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AimGlobal forests and their structural and functional features are shaped by many mechanisms that impact tree vital rates. Although many studies have tried to quantify how specific mechanisms influence vital rates, their relative importance among forests remains unclear. We aimed to assess the patterns of variation in vital rates among species and in space and time across forests to understand and provide a baseline for expectations of the relative importance of the different mechanisms in different contexts.Location21 forest plots worldwide.Time period1981-2021Major taxa studiedWoody plantsMethodsWe developed a conceptual and statistical framework (variance partitioning of multilevel models) that attributes the variability in growth, mortality, and recruitment to variation in species, space, and time, and their interactions, which we refer to asorganising principles(OPs). We applied it to data from 21 forest plots covering more than 2.9 million trees of approximately 6,500 species.ResultsDifferences among species, thespeciesOP, were a major source of variability in tree vital rates, explaining 28-33% of demographic variance alone, and in interaction withspace14-17%, totalling 40-43%. Models with small spatial grain sizes (quadrats at 5 × 5 m) retained most of the spatial OP, but a large proportion of variance remained unexplained (31-55%). The average variability among species declined with species richness across forests, indicating that diverse forests featured smaller interspecific differences in vital rates.Main conclusionsDecomposing variance in vital rates into the proposed OPs showed that taxonomy is crucial to predictions and understanding of tree demography. Our framework has a high potential for identifying the structuring mechanisms of global forest dynamics as it highlights the most promising avenues for future research both in terms of understanding the relative contributions of mechanisms to forest demography and diversity and for improving projections of forest ecosystems.

Lianas are quintessential components of tropical forests competing strongly with trees for resources. Yet, their role in the structure and functioning of forests is rarely studied. Here, we investigate the impact of lianas on the carbon stocks and sink potential of an intact moist tropical forest in Panama using 3D terrestrial laser scanning. We find that liana-infested trees are significantly shorter with smaller crown areas, thereby resulting in a significant liana-induced reduction in stand-level carbon stocks (5.3%) and coarse woody productivity (24.5%). The widely used pantropical allometric model overestimates the carbon stocks by 10% and underestimates the liana impact on woody productivity by 1.5% at the study site with current infestation level. Increasing liana abundance across Neotropics will further worsen this impact. We show that by ignoring liana impact in carbon estimation, we are currently overestimating the potential of nature-based climate solutions to lock up atmospheric carbon.

Stable isotopologues of water are widely used to derive relative root water uptake (RWU) profiles and average RWU depth in lignified plants. Uniform isotope composition of plant xylem water (δxyl) along the stem length of woody plants is a central assumption of the isotope tracing approach which has never been properly evaluated. Here we evaluate whether strong variation in δxyl within woody plants exists using empirical field observations from French Guiana, northwestern China, and Germany. In addition, supported by a mechanistic plant hydraulic model, we test hypotheses on how variation in δxyl can develop through the effects of diurnal variation in RWU, sap flux density, diffusion, and various other soil and plant parameters on the δxyl of woody plants. The hydrogen and oxygen isotope composition of plant xylem water shows strong temporal (i.e., sub-daily) and spatial (i.e., along the stem) variation ranging up to 25.2 ‰ and 6.8 ‰ for δ2H and δ18O, respectively, greatly exceeding the measurement error range in all evaluated datasets. Model explorations predict that significant δxyl variation could arise from diurnal RWU fluctuations and vertical soil water heterogeneity. Moreover, significant differences in δxyl emerge between individuals that differ only in sap flux densities or are monitored at different times or heights. This work shows a complex pattern of δxyl transport in the soil–root–xylem system which can be related to the dynamics of RWU by plants. These dynamics complicate the assessment of RWU when using stable water isotopologues but also open new opportunities to study drought responses to environmental drivers. We propose including the monitoring of sap flow and soil matric potential for more robust estimates of average RWU depth and expansion of attainable insights in plant drought strategies and responses.

