Resolving whole-plant economics from leaf, stem and root traits of 1467 Amazonian tree species
It remains unclear how evolutionary and ecological processes have shaped the wide variety of plant life strategies, especially in highly diverse ecosystems like tropical forests. Some evidence suggests that species have diversified across a gradient of ecological strategies, with different plant tissues converging to optimize resource use across environmental gradients. Alternative hypotheses propose that species have diversified following independent selection on different tissues, resulting in a decoupling of trait syndromes across organs. To shed light on the subject, we assembled an unprecedented dataset combining 19 leaf, stem and root traits for 1467 tropical tree species inventoried across 71 0.1-ha plots spanning broad environmental gradients in French Guiana. Nearly 50% of the overall functional heterogeneity was expressed along four orthogonal dimensions, after accounting for phylogenetic dependences among species. The first dimension related to fine root functioning, while the second and third dimensions depicted two decoupled leaf economics spectra, and the fourth dimension encompassed a wood economics spectrum. Traits involved in orthogonal functional strategies, five leaf traits in particular but also trunk bark thickness, were consistently associated with a same gradient of soil texture and nutrient availability. Root traits did not show any significant association with edaphic variation, possibly because of the prevailing influence of other factors (mycorrhizal symbiosis, phylogenetic constraints). Our study emphasises the existence of multiple functional dimensions that allow tropical tree species to optimize their performance in a given environment, bringing new insights into the debate around the presence of a whole plant economic spectrum in tropical forest tree communities. It also emphasizes the key role that soil heterogeneity plays in shaping tree species assembly. The extent to which different organs are decoupled and respond to environmental gradients may also help to improve our predictions of species distribution changes in responses to habitat modification and environmental changes.
Environmental factors predict community functional composition in Amazonian forests
1. The consequences of biodiversity loss for ecosystem services largely depend on the functional identities of extirpated species. However, poor descriptions of spatial patterns of community functional composition across landscapes hamper accurate predictions, particularly in highly diverse tropical regions. Therefore, understanding how community functional composition varies across environmental gradients remains an important challenge.
2. We sampled 15 functional traits in 800 Neotropical tree species across 13 forest plots representative of the broad climatic and soil gradients encompassed by three widespread lowland forest habitats (terra firme forests on clay-rich soils, seasonally flooded forests and white-sand forests) at opposite ends of Amazonia (Peru and French Guiana). We combined univariate and multivariate approaches to test the magnitude and predictability of environmental filtering on community leaf and wood functional composition.
3. Directional shifts in community functional composition correlated with environmental changes across the 13 plots, with denser leaves, stems and roots in forests occurring in environments with limited water and soil-nutrient availability. Critically, these relationships allowed us to accurately predict the functional composition of 61 additional forest plots from environmental data alone.
4. Synthesis. Environmental filtering consistently shapes the functional composition of highly diverse tropical forests at large scales across the terra firme, seasonally flooded and white-sand forests of lowland Amazonia. Environmental factors drive and allow the prediction of variation in community functional composition among habitat types in Amazonian forests.
Biogeographic history and habitat specialization shape floristic and phylogenetic composition across Amazonian forests
A major challenge remains to understand the relative contributions of history, dispersal, and environmental filtering to the assembly of hyperdiverse communities across spatial scales. Here, we examine the extent to which biogeographical history and habitat specialization have generated turnover among and within lineages of Amazonian trees across broad geographic and environmental gradients. We replicated standardized tree inventories in 102 0.1-ha plots located in two distant regions-the western Amazon and the eastern Guiana shield. Within each region, we used a nested design to replicate plots on contrasted habitats: white-sand, terra firme, and seasonally flooded forests. Our plot network encompassed 26,386 trees that together represented 2,745 distinct taxa, which we standardized across all plots and regions. We combined taxonomic and phylogenetic data with detailed soil measurements and climatic data to: (1) test whether patterns of taxonomic and phylogenetic composition are consistent with recent or historical processes, (2) disentangle the relative effects of habitat, environment, and geographic distance on taxonomic and phylogenetic turnover among plots, and (3) contrast the proportion of habitat specialists among species from each region. We found substantial species turnover between Peru and French Guiana, with only 8.8% of species shared across regions; genus composition remained differentiated across habitats and regions, whereas turnover at higher taxonomic levels (family, order) was much lower. Species turnover across plots was explained primarily by regions, but also substantially by habitat differences and to a lesser extent by spatial distance within regions. Conversely, the composition of higher taxonomic levels was better explained by habitats (especially comparing white-sand forests to other habitats) than spatial distance. White-sand forests harbored most of the habitat specialists in both regions, with stronger habitat specialization in Peru than in French Guiana. Our results suggest that recent diversification events have resulted in extremely high turnover in species and genus composition with relatively little change in the composition of higher lineages. Our results also emphasize the contributions of rare habitats, such as white-sand forests, to the extraordinary diversity of the Amazon and underline their importance as conservation priorities.
Phylogenetic density dependence and environmental filtering predict seedling mortality in a tropical forest
Negative density dependence (NDD) and environmental filtering (EF) shape community assembly, but their relative importance is poorly understood. Recent studies have shown that seedling's mortality risk is positively related to the phylogenetic relatedness of neighbours. However, natural enemies, whose depredations often cause NDD, respond to functional traits of hosts rather than phylogenetic relatedness per se. To understand the roles of NDD and EF in community assembly, we assessed the effects on seedling mortality of functional similarity, phylogenetic relatedness and stem density of neighbouring seedlings and adults in a species?rich tropical forest. Mortality risks increased for common species when their functional traits departed substantially from the neighbourhood mean, and for all species when surrounded by close relatives. This indicates that NDD affects community assembly more broadly than does EF, and leads to the tentative conclusion that natural enemies respond to phylogenetically correlated traits. Our results affirm the prominence of NDD in structuring species-rich communities.
