Green deserts, but not always: A global synthesis of native woody species regeneration under tropical tree monocultures.

Tree monocultures constitute an increasing fraction of the global tree cover and are the dominant tree-growing strategy of forest landscape restoration commitments. Their advantages to produce timber are well known, but their value for biodiversity is highly controversial and context dependent. Therefore, understanding whether, and in which conditions, they can harbor native species regeneration is crucial. Here, we conducted meta-analyses based on a global survey of the literature and on a database created with local, unpublished studies throughout Brazil to evaluate the regeneration potential of native species under tree monocultures and the way management influences this regeneration. Native woody species regeneration under tree monocultures harbors a substantial fraction of the diversity (on average 40% and 68% in the global and Brazilian surveys, respectively) and abundance (on average 25% and 60% in the global and Brazilian surveys, respectively) of regeneration observed in natural forests. Plantations with longer rotation lengths, composed of native tree species, and located adjacent to forest remnants harbor more species. Pine plantations harbor more native individuals than eucalypt plantations, and the abundance of regenerating trees is higher in sites with higher mean temperatures. Species-area curves revealed that the number of woody species under pine and eucalypt plantations in Brazil is 606 and 598 species, respectively, over an aggregated sampled area of ca. 12 ha. We highlight that the understory of tree monocultures can harbor a considerable diversity of regenerating native species at the landscape and regional scales, but this diversity strongly depends on management. Long-rotation length and favorable location are key factors for woody regeneration success under tropical tree monocultures. Therefore, tree monocultures can play a role in forest landscape restoration and conservation, but only if they are planned and managed for achieving this purpose.

Plant hydraulics at the heart of plant, crops and ecosystem functions in the face of climate change.

Plant hydraulics is crucial for assessing the plants' capacity to extract and transport water from the soil up to their aerial organs. Along with their capacity to exchange water between plant compartments and regulate evaporation, hydraulic properties determine plant water relations, water status and susceptibility to pathogen attacks. Consequently, any variation in the hydraulic characteristics of plants is likely to significantly impact various mechanisms and processes related to plant growth, survival and production, as well as the risk of biotic attacks and forest fire behaviour. However, the integration of hydraulic traits into disciplines such as plant pathology, entomology, fire ecology or agriculture can be significantly improved. This review examines how plant hydraulics can provide new insights into our understanding of these processes, including modelling processes of vegetation dynamics, illuminating numerous perspectives for assessing the consequences of climate change on forest and agronomic systems, and addressing unanswered questions across multiple areas of knowledge.

Rethinking the nature of intraspecific variability and its consequences on species coexistence.

Intraspecific variability (IV) has been proposed to explain species coexistence in diverse communities. Assuming, sometimes implicitly, that conspecific individuals can perform differently in the same environment and that IV increases niche overlap, previous studies have found contrasting results regarding the effect of IV on species coexistence. We aim at showing that the large IV observed in data does not mean that conspecific individuals are necessarily different in their response to the environment and that the role of high-dimensional environmental variation in determining IV has largely remained unexplored in forest plant communities. We first used a simulation experiment where an individual attribute is derived from a high-dimensional model, representing "perfect knowledge" of individual response to the environment, to illustrate how large observed IV can result from "imperfect knowledge" of the environment. Second, using growth data from clonal Eucalyptus plantations in Brazil, we estimated a major contribution of the environment in determining individual growth. Third, using tree growth data from long-term tropical forest inventories in French Guiana, Panama and India, we showed that tree growth in tropical forests is structured spatially and that despite a large observed IV at the population level, conspecific individuals perform more similarly locally than compared with heterospecific individuals. As the number of environmental dimensions that are well quantified at fine scale is generally lower than the actual number of dimensions influencing individual attributes, a great part of observed IV might be represented as random variation across individuals when in fact it is environmentally driven. This mis-representation has important consequences for inference about community dynamics. We emphasize that observed IV does not necessarily impact species coexistence per se but can reveal species response to high-dimensional environment, which is consistent with niche theory and the observation of the many differences between species in nature.

Small and slow is safe: On the drought tolerance of tropical tree species.