Regression dilution is a statistical inference bias that causes underestimation of the strength of dependency between two variables when the predictors are error-prone proxies (EPPs). EPPs are widely used in plant community studies focused on negative density-dependence (NDD) to quantify competitive interactions. Because of the nature of the bias, conspecific NDD is often overestimated in recruitment analyses, and in some cases, can be erroneously detected when absent. In contrast, for survival analyses, EPPs typically cause NDD to be underestimated, but underestimation is more severe for abundant species and for heterospecific effects, thereby generating spurious negative relationships between the strength of NDD and the abundances of con- and heterospecifics. This can explain why many studies observed rare species to suffer more severely from conspecific NDD, and heterospecific effects to be disproportionally smaller than conspecific effects. In general, such species-dependent bias is often related to traits associated with likely mechanisms of NDD, which creates false patterns and complicates the ecological interpretation of the analyses. Classic examples taken from literature and simulations demonstrate that this bias has been pervasive, which calls into question the emerging paradigm that intraspecific competition has been demonstrated by direct field measurements to be generally stronger than interspecific competition.

Many empirical studies motivated by an interest in stable coexistence have quantified negative density dependence, negative frequency dependence, or negative plant-soil feedback, but the links between these empirical results and ecological theory are not straightforward. Here, we relate these analyses to theoretical conditions for stabilisation and stable coexistence in classical competition models. By stabilisation, we mean an excess of intraspecific competition relative to interspecific competition that inherently slows or even prevents competitive exclusion. We show that most, though not all, tests demonstrating negative density dependence, negative frequency dependence, and negative plant-soil feedback constitute sufficient conditions for stabilisation of two-species interactions if applied to data for per capita population growth rates of pairs of species, but none are necessary or sufficient conditions for stable coexistence of two species. Potential inferences are even more limited when communities involve more than two species, and when performance is measured at a single life stage or vital rate. We then discuss two approaches that enable stronger tests for stable coexistence-invasibility experiments and model parameterisation. The model parameterisation approach can be applied to typical density-dependence, frequency-dependence, and plant-soil feedback data sets, and generally enables better links with mechanisms and greater insights, as demonstrated by recent studies.

The vertical distribution of absorbing roots is one of the most influential plant traits determining plant strategy to access below ground resources. Yet little is known of natural variability in root distribution since collecting field data is challenging and labor-intensive. Studying stable water isotope compositions in plants could offer a cost-effective and practical solution to estimate the absorbing root surfaces distribution. However, such an approach requires developing realistic inverse modeling techniques that enable robust estimation of rooting distributions and associated uncertainty from xylem water isotopic composition observations. This study introduces an inverse modeling method that supports the assessment of the root allocation parameter (β) that defines the exponential vertical decay of a plants’ absorbing root surfaces distribution with soil depth. The method requires measurements obtained from xylem and soil water isotope composition, soil water potentials, and sap flow velocities when plants’ xylem water is sampled at a certain height above the rooting point. In a simulation study, we show that the approach can provide unbiased estimates of β and its associated uncertainty due to measuring errors and unmeasured environmental factors that can impact the xylem water isotopic data. We also recommend improving the accuracy and power of β estimation, highlighting the need for considering accurate soil water potential and sap flow monitoring. Finally, we apply the inverse modeling method to xylem water isotope data of lianas and trees collected in French Guiana. Our work shows that the inverse modeling procedure provides a robust analytical and statistical framework to estimate β. The method accounts for potential bias due to extraction errors and unmeasured environmental factors, which improves the viability of using stable water isotope compositions to estimate the distribution of absorbing root surfaces complementary to the assessment of relative root water uptake profiles.

ForestGEO is a network of scientists and long-term forest dynamics plots (FDPs) spanning the Earth's major forest types. ForestGEO's mission is to advance understanding of the diversity and dynamics of forests and to strengthen global capacity for forest science research. ForestGEO is unique among forest plot networks in its large-scale plot dimensions, censusing of all stems ≥1 cm in diameter, inclusion of tropical, temperate and boreal forests, and investigation of additional biotic (e.g., arthropods) and abiotic (e.g., soils) drivers, which together provide a holistic view of forest functioning. The 71 FDPs in 27 countries include approximately 7.33 million living trees and about 12,000 species, representing 20% of the world's known tree diversity. With >1300 published papers, ForestGEO researchers have made significant contributions in two fundamental areas: species coexistence and diversity, and ecosystem functioning. Specifically, defining the major biotic and abiotic controls on the distribution and coexistence of species and functional types and on variation in species' demography has led to improved understanding of how the multiple dimensions of forest diversity are structured across space and time and how this diversity relates to the processes controlling the role of forests in the Earth system. Nevertheless, knowledge gaps remain that impede our ability to predict how forest diversity and function will respond to climate change and other stressors. Meeting these global research challenges requires major advances in standardizing taxonomy of tropical species, resolving the main drivers of forest dynamics, and integrating plot-based ground and remote sensing observations to scale up estimates of forest diversity and function, coupled with improved predictive models. However, they cannot be met without greater financial commitment to sustain the long-term research of ForestGEO and other forest plot networks, greatly expanded scientific capacity across the world's forested nations, and increased collaboration and integration among research networks and disciplines addressing forest science.

