The links between wood traits and species demography change during tree development in a lowland tropical rainforest.

One foundational assumption of trait-based ecology is that traits can predict species demography. However, the links between traits and demographic rates are, in general, not as strong as expected. These weak associations may be due to the use of traits that are distantly related to performance, and/or the lack of consideration of size-related variations in both traits and demographic rates. Here, we examined how wood traits were related to demographic rates in 19 tree species from a lowland forest in eastern Amazonia. We measured 11 wood traits (i.e. structural, anatomical and chemical traits) in sapling, juvenile and adult wood; and related them to growth and mortality rates (MR) at different ontogenetic stages. The links between wood traits and demographic rates changed during tree development. At the sapling stage, relative growth rates (RGR) were negatively related to wood specific gravity (WSG) and total parenchyma fractions, while MR decreased with radial parenchyma fractions, but increased with vessel lumen area (VA). Juvenile RGR were unrelated to wood traits, whereas juvenile MR were negatively related to WSG and axial parenchyma fractions. At the adult stage, RGR scaled with VA and wood potassium concentrations. Adult MR were not predicted by any trait. Overall, the strength of the trait-demography associations decreased at later ontogenetic stages. Our results indicate that the associations between traits and demographic rates can change as trees age. Also, wood chemical or anatomical traits may be better predictors of growth and MR than WSG. Our findings are important to expand our knowledge on tree life-history variations and community dynamics in tropical forests, by broadening our understanding on the links between wood traits and demography during tree development.

High intraspecific growth variability despite strong evolutionary legacy in an Amazonian forest.

Tree growth is key to species performance. However, individual growth variability within species remains underexplored for a whole community, and the role of species evolutionary legacy and local environments remains unquantified. Based on 36 years of diameter records for 7961 trees from 138 species, we assessed individual growth across an Amazonian forest. We related individual growth to taxonomy, topography and neighbourhood, before exploring species growth link to functional traits and distribution along the phylogeny. We found most variation in growth among individuals within species, even though taxonomy explained a third of the variation. Species growth was phylogenetically conserved up to the genus. Traits of roots, wood and leaves were good predictors of growth, suggesting their joint selection during convergent evolutions. Neighbourhood crowding significantly decreased individual growth, although much of inter-individual variation remains unexplained. The high intraspecific variation observed could allow individuals to respond to the heterogeneous environments of Amazonian forests.

Thresholds for persistent leaf photochemical damage predict plant drought resilience in a tropical rainforest.

Water stress can cause declines in plant function that persist after rehydration. Recent work has defined 'resilience' traits characterizing leaf resistance to persistent damage from drought, but whether these traits predict resilience in whole-plant function is unknown. It is also unknown whether the coordination between resilience and 'resistance' - the ability to maintain function during drought - observed globally occurs within ecosystems. For eight rainforest species, we dehydrated and subsequently rehydrated leaves, and measured water stress thresholds for declines in rehydration capacity and maximum quantum yield of photosystem II (Fv /Fm ). We tested correlations with embolism resistance and dry season water potentials (ΨMD ), and calculated safety margins for damage (ΨMD - thresholds) and tested correlations with drought resilience in sap flow and growth. Ψ thresholds for persistent declines in Fv /Fm , indicating resilience, were positively correlated with ΨMD and thresholds for leaf vein embolism. Safety margins for persistent declines in Fv /Fm , but not rehydration capacity, were positively correlated with drought resilience in sap flow. Correlations between resistance and resilience suggest that species' differences in performance during drought are perpetuated after drought, potentially accelerating shifts in forest composition. Resilience to photochemical damage emerged as a promising functional trait to characterize whole-plant drought resilience.

Soil resistance and recovery during neotropical forest succession.

