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Tropical forests are global centres of biodiversity and carbon storage.Many tropical countries aspire to protect forest to fulfil biodiversity and climate mitigation policy targets, but the conservation strategies needed to achieve these two functions depend critically on the tropical forest tree diversity-carbon storage relationship. Assessing this relationship is challenging due to the scarcity of inventories where carbon stocks in aboveground biomass and species identifications have been simultaneously and robustly quantified.Here, we compile a unique pan-tropical dataset of 360 plots located in structurally intact old-growth closed-canopy forest, surveyed using standardised methods, allowing a multi-scale evaluation of diversity-carbon relationships in tropical forests. Diversity-carbon relationships among all plots at 1 ha scale across the tropics are absent, and within continents are either weak (Asia) or absent (Amazonia, Africa). A weak positive relationship is detectable within 1 ha plots, indicating that diversity effects in tropical forests may be scale dependent.The absence of clear diversity-carbon relationships at scales relevant to conservation planning means that carbon-centred conservation strategies will inevitably miss many high diversity ecosystems. As tropical forests can have any combination of tree diversity and carbon stocks both require explicit consideration when optimising policies to manage tropical carbon and biodiversity.

Tree mortality rates appear to be increasing in moist tropical forests (MTFs) with significant carbon cycle consequences. Here, we review the state of knowledge regarding MTF tree mortality, create a conceptual framework with testable hypotheses regarding the drivers, mechanisms and interactions that may underlie increasing MTF mortality rates, and identify the next steps for improved understanding and reduced prediction. Increasing mortality rates are associated with rising temperature and vapor pressure deficit, liana abundance, drought, wind events, fire and, possibly, CO2 fertilization-induced increases in stand thinning or acceleration of trees reaching larger, more vulnerable heights. The majority of these mortality drivers may kill trees in part through carbon starvation and hydraulic failure. The relative importance of each driver is unknown. High species diversity may buffer MTFs against large-scale mortality events, but recent and expected trends in mortality drivers give reason for concern regarding increasing mortality within MTFs. Models of tropical tree mortality are advancing the representation of hydraulics, carbon and demography, but require more empirical knowledge regarding the most common drivers and their subsequent mechanisms. We outline critical datasets and model developments required to test hypotheses regarding the underlying causes of increasing MTF mortality rates, and improve prediction of future mortality under climate change.

Tropical ecosystems have the highest levels of biodiversity, cycle more water and absorb more carbon than any other terrestrial ecosystem on Earth. Consequently, these ecosystems are extremely important components of Earth’s climatic system and biogeochemical cycles. Plant hydraulics is an essential discipline to understand and predict the dynamics of tropical vegetation in scenarios of changing water availability. Using published plant hydraulic data we show that the trade-off between drought avoidance (expressed as deep-rooting, deciduousness and capacitance) and hydraulic safety (P50 – the water potential when plants lose 50% of their maximum hydraulic conductivity) is a major axis of physiological variation across tropical ecosystems. We also propose a novel and independent axis of hydraulic trait variation linking vulnerability to hydraulic failure (expressed as the hydraulic safety margin (HSM)) and growth, where inherent fast-growing plants have lower HSM compared to slow-growing plants. We surmise that soil nutrients are fundamental drivers of tropical community assembly determining the distribution and abundance of the slow-safe/fast-risky strategies. We conclude showing that including either the growth-HSM or the resistance-avoidance trade-off in models can make simulated tropical rainforest communities substantially more vulnerable to drought than similar communities without the trade-off. These results suggest that vegetation models need to represent hydraulic trade-off axes to accurately project the functioning and distribution of tropical ecosystems.

Aim: Large trees (d.b.h. ≥70 cm) store large amounts of biomass. Several studies suggest that large trees may be vulnerable to changing climate, potentially leading to declining forest biomass storage. Here we determine the importance of large trees for tropical forest biomass storage and explore which intrinsic (species trait) and extrinsic (environment) variables are associated with the density of large trees and forest biomass at continental and pan-tropical scales. Location: Pan-tropical. Methods: Aboveground biomass (AGB) was calculated for 120 intact lowland moist forest locations. Linear regression was used to calculate variation in AGB explained by the density of large trees. Akaike information criterion weights (AICcwi) were used to calculate averaged correlation coefficients for all possible multiple regression models between AGB/density of large trees and environmental and species trait variables correcting for spatial autocorrelation. Results: Density of large trees explained c. 70% of the variation in pan-tropical AGB and was also responsible for significantly lower AGB in Neotropical [287.8 (mean) ± 105.0 (SD) Mg ha⁻¹] versus Palaeotropical forests (Africa 418.3 ± 91.8 Mg ha⁻¹; Asia 393.3 ± 109.3 Mg ha⁻¹). Pan-tropical variation in density of large trees and AGB was associated with soil coarseness (negative), soil fertility (positive), community wood density (positive) and dominance of wind dispersed species (positive), temperature in the coldest month (negative), temperature in the warmest month (negative) and rainfall in the wettest month (positive), but results were not always consistent among continents. Main conclusions: Density of large trees and AGB were significantly associated with climatic variables, indicating that climate change will affect tropical forest biomass storage. Species trait composition will interact with these future biomass changes as they are also affected by a warmer climate. Given the importance of large trees for variation in AGB across the tropics, and their sensitivity to climate change, we emphasize the need for in-depth analyses of the community dynamics of large trees.

