Tropical tree growth sensitivity to climate is driven by species intrinsic growth rate and leaf traits.

A better understanding of how climate affects growth in tree species is essential for improved predictions of forest dynamics under climate change. Long-term climate averages (mean climate) drive spatial variations in species' baseline growth rates, whereas deviations from these averages over time (anomalies) can create growth variation around the local baseline. However, the rarity of long-term tree census data spanning climatic gradients has so far limited our understanding of their respective role, especially in tropical systems. Furthermore, tree growth sensitivity to climate is likely to vary widely among species, and the ecological strategies underlying these differences remain poorly understood. Here, we utilize an exceptional dataset of 49 years of growth data for 509 tree species across 23 tropical rainforest plots along a climatic gradient to examine how multiannual tree growth responds to both climate means and anomalies, and how species' functional traits mediate these growth responses to climate. We show that anomalous increases in atmospheric evaporative demand and solar radiation consistently reduced tree growth. Drier forests and fast-growing species were more sensitive to water stress anomalies. In addition, species traits related to water use and photosynthesis partly explained differences in growth sensitivity to both climate means and anomalies. Our study demonstrates that both climate means and anomalies shape tree growth in tropical forests and that species traits can provide insights into understanding these demographic responses to climate change, offering a promising way forward to forecast tropical forest dynamics under different climate trajectories.

Fine root dynamics across pantropical rainforest ecosystems.

Fine roots constitute a significant component of the net primary productivity (NPP) of forest ecosystems but are much less studied than aboveground NPP. Comparisons across sites and regions are also hampered by inconsistent methodologies, especially in tropical areas. Here, we present a novel dataset of fine root biomass, productivity, residence time, and allocation in tropical old-growth rainforest sites worldwide, measured using consistent methods, and examine how these variables are related to consistently determined soil and climatic characteristics. Our pantropical dataset spans intensive monitoring plots in lowland (wet, semi-deciduous, and deciduous) and montane tropical forests in South America, Africa, and Southeast Asia (n = 47). Large spatial variation in fine root dynamics was observed across montane and lowland forest types. In lowland forests, we found a strong positive linear relationship between fine root productivity and sand content, this relationship was even stronger when we considered the fractional allocation of total NPP to fine roots, demonstrating that understanding allocation adds explanatory power to understanding fine root productivity and total NPP. Fine root residence time was a function of multiple factors: soil sand content, soil pH, and maximum water deficit, with longest residence times in acidic, sandy, and water-stressed soils. In tropical montane forests, on the other hand, a different set of relationships prevailed, highlighting the very different nature of montane and lowland forest biomes. Root productivity was a strong positive linear function of mean annual temperature, root residence time was a strong positive function of soil nitrogen content in montane forests, and lastly decreasing soil P content increased allocation of productivity to fine roots. In contrast to the lowlands, environmental conditions were a better predictor for fine root productivity than for fractional allocation of total NPP to fine roots, suggesting that root productivity is a particularly strong driver of NPP allocation in tropical mountain regions.

On the importance of root traits in seedlings of tropical tree species.

Plant biomass allocation may be optimized to acquire and conserve resources. How trade-offs in the allocation of tropical tree seedlings depend on different stressors remains poorly understood. Here we test whether above- and below-ground traits of tropical tree seedlings could explain observed occurrence along gradients of resources (light, water) and defoliation (fire, herbivory). We grew 24 tree species occurring in five African vegetation types, varying from dry savanna to moist forest, in a glasshouse for 6 months, and measured traits associated with biomass allocation. Classification based on above-ground traits resulted in clusters representing savanna and forest species, with low and high shoot investment, respectively. Classification based on root traits resulted in four clusters representing dry savanna, humid savanna, dry forest and moist forest, characterized by a deep mean rooting depth, root starch investment, high specific root length in deeper soil layers, and high specific root length in the top soil layer, respectively. In conclusion, tree seedlings in this study show root trait syndromes, which vary along gradients of resources and defoliation: seedlings from dry areas invest in deep roots, seedlings from shaded environments optimize shoot investment, and seedlings experiencing frequent defoliation store resources in the roots.