The recovery of soil conditions is crucial for successful ecosystem restoration and, hence, for achieving the goals of the UN Decade on Ecosystem Restoration. Here, we assess how soils resist forest conversion and agricultural land use, and how soils recover during subsequent tropical forest succession on abandoned agricultural fields. Our overarching question is how soil resistance and recovery depend on local conditions such as climate, soil type and land-use history. For 300 plots in 21 sites across the Neotropics, we used a chronosequence approach in which we sampled soils from two depths in old-growth forests, agricultural fields (i.e. crop fields and pastures), and secondary forests that differ in age (1-95 years) since abandonment. We measured six soil properties using a standardized sampling design and laboratory analyses. Soil resistance strongly depended on local conditions. Croplands and sites on high-activity clay (i.e. high fertility) show strong increases in bulk density and decreases in pH, carbon (C) and nitrogen (N) during deforestation and subsequent agricultural use. Resistance is lower in such sites probably because of a sharp decline in fine root biomass in croplands in the upper soil layers, and a decline in litter input from formerly productive old-growth forest (on high-activity clays). Soil recovery also strongly depended on local conditions. During forest succession, high-activity clays and croplands decreased most strongly in bulk density and increased in C and N, possibly because of strongly compacted soils with low C and N after cropland abandonment, and because of rapid vegetation recovery in high-activity clays leading to greater fine root growth and litter input. Furthermore, sites at low precipitation decreased in pH, whereas sites at high precipitation increased in N and decreased in C : N ratio. Extractable phosphorus (P) did not recover during succession, suggesting increased P limitation as forests age. These results indicate that no single solution exists for effective soil restoration and that local site conditions should determine the restoration strategies. This article is part of the theme issue 'Understanding forest landscape restoration: reinforcing scientific foundations for the UN Decade on Ecosystem Restoration'.

Defaunation changes leaf trait composition of recruit communities in tropical forests in French Guiana.

Hunting impacts tropical vertebrate populations, causing declines of species that function as seed dispersers and predators, or that browse seedlings and saplings. Whether and how the resulting reductions in seed dispersal, seed predation, and browsing translate to changes in the tree composition is poorly understood. Here, we assess the effect of defaunation on the functional composition of communities of tree recruits in tropical rainforests in French Guiana. We selected eight sites along a gradient of defaunation, caused by differences in hunting pressure, in otherwise intact old-growth forests in French Guiana. We measured shifts in functional composition by comparing leaf and fruit traits and wood density between tree recruits (up to 5 cm diameter at breast height) and adults, and tested whether and how these compositional shifts related to defaunation. We found a positive relationship with defaunation for shifts in specific leaf area, a negative relationship for shifts of leaf toughness and wood density, and a weak relationship for shifts in fruit traits. Our results suggest that the loss of vertebrates affects ecological processes such as seed dispersal and browsing, of which browsing remains understudied. Even though these changes sometimes seem minor, together they result in major shifts in forest composition. These changes have long-term ramifications that may alter forest dynamics for generations.

Strong floristic distinctiveness across Neotropical successional forests.

Forests that regrow naturally on abandoned fields are important for restoring biodiversity and ecosystem services, but can they also preserve the distinct regional tree floras? Using the floristic composition of 1215 early successional forests (≤20 years) in 75 human-modified landscapes across the Neotropic realm, we identified 14 distinct floristic groups, with a between-group dissimilarity of 0.97. Floristic groups were associated with location, bioregions, soil pH, temperature seasonality, and water availability. Hence, there is large continental-scale variation in the species composition of early successional forests, which is mainly associated with biogeographic and environmental factors but not with human disturbance indicators. This floristic distinctiveness is partially driven by regionally restricted species belonging to widespread genera. Early secondary forests contribute therefore to restoring and conserving the distinctiveness of bioregions across the Neotropical realm, and forest restoration initiatives should use local species to assure that these distinct floras are maintained.

Multidimensional tropical forest recovery.

Tropical forests disappear rapidly because of deforestation, yet they have the potential to regrow naturally on abandoned lands. We analyze how 12 forest attributes recover during secondary succession and how their recovery is interrelated using 77 sites across the tropics. Tropical forests are highly resilient to low-intensity land use; after 20 years, forest attributes attain 78% (33 to 100%) of their old-growth values. Recovery to 90% of old-growth values is fastest for soil (<1 decade) and plant functioning (<2.5 decades), intermediate for structure and species diversity (2.5 to 6 decades), and slowest for biomass and species composition (>12 decades). Network analysis shows three independent clusters of attribute recovery, related to structure, species diversity, and species composition. Secondary forests should be embraced as a low-cost, natural solution for ecosystem restoration, climate change mitigation, and biodiversity conservation.

