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Global warming may force montane species to shift upward to keep pace with their shifting climate niche. How species differences in such distribution shifts depend on their elevational positions, elevation-dependent warming rates, and other environmental constraints, or plant functional traits is poorly understood. Here, we analyzed for 137 Himalayan tree species how distribution shifts vary with elevational niche positions, environmental constraints, and their functional traits. We developed ecological niche models using MaxEnt by combining species survey and botanical collections data with 19 environmental predictors. Species distributions were projected to 1985 and 2050 conditions, and elevational range parameters and distribution areas were derived. Under the worst-case RCP 8.5 scenario, species are predicted to shift, on average, 3 m/year in optimum elevation, and have 33% increase in distribution area. Highland species showed faster predicted elevational shifts than lowland species. Lowland and highland species are predicted to expand in distribution area in contrast to mid-elevation species. Tree species for which species distribution models are driven by responses to temperature, aridity, or soil clay content showed the strongest predicted upslope shifts. Tree species with conservative trait values that enable them to survive resource poor conditions (i.e., narrow conduits) showed larger predicted upslope shifts than species with wide conduits. The predicted average upslope shift in maximum elevation (8 m/year) is >2 times faster than the current observations indicating that many species will not be able to track climate change and potentially go extinct, unless they are supported by active conservation measures, such as assisted migration. Abstract in Nepali is available with online material.

High demands on forest for carbon storage and provision of timber and biofuel require precise and reliable estimates of the biomass, carbon and nutrient stocks in different tree compartments. Whether the fraction of biomass distributed in aboveground tree compartments and the carbon and nutrient concentrations varies systematically across trees in different canopy positions remains unclear despite its importance for understanding forest ecology. Here, we compared the distribution of biomass, carbon and nutrients from underlying carbon and nutrient concentrations between different aboveground tree compartments for 15 mature trees of European beech (Fagus sylvatica), Douglas fir (Pseudotsuga menziesii) and Scots pine (Pinus sylvestris) with dominant, intermediate and suppressed canopy position. We show that carbon concentrations were relatively constant across tree compartments while nutrient concentrations increased from stem, bark, branches towards needles. Canopy position had only minor effects on carbon and nutrient concentrations and on the distribution of biomass, carbon and nutrients between aboveground tree components. Nutrient concentrations and stochiometric results confirm that the forests were affected by high N deposition and low availability of P and base cations. Our results imply that predictions from allometric scaling theory better apply to aboveground tree components than from functional equilibrium theory. Models aiming for estimating tree and forest biomass and carbon and nutrient stocks can apply equal biomass, carbon and nutrient stocks for trees independent of canopy position as a valid assumption but testing this assumption for a broader range of species and site conditions remains recommended.

Conifers face increased drought mortality risks because of drought-induced embolism in their vascular system. Variation in embolism resistance may result from species differences in pit structure and function, as pits control the air seeding between water-transporting conduits. This study quantifies variation in embolism resistance and hydraulic conductivity for 28 conifer species grown in a 50-year-old common garden experiment and assesses the underlying mechanisms. Conifer species with a small pit aperture, high pit aperture resistance, and large valve effect were more resistant to embolism, as they all may reduce air seeding. Surprisingly, hydraulic conductivity was only negatively correlated with tracheid cell wall thickness. Embolism resistance and its underlying pit traits related to pit size and sealing were more strongly phylogenetically controlled than hydraulic conductivity and anatomical tracheid traits. Conifers differed in hydraulic safety and hydraulic efficiency, but there was no trade-off between safety and efficiency because they are driven by different xylem anatomical traits that are under different phylogenetic control., We used a common garden experiment to assess the mechanisms underlying embolism resistance and hydraulic conductivity of conifer species. Pit size and sealing explain embolism resistance, while hydraulic conductivity is only related to wall thickness.

Funder: Agua Salud Project, Funder: U.S. Department of Energy; Id: http://dx.doi.org/10.13039/100000015, Funder: CAPES; Id: http://dx.doi.org/10.13039/501100002322, Data capturing multiple axes of tree size and shape, such as a tree's stem diameter, height and crown size, underpin a wide range of ecological research-from developing and testing theory on forest structure and dynamics, to estimating forest carbon stocks and their uncertainties, and integrating remote sensing imagery into forest monitoring programmes. However, these data can be surprisingly hard to come by, particularly for certain regions of the world and for specific taxonomic groups, posing a real barrier to progress in these fields. To overcome this challenge, we developed the Tallo database, a collection of 498,838 georeferenced and taxonomically standardized records of individual trees for which stem diameter, height and/or crown radius have been measured. These data were collected at 61,856 globally distributed sites, spanning all major forested and non-forested biomes. The majority of trees in the database are identified to species (88%), and collectively Tallo includes data for 5163 species distributed across 1453 genera and 187 plant families. The database is publicly archived under a CC-BY 4.0 licence and can be access from: https://doi.org/10.5281/zenodo.6637599. To demonstrate its value, here we present three case studies that highlight how the Tallo database can be used to address a range of theoretical and applied questions in ecology-from testing the predictions of metabolic scaling theory, to exploring the limits of tree allometric plasticity along environmental gradients and modelling global variation in maximum attainable tree height. In doing so, we provide a key resource for field ecologists, remote sensing researchers and the modelling community working together to better understand the role that trees play in regulating the terrestrial carbon cycle.

Lianas are increasing in relative abundance and biomass in many tropical forests. We tested the hypothesis that lianas conform to the fast and acquisitive end of the continuum of plant strategies, allowing lianas to acquire resources faster than trees. We assessed functional traits representative of the leaf (LES) and wood economics spectrum (WES), including plant hydraulics, in 16 liana and 16 tree species in the canopy of two tropical forests at the extremes of the climatic and geological gradient across the Isthmus of Panama. For both forests, we observed a trade-off between the construction of more productive leaves with rapid turnover and expensive leaves with slower turnover. We also found trait variation associated with wood and hydraulic traits. These two axes were orthogonal, suggesting that trade-offs at the leaf and stem, including plant hydraulics, operate independently. For the dry forest, lianas had cheaper and more efficient leaves than trees. For the wet forest, lianas and trees overlapped in leaf and stem characteristics. Moreover, the duration of green foliage highly explained the variation between dry forest species, reflecting different adaptations to drought. In the wet forest, fast-growing species benefited from a higher return on investments of leaf vascular tissues than slow-growing species and they had a higher capacity to transport water through the leaf. A higher capacity to construct more productive leaves and display leaves with lower costs may favour lianas over trees in light interception, photosynthetic rates, and growth under high light and nutrient availability in dry forests. Synthesis. Lianas in a dry tropical forest had a more acquisitive strategy than trees, characterized by more productive leaves and more efficient display for light interception. In dry environments, lianas appear to benefit from high-light and nutrient-rich soils and thus take advantage of higher resource conditions compared to trees. By contrast, in a wet tropical forest, lianas and trees overlapped in leaf and stem characteristics and lianas were not more acquisitive than trees. In wet environments, low light availability and nutrient-poor soils in a context of low water limitation may constrain variation in resource acquisition strategies between lianas and trees.

