Leaf Functional Diversity Research Articles

Diversity-enhanced canopy space occupation and leaf functional diversity jointly promote overyielding in tropical tree communities

Understanding the mechanisms that drive biodiversity-productivity relationships is critical for guiding forest restoration. Although complementarity among trees in the canopy space has been suggested as a key mechanism for greater productivity in mixed-species tree communities, empirical evidence remains limited. Here, we used data from a tropical tree diversity experiment to disentangle the effects of tree species richness and community functional characteristics (community-weighted mean and functional diversity of leaf traits) on canopy space filling, and how these effects are related to overyielding. We found that canopy space filling was largely explained by species identity effects rather than tree diversity effects. Communities with a high abundance of species with a conservative resource-use strategy were those with most densely packed canopies. Across monocultures and mixtures, a higher canopy space filling translated into an enhanced wood productivity. Importantly, most communities (83 %) produced more wood volume than the average of their constituent species in monoculture (i.e. most communities overyielded). Our results show that overyielding increased with leaf functional diversity and positive net biodiversity effects on canopy space filling, which mainly arose due to a high taxonomic diversity. These findings suggest that both taxonomic diversity-enhanced canopy space filling and canopy leaf diversity are important drivers for overyielding in mixed-species forests. Consequently, restoration initiatives should promote stands with functionally diverse canopies by selecting tree species with large interspecific differences in leaf nutrition, as well as leaf and branch morphology to optimize carbon capture in young forest stands.

Leaf functional diversity and environmental filtering in a tropical dry forest: Comparison between two geological substrates.

The role of geological substrate in shaping plant community functional diversity remains poorly understood. Considering the involvement of leaves in the energy, water, and nutrient economics of plants, we hypothesized that leaves experience geology-related filtering, which in turn shapes their functional attributes and community leaf functional diversity on different substrates. We studied tropical dry forest communities on limestone and siliciclastic phyllite-derived soils, comparing their functional diversity and soil physico-chemical properties. We predicted the most benign habitat (less severe filter) to be associated with higher leaf functional diversity and an acquisitive strategy prevalence, while the more stressful habitat should show conservative leaf traits and lower leaf functional diversity. We measured six traits in 31 common tree species (representing ~80% of community crown cover): leaf area, specific leaf area, leaf thickness, leaf dry matter content, petiole length, and leaf blade narrowness. Leaf functional diversity was assessed through the functional trait dispersion metric. Intraspecific functional variation was examined in 25 species shared between substrates. The limestone substrate was more fertile (higher phosphorous) with higher water retention, while phyllite had higher nitrogen and lower humidity. Principal component analysis segregated plots by substrate, with limestone plots being more clustered. Community leaf functional diversity was higher in the limestone forest. Most species examined showed inter-substrate trait differences in at least one leaf functional trait. The two substrates constituted distinct growth environments, with the more benign substrate associated with higher community leaf functional diversity. The intraspecific analysis revealed the prevalence of acquisitive traits in the more benign and more conservative traits in the more stressful habitat. This study advances our understanding of the role of geological substrate as an environmental filter in tropical dry forests, influencing leaf functional responses and emphasizing the importance of intraspecific functional variation.

Biogeography of global drylands.

Despite their extent and socio-ecological importance, a comprehensive biogeographical synthesis of drylands is lacking. Here we synthesize the biogeography of key organisms (vascular and nonvascular vegetation and soil microorganisms), attributes (functional traits, spatial patterns, plant-plant and plant-soil interactions) and processes (productivity and land cover) across global drylands. These areas have a long evolutionary history, are centers of diversification for many plant lineages and include important plant diversity hotspots. This diversity captures a strikingly high portion of the variation in leaf functional diversity observed globally. Part of this functional diversity is associated with the large variation in response and effect traits in the shrubs encroaching dryland grasslands. Aridity and its interplay with the traits of interacting plant species largely shape biogeographical patterns in plant-plant and plant-soil interactions, and in plant spatial patterns. Aridity also drives the composition of biocrust communities and vegetation productivity, which shows large geographical variation. We finish our review by discussing major research gaps, which include: studying regular vegetation spatial patterns; establishing large-scale plant and biocrust field surveys assessing individual-level trait measurements; knowing whether the impacts of plant-plant and plant-soil interactions on biodiversity are predictable; and assessing how elevated CO2 modulates future aridity conditions and plant productivity.

How does leaf functional diversity affect the light environment in forest canopies? An in-silico biodiversity experiment

The interaction of shortwave radiation with vegetation drives basic processes of the biosphere, such as primary productivity, species interactions through light competition, and energy fluxes between the atmosphere, vegetation, and soil. Here, we aim to understand the effects of leaf functional trait diversity on canopy light absorption. We focus on the diversity of three key functional traits that influence the light-canopy interaction: leaf area index (LAI), leaf angle distribution (LAD) and leaf optical properties (LOP). We used a 3D radiative transfer model to perform an in-silico biodiversity experiment to study the effects of leaf functional diversity on a light proxy for productivity (the fraction of absorbed photosynthetically active radiation (FAPAR)) and net radiation (shortwave albedo). We found that diverse canopies had lower albedo and higher FAPAR than the average of the corresponding monoculture values. In mixtures, FAPAR was unequally re-distributed between trees with distinct traits: some plant functional types absorbed more light and some plant functional types absorbed less than in monocultures. The net biodiversity effect on absorptance was greater when combining plant functional types with more distinct leaf traits. Our results support the mechanistic understanding of overyielding effects in functionally diverse canopies and may partially explain some of the growth-promoting mechanisms in biodiversity-ecosystem functioning experiments. They can further help to account for biodiversity effects in climate models.

Leaf traits and canopy structure together explain canopy functional diversity: an airborne remote sensing approach.

