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Leaf area index estimates in dense evergreen tropical moist forest almost exclusively rest on indirect methods most of which being of limited accuracy or spatial resolution. In this study we examine the potential of full waveform Aerial Laser Scanning (ALS) to derive accurate spatially explicit estimates of Plant Area Index (PAI). A discrete representation of the forest canopy is introduced in the form of a 3D voxelized space. For each voxel (elementary volume, typically one cubic m) a first estimate of local transmittance of vegetation is computed as the ratio of the sum of energy exiting a voxel to the sum of energy entering the same voxel. A spatially hierarchical model is subsequently applied to refine estimates of individual voxel transmittance. Plant area density (PAD) profiles are then computed from the local transmittance values by applying Beer Lambert's turbid medium approximation. PAI values are obtained from vertical integration of PAD profiles. The model is shown to be robust to low sampling intensity and high occlusion rates. We further compared simulated values of gap fraction obtained by ray tracing for 5 angular sectors with in situ LAI2200 measurements taken at 135 positions in a 0.5 ha forest plot located in the center of the scene. The overall patterns of simulated and measured values (average value per inclination and pattern of variation along a 70 m transect line) were highly consistent. A slight but systematic discrepancy was observed along the inclination gradient, gap fractions derived from ray tracing in the voxelized scene being slightly lower than the measured values. This difference might be the consequence of multiple reflections which have been found to bias gap fractions estimates produced by LAI2200. PAI estimates derived from LAI2200 measurements (either simulated 6.8 or observed 5.9) are much lower than the PAI derived from vertical integration of local PAD (13.6). This large difference reflects the fact that distribution of foliage is strongly spatially structured and that this structural information is not properly accounted for in PAI estimates derived from mean gap fraction per elevation angle. After adjusting local transmittance to match mean LAI 2200 profiles the PAI at plot level was found to be 13.2 m2·m− 2. We conclude that Aerial Laser Scanning can produce accurate maps of Plant Area Index over large areas with unmatched efficacy, accuracy and ease. This should be of major relevance for many forest ecological studies.

International audience; Background and Aims: Morphological variation in light-foraging strategies potentially plays important roles in efficient light utilization and carbon assimilation in spatially and temporally heterogeneous environments such as tropical moist forest understorey. By considering a suite of morphological traits at various hierarchical scales, we examined the functional significance of crown shape diversity and plasticity in response to canopy openness.Methods: We conducted a field comparative study in French Guiana among tree saplings of 14 co-occurring species differing in light-niche optimum and breadth. Each leaf, axis or crown functional trait was characterized by a median value and a degree of plasticity expressed under contrasting light regimes.Key Results: We found divergent patterns between shade-tolerant and heliophilic species on the one hand and between shade and sun plants on the other. Across species, multiple regression analysis showed that relative crown depth was positively correlated with leaf lifespan and not correlated with crown vertical growth rate. Within species displaying a reduction in crown depth in the shade, we observed that crown depth was limited by reduced crown vertical growth rate and not by accelerated leaf or branch shedding. In addition, the study provides contrasting examples of morphological multilevel plastic responses, which allow the maintenance of efficient foliage and enable effective whole-plant light capture in shaded conditions under a moderate vertical light gradient.Conclusions: This result suggests that plastic adjustment of relative crown depth does not reflect a strategy maximizing light capture efficiency. Integrating and scaling-up leaf-level dynamics to shoot- and crown-level helps to interpret in functional and adaptive terms inter- and intraspecific patterns of crown traits and to better understand the mechanism of shade tolerance.

International audience; Differential growth response to light level is widely accepted as a potential mechanism for maintaining tree species richness in tropical forests. The position of tree species in the hierarchy of the canopy is considered an important indicator of species light capture and growth strategy. Paradoxically, the relative importance of species identity and competition for light in determining individual tree growth is poorly documented at the adult stage. In this study, we used a hierarchical Bayesian model to quantify the overall importance of species identity, light and belowground competition as determinants of tree growth in French Guiana tropical forest. Light competitive status is assessed by a crown exposure score and below ground competition is estimated from local crowding. We examined species sensitivity to both types of competition in relation to adult stature.Our results are based on annual diameter increments of more than 13,510 stems from 282 species monitored over 10 years. Mean annual growth rate was 0.11 cm y−1 with species identity explaining 35% of the individual variation in growth rate. Crown exposure and local crowding explained 3.5% and 2.4% of the variation in growth rate, respectively. Predicted changes in growth rate as crown exposure (resp. local crowding) index changed from lower to upper interquartile level was 0.03 cm y−1 (resp. 0.02 cm y−1). Species sensitivity to crown exposure and to local crowding were positively correlated (i) with predicted growth rate at high-light standardized conditions and (ii) with adult stature.This vertical niche partitioning is invoked to explain the limited contribution made by level of light competition for predicting individual tropical tree growth as the community-level response is dominated by the abundance of small-statured species with low sensitivity to light level.Light appears to drive the stem growth rate of tropical trees through species differentiation more than through individual tree growth limitation. This vertical stratification complements the previously reported regeneration niche and together these provide evidence for light niche partitioning in the three-dimensional space of tropical forests.