Tropical montane systems are characterized by a high plant species diversity and complex environmental gradients. Climate warming may force species to track suitable climatic conditions and shift their distribution upward, which may be particularly problematic for species with narrow elevational ranges. To better understand the fate of montane plant species in the face of climate change, we evaluated a) which environmental factors best predict the distribution of 277 plant species along the Himalayan elevational gradient in Nepal, and b) whether species elevational ranges increase with increasing elevation. To this end, we developed ecological niche models using MaxEnt by combining species survey and presence data with 19 environmental predictors. Key environmental factors that best predicted the distribution of Himalayan plant species were mean annual temperature (for 54.5% of the species) followed by soil clay content (10.2%) and slope (9.4%). Although temperature is the best predictor, it is associated with many other covariates that may explain species distribution, such as irradiance and potential evapotranspiration. Species at both ends of the Himalayan elevational gradient had narrower elevational ranges than species in the middle. Our results suggest that with further global warming, most Himalayan plant species have to migrate upward, which is especially critical for upland species with narrow distribution ranges.

Among tropical forests, lianas are predicted to have a growth advantage over trees during seasonal drought, with substantial implications for tree and forest dynamics. We tested the hypotheses that lianas maintain higher water status than trees during seasonal drought and that lianas maximize leaf cover to match high, dry-season light conditions, while trees are more limited by moisture availability during the dry season. We monitored the seasonal dynamics of predawn and midday leaf water potentials and leaf phenology for branches of 16 liana and 16 tree species in the canopies of two lowland tropical forests with contrasting rainfall regimes in Panama. In a wet, weakly seasonal forest, lianas maintained higher water balance than trees and maximized their leaf cover during dry-season conditions, when light availability was high, while trees experienced drought stress. In a drier, strongly seasonal forest, lianas and trees displayed similar dry season reductions in leaf cover following strong decreases in soil water availability. Greater soil moisture availability and a higher capacity to maintain water status allow lianas to maintain the turgor potentials that are critical for plant growth in a wet and weakly seasonal forest but not in a dry and strongly seasonal forest.

Large lianas and trees in the forest canopy are challenged by hydraulic and mechanical failures and need to balance hydraulic conductivity, hydraulic safety and mechanical safety. Our study integrates these functions in canopy branches to understand the performance of canopy trees and lianas, and their difference. We sampled and measured branches from 22 species at a canopy crane in the tropical forest at Xishuangbanna, SW China. We quantified the hydraulic conductivity from the xylem-specific hydraulic conductivity (KS), hydraulic safety from the cavitation resistance (P50) and mechanical safety from the modulus of rupture (MOR) to evaluate trade-offs and differences between lianas and trees. We also measured a number of anatomical features that may influence these three functional traits. Our results suggest the following: trade-offs between hydraulic conductivity, hydraulic safety and mechanical safety are weak or absent; liana branches better resist external mechanical forces (higher MOR) than tree branches; and liana and tree branches were similar in hydraulic performance (KS and P50). The anatomical features underlying KS, P50 and MOR may differ between lianas and trees. We conclude that canopy branches of lianas and trees diverged in mechanical design due to fundamental differences in wood formation, but converged in hydraulic design.

Xylem conduit diameter (D max ) of woody angiosperm adults scales with plant size and widens from the stem apex downwards. We hypothesized that, notwithstanding relative growth rate (RGR), growth form or leaf habit, woody seedling conduit D max scales linearly with plant size across species; this scaling should be applicable to all vegetative organs, with consistent conduit widening from leaf via stem to main root and coupling with whole-leaf area and whole-stem xylem area. To test these hypotheses, organ-specific xylem anatomy traits and size-related traits in laboratory-grown seedlings were analyzed across 55 woody European species from cool-temperate and Mediterranean climates. As hypothesized, conduit D max of each organ showed similar scaling with plant size and consistent basipetal widening from the leaf midvein via the stem to the main root across species, independently of growth form, RGR and leaf habit. We also found a strong correlation between D max and average leaf area, and between stem xylem area and whole-plant leaf area. We conclude that seedlings of ecologically wide-ranging woody species converge in their allometric scaling of conduit diameters within and across plant organs. These relationships will contribute to modeling of water transport in woody vegetation that accounts for the whole life history from the trees’ regeneration phase to adulthood.

Reactivation of axial water flow in ring-porous species is a complex process related to stem water content and developmental stage of both earlywood-vessel and leaf formation. Yet empirical evidence with non-destructive methods on the dynamics of water flow resumption in relation to these mechanisms is lacking. Here we combined in vivo magnetic resonance imaging and wood-anatomical observations to monitor the dynamic changes in stem water content and flow during spring reactivation in 4-year-old pedunculate oaks (Quercus robur L.) saplings. We found that previous year latewood vessels and current year developing earlywood vessels form a functional unit for water flow during growth resumption. During spring reactivation, water flow shifted from latewood towards the new earlywood, paralleling the formation of earlywood vessels and leaves. At leaves' full expansion, volumetric water content of previous rings drastically decreased due to the near-absence of water in fibre tissue. We conclude (i) that in ring-porous oak, latewood vessels play an important hydraulic role for bridging the transition between old and new water-conducting vessels and (ii) that fibre and parenchyma provides a place for water storage.

Lianas are a distinctive component of tropical forests and their increase in abundance may have profound ramifications for forest composition and ecosystem functioning. However, so far, the current view considers lianas as a single, functional plant type and, therefore, ignores the life history differences among species resulting from their climbing mechanisms. Here, we integrate data from the main functional traits used to characterize plant form and function with global abundance data to reveal that lianas consistently have differences associated with their main climbing mechanisms. Overall, lianas with active climbing were characterized by an acquisitive strategy, showing higher specific leaf area, foliar nitrogen and slightly higher (marginally significant) maximum photosynthetic rates compared to lianas with passive climbing. Using structural equation modeling, we show that across the tropics the abundance of lianas with active climbing is mainly determined by forest structure (stem size distribution). Our study shows that active and passive climbing lianas clearly diverge in their functional traits and factors affecting their distribution. We conclude that a shift in the current view, where lianas are considered a single, functional plant type, is urgently needed to increase our predictability of their effects on tropical forests in the future.

Hydraulic failure caused by severe drought contributes to aboveground dieback and whole-plant death. The extent to which dieback or whole-plant death can be predicted by plant hydraulic traits has rarely been tested among species with different leaf habits and/or growth forms. We investigated 19 hydraulic traits in 40 woody species in a tropical savanna and their potential correlations with drought response during an extreme drought event during the El Niño–Southern Oscillation in 2015. Plant hydraulic trait variation was partitioned substantially by leaf habit but not growth form along a trade-off axis between traits that support drought tolerance versus avoidance. Semi-deciduous species and shrubs had the highest branch dieback and top-kill (complete aboveground death) among the leaf habits or growth forms. Dieback and top-kill were well explained by combining hydraulic traits with leaf habit and growth form, suggesting integrating life history traits with hydraulic traits will yield better predictions.

A central paradigm in comparative ecology is that species sort out along a global economic strategy spectrum, ranging from slow to fast growth. Many studies evaluated plant strategy spectra for leaf traits, b u t few studies evaluated stem strategy spectra using a comprehensive set of anatomical, chemical and morphological traits, addressing key stem functions of different stem compartments (inner wood, outer wood and bark). This study evaluates how stem traits vary in the wood and bark of temperate tree species, and whether a slow-fast growth strategy spectrum exists and what traits make up this plant strategy spectrum. For 14 temperate gymnosperm and angiosperm species, 20 traits belonging to six key stem functions were measured for three stem compartments. Both across and within gymnosperms and angiosperms, a slow-fast stem strategy spectrum is found. Gymnosperms have slow traits and showed converging stem strategies because of their uniform tracheids. Angiosperms have fast traits and showed diverging stem strategies because of a wider array of tissues (vessels, parenchyma and fibers) and vessel size and arrangements (ring-porous versus diffuse porous). Gymnosperms showed a main trade-off between hydraulic efficiency and safety, and angiosperms showed a main trade-off between ‘slow’ diffuse porous species and ‘fast’ ring porous species. The slow traits of gymnosperms allow for a high hydraulic safety, an evergreen leaf habit and steady but slow growth makes them successful in unproductive habitats whereas the fast traits of angiosperms allow for high conductivity, a deciduous leaf habit and fast growth which makes them successful in productive habitats.