Forest albedo changes with vegetation dynamics, ecosystem demography and succession as different plant species can be characterized by contrasting leaf traits, as well as different allocation strategies. In tropical ecosystems, lianas (woody vines) strongly impact the forest biogeochemical cycles by competing with co-occurring trees for above- and below-ground resources. In addition to the particular location of their foliage (often at the top of the canopy), lianas were shown to contrast with tropical trees in terms of biochemical and structural leaf properties, as well as allocation strategies. As a consequence, liana spectral signature differs in several regions of the leaf spectrum (visible, near infrared and shortwave infrared) from tree’s one. At larger scale, the forest canopy reflectance spectrum is also affected by the relative abundance of lianas.To evaluate the impact of lianas on the radiative transfer and albedo of tropical forests, we collected all published reflectance spectra of liana and co-occurring tree leaves as well as the canopy reflectance spectra characterized by high and low liana coverage. We then calibrated both a leaf (PROSPECT-5) and a canopy (ED2-RTM) radiative transfer model on those data to reproduce the spectral signatures together with their differences, for each single study and site. The Bayesian framework that we used generated leaf biochemical and structural trait distributions, as well as allocation pattern strategies that could be compared between both growth forms.Collected spectra could be fairly well reproduced at both leaf and canopy levels by the selected mechanistic models. In most studies, calibration led to significantly lower chlorophyll and carotenoid contents, higher relative water content, and larger specific leaf areas for liana leaves, which allowed reproducing observed higher leaf reflectance values in the visible and shortwave infrared and the lower ones in the near infrared. We then validated our findings with independent field data on leaf chemistry and structure from the same studies.These calibrated radiative transfer models are tools that can be used in future research to investigate the liana impact on the global energy budget of tropical forests, as well as to monitor spatial and temporal liana abundance.

1. Stable isotopologues of water are a widely used tool to derive the depth of root water uptake (RWU) in lignified plants. Uniform isotope composition of plant xylem water (i-H2O-xyl) along the stem length is a central assumption, which has never been properly evaluated. 2. We studied the effects of diurnal variation in RWU, sap flux density and various other soil and plant parameters on i-H2O-xyl within a plant using a mechanistic plant hydraulic model and empirical field observations from French Guiana and northwestern China. 3. Our model predicts significant i-H2O-xyl variation arising from diurnal RWU fluctuations and vertical soil water heterogeneity. Moreover, significant differences in i-H2O-xyl emerge between individuals with different sap flux densities. In line with model predictions, field data show excessive i-H2O-xyl variation during the day or along stem length ranging up to 25.2 ‰ in δ2H and 6.8 ‰ in δ18O, largely exceeding the measurement error range. 4. Our work show that the fundamental assumption of uniform i-H2O-xyl is violated both theoretically and empirically and therefore a real danger exists of significant biases when using stable water isotopologues to assess RWU. We propose to include monitoring of sap flow and soil water potential for more robust RWU depth estimates.