Plant functional diversity is strongly connected to photosynthetic carbon assimilation in terrestrial ecosystems. However, many of the plant functional traits that regulate photosynthetic capacity, including foliar nitrogen concentration and leaf mass per area, vary significantly between and within plant functional types and vertically through forest canopies, resulting in considerable landscape-scale heterogeneity in three dimensions. Hyperspectral imagery has been used extensively to quantify functional traits across a range of ecosystems but is generally limited to providing information for top of canopy leaves only. On the other hand, lidar data can be used to retrieve the vertical structure of forest canopies. Because these data are rarely collected at the same time, there are unanswered questions about the effect of forest structure on the three -dimensional spatial patterns of functional traits across ecosystems. In the United States, the National Ecological Observatory Network's Airborne Observation Platform (NEON AOP) provides an opportunity to address this structure-function relationship by collecting lidar and hyperspectral data together across a variety of ecoregions. With a fusion of hyperspectral and lidar data from the NEON AOP and field-collected foliar trait data, we assessed the impacts of forest structure on spatial patterns of N. In addition, we examine the influence of abiotic gradients and management regimes on top-of-canopy percent N and total canopy N (i.e., the total amount of N [g/m2 ] within a forest canopy) at a NEON site consisting of a mosaic of open longleaf pine and dense broadleaf deciduous forests. Our resulting maps suggest that, in contrast to top of canopy values, total canopy N variation is dampened across this landscape resulting in relatively homogeneous spatial patterns. At the same time, we found that leaf functional diversity and canopy structural diversity showed distinct dendritic patterns related to the spatial distribution of plant functional types.

Expanding our understanding of leaf functional syndromes in savanna systems: the role of plant growth form.

The assessment of leaf strategies has been a common theme in ecology, especially where multiple sources of environmental constraints (fire, seasonal drought, nutrient-poor soils) impose a strong selection pressure towards leaf functional diversity, leading to inevitable tradeoffs among leaf traits, and ultimately to niche segregation among coexisting species. As diversification on leaf functional strategies is dependent on integration at whole plant level, we hypothesized that regardless of phylogenetic relatedness, leaf trait functional syndromes in a multivariate space would be associated with the type of growth form. We measured traits related to leaf gas exchange, structure and nutrient status in 57 coexisting species encompassing all Angiosperms major clades, in a wide array of plant morphologies (trees, shrubs, sub-shrubs, herbs, grasses and palms) in a savanna of Central Brazil. Growth forms differed in mean values for the studied functional leaf traits. We extracted 4 groups of functional typologies: grasses (elevated leaf dark respiration, light-saturated photosynthesis on a leaf mass and area basis, lower values of leaf Ca and Mg), herbs (high values of SLA, leaf N and leaf Fe), palms (high values of stomatal conductance, leaf transpiration and leaf K) and woody eudicots (sub-shrubs, shrubs and trees; low SLA and high leaf Ca and Mg). Despite the large range of variation among species for each individual trait and the independent evolutionary trajectory of individual species, growth forms were strongly associated with particular leaf trait combinations, suggesting clear evolutionary constraints on leaf function for morphologically similar species in savanna ecosystems.

Tree diversity promotes predator but not omnivore ants in a subtropical Chinese forest

1. Epigeic ants are functionally important arthropods in tropical and subtropical forests, particularly by acting as predators. High predation pressure has been hypothesised to be a mechanism facilitating high diversity across trophic levels.2. In this study, standardised pitfall traps were used in a highly diverse subtropical forest to test if and how ant species richness is related to tree species richness and a comprehensive set of other environmental variables such as successional age, soil properties or elevation.3. A total of 13 441 ant individuals belonging to 3839 species occurrences and 71 species were collected, of which 26 species were exclusive predators and 45 species were omnivores.4. Occurrence and species richness of total and omnivore ants were positively related to soil pH. Predator ant occurrence was unrelated to all environmental variables tested.5. The species richness of predator ants increased with tree species richness but decreased with leaf functional diversity and shrub cover. Elevation negatively influenced only total ant species richness.6. The evenness of predators increased with tree species richness, while the evenness of all ants decreased with shrub cover. Omnivore ant evenness decreased with tree evenness, but increased with successional age.7. The results highlight the value of diverse forests in maintaining species richness and community evenness of a functionally important predator group. Moreover, the results stress the importance of analysing trophic groups separately when investigating biodiversity effects.

Trait-mediated assembly processes predict successional changes in community diversity of tropical forests

Interspecific differences in relative fitness can cause local dominance by a single species. However, stabilizing interspecific niche differences can promote local diversity. Understanding these mechanisms requires that we simultaneously quantify their effects on demography and link these effects to community dynamics. Successional forests are ideal systems for testing assembly theory because they exhibit rapid community assembly. Here, we leverage functional trait and long-term demographic data to build spatially explicit models of successional community dynamics of lowland rainforests in Costa Rica. First, we ask what the effects and relative importance of four trait-mediated community assembly processes are on tree survival, a major component of fitness. We model trait correlations with relative fitness differences that are both density-independent and -dependent in addition to trait correlations with stabilizing niche differences. Second, we ask how the relative importance of these trait-mediated processes relates to successional changes in functional diversity. Tree dynamics were more strongly influenced by trait-related interspecific variation in average survival than trait-related responses to neighbors, with wood specific gravity (WSG) positively correlated with greater survival. Our findings also suggest that competition was mediated by stabilizing niche differences associated with specific leaf area (SLA) and leaf dry matter content (LDMC). These drivers of individual-level survival were reflected in successional shifts to higher SLA and LDMC diversity but lower WSG diversity. Our study makes significant advances to identifying the links between individual tree performance, species functional traits, and mechanisms of tropical forest succession.