Background and Aims Conifers are key components of many temperate and boreal forests and are important for forestry, but species differences in stem growth responses to climate are still poorly understood and may hinder effective management of these forests in a warmer and drier future. Methods We studied 19 Northern Hemisphere conifer species planted in a 50-year-old common garden experiment in the Netherlands to (1) assess the effect of temporal dynamics in climate on stem growth, (2) test for a possible positive relationship between the growth potential and climatic growth sensitivity across species, and (3) evaluate the extent to which stem growth is controlled by phylogeny. Key results Eighty-nine per cent of the species showed a significant reduction in stem growth to summer drought, 37 % responded negatively to spring frost and 32 % responded positively to higher winter temperatures. Species differed largely in their growth sensitivity to climatic variation and showed, for example, a four-fold difference in growth reduction to summer drought. Remarkably, we did not find a positive relationship between productivity and climatic sensitivity, but instead observed that some species combined a low growth sensitivity to summer drought with high growth potential. Both growth sensitivity to climate and growth potential were partly phylogenetically controlled. Conclusions A warmer and drier future climate is likely to reduce the productivity of most conifer species. We did not find a relationship between growth potential and growth sensitivity to climate; instead, some species combined high growth potential with low sensitivity to summer drought. This may help forest managers to select productive species that are able to cope with a warmer and drier future.

Altres ajuts: acords transformatius de la UAB A major foundation of trait-based ecology is that traits have an impact on individual performance. However, trait-growth relationships have not been extensively assessed in trees, especially outside tropical ecosystems. In addition, measuring traits directly related to physiological processes remains difficult and the differences between inter- and intraspecific relationships are seldom explored. Here, we use individual-level data on a set of hydraulic, leaf and stem traits to assess their ability to predict basal area increment (BAI) and growth efficiency (BAI per unit of tree leaf area, GE) among and within species for six dominant tree species along a water availability gradient under Mediterranean climate (Catalonia, NE Spain). Measured traits include: leaf mass per area (LMA), leaf nitrogen concentration (N), leaf C isotopic composition (δC), the leaf water potential at turgor loss (P), stem wood density (WD) and branch-level estimates of the Huber value (Hv), sapwood- and leaf-specific hydraulic conductivity (K and K) and resistance to xylem embolism (P). Trait-growth associations were generally weak, particularly for BAI and within species. High values of both growth metrics were associated with 'conservative' leaf and hydraulic traits. In particular, BAI was negatively associated with K (and wood density), while GE increased with LMA, allocation to sapwood relative to leaves (Hv) and resistance to xylem embolism (P). Climate effects on BAI and GE were indirectly mediated by changes in traits, stand structure and tree basal area. Overall, these results suggest that maintaining functionality over extended periods of time may be more important than maximum gas exchange or hydraulic capacity to achieve high radial growth under Mediterranean climates. Our study reveals that widely used 'functional traits' may be poor predictors of tree growth variability along environmental gradients. Moreover, trait effects (when present) do not necessarily conform to simple hypotheses based on our understanding of organ-level processes. An improved understanding of trait coordination along common axes of variation together with a revaluation of the variables that better reflect whole-tree performance can greatly improve our understanding of trait-growth relationships. A free Plain Language Summary can be found within the Supporting Information of this article.

The timing of diel stem growth of mature forest trees is still largely unknown, as empirical data with high temporal resolution have not been available so far. Consequently, the effects of day–night conditions on tree growth remained uncertain. - Here we present the first comprehensive field study of hourly-resolved radial stem growth of seven temperate tree species, based on 57 million underlying data points over a period of up to 8 yr. - We show that trees grow mainly at night, with a peak after midnight, when the vapour pressure deficit (VPD) is among the lowest. A high VPD strictly limits radial stem growth and allows little growth during daylight hours, except in the early morning. Surprisingly, trees also grow in moderately dry soil when the VPD is low. Species-specific differences in diel growth dynamics show that species able to grow earlier during the night are associated with the highest number of hours with growth per year and the largest annual growth increment. - We conclude that species with the ability to overcome daily water deficits faster have greater growth potential. Furthermore, we conclude that growth is more sensitive than carbon uptake to dry air, as growth stops before stomata are known to close. ISSN:0028-646X ISSN:1469-8137

Plant functional traits and strategies hold the promise to explain species distribution, but few studies have linked multiple traits to multiple niche dimensions (i.e., light, water, and nutrients). Here, we analyzed for 29 liana species in a Chinese tropical seasonal rainforest how: (1) trait associations and trade-offs lead to different plant strategies; and (2) how these traits shape species’ niche dimensions. Eighteen functional traits related to light, water, and nutrient use were measured and species niche dimensions were quantified using species distribution in a 20-ha plot combined with data on canopy gaps, topographic water availability, and soil nutrients. We found a tissue toughness spectrum ranging from soft to hard tissues along which species also varied from acquisitive to conservative water use, and a resource acquisition spectrum ranging from low to high light capture and nutrient use. Intriguingly, each spectrum partly reflected the conservative–acquisitive paradigm, but at the same time, the tissue toughness and the resource acquisition spectrum were uncoupled. Resource niche dimensions were better predicted by individual traits than by multivariate plant strategies. This suggests that trait components that underlie multivariate strategy axes, rather than the plant strategies themselves determine species distributions. Different traits were important for different niche dimensions. In conclusion, plant functional traits and strategies can indeed explain species distributions, but not in a simple and straight forward way. Although the identification of global plant strategies has significantly advanced the field, this research shows that global, multivariate generalizations are difficult to translate to local conditions, as different components of these strategies are important under different local conditions. Supplementary Information The online version contains supplementary material available at 10.1007/s00442-021-04937-4.

Plant functional traits determine plant performance and have therefore the potential to shape and predict species distributions along environmental gradients. This study analyses how traits affect tree species distribution along an elevational gradient in the Himalayas, Nepal. We addressed three questions: (a) what plant strategies can be distinguished among tree species? (b) how are plant traits and strategies associated with elevation? and (c) what plant traits are the best predictors of species positions along elevational gradient? We quantified for 31 tree species a set of 39 plant traits related to resource uptake, use and conservation. We analysed how traits cluster into separate functions using a cluster analysis, and how traits and clusters associate into distinct plant strategies using a principal component analysis. The cluster analysis showed five clusters of traits, reflecting (a) efficiency in vertical expansion, (b) efficiency in horizontal expansion, (c) efficiency in metabolism, (d) physical defence and (e) tree functional groups (conifers vs. broadleaf species). The first PCA axis reflects trade-offs in traits and clusters linked to elevation: highland species had trait values that increase safety against freezing induced cavitation, high solar radiation and strong wind, and that increase resource conservation. In contrast, lowland species had trait values that increase efficiency of resource acquisition, metabolism and expansion. Additionally, a bivariate analysis showed that the first PCA axis is more strongly related to elevation than the individual traits, indicating that the combination of traits is more important than the individual traits. An all subsets regression analysis showed that a small set of traits best explain species distribution: highland species had smaller size (low basal area), safer hydraulics (low conduit diameter) and lower leaf and branch display efficiency (low leaf area per xylem area, specific branch length) that increase persistence under harsh conditions. Remarkably, leaf traits were poor predictors of species' elevational positions. Synthesis. Multiple trade-offs in plant size, hydraulics and light competitiveness shape species distribution along the elevational gradient. Along this extreme environmental gradient, stem and branch traits that integrate multiple plant organs and functions are better predictors of species' elevational distributions than leaf traits.

