New tree height allometries derived from terrestrial laser scanning reveal substantial discrepancies with forest inventory methods in tropical rainforests.

Tree allometric models, essential for monitoring and predicting terrestrial carbon stocks, are traditionally built on global databases with forest inventory measurements of stem diameter (D) and tree height (H). However, these databases often combine H measurements obtained through various measurement methods, each with distinct error patterns, affecting the resulting H:D allometries. In recent decades, terrestrial laser scanning (TLS) has emerged as a widely accepted method for accurate, non-destructive tree structural measurements. This study used TLS data to evaluate the prediction accuracy of forest inventory-based H:D allometries and to develop more accurate pantropical allometries. We considered 19 tropical rainforest plots across four continents. Eleven plots had forest inventory and RIEGL VZ-400(i) TLS-based D and H data, allowing accuracy assessment of local forest inventory-based H:D allometries. Additionally, TLS-based data from 1951 trees from all 19 plots were used to create new pantropical H:D allometries for tropical rainforests. Our findings reveal that in most plots, forest inventory-based H:D allometries underestimated H compared with TLS-based allometries. For 30-metre-tall trees, these underestimations varied from -1.6 m (-5.3%) to -7.5 m (-25.4%). In the Malaysian plot with trees reaching up to 77 m in height, the underestimation was as much as -31.7 m (-41.3%). We propose a TLS-based pantropical H:D allometry, incorporating maximum climatological water deficit for site effects, with a mean uncertainty of 19.1% and a mean bias of -4.8%. While the mean uncertainty is roughly 2.3% greater than that of the Chave2014 model, this model demonstrates more consistent uncertainties across tree size and delivers less biased estimates of H (with a reduction of 8.23%). In summary, recognizing the errors in H measurements from forest inventory methods is vital, as they can propagate into the allometries they inform. This study underscores the potential of TLS for accurate H and D measurements in tropical rainforests, essential for refining tree allometries.

The role of height-driven constraints and compensations on tree vulnerability to drought.

Frequent observations of higher mortality in larger trees than in smaller ones during droughts have sparked an increasing interest in size-dependent drought-induced mortality. However, the underlying physiological mechanisms are not well understood, with height-associated hydraulic constraints often being implied as the potential mechanism driving increased drought vulnerability. We performed a quantitative synthesis on how key traits that drive plant water and carbon economy change with tree height within species and assessed the implications that the different constraints and compensations may have on the interacting mechanisms (hydraulic failure, carbon starvation and/or biotic-agent attacks) affecting tree vulnerability to drought. While xylem tension increases with tree height, taller trees present a range of structural and functional adjustments, including more efficient water use and transport and greater water uptake and storage capacity, that mitigate the path-length-associated drop in water potential. These adaptations allow taller trees to withstand episodic water stress. Conclusive evidence for height-dependent increased vulnerability to hydraulic failure and carbon starvation, and their coupling to defence mechanisms and pest and pathogen dynamics, is still lacking. Further research is needed, particularly at the intraspecific level, to ascertain the specific conditions and thresholds above which height hinders tree survival under drought.

Vapour pressure deficit modulates hydraulic function and structure of tropical rainforests under nonlimiting soil water supply.

Atmospheric conditions are expected to become warmer and drier in the future, but little is known about how evaporative demand influences forest structure and function independently from soil moisture availability, and how fast-response variables (such as canopy water potential and stomatal conductance) may mediate longer-term changes in forest structure and function in response to climate change. We used two tropical rainforest sites with different temperatures and vapour pressure deficits (VPD), but nonlimiting soil water supply, to assess the impact of evaporative demand on ecophysiological function and forest structure. Common species between sites allowed us to test the extent to which species composition, relative abundance and intraspecific variability contributed to site-level differences. The highest VPD site had lower midday canopy water potentials, canopy conductance (gc ), annual transpiration, forest stature, and biomass, while the transpiration rate was less sensitive to changes in VPD; it also had different height-diameter allometry (accounting for 51% of the difference in biomass between sites) and higher plot-level wood density. Our findings suggest that increases in VPD, even in the absence of soil water limitation, influence fast-response variables, such as canopy water potentials and gc , potentially leading to longer-term changes in forest stature resulting in reductions in biomass.

Automatic Tree Height Measurement Based on Three-Dimensional Reconstruction Using Smartphone.

Tree height is a crucial structural parameter in forest inventory as it provides a basis for evaluating stock volume and growth status. In recent years, close-range photogrammetry based on smartphone has attracted attention from researchers due to its low cost and non-destructive characteristics. However, such methods have specific requirements for camera angle and distance during shooting, and pre-shooting operations such as camera calibration and placement of calibration boards are necessary, which could be inconvenient to operate in complex natural environments. We propose a tree height measurement method based on three-dimensional (3D) reconstruction. Firstly, an absolute depth map was obtained by combining ARCore and MidasNet. Secondly, Attention-UNet was improved by adding depth maps as network input to obtain tree mask. Thirdly, the color image and depth map were fused to obtain the 3D point cloud of the scene. Then, the tree point cloud was extracted using the tree mask. Finally, the tree height was measured by extracting the axis-aligned bounding box of the tree point cloud. We built the method into an Android app, demonstrating its efficiency and automation. Our approach achieves an average relative error of 3.20% within a shooting distance range of 2-17 m, meeting the accuracy requirements of forest survey.