Drier tropical forests are susceptible to functional changes in response to a long-term drought.

Climatic changes have profound effects on the distribution of biodiversity, but untangling the links between climatic change and ecosystem functioning is challenging, particularly in high diversity systems such as tropical forests. Tropical forests may also show different responses to a changing climate, with baseline climatic conditions potentially inducing differences in the strength and timing of responses to droughts. Trait-based approaches provide an opportunity to link functional composition, ecosystem function and environmental changes. We demonstrate the power of such approaches by presenting a novel analysis of long-term responses of different tropical forest to climatic changes along a rainfall gradient. We explore how key ecosystem's biogeochemical properties have shifted over time as a consequence of multi-decadal drying. Notably, we find that drier tropical forests have increased their deciduous species abundance and generally changed more functionally than forests growing in wetter conditions, suggesting an enhanced ability to adapt ecologically to a drying environment.

Small-scale indirect plant responses to insect herbivory could have major impacts on canopy photosynthesis and isoprene emission.

Insect herbivores cause substantial changes in the leaves they attack, but their effects on the ecophysiology of neighbouring, nondamaged leaves have never been quantified in natural canopies. We studied how winter moth (Operophtera brumata), a common herbivore in temperate forests, affects the photosynthetic and isoprene emission rates of its host plant, the pedunculate oak (Quercus robur). Through a manipulative experiment, we measured leaves on shoots damaged by caterpillars or mechanically by cutting, or left completely intact. To quantify the effects at the canopy scale, we surveyed the extent and patterns of leaf area loss in the canopy. Herbivory reduced photosynthesis both in damaged leaves and in their intact neighbours. Isoprene emission rates significantly increased after mechanical leaf damage. When scaled up to canopy-level, herbivory reduced photosynthesis by 48 ± 10%. The indirect effects of herbivory on photosynthesis in undamaged leaves (40%) were much more important than the direct effects of leaf area loss (6%). If widespread across other plant-herbivore systems, these findings suggest that insect herbivory has major and previously underappreciated influences in modifying ecosystem carbon cycling, with potential effects on atmospheric chemistry.

On the relationship between fire regime and vegetation structure in the tropics.

We assessed data from 11 experiments examining the effects of the timing and/or frequency of fire on tropical forest and/or savanna vegetation structure over one decade or more. The initial 'control treatment' in many such cases consisted of previously cleared land. This is as opposed to natural vegetation subject to some sort of endogenous fire regime before the imposition of fire treatments. Effects of fire on fractional foliar cover are up to 10-fold greater when clearing pre-treatments are imposed. Moreover, because many of the 'classic' fire trials were initialised with applied management questions in mind, most have also used burning regimes much more frequent and/or severe than those occurring in the absence of human activity. Once these factors are taken into account, our modelling analysis shows that nonanthropogenic fire regimes serve to reduce canopy vegetative cover to a much lower extent than has previously been argued to be the case. These results call into question the notion that fire effects on tropical vegetation can be of a sufficient magnitude to maintain open-type savanna ecosystems under climatic/soil regimes otherwise sufficient to give rise to a more luxurious forest-type vegetation cover.

Forest biomass, productivity and carbon cycling along a rainfall gradient in West Africa.

Net Primary Productivity (NPP) is one of the most important parameters in describing the functioning of any ecosystem and yet it arguably remains a poorly quantified and understood component of carbon cycling in tropical forests, especially outside of the Americas. We provide the first comprehensive analysis of NPP and its carbon allocation to woody, canopy and root growth components at contrasting lowland West African forests spanning a rainfall gradient. Using a standardized methodology to study evergreen (EF), semi-deciduous (SDF), dry forests (DF) and woody savanna (WS), we find that (i) climate is more closely related with above and belowground C stocks than with NPP (ii) total NPP is highest in the SDF site, then the EF followed by the DF and WS and that (iii) different forest types have distinct carbon allocation patterns whereby SDF allocate in excess of 50% to canopy production and the DF and WS sites allocate 40%-50% to woody production. Furthermore, we find that (iv) compared with canopy and root growth rates the woody growth rate of these forests is a poor proxy for their overall productivity and that (v) residence time is the primary driver in the productivity-allocation-turnover chain for the observed spatial differences in woody, leaf and root biomass across the rainfall gradient. Through a systematic assessment of forest productivity we demonstrate the importance of directly measuring the main components of above and belowground NPP and encourage the establishment of more permanent carbon intensive monitoring plots across the tropics.