Topography shapes the local coexistence of tree species within species complexes of Neotropical forests.

Forest inventories in Amazonia include around 5000 described tree species belonging to more than 800 genera. Numerous species-rich genera share genetic variation among species because of recent speciation and/or recurrent hybridisation, forming species complexes. Despite the key role that tree species complexes play in understanding Neotropical diversification, and their need to exploit a diversity of niches, little is known about the mechanisms that allow local coexistence of tree species complexes and their species in sympatry. In this study, we explored the fine-scale distribution of five tree species complexes and 22 species within these complexes. Combining forest inventories, botanical determination, and LiDAR-derived topographic data over 120 ha of permanent plots in French Guiana, we used a Bayesian modelling framework to test the role of fine-scale topographic wetness and tree neighbourhood on the occurrence of species complexes and the relative distribution of species within complexes. Species complexes of Neotropical trees were widely spread across the topographic wetness gradient at the local scale. Species within complexes showed pervasive niche differentiation along with topographic wetness and competition gradients. Similar patterns of species-specific habitat preferences were observed within several species complexes: species more tolerant to competition for resources grow in drier and less fertile plateaus and slopes. If supported by partial reproductive isolation of species and adaptive introgression at the species complex level, our results suggest that both species-specific habitat specialisation within species complexes and the broad ecological distribution of species complexes might explain the success of these species complexes at the regional scale.

Impacts of Degradation on Water, Energy, and Carbon Cycling of the Amazon Tropical Forests.

Selective logging, fragmentation, and understory fires directly degrade forest structure and composition. However, studies addressing the effects of forest degradation on carbon, water, and energy cycles are scarce. Here, we integrate field observations and high-resolution remote sensing from airborne lidar to provide realistic initial conditions to the Ecosystem Demography Model (ED-2.2) and investigate how disturbances from forest degradation affect gross primary production (GPP), evapotranspiration (ET), and sensible heat flux (H). We used forest structural information retrieved from airborne lidar samples (13,500 ha) and calibrated with 817 inventory plots (0.25 ha) across precipitation and degradation gradients in the eastern Amazon as initial conditions to ED-2.2 model. Our results show that the magnitude and seasonality of fluxes were modulated by changes in forest structure caused by degradation. During the dry season and under typical conditions, severely degraded forests (biomass loss ≥66%) experienced water stress with declines in ET (up to 34%) and GPP (up to 35%) and increases of H (up to 43%) and daily mean ground temperatures (up to 6.5°C) relative to intact forests. In contrast, the relative impact of forest degradation on energy, water, and carbon cycles markedly diminishes under extreme, multiyear droughts, as a consequence of severe stress experienced by intact forests. Our results highlight that the water and energy cycles in the Amazon are driven by not only climate and deforestation but also the past disturbance and changes of forest structure from degradation, suggesting a much broader influence of human land use activities on the tropical ecosystems.

Slow rate of secondary forest carbon accumulation in the Guianas compared with the rest of the Neotropics.

Secondary forests are a prominent component of tropical landscapes, and they constitute a major atmospheric carbon sink. Rates of carbon accumulation are usually inferred from chronosequence studies, but direct estimates of carbon accumulation based on long-term monitoring of stands are rarely reported. Recent compilations on secondary forest carbon accumulation in the Neotropics are heavily biased geographically as they do not include estimates from the Guiana Shield. We analysed the temporal trajectory of aboveground carbon accumulation and floristic composition at one 25-ha secondary forest site in French Guiana. The site was clear-cut in 1976, abandoned thereafter, and one large plot (6.25 ha) has been monitored continuously since. We used Bayesian modeling to assimilate inventory data and simulate the long-term carbon accumulation trajectory. Canopy change was monitored using two aerial lidar surveys conducted in 2009 and 2017. We compared the dynamics of this site with that of a surrounding old-growth forest. Finally, we compared our results with that from secondary forests in Costa Rica, which is one of the rare long-term monitoring programs reaching a duration comparable to our study. Twenty years after abandonment, aboveground carbon stock was 64.2 (95% credibility interval 46.4, 89.0) Mg C/ha, and this stock increased to 101.3 (78.7, 128.5) Mg C/ha 20 yr later. The time to accumulate one-half of the mean aboveground carbon stored in the nearby old-growth forest (185.6 [155.9, 200.2] Mg C/ha) was estimated at 35.0 [20.9, 55.9] yr. During the first 40 yr, the contribution of the long-lived pioneer species Xylopia nitida, Goupia glabra, and Laetia procera to the aboveground carbon stock increased continuously. Secondary forest mean-canopy height measured by lidar increased by 1.14 m in 8 yr, a canopy-height increase consistent with an aboveground carbon accumulation of 7.1 Mg C/ha (or 0.89 Mg C·ha-1 ·yr-1 ) during this period. Long-term AGC accumulation rate in Costa Rica was almost twice as fast as at our site in French Guiana. This may reflect higher fertility of Central American forest communities or a better adaptation of the forest tree community to intense and frequent disturbances. This finding may have important consequences for scaling-up carbon uptake estimates to continental scales.