As a response to the increased pressure of global climate change on most ecosystems, national and international agreements aim at creating forests that are productive, resilient to climate change, and that store carbon to mitigate global warming. However, these aims are being challenged by increased tree mortality rates and decreased tree growth rates in response to increased incidence of extreme drought events. These phenomena make us aware of a lack of crucial insights into the effects of forest management on the growth and survival of trees, and on carbon storage in both trees and forest soils under increased incidence of drought. Here we compile current knowledge on how forest management and drought impact on tree growth and survival, and above- and belowground carbon storage in forest ecosystems. Based on this, we propose that climate-smart forestry may benefit from controlling stand density at intermediate levels (>60%, e.g.~80%) by applying low levels of tree harvest intensity on a regular base. Furthermore, we propose that the actual optimal density will depend on the tree species, site conditions and management history. As a next step, studies are needed that take an above- and belowground approach and combine forest experiments with mechanistic models on water, carbon and nutrient flows in trees and soils within forests in order to transform current results, which focus on either soil or trees and are often highly-context dependent, to a more generic forest framework. Such a generic framework would be needed to enhance understanding across forest ecosystems on how forest management may promote forest resilience, productivity and carbon storage with increasing drought.

Questions Lianas are a conspicuous element of tropical forests but have largely been ignored in species‐level vegetation surveys. As a result, there is limited understanding of how environmental factors structure liana communities. Location A 20‐ha forest dynamics plot in Xishuangbanna National Nature Reserve, southwestern China. Methods We evaluated the distribution of the 50 most abundant liana species, comprising >18,000 individuals, in the 20‐ha forest plot. Ordination analysis and generalized linear mixed models were used to evaluate how species distribution and abundance are associated with soil pH, soil phosphorus (P), soil nitrogen (N), and soil potassium (K), canopy gaps and topography. We calculated the average weighted distribution as a proxy for the optimum resource condition for each species. Results The first two axes of a canonical correspondence analysis explained 65% of the variation in liana species composition, with pH and P being the strongest drivers and highly correlated with each other. We modelled the responses of liana species to soil nutrients, and found a negative, unimodal or positive response of liana abundance with increasing soil nutrient concentrations. Forty‐six of the 50 species occurred under significantly higher or lower soil nutrient conditions than expected at random. Lianas mainly separated along the P gradient, whereas for N and K most liana species tended to occupy locations with high nutrient concentrations. Conclusions Although lianas are thought to be notoriously light‐demanding, soil conditions were stronger drivers of liana species distribution than gaps. Species differences in distributions were mainly driven by soil gradients in pH and P, highlighting the importance of soil nutrient status for liana niche partitioning in wet tropical forests on highly weathered soils. Most liana species had high resource requirements for N, K and light, which come along with their fast growth and acquisitive resource use strategy. Hence, below‐ground resource availability plays an important role in shaping the assembly of liana communities.

Severe droughts have the potential to reduce forest productivity and trigger tree mortality. Most trees face several drought events during their life and therefore resilience to dry conditions may be crucial to long-term survival. We assessed how growth resilience to severe droughts, including its components resistance and recovery, is related to the ability to survive future droughts by using a tree-ring database of surviving and now-dead trees from 118 sites (22 species, >3,500 trees). We found that, across the variety of regions and species sampled, trees that died during water shortages were less resilient to previous non-lethal droughts, relative to coexisting surviving trees of the same species. In angiosperms, drought-related mortality risk is associated with lower resistance (low capacity to reduce impact of the initial drought), while it is related to reduced recovery (low capacity to attain pre-drought growth rates) in gymnosperms. The different resilience strategies in these two taxonomic groups open new avenues to improve our understanding and prediction of drought-induced mortality., Resilience to drought is crucial for tree survival under climate change. Here, DeSoto et al. show that trees that died during drought were less resilient to previous dry events compared to surviving conspecifics, but the resilience strategies differ between angiosperms and gymnosperms.

1. The root economics spectrum (RES) hypothesis predicts that fast-growing tree species have short-lived roots with high specific root length (SRL) to allow rapid resource uptake, and opposite trait expressions for slow-growing species. Yet, the mixed support for this hypothesis suggests that trees can adopt alternative strategies to increase resource uptake, besides an increase in SRL. 2. We combined a novel mechanistic whole-tree model and empirical fine-root data of ten tree species to test the effects of one of these alternative strategies, notably increasing fine-root mass, on the tree’s net C gain (used here as a proxy for tree performance), and to assess how fine-root lifespan influences the relative importance of SRL and fine-root mass for the C balance of trees. 3. Our results indicate that accounting for the short lifespan of high-SRL roots has important implications for explaining tree performance and the role of roots herein. Without considering their faster turnover, high-SRL roots and low fine-root mass resulted in the highest performance as predicted from the RES. Yet, when their higher turnover rates were accounted for, a high fine-root mass and low SRL lead to the highest performance. Both our model outcomes and field data further show a negative relationship between SRL and fine-root mass through which species aim to realise a similar root length density. This trade-off further indicates how high a SRL and low fine-root mass as well as opposite trait values can both lead to a positive C balance in a similar environment. 4. Our study may explain why high-SRL roots do not necessarily lead to the fastest tree growth as often hypothesised and demonstrates the importance of fine-root mass in combination with fine-root lifespan for explaining interspecific differences in tree performance. More generally, our work demonstrates the value of identifying and investigating different belowground strategies across species from a whole-plant modelling perspective, and identifies the relationship between SRL, fine-root biomass and lifespan as an important functional dimension to variation in species’ performance.

Drought limits the long-distance transport of water in the xylem due to the reduced leaf-to-soil water potential difference and possible embolism-related losses of conductance and of sugars in the phloem due to the higher viscosity of the dehydrated sugary solution. This condition can have cascading effects in water and carbon (C) fluxes that may ultimately cause tree death. We hypothesize that the maintenance of xylem and phloem conductances is fundamental for survival also under reduced resource availability, when trees may produce effective and low C cost anatomical adjustments in the xylem and phloem close to the treetop where most of the hydraulic resistance is concentrated. We analyzed the treetop xylem and phloem anatomical characteristics in coexisting Scots pine trees, symptomatic and non-symptomatic of drought-induced dieback. We selected the topmost 55 cm of the main stem and selected several sampling positions at different distances from the stem apex to test for differences in the axial patterns between the two groups of trees. We measured the annual ring area, the tracheid hydraulic diameter (Dh) and cell wall thickness (CWT), the conductive phloem area and the average lumen diameter of the 20 largest phloem sieve cells (Dph). Declining trees grew less than the non-declining ones, and despite the similar axial scaling of anatomical traits, had larger Dh and lower CWT. Moreover, declining trees had wider Dph. Our results demonstrate that even under drought stress, maintenance of xylem and phloem efficiencies is of primary importance for survival, even if producing fewer larger tracheids may lead to a xylem more vulnerable to embolism formation.