Divergence of hydraulic traits among tropical forest trees across topographic and vertical environment gradients in Borneo.

Fine-scale topographic-edaphic gradients are common in tropical forests and drive species spatial turnover and marked changes in forest structure and function. We evaluate how hydraulic traits of tropical tree species relate to vertical and horizontal spatial niche specialization along such a gradient. Along a topographic-edaphic gradient with uniform climate in Borneo, we measured six key hydraulic traits in 156 individuals of differing heights in 13 species of Dipterocarpaceae. We investigated how hydraulic traits relate to habitat, tree height and their interaction on this gradient. Embolism resistance increased in trees on sandy soils but did not vary with tree height. By contrast, water transport capacity increased on sandier soils and with increasing tree height. Habitat and height only interact for hydraulic efficiency, with slope for height changing from positive to negative from the clay-rich to the sandier soil. Habitat type influenced trait-trait relationships for all traits except wood density. Our data reveal that variation in the hydraulic traits of dipterocarps is driven by a combination of topographic-edaphic conditions, tree height and taxonomic identity. Our work indicates that hydraulic traits play a significant role in shaping forest structure across topographic-edaphic and vertical gradients and may contribute to niche specialization among dipterocarp species.

Tallo: A global tree allometry and crown architecture database.

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.

Fisheye Image Detection of Trees Using Improved YOLOX for Tree Height Estimation.

Tree height is an essential indicator in forestry research. This indicator is difficult to measure directly, as well as wind disturbance adds to the measurement difficulty. Therefore, tree height measurement has always been an issue that experts and scholars strive to improve. We propose a tree height measurement method based on tree fisheye images to improve the accuracy of tree height measurements. Our aim is to extract tree height extreme points in fisheye images by proposing an improved lightweight target detection network YOLOX-tiny. We added CBAM attention mechanism, transfer learning, and data enhancement methods to improve the recall rate, F1 score, AP, and other indicators of YOLOX-tiny. This study improves the detection performance of YOLOX-tiny. The use of deep learning can improve measurement efficiency while ensuring measurement accuracy and stability. The results showed that the highest relative error of tree measurements was 4.06% and the average relative error was 1.62%. The analysis showed that the method performed better at all stages than in previous studies.

Optimization and Evaluation of Sensor Angles for Precise Assessment of Architectural Traits in Peach Trees.

Fruit industries play a significant role in many aspects of global food security. They provide recognized vitamins, antioxidants, and other nutritional supplements packed in fresh fruits and other processed commodities such as juices, jams, pies, and other products. However, many fruit crops including peaches (Prunus persica (L.) Batsch) are perennial trees requiring dedicated orchard management. The architectural and morphological traits of peach trees, notably tree height, canopy area, and canopy crown volume, help to determine yield potential and precise orchard management. Thus, the use of unmanned aerial vehicles (UAVs) coupled with RGB sensors can play an important role in the high-throughput acquisition of data for evaluating architectural traits. One of the main factors that define data quality are sensor imaging angles, which are important for extracting architectural characteristics from the trees. In this study, the goal was to optimize the sensor imaging angles to extract the precise architectural trait information by evaluating the integration of nadir and oblique images. A UAV integrated with an RGB imaging sensor at three different angles (90°, 65°, and 45°) and a 3D light detection and ranging (LiDAR) system was used to acquire images of peach trees located at the Washington State University's Tukey Horticultural Orchard, Pullman, WA, USA. A total of four approaches, comprising the use of 2D data (from UAV) and 3D point cloud (from UAV and LiDAR), were utilized to segment and measure the individual tree height and canopy crown volume. Overall, the features extracted from the images acquired at 45° and integrated nadir and oblique images showed a strong correlation with the ground reference tree height data, while the latter was highly correlated with canopy crown volume. Thus, selection of the sensor angle during UAV flight is critical for improving the accuracy of extracting architectural traits and may be useful for further precision orchard management.

Climate and functional traits jointly mediate tree water-use strategies.

Tree water use is central to plant function and ecosystem fluxes. However, it is still unknown how organ-level water-relations traits are coordinated to determine whole-tree water-use strategies in response to drought, and whether this coordination depends on climate. Here we used a global sap flow database (SAPFLUXNET) to study the response of water use, in terms of whole-tree canopy conductance (G), to vapour pressure deficit (VPD) and to soil water content (SWC) for 142 tree species. We investigated the individual and coordinated effect of six water-relations traits (vulnerability to embolism, Huber value, hydraulic conductivity, turgor-loss point, rooting depth and leaf size) on water-use parameters, also accounting for the effect of tree height and climate (mean annual precipitation, MAP). Reference G and its sensitivity to VPD were tightly coordinated with water-relations traits rather than with MAP. Species with efficient xylem transport had higher canopy conductance but also higher sensitivity to VPD. Moreover, we found that angiosperms had higher reference G and higher sensitivity to VPD than did gymnosperms. Our results highlight the need to consider trait integration and reveal the complications and challenges of defining a single, whole-plant resource use spectrum ranging from 'acquisitive' to 'conservative'.

