Claire Fortunel

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 seedlings 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.

Environmental factors predict community functional composition in Amazonian forests

Summary 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.

Rare species contribute disproportionately to the functional structure of species assemblages.

There is broad consensus that the diversity of functional traits within species assemblages drives several ecological processes. It is also widely recognized that rare species are the first to become extinct following human-induced disturbances. Surprisingly, however, the functional importance of rare species is still poorly understood, particularly in tropical species-rich assemblages where the majority of species are rare, and the rate of species extinction can be high. Here, we investigated the consequences of local and regional extinctions on the functional structure of species assemblages. We used three extensive datasets (stream fish from the Brazilian Amazon, rainforest trees from French Guiana, and birds from the Australian Wet Tropics) and built an integrative measure of species rarity versus commonness, combining local abundance, geographical range, and habitat breadth. Using different scenarios of species loss, we found a disproportionate impact of rare species extinction for the three groups, with significant reductions in levels of functional richness, specialization, and originality of assemblages, which may severely undermine the integrity of ecological processes. The whole breadth of functional abilities within species assemblages, which is disproportionately supported by rare species, is certainly critical in maintaining ecosystems particularly under the ongoing rapid environmental transitions.

Globally, functional traits are weak predictors of juvenile tree growth, and we do not know why

Additional co-authors: Herve Jactel, Xuefei Li, Kaoru Kitajima, Julia Koricheva, Cristina Martinez-Garza, Christian Messier, Alain Paquette, Christopher Philipson, Daniel Piotto, Lourens Poorter, Juan M. Posada, Catherine Potvin, Kalle Rainio, Sabrina E. Russo, Mariacarmen Ruiz-Jaen, Michael Scherer-Lorenzen, Campbell O. Webb, S. Joseph Wright, Rakan A. Zahawi, and Andy Hector

Herbivory, growth rates, and habitat specialization in tropical tree lineages: implications for Amazonian beta-diversity

Tropical plant diversity is extraordinarily high at both local and regional scales. Many studies have demonstrated that natural enemies maintain local diversity via negative density dependence, but we know little about how natural enemies influence beta-diversity across habitats and/or regions. One way herbivores could influence plant beta-diversity is by driving allocation trade-offs that promote habitat specialization across resource gradients. We therefore predicted that increasing resource availability should be accompanied by increasing herbivory rates and decreasing plant allocation to defense. Second, relative abundances within plant lineages are predicted to reflect patterns of habitat specialization and allocation trade- offs. A phylogenetic context is vital not only to compare homologous plant traits (including defense strategies) across habitat types, but also to connect evolutionary trade-offs to patterns of species diversification in each phylogenetic lineage. We tested these predictions for trees in white-sand, clay terra firme, and seasonally flooded forests in Peru and French Guiana that represent the range of soil fertility, forest structure, and floristic compositions found throughout the Amazon region. We established 74 0.5-ha plots in these habitats and sampled all trees. Within 12 representative plots we marked newly expanding leaves of 394 saplings representing 68 species, including the most abundant species in each plot in addition to species from five focal lineages:SwartziaandInga(Fabaceae), Protieae (Burseracaeae), Bombacoideae (Malvaceae), andMicropholis(Sapotaceae). We measured total leaf production rates for each sapling and calculated relative herbivory impact as the ratio between herbivory rate and leaf production rate. Herbivory rates averaged 2.1%per month, did not correlate with leaf production rate, and were similar across habitats. Relative herbivore impacts exceeded leaf production rates for most species. Leaf production rate averaged 2.8%, was significantly higher in seasonally flooded forests than the other two habitats, and exhibited significant correlations with specific leaf area. Species with high herbivory rates exhibited significantly lower relative abundances in terra firme forests. Overall, focal species within lineages present contrasting patterns regarding their herbivory rates and leaf production rate within habitats. These results highlight why a lineage-based approach is necessary when attempting to connect hypotheses regarding evolutionary trade-offs to community assembly patterns.

Wood specific gravity and anatomy of branches and roots in 113 Amazonian rainforest tree species across environmental gradients

Wood specific gravity (WSG) is a strong predictor of tree performance across environmental gradients. Yet it remains unclear how anatomical elements linked to different wood functions contribute to variation in WSG in branches and roots across tropical forests. We examined WSG and wood anatomy in white sand, clay terra firme and seasonally flooded forests in French Guiana, spanning broad environmental gradients found throughout Amazonia. We measured 15 traits relating to branches and small woody roots in 113 species representing the 15 most abundant species in each habitat and representative species from seven monophyletic lineages occurring in all habitats. Fiber traits appear to be major determinants of WSG, independent of vessel traits, in branches and roots. Fiber traits and branch and root WSG increased from seasonally flooded species to clay terra firme species and lastly to white sand species. Branch and root wood traits were strongly phylogenetically constrained. Lineages differed in wood design, but exhibited similar variation in wood structure across habitats. We conclude that tropical trees can invest differently in support and transport to respond to environmental conditions. Wind disturbance and drought stress represent significant filters driving tree distribution of Amazonian forests; hence we suggest that biophysical explanations should receive more attention.

Functional trait differences influence neighbourhood interactions in a hyperdiverse Amazonian forest

As distinct community assembly processes can produce similar community patterns, assessing the ecological mechanisms promoting coexistence in hyperdiverse rainforests remains a considerable challenge. We use spatially explicit neighbourhood models of tree growth to quantify how functional trait and phylogenetic similarities predict variation in growth and crowding effects for the 315 most abundant tree species in a 25-ha lowland rainforest plot in Ecuador. We find that functional trait differences reflect variation in (1) species maximum potential growth, (2) the intensity of interspecific interactions for some species, and (3) species sensitivity to neighbours. We find that neighbours influenced tree growth in 28% of the 315 focal tree species. Neighbourhood effects are not detected in the remaining 72%, which may reflect the low statistical power to model rare taxa and/or species insensitivity to neighbours. Our results highlight the spectrum of ways in which functional trait differences can shape community dynamics in highly diverse rainforests.

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.

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.

Geographical Variation in Community Divergence: Insights from Tropical Forest Monodominance by Ectomycorrhizal Trees*

Convergence occurs in both species traits and community structure, but how convergence at the two scales influences each other remains unclear. To address this question, we focus on tropical forest monodominance, in which a single, often ectomycorrhizal (EM) tree species occasionally dominates forest stands within a landscape otherwise characterized by diverse communities of arbuscular mycorrhizal (AM) trees. Such monodominance is a striking potential example of community divergence resulting in alternative stable states. However, it is observed only in some tropical regions. A diverse suite of AM and EM trees locally codominate forest stands elsewhere. We develop a hypothesis to explain this geographical difference using a simulation model of plant community assembly. Simulation results suggest that in a region with a few EM species (e.g., South America), EM trees experience strong selection for convergent traits that match the abiotic conditions of the environment. Consequently, EM species successfully compete against other species to form monodominant stands via positive plant-soil feedbacks. By contrast, in a region with many EM species (e.g., Southeast Asia), species maintain divergent traits because of complex plant-soil feedbacks, with no species having traits that enable monodominance. An analysis of plant trait data from Borneo and Peruvian Amazon was inconclusive. Overall, this work highlights the utility of geographical comparison in understanding the relationship between trait convergence and community convergence.