Niche differentiation is a key issue in the current debate on community assembly mechanisms. In highly diverse moist tropical forests, tree species sensitivity to canopy openness is thought to be a major axis in niche differentiation. In the past, the syndrome of traits driving the demographic trade-off involved in the niche-based theory of coexistence has always been established among species situated at the two extremities of the shade-tolerance gradient, even though most tropical tree species have intermediate light requirements. In addition, trait plasticity has seldom been linked to tropical tree species distribution along environmental gradients. This article examines covariations between leaf traits, whole-plant traits and niche parameters among 14 tree species with intermediate light requirements in French Guiana and across a range of canopy openness. Each functional trait measured under field conditions was characterized by a median value and a degree of plasticity expressed under contrasting light regimes. Niche differentiation was characterized in terms of spatial light gradient. We first examined covariations between functional traits then explored to what degree the median value and plasticity in functional traits could predict light niche characteristics at the sapling stage and the ontogenetic change in light availability estimated by adult stature. Leaf mass per area (LMA) was positively correlated with leaf life span (LLS); species with higher LMA and higher LLS displayed lower diameter growth rates (GRs) and lower responsiveness to canopy gap at both whole-plant and population levels. This proved that the relationships previously established over a broader range of species held true within the narrow range of the light requirements covered. Height GR plasticity accounted for 49% of the variation in light niche optimum. LMA plasticity, unlike LLS plasticity, was significantly correlated with light niche breadth and adult stature. Synthesis. This study demonstrates the relevance of considering the phenotypic plasticity in functional traits in community ecology, particularly for quantifying breadth of species distribution over environmental gradients. Our findings did not support Hubbell's hypothesis of functional equivalence and suggest that even a rather subtle variation in forest canopy disturbance promotes the coexistence of tropical tree species. (Resume d'auteur)

UMR AMAP, Equipe 3; International audience; In agroforestry systems, the distribution of light transmitted under tree canopies can be a limiting factor for the development of intercrops. The light available for intercrops depends on the quantity of light intercepted by tree canopies and, consequently, on the architecture of the tree species present. The influence of tree architecture on light transmission was analysed using dynamic 3D architectural models. The architectural analysis of Acacia mangium and Tectona grandis was performed in Indonesian agroforestry systems with trees aged from 1 to 3 years. 3D virtual trees were then generated with the AmapSim simulation software and 3D virtual experiments in which tree age, planting density, planting pattern and pruning intensity varied were reconstructed in order to simulate light available for the crop. Canopy closure of trees was more rapid in A. mangium than in T. grandis agroforestry systems; after 3 years the quantity of light available for A. mangium intercrops was three times lower than under T. grandis. Simulations with A. mangium showed that practices such as pruning and widening tree spacing enable to increase the total transmitted light within the stand. On T. grandis, modification of the tree row azimuth resulted in changes in the spatial and seasonal distribution of light available for the intercrops. These results are discussed in terms of agroforestry system management.

Correspondance: malezieux@cirad.fr; International audience; The evolution of natural ecosystems is controled by a high level of biodiversity, In sharp contrast, intensive agricultural systems involve monocultures associated with high input of chemical fertilisers and pesticides. Intensive agricultural systems have clearly negative impacts on soil and water quality and on biodiversity conservation. Alternatively, cropping systems based on carefully designed species mixtures reveal many potential advantages under various conditions, both in temperate and tropical agriculture. This article reviews those potential advantages by addressing the reasons for mixing plant species; the concepts and tools required for understanding and designing cropping systems with mixed species; and the ways of simulating multispecies cropping systems with models. Multispecies systems are diverse and may include annual and perennial crops on a gradient of complexity from 2 to n species. A literature survey shows potential advantages such as (1) higher overall productivity, (2) better control of pests and diseases, (3) enhanced ecological services and (4) greater economic profitability. Agronomic and ecological conceptual frameworks are examined for a clearer understanding of cropping systems, including the concepts of competition and facilitation, above- and belowground interactions and the types of biological interactions between species that enable better pest management in the system. After a review of existing models, future directions in modelling plant mixtures are proposed. We conclude on the need to enhance agricultural research on these multispecies systems, combining both agronomic and ecological concepts and tools.