in NSC, starch or soluble sugar concentrations between infested and non-infested trees, in either site. This result suggests that negative liana impact on trees may be mediated through mechanisms other than depletion of host tree NSC concentrations. We found lianas have higher stem NSC and starch than trees in both sites. The consistent differences in starch concentrations, a long term NSC reserve, between life forms across sites reflect differences in carbon gain and use of lianas and trees. Soluble sugar concentrations were higher in lianas than in trees in the moist site but indistinguishable between life forms in the dry site. The lack of difference in soluble sugars between trees and lianas in the dry site emphasize the importance of this NSC fraction for plant metabolism of plants occurring in water limited environments. Abstract in Portuguese and Spanish are available in the supplementary material.

Functional susceptibility of tropical forests to climate change.

Tropical forests are some of the most biodiverse ecosystems in the world, yet their functioning is threatened by anthropogenic disturbances and climate change. Global actions to conserve tropical forests could be enhanced by having local knowledge on the forests' functional diversity and functional redundancy as proxies for their capacity to respond to global environmental change. Here we create estimates of plant functional diversity and redundancy across the tropics by combining a dataset of 16 morphological, chemical and photosynthetic plant traits sampled from 2,461 individual trees from 74 sites distributed across four continents together with local climate data for the past half century. Our findings suggest a strong link between climate and functional diversity and redundancy with the three trait groups responding similarly across the tropics and climate gradient. We show that drier tropical forests are overall less functionally diverse than wetter forests and that functional redundancy declines with increasing soil water and vapour pressure deficits. Areas with high functional diversity and high functional redundancy tend to better maintain ecosystem functioning, such as aboveground biomass, after extreme weather events. Our predictions suggest that the lower functional diversity and lower functional redundancy of drier tropical forests, in comparison with wetter forests, may leave them more at risk of shifting towards alternative states in face of further declines in water availability across tropical regions.

Examining variation in the leaf mass per area of dominant species across two contrasting tropical gradients in light of community assembly.

Understanding variation in key functional traits across gradients in high diversity systems and the ecology of community changes along gradients in these systems is crucial in light of conservation and climate change. We examined inter- and intraspecific variation in leaf mass per area (LMA) of sun and shade leaves along a 3330-m elevation gradient in Peru, and in sun leaves across a forest-savanna vegetation gradient in Brazil. We also compared LMA variance ratios (T-statistics metrics) to null models to explore internal (i.e., abiotic) and environmental filtering on community structure along the gradients. Community-weighted LMA increased with decreasing forest cover in Brazil, likely due to increased light availability and water stress, and increased with elevation in Peru, consistent with the leaf economic spectrum strategy expected in colder, less productive environments. A very high species turnover was observed along both environmental gradients, and consequently, the first source of variation in LMA was species turnover. Variation in LMA at the genus or family levels was greater in Peru than in Brazil. Using dominant trees to examine possible filters on community assembly, we found that in Brazil, internal filtering was strongest in the forest, while environmental filtering was observed in the dry savanna. In Peru, internal filtering was observed along 80% of the gradient, perhaps due to variation in taxa or interspecific competition. Environmental filtering was observed at cloud zone edges and in lowlands, possibly due to water and nutrient availability, respectively. These results related to variation in LMA indicate that biodiversity in species rich tropical assemblages may be structured by differential niche-based processes. In the future, specific mechanisms generating these patterns of variation in leaf functional traits across tropical environmental gradients should be explored.

Resilience of savanna forest after clear-cutting in the cerrado-amazon transition zone / Resiliência de cerradão após corte raso da vegetação na transição cerrado-amazônia

The dynamics of the natural process of recuperation of the structure and diversity of native vegetation following anthropogenic disturbance has been the subject of a great deal of controversy in restoration ecology research. The present study evaluates the natural regeneration of savanna forest (cerradão) 32 and 36 years after the clearcutting of the vegetation. We compared species diversity, and the structure and dynamics of the vegetation in two communities, one representing preserved cerradão (PC), and the other, the regenerating cerradão (RC), which was clearcut in 1976. Surveys were conducted in 2008 and 2012, 32 and 36 years after clear-cutting, respectively. In 2008, we demarcated 81 permanent 10 m x 10 m plots, 50 in the RC and 31 in the PC, and measured all live plants with a diameter at soil level > 5 cm. In 2012, the plots were resampled, including the original plants and all the recruits. The species were classified as specialists in savanna (SA) or forest habitat (FO), or as generalists (SA/FO). The RC presented the highest species richness and diversity, density, annual increment, and mortality rates. However, no significant differences were found between communities in the distribution of specialist or generalist species, or between years (2008 and 2012) in basal area or recruitment rates. While the species composition of the two communities is highly similar, the RC was characterized by a higher frequency of SA species, and was more similar to nearby savanna communities (cerrado sensu stricto). Trees in the RC were smaller and suffered higher rates of mortality than those in the PC, but also higher annual increments. While the RC demonstrated a high degree of resilience following clear-cutting, it was still found to be at an intermediate stage of succession, even after almost four decades, indicating that regeneration is a slow process.

A dinâmica de recuperação da diversidade e estrutura da vegetação nativa, a partir de processos naturais, após distúrbios antrópicos, tem sido objeto de controvérsias em estudos de ecologia da restauração. Nosso objetivo foi avaliar a regeneração natural de cerradão em 32 e 36 anos, após corte raso da vegetação. Comparamos os parâmetros florísticos e estruturais e a dinâmica da vegetação (entre 2008 e 2012) de duas comunidades de cerradão, sendo uma de cerradão preservado (CP) e outra em regeneração desde 1976 (CR). Demarcamos em 2008, 81 parcelas permanentes (10 x 10 m), sendo 50 no CR e 31 no CP e medimos todas as plantas vivas com diâmetro a altura do solo > 5 cm. Em 2012, reamostramos todas as plantas e incluímos os recrutas. Classificamos as espécies como especialistas em habitats savânicos (SA), florestal (FO) ou generalistas em habitats savânicos e florestais (SA/FO). O CR apresentou maior riqueza e diversidade de espécies, densidade de indivíduos, incremento periódico anual e taxa de mortalidade. Porém, as duas áreas não apresentaram diferenças na distribuição das espécies entre os habitats (SA, FO, SA/FO) e área basal (em 2008 e 2012) e taxa de recrutamento (entre 2008 e 2012). Apesar do CR e CP apresentarem elevada similaridade florística, o CR apresentou maior frequência de indivíduos de espécies com habitat SA bem como maior similaridade com duas outras comunidades savânicas (cerrado sentido restrito) próximas. Ainda, o CR possui indivíduos com menores diâmetros e consequentemente maiores taxas de mortalidade e incremento periódico anual. Apesar da resiliência do cerradão em relação ao corte raso esse ainda se encontra em estágio intermediário de sucessão, mesmo quase quatro décadas após o distúrbio, indicando que a regeneração é um processo lento.