Amazon tree dominance across forest strata
The forests of Amazonia are among the most biodiverse plant communities on Earth. Given the immediate threats posed by climate and land-use change, an improved understanding of how this extraordinary biodiversity is spatially organized is urgently required to develop effective conservation strategies. Most Amazonian tree species are extremely rare but a few are common across the region. Indeed, just 227 'hyperdominant' species account for >50% of all individuals >10 cm diameter at 1.3 m in height. Yet, the degree to which the phenomenon of hyperdominance is sensitive to tree size, the extent to which the composition of dominant species changes with size class and how evolutionary history constrains tree hyperdominance, all remain unknown. Here, we use a large floristic dataset to show that, while hyperdominance is a universal phenomenon across forest strata, different species dominate the forest understory, midstory and canopy. We further find that, although species belonging to a range of phylogenetically dispersed lineages have become hyperdominant in small size classes, hyperdominants in large size classes are restricted to a few lineages. Our results demonstrate that it is essential to consider all forest strata to understand regional patterns of dominance and composition in Amazonia. More generally, through the lens of 654 hyperdominant species, we outline a tractable pathway for understanding the functioning of half of Amazonian forests across vertical strata and geographical locations.
Globally, functional traits are weak predictors of juvenile tree growth, and we do not know why
1. Plant functional traits, in particular specific leaf area (SLA), wood density and seed mass, are often good predictors of individual tree growth rates within communities. Individuals and species with high SLA, low wood density and small seeds tend to have faster growth rates. 2. If community-level relationships between traits and growth have general predictive value, then similar relationships should also be observed in analyses that integrate across taxa, biogeographic regions and environments. Such global consistency would imply that traits could serve as valuable proxies for the complex suite of factors that determine growth rate, and, therefore, could underpin anew generation of robust dynamic vegetation models. Alternatively, growth rates may depend more strongly on the local environment or growth-trait relationships may vary along environmental gradients. 3. We tested these alternative hypotheses using data on 27 352 juvenile trees, representing 278 species from 27 sites on all forested continents, and extensive functional trait data, 38% of which were obtained at the same sites at which growth was assessed. Data on potential evapotranspiration (PET), which summarizes the joint ecological effects of temperature and precipitation, were obtained from a global data base. 4. We estimated size-standardized relative height growth rates (SGR) for all species, then related them to functional traits and PET using mixed-effect models for the fastest growing species and for all species together. 5. Both the mean and 95th percentile SGR were more strongly associated with functional traits than with PET. PET was unrelated to SGR at the global scale. SGR increased with increasing SLA and decreased with increasing wood density and seed mass, but these traits explained only 3.1% of the variation in SGR. SGR-trait relationships were consistently weak across families and biogeographic zones, and over a range of tree statures. Thus, the most widely studied functional traits in plant ecology were poor predictors of tree growth over large scales. 6. Synthesis.We conclude that these functional traits alone may be unsuitable for predicting growth of trees over broad scales. Determining the functional traits that predict vital rates under specific environmental conditions may generate more insight than a monolithic global relationship can offer.
Environmental factors predict community functional composition in Amazonian forests
1. The consequences of biodiversity loss for ecosystem services largely depend on the functional identities of extirpated species. However, poor descriptions of spatial patterns of community functional composition across landscapes hamper accurate predictions, particularly in highly diverse tropical regions. Therefore, understanding how community functional composition varies across environmental gradients remains an important challenge. 2. We sampled 15 functional traits in 800 Neotropical tree species across 13 forest plots representative of the broad climatic and soil gradients encompassed by three widespread lowland forest habitats (terra firme forests on clay-rich soils, seasonally flooded forests and white-sand forests) at opposite ends of Amazonia (Peru and French Guiana). We combined univariate and multivariate approaches to test the magnitude and predictability of environmental filtering on community leaf and wood functional composition. 3. Directional shifts in community functional composition correlated with environmental changes across the 13 plots, with denser leaves, stems and roots in forests occurring in environments with limited water and soil-nutrient availability. Critically, these relationships allowed us to accurately predict the functional composition of 61 additional forest plots from environmental data alone. 4. Synthesis. Environmental filtering consistently shapes the functional composition of highly diverse tropical forests at large scales across the terra firme, seasonally flooded and white-sand forests of lowland Amazonia. Environmental factors drive and allow the prediction of variation in community functional composition among habitat types in Amazonian forests.
There's no place like home: seedling mortality contributes to the habitat specialisation of tree species across Amazonia
Understanding the mechanisms generating species distributions remains a challenge, especially in hyperdiverse tropical forests. We evaluated the role of rainfall variation, soil gradients and herbivory on seedling mortality, and how variation in seedling performance along these gradients contributes to habitat specialisation. In a 4-year experiment, replicated at the two extremes of the Amazon basin, we reciprocally transplanted 4638 tree seedlings of 41 habitat-specialist species from seven phylogenetic lineages among the three most important forest habitats of lowland Amazonia. Rainfall variation, flooding and soil gradients strongly influenced seedling mortality, whereas herbivory had negligible impact. Seedling mortality varied strongly among habitats, consistent with predictions for habitat specialists in most lineages. This suggests that seedling performance is a primary determinant of the habitat associations of adult trees across Amazonia. It further suggests that tree diversity, currently mostly harboured in terra firme forests, may be strongly impacted by the predicted climate changes in Amazonia.