Understanding how evolutionary history and the coordination between trait trade-off axes shape the drought tolerance of trees is crucial to predict forest dynamics under climate change. Here, we compiled traits related to drought tolerance and the fast-slow and stature-recruitment trade-off axes in 601 tropical woody species to explore their covariations and phylogenetic signals. We found that xylem resistance to embolism (P50) determines the risk of hydraulic failure, while the functional significance of leaf turgor loss point (TLP) relies on its coordination with water use strategies. P50 and TLP exhibit weak phylogenetic signals and substantial variation within genera. TLP is closely associated with the fast-slow trait axis: slow species maintain leaf functioning under higher water stress. P50 is associated with both the fast-slow and stature-recruitment trait axes: slow and small species exhibit more resistant xylem. Lower leaf phosphorus concentration is associated with more resistant xylem, which suggests a (nutrient and drought) stress-tolerance syndrome in the tropics. Overall, our results imply that (1) drought tolerance is under strong selective pressure in tropical forests, and TLP and P50 result from the repeated evolutionary adaptation of closely related taxa, and (2) drought tolerance is coordinated with the ecological strategies governing tropical forest demography. These findings provide a physiological basis to interpret the drought-induced shift toward slow-growing, smaller, denser-wooded trees observed in the tropics, with implications for forest restoration programmes.

Increased hydraulic constraints in Eucalyptus plantations fertilized with potassium.

Fertilization is commonly used to increase growth in forest plantations, but it may also affect tree water relations and responses to drought. Here, we measured changes in biomass, transpiration, sapwood-to-leaf area ratio (As :Al ) and sap flow driving force (ΔΨ) during the 6-year rotation of tropical plantations of Eucalyptus grandis under controlled conditions for throughfall and potassium (K) fertilization. K fertilization increased final tree height by 8 m. Throughfall exclusion scarcely affected tree functioning because of deep soil water uptake. Tree growth increased in K-supplied plots and remained stable in K-depleted plots as tree height increased, while growth per unit leaf area increased in all plots. Stand transpiration and hydraulic conductance standardized per leaf area increased with height in K-depleted plots, but remained stable or decreased in K-supplied plots. Greater Al in K-supplied plots increased the hydraulic constraints on water use. This involved a direct mechanism through halved As :Al in K-supplied plots relative to K-depleted plots, and an indirect mechanism through deteriorated water status in K-supplied plots, which prevented the increase in ΔΨ with tree height. K fertilization in tropical plantations reduces the hydraulic compensation to growth, which could increase the risk of drought-induced dieback under climate change.

Neighbourhood-mediated shifts in tree biomass allocation drive overyielding in tropical species mixtures.

Variations in crown forms promote canopy space-use and productivity in mixed-species forests. However, we have a limited understanding on how this response is mediated by changes in within-tree biomass allocation. Here, we explored the role of changes in tree allometry, biomass allocation and architecture in shaping diversity-productivity relationships (DPRs) in the oldest tropical tree diversity experiment. We conducted whole-tree destructive biomass measurements and terrestrial laser scanning. Spatially explicit models were built at the tree level to investigate the effects of tree size and local neighbourhood conditions. Results were then upscaled to the stand level, and mixture effects were explored using a bootstrapping procedure. Biomass allocation and architecture substantially changed in mixtures, which resulted from both tree-size effects and neighbourhood-mediated plasticity. Shifts in biomass allocation among branch orders explained substantial shares of the observed overyielding. By contrast, root-to-shoot ratios, as well as the allometric relationships between tree basal area and aboveground biomass, were little affected by the local neighbourhood. Our results suggest that generic allometric equations can be used to estimate forest aboveground biomass overyielding from diameter inventory data. Overall, we demonstrate that shifts in tree biomass allocation are mediated by the local neighbourhood and promote DPRs in tropical forests.

Intensive silviculture enhances biomass accumulation and tree diversity recovery in tropical forest restoration.

Maximizing initial aboveground woody biomass (AGB) accumulation in order to obtain early payments for carbon stocking is essential for the financial viability of reforestation programs fostered by climate mitigation efforts. Intensive silviculture, i.e., silviculture traditionally used in commercial forestry to maximize productivity and gains, has recently been advocated as a promising approach to enhance AGB accumulation in restoration plantations. However, this approach may hamper natural forest regeneration and ecological succession due to high competition between colonizing plants and planted trees. We investigated the impacts of different silvicultural treatments applied to restoration plantations with 20 native tree species on AGB accumulation and spontaneous regeneration of native woody species in an experiment set up in the Atlantic Forest of Brazil. Intensive silviculture demonstrated a remarkable potential to enhance AGB accumulation in restoration plantations by increasing up to three times the AGB of tree stands (from ~25 to 75 Mg/ha in the 12th year). Intensive fertilization/weed control enhanced AGB accumulation, while higher tree density and the proportion of pioneers did not have a significant effect on AGB over the time. In spite of higher costs (cost increase of 13-19%), the cost-effectiveness for AGB accumulation of intensive silviculture was comparable to that of traditional silviculture applied to restoration (US$50-100/Mg AGB for 3 × 2 m spacing). Contrary to our expectations, we did not find a trade-off between AGB accumulation by planted trees and the spontaneous regeneration of tree species, since intensive silviculture enhanced the regeneration of both planted (total of 12 species) and colonizing woody species (total of 30 species) in the plantation understory. Specifically, a strong association was found between AGB stocks and the abundance and richness of colonizing species, a vast majority of which (90% of species and 95% of individuals) were dispersed by animals. We report a case of positive correlation between AGB stocking and woody species regeneration in the restoration of the Atlantic Forest. Fostering the establishment and maintenance of restoration tree plantations can, in some cases, be a win-win strategy for climate mitigation and biodiversity conservation in human-modified tropical landscapes.