Fire effects and ecological recovery pathways of tropical montane cloud forests along a time chronosequence.

Tropical montane cloud forests (TMCFs) harbour high levels of biodiversity and large carbon stocks. Their location at high elevations make them especially sensitive to climate change, because a warming climate is enhancing upslope species migration, but human disturbance (especially fire) may in many cases be pushing the treeline downslope. TMCFs are increasingly being affected by fire, and the long-term effects of fire are still unknown. Here, we present a 28-year chronosequence to assess the effects of fire and recovery pathways of burned TMCFs, with a detailed analysis of carbon stocks, forest structure and diversity. We assessed rates of change of carbon (C) stock pools, forest structure and tree-size distribution pathways and tested several hypotheses regarding metabolic scaling theory (MST), C recovery and biodiversity. We found four different C stock recovery pathways depending on the selected C pool and time since last fire, with a recovery of total C stocks but not of aboveground C stocks. In terms of forest structure, there was an increase in the number of small stems in the burned forests up to 5-9 years after fire because of regeneration patterns, but no differences on larger trees between burned and unburned plots in the long term. In support of MST, after fire, forest structure appears to approximate steady-state size distribution in less than 30 years. However, our results also provide new evidence that the species recovery of TMCF after fire is idiosyncratic and follows multiple pathways. While fire increased species richness, it also enhanced species dissimilarity with geographical distance. This is the first study to report a long-term chronosequence of recovery pathways to fire suggesting faster recovery rates than previously reported, but at the expense of biodiversity and aboveground C stocks.

Revitalising Indigenous cultural fire practice: benefits and partnerships.

Indigenous cultural fire practitioners proactively revitalise their stewardship/custodianship of their traditional territories to generate diverse social, cultural, economic, self-determination, and ecological benefits. Government, researchers, and natural resource managers can overcome ongoing colonial legacies by enabling Indigenous leadership, providing ongoing investment and removing imposed barriers that restrict cultural fire practices.

Branching principles of animal and plant networks identified by combining extensive data, machine learning and modelling.

Branching in vascular networks and in overall organismic form is one of the most common and ancient features of multicellular plants, fungi and animals. By combining machine-learning techniques with new theory that relates vascular form to metabolic function, we enable novel classification of diverse branching networks-mouse lung, human head and torso, angiosperm and gymnosperm plants. We find that ratios of limb radii-which dictate essential biologic functions related to resource transport and supply-are best at distinguishing branching networks. We also show how variation in vascular and branching geometry persists despite observing a convergent relationship across organisms for how metabolic rate depends on body mass.

Long-term droughts may drive drier tropical forests towards increased functional, taxonomic and phylogenetic homogeneity.

Tropical ecosystems adapted to high water availability may be highly impacted by climatic changes that increase soil and atmospheric moisture deficits. Many tropical regions are experiencing significant changes in climatic conditions, which may induce strong shifts in taxonomic, functional and phylogenetic diversity of forest communities. However, it remains unclear if and to what extent tropical forests are shifting in these facets of diversity along climatic gradients in response to climate change. Here, we show that changes in climate affected all three facets of diversity in West Africa in recent decades. Taxonomic and functional diversity increased in wetter forests but tended to decrease in forests with drier climate. Phylogenetic diversity showed a large decrease along a wet-dry climatic gradient. Notably, we find that all three facets of diversity tended to be higher in wetter forests. Drier forests showed functional, taxonomic and phylogenetic homogenization. Understanding how different facets of diversity respond to a changing environment across climatic gradients is essential for effective long-term conservation of tropical forest ecosystems.