Height–diameter allometric (H-D) relationships play an important role in the light capture and stability of a tree, and it is poorly understood whether the relationships changes with growth stages or functional groups along a large scale environmental gradients. We present a comparative study of H-D relationships of 6810 seedlings and 19,707 trees from about 1000 species in 9 different sites across China. We showed that allometric exponents for seedlings and trees differ between different sites and do not represent a single, constant theoretical value (e.g. 2/3, 1/2 or 1). Seedlings had greater exponents than trees in most sites. The exponents varied between canopy and understory trees in 5 sites, while were indistinguishable from each other in the remaining 4 sites. Canopy-tree seedlings had smaller exponents than understory-tree seedlings in 3 sites but had indistinguishable exponents in the other 6 sites. Gymnosperm trees had exponents greater than or indistinguishable from angiosperm trees. Elevation alone, or together with mean temperature of the warmest month explained variation of tree allometries for canopy-tree and understory-tree seedlings. The exponents of canopy trees decreased with mean annual precipitation. Our results do not support the predictions from metabolic scaling theory or biomechanical models that height–diameter allometries are invariant. Our study provides insight into how ontogeny, adult stature, phyletic affiliations and environmental limitations affect height–diameter allometric relationships at biogeographical scales.

Some temperate tree species mitigate the negative impacts of frost-induced xylem cavitation by restoring impaired hydraulic function via positive pressures, and may therefore be more resistant to frost fatigue (the phenomenon that post-freezing xylem becomes more susceptible to hydraulic dysfunction) than nonpressure-generating species. We test this hypothesis and investigate underlying anatomical/physiological mechanisms. Using a common garden experiment, we studied key hydraulic traits and detailed xylem anatomical characteristics of 18 sympatric tree species. These species belong to three functional groups, that is, one generating both root and stem pressures (RSP), one generating only root pressure (RP), and one unable to generate such pressures (NP). The three functional groups diverged substantially in hydraulic efficiency, resistance to drought-induced cavitation, and frost fatigue resistance. Most notably, RSP and RP were more resistant to frost fatigue than NP, but this was at the cost of reduced hydraulic conductivity for RSP and reduced resistance to drought-induced cavitation for RP. Our results show that, in environments with strong frost stress: these groups diverge in hydraulic functioning following multiple trade-offs between hydraulic efficiency, resistance to drought and resistance to frost fatigue; and how differences in anatomical characteristics drive such divergence across species.

Recent studies show that mixed species forests sometimes have higher stand productivity than monospecific forests, which we refer to as overyielding. Yet, results for temperate forests are ambiguous, possibly because forests differ in local site conditions, thinning history and forest age. In line with the niche complementarity hypothesis, we expect stronger overyielding for forests with species differing in both leaf phenology (evergreen or deciduous) and shade tolerance. We also hypothesize that overyielding will decrease with stand development because of decreasing resource availability. We compared 4 two-species mixtures with their corresponding monospecific stands from long-term field measurements in the Netherlands. The mixtures were Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco)–common beech (Fagus sylvatica L.), Scots pine (Pinus sylvestris L.)–common oak (Quercus robur L.), common oak–common beech, and common oak–silver birch (Betula pendula Roth). Overyielding was observed in 2 of the 4 mixtures: Douglas-fir–common beech mixtures had 35.9% and 36.7% higher volume growth relative to Douglas-fir and common beech monocultures, respectively; Scots pine–common oak mixtures had 20.3% and 31.2% higher volume growth relative to Scots pine and common oak monocultures, respectively, on average over time. Furthermore, overyielding was relatively constant for the two mixtures through stand development. This result was robust after accounting for possible effects of site quality and thinning history, where site quality contributed independently to stand productivity and thinning history had no effect. No significant overyielding effects were observed for the two deciduous mixed stands, i.e. common oak–common beech and common oak–silver birch. Mixing tree species in temperate forests resulted in overyielding for evergreen–deciduous species mixtures, but not for deciduous–deciduous species mixtures. This indicates that leaf phenology contributes to overyielding effects. Overyielding was higher in the Douglas-fir–common beech mixtures than the Scots pine–common oak mixtures, which coincides with a stronger contrast in shade tolerance between Douglas-fir and common beech and thus stronger complementarity. Our results support the complementarity hypothesis and imply that such mechanisms are maintained with stand development. It therefore appears that mixing evergreen with deciduous species with contrasting shade tolerance is a valid management strategy for increasing diversity and productivity of temperate forests in the Netherlands.

Plasticity of biomass allocation is a key to growth and survival of trees exposed to variable levels of stress in their lifetime. Most of our understanding of dynamic biomass allocation comes from seedling studies, but plasticity may be different in mature trees. We used stem analysis to reconstruct whole-tree growth and biomass allocation patterns in Quercus pubescens trees harvested from a dry woodland in Valais, Switzerland. We identified three distinct growth phases. In phase I, a primary root developed but the aboveground structure did not persist. In phase II, height growth occurred and secondary roots developed. In phase III, height growth ceased and stems and roots only grew radially. Reference trees harvested from a less dry site nearby only showed phase II-type growth. In line with our hypothesis, drought-stressed trees maintained more biomass in roots and less in aboveground woody parts than reference trees. Contrary to our expectation, stressed trees allocated proportionally more resources to leaves and less to roots in the growing season before harvest than reference trees. It appears that sub-seasonal wood anatomical adjustments to water availability minimize hydraulic failure, thus enabling these dry woodland trees to invest preferentially in leaves. Wet years did not see preferential investment in aboveground tissues, suggesting more restricted plasticity in biomass allocation in these mature trees than in seedlings. It is concluded that trees beyond seedling stage show different responses to variation in drought than the better-studied seedlings.

Trees scale leaf (A(L)) and xylem (A(X)) areas to couple leaf transpiration and carbon gain with xylem water transport. Some species are known to acclimate in A(L) : A(X) balance in response to climate conditions, but whether trees of different species acclimate in A(L) : A(X) in similar ways over their entire (continental) distributions is unknown. We analyzed the species and climate effects on the scaling of A(L) vs A(X) in branches of conifers (Pinus sylvestris, Picea abies) and broadleaved (Betula pendula, Populus tremula) sampled across a continental wide transect in Europe. Along the branch axis, A(L) and A(X) change in equal proportion (isometric scaling: b similar to 1) as for trees. Branches of similar length converged in the scaling of A(L) vs A(X) with an exponent of b = 0.58 across European climates irrespective of species. Branches of slow- growing trees from Northern and Southern regions preferentially allocated into new leaf rather than xylem area, with older xylem rings contributing to maintaining total xylem conductivity. In conclusion, trees in contrasting climates adjust their functional balance between water transport and leaf transpiration by maintaining biomass allocation to leaves, and adjusting their growth rate and xylem production to maintain xylem conductance.

Tree species largely differ in the amount of annual stem growth explained by current-year conditions. Historic conditions have been shown to additionally explain a significant fraction of the unexplained variance. So far there is no mechanism described explaining why species differ in such legacy effects, obscuring our understanding of species differences in annual stem growth responses to climate. We present a generic conceptual view on key processes determining stem growth. We link current and historic conditions and their impacts on growth by considering the lifetime of functional organs (leaves and sapwood) and reserves (carbon pool) as a way to quantify legacy effects. We propose how tree species with long organ lifetimes are determined by longer periods of historic conditions than species with short organ lifetimes, and why these species differ in their responses to current conditions. We investigated the hypothesis that including lifetime as variable in a process-based tree model allows for explaining different growth responses to current-year conditions. We show that species with short organ and reserve lifetimes are more sensitive to—and better track—current environmental conditions and therefore respond more strongly to current conditions than species with long lifetimes. Instead, the species with longer organ lifetimes respond more strongly to historic conditions and thus buffer their growth responses to current conditions. We propose the impact of historic environmental conditions being controlled by organ and reserve lifetimes and partially explaining the strength of legacy effects and the explanatory power of current-year environmental conditions on stem growth of different species.