In the past decade, plant root water uptake (RWU) has been a major focus of ecohydrological studies employing water stable isotopes. The interest of the isotopic community for RWU rose concomitantly to the development of open-access multi-source mixing models based on Bayesian inference. Another more general reason was certainly the decrease in analytical cost with the advent of isotope-specific laser absorption spectrometry. The isotopic methodology used to determine relative profiles of RWU works on the premises that (i) RWU does not fractionate stable isotopes in water and (ii) the isotopic composition of water inside the xylem vessel of the last non-evaporating part of the plant (typically the stem) is that of RWU. Following a simple mass balance approach, the isotopic composition of RWU can be linked back by inversion to contributions to RWU (i.e., relative RWU) of a set of potential water sources (of known isotopic compositions) originating from the soil profile.In recent research, the preferred tool for inverting water isotope data was Bayesian models and the literature shows that only a handful of studies complemented isotope analysis with observation of plant water status and flow. Consequently, most of the gathered information on RWU cannot be used to test hypotheses on which are built physically-based soil-root water flow models. The authors have on the other hand initiated an effort within the framework of dual experimental-modeling approaches, where tightly-controlled experiments are thought and prepared in order to validate, parameterize models, or test hypotheses. The present contribution gives an overview of the different attempts at integrating both water and isotope observations types and confronting them to model simulations explicitly accounting for root system architecture and hydraulic properties. It addresses the meaningfulness and limitations of isotope data, especially in the context of labeling experiments when treated with statistical (e.g. Bayesian) models. We finally propose a way forward and present improvements to be achieved on both experimental and modeling sides to increase the reliability and precision of isotope-derived estimates of RWU.

Lianas and other climbing plants are structural parasites of trees, generally reducing host tree survival, growth, and reproduction, yet their influences on the outcome of competition among tree species have remained largely unexplored. We propose that there are three distinct components to liana–tree interactions: prevalence, defined as the proportion of infested trees; load, defined as the mean liana cover on infested trees; and tolerance, defined as the effect of a given level of infestation on tree population growth rates. We introduce a new metric that integrates these components, the liana burden, defined as the total effect of lianas on per capita population growth rates given current prevalence, load, and tolerance. Using these metrics, we quantify variation among 33 co‐occurring tropical tree species in liana–tree interactions and its relation with shade‐tolerance. The focal tree species vary strongly in liana prevalence, load, tolerance, and burden. Interspecific variation in tolerance is the largest contributor to interspecific variation in burden. Species rankings of per capita population growth rates under current liana infestation levels differ somewhat from rankings under liana‐free conditions, and differ strongly from rankings under uniformly high liana infestation. Thus, lianas alter competitive hierarchies to benefit tree species that are relatively tolerant of and/or resistant to lianas. Among the focal tree species, shade‐tolerance is positively correlated with liana tolerance and prevalence, but largely unrelated to load and burden, meaning shade‐tolerance does not predict which species are competitively disadvantaged by lianas. We describe a variety of mechanisms by which lianas may potentially increase or decrease niche differences among tree species, including interactions with spatial and temporal environmental niche partitioning, and potential differences among tree species in relative vulnerability to different liana species. Synthesis. Lianas, like other natural enemies, can in principle alter competitive hierarchies and niche structure of co‐occurring tree species, and our analyses suggest such influences are substantial in our focal tropical tree community and likely many other tree communities as well. Quantifying these effects requires a more comprehensive approach including analyses and modelling of dynamics of liana–tree interactions and their variation with tree and liana species identities.

Life-history theory posits that trade-offs between demographic rates constrain the range of viable life-history strategies. For coexisting tropical tree species, the best established demographic trade-off is the growth-survival trade-off. However, we know surprisingly little about co-variation of growth and survival with measures of reproduction. We analysed demographic rates from seed to adult of 282 co-occurring tropical tree and shrub species, including measures of reproduction and accounting for ontogeny. Besides the well-established fast-slow continuum, we identified a second major dimension of demographic variation: a trade-off between recruitment and seedling performance vs. growth and survival of larger individuals (≥ 1 cm dbh) corresponding to a 'stature-recruitment' axis. The two demographic dimensions were almost perfectly aligned with two independent trait dimensions (shade tolerance and size). Our results complement recent analyses of plant life-history variation at the global scale and reveal that demographic trade-offs along multiple axes act to structure local communities.

Aim To evaluate how wood density relates to tree growth rates in simple models and two tropical forests. Location Barro Colorado Island, Panama; and Pasoh Forest Reserve, Malaysia. Time Period 1986–2010. Major Taxa Studied Trees. Methods We derived expected relationships of wood density with diameter growth at a given diameter under a null hypothesis that aboveground biomass growth is independent of wood density, and an alternative hypothesis that biomass growth scales with crown area, which itself increases with wood density. We tested these assumptions and predictions through analyses of interspecific relationships of wood density with height, crown area and diameter growth at constant diameter in two tropical forests. Results Height was unrelated to wood density, whereas crown areas showed a slightly positive relationship to wood density. Thus, the expected exponent of diameter growth with wood density was equal to minus one under the null hypothesis, and equal to the exponent of crown area with wood density minus one under the alternative hypothesis. Empirical relationships of diameter growth and biomass growth with wood density were broadly consistent with the null hypothesis that biomass growth is unrelated to wood density at both sites, except in trees

Contains fulltext : 184075.pdf (Publisher’s version ) (Open Access) 9 p.