Leaf-level photosynthetic capacity dynamics in relation to soil and foliar nutrients along forest-savanna boundaries in Ghana and Brazil.

Forest-savanna boundaries extend across large parts of the tropics but the variability of photosynthetic capacity in relation to soil and foliar nutrients across these transition zones is poorly understood. For this reason, we compared photosynthetic capacity (maximum rate of carboxylation of Rubisco at 25 C° (Vcmax25), leaf mass, nitrogen (N), phosphorus (P) and potassium (K) per unit leaf area (LMA, Narea, Parea and Karea, respectively), in relation to respective soil nutrients from 89 species at seven sites along forest-savanna ecotones in Ghana and Brazil. Contrary to our expectations, edaphic conditions were not reflected in foliar nutrient concentrations but LMA was slightly higher in lower fertility soils. Overall, each vegetation type within the ecotones demonstrated idiosyncratic and generally weak relationships between Vcmax25 and Narea, Parea and Karea. Species varied significantly in their Vcmax25 ↔ Narea relationship due to reduced investment of total Narea in photosynthetic machinery with increasing LMA. We suggest that studied species in the forest-savanna ecotones do not maximize Vcmax25 per given total Narea due to adaptation to intermittent water availability. Our findings have implications for global modeling of Vcmax25 and forest-savanna ecotone productivity.

ENSO Drives interannual variation of forest woody growth across the tropics.

Meteorological extreme events such as El Niño events are expected to affect tropical forest net primary production (NPP) and woody growth, but there has been no large-scale empirical validation of this expectation. We collected a large high-temporal resolution dataset (for 1-13 years depending upon location) of more than 172 000 stem growth measurements using dendrometer bands from across 14 regions spanning Amazonia, Africa and Borneo in order to test how much month-to-month variation in stand-level woody growth of adult tree stems (NPPstem) can be explained by seasonal variation and interannual meteorological anomalies. A key finding is that woody growth responds differently to meteorological variation between tropical forests with a dry season (where monthly rainfall is less than 100 mm), and aseasonal wet forests lacking a consistent dry season. In seasonal tropical forests, a high degree of variation in woody growth can be predicted from seasonal variation in temperature, vapour pressure deficit, in addition to anomalies of soil water deficit and shortwave radiation. The variation of aseasonal wet forest woody growth is best predicted by the anomalies of vapour pressure deficit, water deficit and shortwave radiation. In total, we predict the total live woody production of the global tropical forest biome to be 2.16 Pg C yr-1, with an interannual range 1.96-2.26 Pg C yr-1 between 1996-2016, and with the sharpest declines during the strong El Niño events of 1997/8 and 2015/6. There is high geographical variation in hotspots of El Niño-associated impacts, with weak impacts in Africa, and strongly negative impacts in parts of Southeast Asia and extensive regions across central and eastern Amazonia. Overall, there is high correlation (r = -0.75) between the annual anomaly of tropical forest woody growth and the annual mean of the El Niño 3.4 index, driven mainly by strong correlations with anomalies of soil water deficit, vapour pressure deficit and shortwave radiation.This article is part of the discussion meeting issue 'The impact of the 2015/2016 El Niño on the terrestrial tropical carbon cycle: patterns, mechanisms and implications'.

Does salt stress constrain spatial distribution of dune building grasses Ammophila arenaria and Elytrichia juncea on the beach?