International audience; Many studies have reported that hydraulic properties vary considerably between tree species, but little is known about their intraspecific variation and, therefore, their capacity to adapt to a warmer and drier climate. Here, we quantify phenotypic divergence and clinal variation for embolism resistance, hydraulic conductivity and branch growth, in four tree species, two angiosperms (Betula pendula, Populus tremula) and two conifers (Picea abies, Pinus sylvestris), across their latitudinal distribution in Europe. Growth and hydraulic efficiency varied widely within species and between populations. The variability of embolism resistance was in general weaker than that of growth and hydraulic efficiency, and very low for all species but Populus tremula. In addition, no and weak support for a safety vs. efficiency trade-off was observed for the angiosperm and conifer species, respectively. The limited variability of embolism resistance observed here for all species except Populus tremula, suggests that forest populations will unlikely be able to adapt hydraulically to drier conditions through the evolution of embolism resistance.

A growing number of studies provides evidence that mixed-species forests often have higher stand productivity than monospecific forests, which is referred to as overyielding. In this study, we explored how the combination of species and soil conditions affect overyielding in terms of periodic annual volume increment (PAIV) in Dutch forests. We studied Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), common beech (Fagus sylvatica L.), Scots pine (Pinus sylvestris L.), pedunculate oak (Quercus robur L.), and silver birch (Betula pendula Roth) growing in four two species combinations (Douglas-fir–common beech, Scots pine–pedunculate oak, pedunculate oak–common beech, and pedunculate oak–silver birch) from 398 long-term permanent field plots all over the Netherlands. We found that the Douglas-fir–common beech and Scots pine–pedunculate oak mixtures always showed overyielding. This overyielding was largely attributed to the Douglas-fir in the former mixture and to the pedunculate oak in the latter mixture, respectively. In both cases, overyielding was stronger at poor soils than at rich soils. The pedunculate oak–common beech mixtures overyielded at poor soils and underyielded at rich soils, which was attributed to the response of the common beech. Overyielding was not observed for the pedunculate oak–silver birch mixtures, irrespective of soil conditions. The results do not support our hypothesis since overyielding was not always driven by fast-growing light-demanding species. Overyielding was stronger for evergreen–deciduous species combinations, suggesting that differences in leaf phenology are a major driver of overyielding. Secondly, our results imply that overyielding is much stronger at poor soils than at rich soils, which is in line with the prediction of the stress-gradient hypothesis. We conclude that the growth of one species benefits from the admixture species, particularly in evergreen–deciduous species mixtures and that soils affect the extent of overyielding as studied in the Netherlands.

The persistence of tropical rainforests, together with their flora and fauna, is highly threatened by anthropogenic disturbances. In this study, we investigate to what extent selective logging influences the structure and composition of a tropical rainforest in Madagascar and subsequently lemur encounter rates and cluster sizes. We quantified forest structure variables and conducted transect surveys of seven sympatric diurnal lemur species in five protected forest sites with different logging histories. We found that DBH, tree height, the interquartile ranges of DBH and tree height (measure of forest heterogeneity), tree species and family richness were relatively high and tree density was relatively low in less disturbed compared to disturbed sites. Although the disturbed forests have not fully recovered to previous conditions, they seem to have recovered from a functional perspective into suitable lemur habitat, as lemur encounter rates and cluster sizes were similar in disturbed and less disturbed sites. We only found slightly higher encounter rates for Varecia variegata (P = 0.078) and lower encounter rates for Eulemur rufifrons (P = 0.059) in less disturbed forests. This is one of the first studies that report the presence of V. variegata, a species characterised by its drastic decline, in previously logged sites. Lemurs travelling between disturbed and less disturbed sites disperse seeds and hereby facilitate forest regeneration. Therefore, we promote the need for better attention to the value of logged forests for biodiversity conservation in Madagascar and suggest that there is considerable potential for regenerating logged forests to support lemur communities.

Many monoculture forests have been converted to mixed-species forests in Europe over the last decades. The main reasons for this conversion were probably to increase productivity, including timber production, and enhance other ecosystem services, such as conservation of biodiversity and other nature values. This study was done by synthesizing results from studies carried out in Dutch mixed forests compared with monoculture stands and evaluating them in the perspective of the current theory. Then we explored possible mechanisms of higher productivity in mixed stands, in relation to the combination of species, stand age and soil fertility, and discussed possible consequences of forest management. The study covered five two-species mixtures and their corresponding monoculture stands from using long-term permanent forest plots over multiple decades as well as two inventories (around 2003 and 2013) across the entire Netherlands. These forest plot data were used together with empirical models at total stand level, species level and tree level. Overyielding in Douglas-fir&ndash, beech and pine&ndash, oak mixtures was maintained over time, probably owing to the intensive thinning and was achieved on the poorer soils. However, this overyielding was not always driven by fast-growing light-demanding species. On individual tree level, intra-specific competition was not necessarily stronger than inter-specific competition and this competitive reduction was less seen at lower soil fertility and dependent on species mixtures. Moreover, size-asymmetric competition for light was more associated with tree basal area growth than size-symmetric competition for soil resources. Overall, this study suggests a substantial potential of species mixing for increasing productivity and implies developing forest management strategies to convert monospecific forests to mixed-species forests that consider the complementarity in resource acquisition of tree species.

Boswellia neglecta S. Moore is a frankincense-producing tree species dominantly found in the dry woodlands of southeastern Ethiopia. Currently, the population of this socio-economically and ecologically important species is threatened by complex anthropogenic and climate change related factors. Evaluation of tree age and its radial growth dynamics in relation to climate variables helps to understand the response of the species to climate change. It is also crucial for sustainable forest resource management and utilization. Dendrochronological and remote-sensing techniques were used to study periodicity of wood formation and leaf phenology and to assess the growth dynamics of B. neglecta. The results show that B. neglecta forms two growth rings per year in the study area. The growth ring structure is characterized by larger vessels at the beginning of each growing season and smaller vessels formed later in the growing season, suggesting adaptation to decreasing soil moisture deficits at the end of the growing season. Seasonality in cambial activity matches with a bimodal leaf phenological pattern. The mean annual radial growth rate of B. neglecta trees is 2.5 mm. Tree age varied between 16 and 28 years, with an average age of 22 years. The young age of these trees indicates recent colonization of B. neglecta in the study region. The growth rate and seasonal canopy greenness (expressed by Normalized Difference Vegetation Index – NDVI) were positively correlated with rainfall, suggesting that rainfall is the main climatic factor controlling growth of B. neglecta. The observed temporal changes in leaf phenology and vessel size across the growth rings indicate that the species is drought tolerant. Therefore, it can be regarded as a key tree species for restoration of moisture-related limited areas across the Horn of Africa.

Bioclimate envelope models have been widely used to illustrate the discrepancy between current species distributions and their potential habitat under climate change. However, the realism and correct interpretation of such projections has been the subject of considerable discussion. Here, we investigate whether climate suitability predictions correlate to tree growth, measured in permanent inventory plots and inferred from tree‐ring records. We use the ensemble classifier RandomForest and species occurrence data from ~200,000 inventory plots to build species distribution models for four important European forestry species: Norway spruce, Scots pine, European beech, and pedunculate oak. We then correlate climate‐based habitat suitability with volume measurements from ~50‐year‐old stands, available from ~11,000 inventory plots. Secondly, habitat projections based on annual historical climate are compared with ring width from ~300 tree‐ring chronologies. Our working hypothesis is that habitat suitability projections from species distribution models should to some degree be associated with temporal or spatial variation in these growth records. We find that the habitat projections are uncorrelated with spatial growth records (inventory plot data), but they do predict interannual variation in tree‐ring width, with an average correlation of .22. Correlation coefficients for individual chronologies range from values as high as .82 or as low as −.31. We conclude that tree responses to projected climate change are highly site‐specific and that local suitability of a species for reforestation is difficult to predict. That said, projected increase or decrease in climatic suitability may be interpreted as an average expectation of increased or reduced growth over larger geographic scales.