1. Successional gradients are ubiquitous in nature, yet few studies have systematically examined the evolutionary origins of taxa that specialize at different successional stages. Here we quantify successional habitat specialization in Neotropical forest trees and evaluate its evolutionary lability along a precipitation gradient. Theoretically, successional habitat specialization should be more evolutionarily conserved in wet forests than in dry forests due to more extreme microenvironmental differentiation between early and late-successional stages in wet forest. 2. We applied a robust multinomial classification model to samples of primary and secondary forest trees from 14 Neotropical lowland forest sites spanning a precipitation gradient from 788 to 4000 mm annual rainfall, identifying species that are old-growth specialists and secondary forest specialists in each site. We constructed phylogenies for the classified taxa at each site and for the entire set of classified taxa and tested whether successional habitat specialization is phylogenetically conserved. We further investigated differences in the functional traits of species specializing in secondary vs. old-growth forest along the precipitation gradient, expecting different trait associations with secondary forest specialists in wet vs. dry forests since water availability is more limiting in dry forests and light availability more limiting in wet forests. 3. Successional habitat specialization is non-randomly distributed in the angiosperm phylogeny, with a tendency towards phylogenetic conservatism overall and a trend towards stronger conservatism in wet forests than in dry forests. However, the specialists come from all the major branches of the angiosperm phylogeny, and very few functional traits showed any consistent relationships with successional habitat specialization in either wet or dry forests. 4. Synthesis. The niche conservatism evident in the habitat specialization of Neotropical trees suggests a role for radiation into different successional habitats in the evolution of species-rich genera, though the diversity of functional traits that lead to success in different successional habitats complicates analyses at the community scale. Examining the distribution of particular lineages with respect to successional gradients may provide more insight into the role of successional habitat specialization in the evolution of species-rich taxa.

Aim: Tropical forests account for a quarter of the global carbon storage and a third of the terrestrial productivity. Few studies have teased apart the relative importance of environmental factors and forest attributes for ecosystem functioning, especially for the tropics. This study aims to relate aboveground biomass (AGB) and biomass dynamics (i.e., net biomass productivity and its underlying demographic drivers: biomass recruitment, growth and mortality) to forest attributes (tree diversity, community-mean traits and stand basal area) and environmental conditions (water availability, soil fertility and disturbance). Location: Neotropics. Methods: We used data from 26 sites, 201 1-ha plots and >92,000 trees distributed across the Neotropics. We quantified for each site water availability and soil total exchangeable bases and for each plot three key community-weighted mean functional traits that are important for biomass stocks and productivity. We used structural equation models to test the hypothesis that all drivers have independent, positive effects on biomass stocks and dynamics. Results: Of the relationships analysed, vegetation attributes were more frequently associated significantly with biomass stocks and dynamics than environmental conditions (in 67 vs. 33% of the relationships). High climatic water availability increased biomass growth and stocks, light disturbance increased biomass growth, and soil bases had no effect. Rarefied tree species richness had consistent positive relationships with biomass stocks and dynamics, probably because of niche complementarity, but was not related to net biomass productivity. Community-mean traits were good predictors of biomass stocks and dynamics. Main conclusions: Water availability has a strong positive effect on biomass stocks and growth, and a future predicted increase in (atmospheric) drought might, therefore, potentially reduce carbon storage. Forest attributes, including species diversity and community-weighted mean traits, have independent and important relationships with AGB stocks, dynamics and ecosystem functioning, not only in relatively simple temperate systems, but also in structurally complex hyper-diverse tropical forests.