Rising sea levels threaten coastal safety by increasing the risk of flooding. Coastal dunes provide a natural form of coastal protection. Understanding drivers that constrain early development of dunes is necessary to assess whether dune development may keep pace with sea-level rise. In this study, we explored to what extent salt stress experienced by dune building plant species constrains their spatial distribution at the Dutch sandy coast. We conducted a field transplantation experiment and a glasshouse experiment with two dune building grasses Ammophila arenaria and Elytrigia juncea. In the field, we measured salinity and monitored growth of transplanted grasses in four vegetation zones: (I) nonvegetated beach, (II) E. juncea occurring, (III) both species co-occurring, and (IV) A. arenaria dominant. In the glasshouse, we subjected the two species to six soil salinity treatments, with and without salt spray. We monitored biomass, photosynthesis, leaf sodium, and nutrient concentrations over a growing season. The vegetation zones were weakly associated with summer soil salinity; zone I and II were significantly more saline than zones III and IV. Ammophila arenaria performed equally (zone II) or better (zones III, IV) than E. juncea, suggesting soil salinity did not limit species performance. Both species showed severe winter mortality. In the glasshouse, A. arenaria biomass decreased linearly with soil salinity, presumably as a result of osmotic stress. Elytrigia juncea showed a nonlinear response to soil salinity with an optimum at 0.75% soil salinity. Our findings suggest that soil salinity stress either takes place in winter, or that development of vegetated dunes is less sensitive to soil salinity than hitherto expected.

Examining variation in the leaf mass per area of dominant species across two contrasting tropical gradients in light of community assembly.

Understanding variation in key functional traits across gradients in high diversity systems and the ecology of community changes along gradients in these systems is crucial in light of conservation and climate change. We examined inter- and intraspecific variation in leaf mass per area (LMA) of sun and shade leaves along a 3330-m elevation gradient in Peru, and in sun leaves across a forest-savanna vegetation gradient in Brazil. We also compared LMA variance ratios (T-statistics metrics) to null models to explore internal (i.e., abiotic) and environmental filtering on community structure along the gradients. Community-weighted LMA increased with decreasing forest cover in Brazil, likely due to increased light availability and water stress, and increased with elevation in Peru, consistent with the leaf economic spectrum strategy expected in colder, less productive environments. A very high species turnover was observed along both environmental gradients, and consequently, the first source of variation in LMA was species turnover. Variation in LMA at the genus or family levels was greater in Peru than in Brazil. Using dominant trees to examine possible filters on community assembly, we found that in Brazil, internal filtering was strongest in the forest, while environmental filtering was observed in the dry savanna. In Peru, internal filtering was observed along 80% of the gradient, perhaps due to variation in taxa or interspecific competition. Environmental filtering was observed at cloud zone edges and in lowlands, possibly due to water and nutrient availability, respectively. These results related to variation in LMA indicate that biodiversity in species rich tropical assemblages may be structured by differential niche-based processes. In the future, specific mechanisms generating these patterns of variation in leaf functional traits across tropical environmental gradients should be explored.

Xylem hydraulic properties of roots and stems of nine Mediterranean woody species.

We studied the hydraulic architecture and water relations of nine co-occurring woody species in a Spanish evergreen oak forest over the course of a dry season. Our main objectives were to: (1) test the existence of a trade-off between hydraulic conductivity and security in the xylem, and (2) establish the safety margins at which the species operated in relation to hydraulic failure, and compare these safety margins between species and tissues (roots vs. stems). Our results showed that the relationship between specific hydraulic conductivity (K s) and resistance to cavitation followed a power function with exponent ≈-2, consistent with the existence of a trade-off between conductivity and security in the xylem, and also consistent with a linear relationship between vessel diameter and the size of inter-vessel pores. The diameter of xylem conduits, K s and vulnerability to xylem embolism were always higher in roots than in stems of the same species. Safety margins from hydraulic failure were narrower in roots than in stems. Among species, the water potential (Ψ) at which 50% of conductivity was lost due to embolism ranged between -0.9 and <-8 MPa for roots and between -2.0 and <-8 MPa for stems. Vulnerability to xylem embolism followed a pattern of: Quercus ilex=Acer monspessulanum=Arbutus unedo=Sorbus torminalis=Cistus laurifolius>Cistus albidus=Ilex aquifolium>Phillyrea latifolia>Juniperus oxycedrus. Gas exchange and seasonal Ψ minima were in general correlated with resistance to xylem embolism. Hydraulic safety margins differed markedly among species, with some of them (J. oxycedrus, I. aquifolium, P. latifolia) showing a xylem overly resistant to cavitation. We hypothesize that this overly resistant xylem may be related to the shape of the relationship between K s and security we have found.