AimThe fine roots of trees may show plastic responses to their resource environment. Several, contrasting hypotheses exist on this plasticity, but empirical evidence for these hypotheses is scattered. This study aims to enhance our understanding of tree root plasticity by examining intra-specific variation in fine-root mass and morphology, fine-root growth and decomposition, and associated mycorrhizal interactions in beech (Fagus sylvatica L.) and spruce (Picea abies (L.) Karst.) forests on soils that differ in resource availability.MethodsWe measured the mass and morphological traits of fine roots (i.e. ≤ 2 mm diameter) sampled to 50 cm depth. Fine-root growth was measured with ingrowth cores, and fine-root decomposition with litter bags. Mycorrhizal fungal biomass was determined using ingrowth mesh bags.ResultsBoth tree species showed more than three times higher fine-root mass, and a ten-fold higher fine-root growth rate on sand than on clay, but no or marginal differences in overall fine-root morphology. Within the fine-root category however, beech stands had relatively more root length of their finest roots on clay than on sand. In the spruce stands, ectomycorrhizal mycelium biomass was larger on sand than on clay.ConclusionsIn temperate beech and spruce forests, fine-root mass and mycorrhizal fungal biomass, rather than fine-root morphology, are changed to ensure uptake under different soil resource conditions. Yet enhancing our mechanistic understanding of fine-root trait plasticity and how it affects tree growth requires more attention to fine-root dynamics, the functional diversity within the fine-roots, and mycorrhizal symbiosis as an important belowground uptake strategy.

Leaf-stem allometry is an important spectrum that linked to biomass allocation and life history strategy in plants, although the determinants and evolutionary significance of leaf-stem allometry remain poorly understood. Leaf and stem architectures - including stem area/mass, petiole area/mass, lamina area/mass, leaf number, specific leaf area (LA), and mass-based leafing intensity (LI) - were measured on the current-year branches for 28 Ficus species growing in a common garden in SW China. The leaf anatomical traits, stem wood density (WD), and stem anatomical and mechanical properties of these species were also measured. We analyzed leaf-stem allometric relationships and their associations with stem hydraulic ad mechanical properties using species-level data and phylogenetically independent contrasts. We found isometric relationship between leaf lamina area/mass and stem area/mass, suggesting that the biomass allocation to leaf was independent to stem size. However, allometric relationship between LA/mass and petiole mass was found, indicating large leaves invest a higher fractional of biomass in petiole than small ones. LI, i.e., leaf numbers per unit of stem mass, was negatively related with leaf and stem size. Species with larger terminal branches tend to have larger vessels and theoretical hydraulic conductivity, but lower WD and mechanical strength. The size of leaf lamina, petiole, and stem was correlated positively with stem theoretical hydraulic conductivity, but negatively with stem WD and mechanical strength. Our results suggest that leaf-stem allometry in Ficus species was shaped by the trade-off between stem hydraulic efficiency and mechanical stability, supporting a functional interpretation of the relationship between leaf and stem dimensions.

Niche differentiation is a major hypothesized determinant of species distributions, but its practical importance is heavily debated and its underlying mechanisms are poorly understood. Trait-based approaches have been used to infer niche differentiation and predict species distributions. For understanding underlying mechanisms, individual traits should be scaled up to whole-plant performance, which has rarely been done. We measured seven key traits that are important for carbon and water balance for 37 tropical tree species. We used a process-based plant physiological model to simulate the carbon budget of saplings along gradients of light and water availability, and quantified the performance of the species in terms of their light compensation points (a proxy for shade tolerance), water compensation points (proxy for drought tolerance), and maximum carbon gain rates (proxy for potential growth rate). We linked species performances to their observed distributions (the realized niches) at two spatial scales in Bolivian lowland forests: along a canopy openness gradient at local scale (~1 km2) and along a rainfall gradient (1100–2200 mm/yr) at regional (~1000 km) scale. We show that the water compensation point was the best predictor of species distributions along water and light resource gradients within and across tropical forests. A sensitivity analysis suggests that the stomatal regulation of minimum leaf water potentials, rather than stem hydraulic traits (sapwood area and specific conductivity), contributed to the species differences in the water compensation point of saplings. The light compensation point and maximum carbon gain, both driven by leaf area index and leaf nitrogen concentration, also contributed to differential species distributions at the local scale, but not or only marginally at the regional scale. Trait-and-physiology-based simulations of whole-plant performance thus help to evaluate the possible roles of individual traits in physiological processes underlying species performance along environmental gradients. The development of such whole-plant concepts will improve our ability to understand responses of plant communities to shifts in resource availability and stress under global change.

Planting provenances originating from southern to northern locations has been discussed as a strategy to speed up species migration and mitigate negative effects of climate change on forest stability and productivity. Especially for drought-susceptible species such as European beech (Fagus sylvatica L.), the introduction of drought-tolerant provenances from the south could be an option. Yet, beech has been found to respond plastically to environmental conditions, suggesting that the climate on the plantation site might be more important for tree growth than the genetic predisposition of potentially drought-adapted provenances. In this study, we compared the radial growth, wood-anatomical traits and leaf phenology of four beech provenances originating from southern (Bulgaria, France) and northern locations (Sweden, the Netherlands) and planted in a provenance trial in the Netherlands. The distribution of precipitation largely differs between the sites of origin. The northern provenances experience a maximum and the southern provenances experience a minimum of rainfall in summer. We compared tree productivity and the anatomy of the water-conducting system for the period from 2000 to 2010, including the drought year 2003. In addition, tree mortality and the timing of leaf unfolding in spring were analysed for the years 2001, 2007 and 2012. Comparison of these traits in the four beech provenances indicates the influence of genetic predisposition and local environmental factors on the performance of these provenances under moderate site conditions. Variation in radial growth was controlled by environment, although the growth level slightly differed due to genetic background. The Bulgarian provenance had an efficient water-conducting system which was moreover unaffected by the drought in 2003, pointing to a high ability of this provenance to cope well with dry conditions. In addition, the Bulgarian provenance showed up as most productive in terms of height and radial growth. Altogether, we conclude that the similarity in ring-width variation among provenances points to environmental control of this trait, whereas the differences encountered in wood-anatomical traits between the well-performing Bulgarian provenance and the other three provenances, as well as the consistent differences in flushing pattern over 3 years under various environmental conditions, support the hypothesis of genetic control of these features.