Increased light-use efficiency sustains net primary productivity of shaded coffee plants in agroforestry system.

In agroforestry systems, shade trees strongly affect the physiology of the undergrown crop. However, a major paradigm is that the reduction in absorbed photosynthetically active radiation is, to a certain extent, compensated by an increase in light-use efficiency, thereby reducing the difference in net primary productivity between shaded and non-shaded plants. Due to the large spatial heterogeneity in agroforestry systems and the lack of appropriate tools, the combined effects of such variables have seldom been analysed, even though they may help understand physiological processes underlying yield dynamics. In this study, we monitored net primary productivity, during two years, on scales ranging from individual coffee plants to the entire plot. Absorbed radiation was mapped with a 3D model (MAESPA). Light-use efficiency and net assimilation rate were derived for each coffee plant individually. We found that although irradiance was reduced by 60% below crowns of shade trees, coffee light-use efficiency increased by 50%, leaving net primary productivity fairly stable across all shade levels. Variability of aboveground net primary productivity of coffee plants was caused primarily by the age of the plants and by intraspecific competition among them (drivers usually overlooked in the agroforestry literature) rather than by the presence of shade trees.

Environmental control of carbon allocation matters for modelling forest growth.

We aimed to evaluate the importance of modulations of within-tree carbon (C) allocation by water and low-temperature stress for the prediction of annual forest growth with a process-based model. A new C allocation scheme was implemented in the CASTANEA model that accounts for lagged and direct environmental controls of C allocation. Different approaches (static vs dynamic) to modelling C allocation were then compared in a model-data fusion procedure, using satellite-derived leaf production estimates and biometric measurements at c. 104 sites. The modelling of the environmental control of C allocation significantly improved the ability of CASTANEA to predict the spatial and year-to-year variability of aboveground forest growth along regional gradients. A significant effect of the previous year's water stress on the C allocation to leaves and wood was reported. Our results also are consistent with a prominent role of the environmental modulation of sink demand in the wood growth of the studied species. Data available at large scales can inform forest models about the processes driving annual and seasonal C allocation. Our results call for a greater consideration of C allocation drivers, especially sink-demand fluctuations, for the simulations of current and future forest productivity with process-based models.

Assessing the effects of management on forest growth across France: insights from a new functional-structural model.

BACKGROUND AND AIMS: The structure of a forest stand, i.e. the distribution of tree size features, has strong effects on its functioning. The management of the structure is therefore an important tool in mitigating the impact of predicted changes in climate on forests, especially with respect to drought. Here, a new functional-structural model is presented and is used to assess the effects of management on forest functioning at a national scale. METHODS: The stand process-based model (PBM) CASTANEA was coupled to a stand structure module (SSM) based on empirical tree-to-tree competition rules. The calibration of the SSM was based on a thorough analysis of intersite and interannual variability of competition asymmetry. The coupled CASTANEA-SSM model was evaluated across France using forest inventory data, and used to compare the effect of contrasted silvicultural practices on simulated stand carbon fluxes and growth. KEY RESULTS: The asymmetry of competition varied consistently with stand productivity at both spatial and temporal scales. The modelling of the competition rules enabled efficient prediction of changes in stand structure within the CASTANEA PBM. The coupled model predicted an increase in net primary productivity (NPP) with management intensity, resulting in higher growth. This positive effect of management was found to vary at a national scale across France: the highest increases in NPP were attained in forests facing moderate to high water stress; however, the absolute effect of management on simulated stand growth remained moderate to low because stand thinning involved changes in carbon allocation at the tree scale. CONCLUSIONS: This modelling approach helps to identify the areas where management efforts should be concentrated in order to mitigate near-future drought impact on national forest productivity. Around a quarter of the French temperate oak and beech forests are currently in zones of high vulnerability, where management could thus mitigate the influence of climate change on forest yield.