The datasets were collected with funding from the National Science Foundation (DEB 0453445; 0453665; 0613666, 0845071, 1019436 & 1558093), Netherlands Organization for Scientific Research (NWO-ALW 801-01-009), the Smithsonian Tropical Research Institute, the Centre for Tropical Forest Science, the John D. and Catherine T. MacArthur Foundation, the Mellon Foundation and the Small World Institute Fund.

This study was supported by the Netherlands Organization for Scientific Research (NWO-ALW 801-01-009; M.D.V.), the Smithsonian Tropical Research Institute (M.D.V.) and the HSBC Climate Partnership (H.C.M.). The datasets were collected with funding from the National Science Foundation (DEB 0425651 & 0948585 to S.P.H.; 0453445 to H.C.M.; 0453665 to S.J.W.; 0613666, 0845071, 1019436 & 1558093 to S.A.S., and 1242622 & 1464389 to L.S.C.), the Smithsonian Tropical Research Institute, the Centre for Tropical Forest Science, the John D. and Catherine T. MacArthur Foundation, the Mellon Foundation and the Small World Institute Fund.

Dioecy has a demographic disadvantage compared with hermaphroditism: only about half of reproductive adults produce seeds. Dioecious species must therefore have fitness advantages to compensate for this cost through increased survival, growth, and/or reproduction. We used a full life cycle approach to quantify the demographic costs and benefits associated with dioecy while controlling for demographic differences between dioecious and hermaphroditic species related to other functional traits. The advantage of this novel approach is that we can focus on the effect of breeding system across a diverse tree community. We built a composite integral projection model for hermaphroditic and dioecious tree populations from Barro Colorado Island, Panama, using long-term demographic and newly collected reproductive data. Integration of all costs and benefits showed that compensation was realized through increased seed production, resulting in no net costs of dioecy. Compensation was also facilitated by the low contribution of reproduction to population growth. Estimated positive effects of dioecy on tree growth and survival were small and insignificant for population growth rates. Our model revealed that, for long-lived organisms, the cost of having males is smaller than generally expected. Hence, little compensation is required for dioecious species to maintain population growth rates similar to those of hermaphroditic species.

Summary The ‘functional traits’ of species have been heralded as promising predictors for species’ demographic rates and life history. Multiple studies have linked plant species’ demographic rates to commonly measured traits. However, predictive power is usually low – raising questions about the practical usefulness of traits – and analyses have been limited to size-independent univariate approaches restricted to a particular life stage. Here we directly evaluated the predictive power of multiple traits simultaneously across the entire life cycle of 136 tropical tree species from central Panama. Using a model-averaging approach, we related wood density, seed mass, leaf mass per area and adult stature (maximum diameter) to onset of reproduction, seed production, seedling establishment, and growth and survival at seedling, sapling and adult stages. Three of the four traits analysed here (wood density, seed mass and adult stature) typically explained 20–60% of interspecific variation at a given vital rate and life stage. There were strong shifts in the importance of different traits throughout the life cycle of trees, with seed mass and adult stature being most important early in life, and wood density becoming most important after establishment. Every trait had opposing effects on different vital rates or at different life stages; for example, seed mass was associated with higher seedling establishment and lower initial survival, wood density with higher survival and lower growth, and adult stature with decreased juvenile but increased adult growth and survival. Forest dynamics are driven by the combined effects of all demographic processes across the full life cycle. Application of a multitrait and full-life cycle approach revealed the full role of key traits, and illuminated how trait effects on demography change through the life cycle. The effects of traits on one life stage or vital rate were sometimes offset by opposing effects at another stage, revealing the danger of drawing broad conclusions about functional trait–demography relationships from analysis of a single life stage or vital rate. Robust ecological and evolutionary conclusions about the roles of functional traits rely on an understanding of the relationships of traits to vital rates across all life stages.