Tree architecture, growth, and mortality change with increasing tree size and associated light conditions. To date, few studies have quantified how size-dependent changes in growth and mortality rates co-vary with architectural traits, and how such size-dependent changes differ across species and possible light capture strategies. We applied a hierarchical Bayesian model to quantify size-dependent changes in demographic rates and correlated demographic rates and architectural traits for 145 co-occurring Malaysian rain-forest tree species covering a wide range of tree sizes. Demographic rates were estimated using relative growth rate in stem diameter (RGR) and mortality rate as a function of stem diameter. Architectural traits examined were adult stature measured as the 95-percentile of the maximum stem diameter (upper diameter), wood density, and three tree architectural variables: tree height, foliage height, and crown width. Correlations between demographic rates and architectural traits were examined for stem diameters ranging from 1 to 47 cm. As a result, RGR and mortality varied significantly with increasing stem diameter across species. At smaller stem diameters, RGR was higher for tall trees with wide crowns, large upper diameter, and low wood density. Increased mortality was associated with low wood density at small diameters, and associated with small upper diameter and wide crowns over a wide range of stem diameters. Positive correlations between RGR and mortality were found over the whole range of stem diameters, but they were significant only at small stem diameters. Associations between architectural traits and demographic rates were strongest at small stem diameters. In the dark understory of tropical rain forests, the limiting amount of light is likely to make the interspecific difference in the effects of functional traits on demography more clear. Demographic performance is therefore tightly linked with architectural traits such as adult stature, wood density, and capacity for horizontal crown expansion. The enhancement of a demographic trade-off due to interspecific variation in functional traits in the understory helps to explain species coexistence in diverse rain forests.

Aims Arbuscular mycorrhizal fungi can have a substantial effect on the water and nutrient uptake by plants and the competition between plants in harsh environments where resource availability comes in pulses. In this study we focus on interspecific competition between Acaia etbaica and Boswellia papyrifera that have distinctive resource acquisition strategies. We compared the extent of interspecific competition with that of intraspecific competition. Methods In a greenhouse study we examined the influence of Arbuscular Mycorrhiza (AM) and pulsed water availability on competitive interactions between seedlings of the rapidly growing species A. etbaica and the slowly growing species B. papyrifera. A factorial experimental design was used. The factors were AM, two water levels and five species combinations Important Findings Seedlings of both species benefitted from AM when grown alone, and the positive growth response to pulsed water availability in B. papyrifera seedlings was in contrast with the negative growth response for A. etbaica seedlings. AM also affected the competitive performance of both species. B. papyrifera was not affected by intraspecific competition, whereas A. etbaica was negatively affected compared to the seedlings grown alone. This effect was stronger in the presence of AM. In interspecific competition, A. etbaica outcompeted B. papyrifera. Mycorrhiza and pulsed water availability did not affect the outcome of interspecific competition, and the aggressivity index of A. etbaica remained unchanged. The extent to which AM influences plant competition in a drought-stressed environment may depend on belowground functional traits of the species. AM and pulsed water availability could modify the balance between intraspecific and interspecific competition. By affecting the balance between intraspecific and interspecific competition, both factors could impact the establishment and survival of seedlings.

Plants can alter wood anatomy to adjust water supply and mechanical stability demands. However, the different ways in which species can adjust rates of water supply through variation in size and number of vessels in the sapwood, and how this variation is related to the trade-off between hydraulic and mechanical functions remain unclear. We tested the hypothesis that plants with higher investment in mechanical support have relatively less margin to change their potential hydraulic conductivity in terms of vascular strategy: the total area of conducting tissue (vessel lumen fraction) and the combination of vessel sizes and numbers (vessel composition). We measured hydraulic and mechanical traits of xylem tissue and compared the relationship between those traits between trees and lianas co-occurring in a semi-deciduous seasonally dry forest (SDF) and an evergreen rainforest (RF). Along the axis of hydraulic-mechanical trait variation, SDF lianas showed a trait combination towards investment in hydraulic conductivity (higher vessel lumen area, percentage of xylem represented by vessels and potential hydraulic conductivity), whereas trees from both forests were characterized by investment in mechanical support (higher wood density, percentage of xylem represented by fibres and number of vessel per area) and RF lianas were intermediate between these spectrum. The main difference between trees and lianas was in vessel lumen fraction and vessel composition, indicating that not only the vessel size but the distribution between the size and number of vessels is important to explain the higher hydraulic conductivity of lianas. Between forests, trees did not differ in wood density (construction costs). Similarly, lianas did not differ in wood density among forests either; therefore, confirming that differences in potential hydraulic conductivity resulted from changes in the distribution between the size and number of vessels (vascular strategy). Our results suggest that vessel lumen fraction and the vessel composition are important dimensions driving variation in construction costs across woody plants. Thus, what makes lianas hydraulically distinctive from trees is the way vessel lumen fraction and vessel composition vary across environments rather than simply having wider vessels.

Studies on climate impacts on tree annual growth are mainly restricted to marginal sites. To date, the climate effects on annual growth of trees in favorable environments remain therefore unclear despite the importance of these sites in terms of forest productivity. Because species respond differently to climate, comparing a multitude of species further enhances our knowledge on climate impacts on tree growth and forest productivity. We present a first study that reveals to what extent radial growth is limited by climate and the groundwater table across 10 temperate tree species growing on a uniform, productive soil (i.e. high nutrient and water contents) in the Netherlands. We ranked our study species according to their shade tolerance, which is associated with species’ resource requirements and growth rates, and examined their annual growth using tree-ring analysis. This allowed us to investigate how these species with diverse ecological backgrounds differ in their growth response to precipitation, temperature, irradiance, potential evapotranspiration (PET), and the groundwater table, when growing under similar and favorable site conditions. Nine out of 10 species had strikingly similar radial growth rates (on average 3.9–4.8 mm yr−1), which contradicts the widely established trade-off between shade tolerance and growth. Populus trichocarpa, the least shade-tolerant of our species, however grew much faster (on average 6.8 mm yr−1). Trees of all species reduced their growth significantly during dry summers, driven by low rainfall and high PET. Yet, the magnitude of their growth responses to climate conditions and especially the groundwater table differed across species. Receding groundwater tables significantly and strongly reduced radial growth of the shade-intolerant species, but not of the shade-tolerant species. The climate impacts on growth were not associated with variation in shade tolerance across our species. Our study demonstrates that even on a productive forest site, summer droughts reduced radial growth across a multitude of common tree species, whereas lower groundwater tables only affected the shade-intolerant species significantly. Thus, the productivity of forests in favorable environments may be seriously affected when summers in north western Europe become hotter and drier as predicted by climate models.

Carbohydrates fixed by photosynthesis are stored in plant organs in the form of starch or sugars. Starch and sugars sum to the total non-structural carbohydrate pool (TNC) and may serve as intermediate pools between assimilation and utilization. We examined the impact of tapping on TNC concentrations in stem-wood, bark and root tissues of the frankincense tree (Boswellia papyrifera (Del.) Hochst) in two natural woodlands of Ethiopia. Two tapping treatments, one without tapping (control) and the other with tapping at 12 incisions, are applied on experimental trees. Trees are tapped in the leafless dry period, diminishing their carbon storage pools. If storage pools are not refilled by assimilation during the wet season, when crowns are in full leaf, tapping may deplete the carbon pool and weaken Boswellia trees. The highest soluble sugar concentrations were in the bark and the highest starch concentrations in the stem-wood. The stem-wood contains 12 times higher starch than soluble sugar concentrations. Hence, the highest TNC concentrations occurred in the stem-wood. Moreover, wood volume was larger than root or bark volumes and, as a result, more TNC was stored in the stem-wood. As predicted, tapping reduced the TNC concentrations and pool sizes in frankincense trees during the dry season. During the wet season, these carbon pools were gradually filled in tapped trees, but never to the size of non-tapped trees. We conclude that TNC is dynamic on a seasonal time scale and offers resilience against stress, highlighting its importance for tree carbon balance. But current resin tapping practices are intensive and may weaken Boswellia populations, jeopardizing future frankincense production.