A central goal of comparative plant ecology is to understand how functional traits vary among species and to what extent this variation has adaptive value. Here we evaluate relationships between four functional traits (seed volume, specific leaf area, wood density, and adult stature) and two demographic attributes (diameter growth and tree mortality) for large trees of 240 tree species from Neotropical forests. We evaluate how these key functional traits are related to survival and growth and whether similar relationships between traits and demography hold across different tropical forests. There was a tendency for a trade-off between growth and survival across rain forest tree species. Wood density, seed volume, and adult stature were significant predictors of growth and/or mortality. Both growth and mortality rates declined with an increase in wood density. This is consistent with greater construction costs and greater resistance to stem damage for denser wood. Growth and mortality rates also declined as seed volume increased. This is consistent with an adaptive syndrome in which species tolerant of low resource availability (in this case shade-tolerant species) have large seeds to establish successfully and low inherent growth and mortality rates. Growth increased and mortality decreased with an increase in adult stature, because taller species have a greater access to light and longer life spans. Specific leaf area was, surprisingly, only modestly informative for the performance of large trees and had ambiguous relationships with growth and survival. Single traits accounted for 9—55% of the interspecific variation in growth and mortality rates at individual sites. Significant correlations with demographic rates tended to be similar across forests and for phylogenetically independent contrasts as well as for cross-species analyses that treated each species as an independent observation. In combination, the morphological traits explained 41% of the variation in growth rate and 54% of the variation in mortality rate, with wood density being the best predictor of growth and mortality. Relationships between functional traits and demographic rates were statistically similar across a wide range of Neotropical forests. The consistency of these results strongly suggests that tropical rain forest species face similar trade-offs in different sites and converge on similar sets of solutions.

1. Essential resources such as water, nutrients and light vary over space and time and plant growth rates are expected to vary accordingly. We examined the effects of climate, soil and logging disturbances on diameter growth rates at the tree and stand level, using 165 1‐ha permanent sample plots distributed across Bolivian tropical lowland forests. 2. We predicted that growth rates would be higher in humid than in dry forests, higher in nutrient‐rich than nutrient‐poor forests and higher in logged than non‐logged forests. 3. Across the 165 plots we found positive basal area increases at the stand level, which agree with the generally reported biomass increases in tropical forests. 4. Multiple regression analysis demonstrated that climate variables, in particular water availability, were the strongest drivers of tree growth. More rainfall, a shorter and less intense dry period and higher temperatures led to higher tree growth rates. 5. Tree growth increased modestly with soil fertility and basal area growth was greatest at intermediate soil fertility. Surprisingly, tree growth showed little or no relationship with total soil nitrogen or plant available soil phosphorus. 6. Growth rates increased in logged plots just after logging, but this effect disappeared after 6 years. 7. Synthesis. Climate is the strongest driver of spatial variation in tree growth, and climate change may therefore have large consequences for forest productivity and carbon sequestration. The negative impact of decreased rainfall and increased rainfall seasonality on tree growth might be partly offset by the positive impact of increased temperature in these forests.

In evergreen tropical forests, the extent, magnitude, and controls on photosynthetic seasonality are poorly resolved and inadequately represented in Earth system models. Combining camera observations with ecosystem carbon dioxide fluxes at forests across rainfall gradients in Amazônia, we show that aggregate canopy phenology, not seasonality of climate drivers, is the primary cause of photosynthetic seasonality in these forests. Specifically, synchronization of new leaf growth with dry season litterfall shifts canopy composition toward younger, more light-use efficient leaves, explaining large seasonal increases (~27%) in ecosystem photosynthesis. Coordinated leaf development and demography thus reconcile seemingly disparate observations at different scales and indicate that accounting for leaf-level phenology is critical for accurately simulating ecosystem-scale responses to climate change.

Aim: To test the extent to which the vertical structure of tropical forests is determined by environment, forest structure or biogeographical history. Location: Pan-tropical. Methods: Using height and diameter data from 20,497 trees in 112 non-contiguous plots, asymptotic maximum height (H AM ) and height—diameter relationships were computed with nonlinear mixed effects (NLME) models to: (1) test for environmental and structural causes of differences among plots, and (2) test if there were continental differences once environment and structure were accounted for; persistence of differences may imply the importance of biogeography for vertical forest structure. NLME analyses for floristic subsets of data (only/excluding Fabaceae and only/excluding Dipterocarpaceae individuals) were used to examine whether family-level patterns revealed biogeographical explanations of cross-continental differences. Results: H AM and allometry were significantly different amongst continents. H AM was greatest in Asian forests (58.3 ± 7.5 m, 95% CI), followed by forests in Africa (45.1 ± 2.6 m), America (35.8 ± 6.0 m) and Australia (35.0 ± 7.4 m), and height—diameter relationships varied similarly; for a given diameter, stems were tallest in Asia, followed by Africa, America and Australia. Precipitation seasonality, basal area, stem density, solar radiation and wood density each explained some variation in allometry and H AM yet continental differences persisted even after these were accounted for. Analyses using floristic subsets showed that significant continental differences in H AM and allometry persisted in all cases. Main conclusions: Tree allometry and maximum height are altered by environmental conditions, forest structure and wood density. Yet, even after accounting for these, tropical forest architecture varies significantly from continent to continent. The greater stature of tropical forests in Asia is not directly determined by the dominance of the family Dipterocarpaceae, as on average non-dipterocarps are equally tall. We hypothesise that dominant large-statured families create conditions in which only tall species can compete, thus perpetuating a forest dominated by tall individuals from diverse families.