The Forest Observation System, building a global reference dataset for remote sensing of forest biomass.

Forest biomass is an essential indicator for monitoring the Earth's ecosystems and climate. It is a critical input to greenhouse gas accounting, estimation of carbon losses and forest degradation, assessment of renewable energy potential, and for developing climate change mitigation policies such as REDD+, among others. Wall-to-wall mapping of aboveground biomass (AGB) is now possible with satellite remote sensing (RS). However, RS methods require extant, up-to-date, reliable, representative and comparable in situ data for calibration and validation. Here, we present the Forest Observation System (FOS) initiative, an international cooperation to establish and maintain a global in situ forest biomass database. AGB and canopy height estimates with their associated uncertainties are derived at a 0.25 ha scale from field measurements made in permanent research plots across the world's forests. All plot estimates are geolocated and have a size that allows for direct comparison with many RS measurements. The FOS offers the potential to improve the accuracy of RS-based biomass products while developing new synergies between the RS and ground-based ecosystem research communities.

Global trait-environment relationships of plant communities.

Plant functional traits directly affect ecosystem functions. At the species level, trait combinations depend on trade-offs representing different ecological strategies, but at the community level trait combinations are expected to be decoupled from these trade-offs because different strategies can facilitate co-existence within communities. A key question is to what extent community-level trait composition is globally filtered and how well it is related to global versus local environmental drivers. Here, we perform a global, plot-level analysis of trait-environment relationships, using a database with more than 1.1 million vegetation plots and 26,632 plant species with trait information. Although we found a strong filtering of 17 functional traits, similar climate and soil conditions support communities differing greatly in mean trait values. The two main community trait axes that capture half of the global trait variation (plant stature and resource acquisitiveness) reflect the trade-offs at the species level but are weakly associated with climate and soil conditions at the global scale. Similarly, within-plot trait variation does not vary systematically with macro-environment. Our results indicate that, at fine spatial grain, macro-environmental drivers are much less important for functional trait composition than has been assumed from floristic analyses restricted to co-occurrence in large grid cells. Instead, trait combinations seem to be predominantly filtered by local-scale factors such as disturbance, fine-scale soil conditions, niche partitioning and biotic interactions.

Contrasting patterns of leaf trait variation among and within species during tropical dry forest succession in Costa Rica.

A coordinated response to environmental drivers amongst individual functional traits is central to the plant strategy concept. However, whether the trait co-ordination observed at the global scale occurs at other ecological scales (especially within species) remains an open question. Here, for sapling communities of two tropical dry forest types in Costa Rica, we show large differences amongst traits in the relative contribution of species turnover and intraspecific variation to their directional changes in response to environmental changes along a successional gradient. We studied the response of functional traits associated with the leaf economics spectrum and drought tolerance using intensive sampling to analyse inter- and intra-specific responses to environmental changes and ontogeny. Although the overall functional composition of the sapling communities changed during succession more through species turnover than through intraspecific trait variation, their relative contributions differed greatly amongst traits. For instance, community mean specific leaf area changed mostly due to intraspecific variation. Traits of the leaf economics spectrum showed decoupled responses to environmental drivers and ontogeny. These findings emphasise how divergent ecological mechanisms combine to cause great differences in changes of individual functional traits over environmental gradients and ecological scales.