Tree height and leaf drought tolerance traits shape growth responses across droughts in a temperate broadleaf forest.

As climate change drives increased drought in many forested regions, mechanistic understanding of the factors conferring drought tolerance in trees is increasingly important. The dendrochronological record provides a window through which we can understand how tree size and traits shape growth responses to droughts. We analyzed tree-ring records for 12 species in a broadleaf deciduous forest in Virginia (USA) to test hypotheses for how tree height, microenvironment characteristics, and species' traits shaped drought responses across the three strongest regional droughts over a 60-yr period. Drought tolerance (resistance, recovery, and resilience) decreased with tree height, which was strongly correlated with exposure to higher solar radiation and evaporative demand. The potentially greater rooting volume of larger trees did not confer a resistance advantage, but marginally increased recovery and resilience, in sites with low topographic wetness index. Drought tolerance was greater among species whose leaves lost turgor (wilted) at more negative water potentials and experienced less shrinkage upon desiccation. The tree-ring record reveals that tree height and leaf drought tolerance traits influenced growth responses during and after significant droughts in the meteorological record. As climate change-induced droughts intensify, tall trees with drought-sensitive leaves will be most vulnerable to immediate and longer-term growth reductions.

Increased hydraulic constraints in Eucalyptus plantations fertilized with potassium.

Fertilization is commonly used to increase growth in forest plantations, but it may also affect tree water relations and responses to drought. Here, we measured changes in biomass, transpiration, sapwood-to-leaf area ratio (As :Al ) and sap flow driving force (ΔΨ) during the 6-year rotation of tropical plantations of Eucalyptus grandis under controlled conditions for throughfall and potassium (K) fertilization. K fertilization increased final tree height by 8 m. Throughfall exclusion scarcely affected tree functioning because of deep soil water uptake. Tree growth increased in K-supplied plots and remained stable in K-depleted plots as tree height increased, while growth per unit leaf area increased in all plots. Stand transpiration and hydraulic conductance standardized per leaf area increased with height in K-depleted plots, but remained stable or decreased in K-supplied plots. Greater Al in K-supplied plots increased the hydraulic constraints on water use. This involved a direct mechanism through halved As :Al in K-supplied plots relative to K-depleted plots, and an indirect mechanism through deteriorated water status in K-supplied plots, which prevented the increase in ΔΨ with tree height. K fertilization in tropical plantations reduces the hydraulic compensation to growth, which could increase the risk of drought-induced dieback under climate change.

Range margin populations show high climate adaptation lags in European trees.

How populations of long-living species respond to climate change depends on phenotypic plasticity and local adaptation processes. Marginal populations are expected to have lags in adaptation (i.e. differences between the climatic optimum that maximizes population fitness and the local climate) because they receive pre-adapted alleles from core populations preventing them from reaching a local optimum in their climatically marginal habitat. Yet, whether adaptation lags in marginal populations are a common feature across phylogenetically and ecologically different species and how lags can change with climate change remain unexplored. To test for range-wide patterns of phenotypic variation and adaptation lags of populations to climate, we (a) built model ensembles of tree height accounting for the climate of population origin and the climate of the site for 706 populations monitored in 97 common garden experiments covering the range of six European forest tree species; (b) estimated populations' adaptation lags as the differences between the climatic optimum that maximizes tree height and the climate of the origin of each population; (c) identified adaptation lag patterns for populations coming from the warm/dry and cold/wet margins and from the distribution core of each species range. We found that (a) phenotypic variation is driven by either temperature or precipitation; (b) adaptation lags are consistently higher in climatic margin populations (cold/warm, dry/wet) than in core populations; (c) predictions for future warmer climates suggest adaptation lags would decrease in cold margin populations, slightly increasing tree height, while adaptation lags would increase in core and warm margin populations, sharply decreasing tree height. Our results suggest that warm margin populations are the most vulnerable to climate change, but understanding how these populations can cope with future climates depend on whether other fitness-related traits could show similar adaptation lag patterns.

Biomass increases attributed to both faster tree growth and altered allometric relationships under long-term carbon dioxide enrichment at a temperate forest.

Increases in atmospheric carbon dioxide (CO2 ) concentrations are expected to lead to increases in the rate of tree biomass accumulation, at least temporarily. On the one hand, trees may simply grow faster under higher CO2 concentrations, preserving the allometric relations that prevailed under lower CO2 concentrations. Alternatively, the allometric relations themselves may change. In this study, the effects of elevated CO2 (eCO2 ) on tree biomass and allometric relations were jointly assessed. Over 100 trees, grown at Duke Forest, NC, USA, were harvested from eight plots. Half of the plots had been subjected to CO2 enrichment from 1996 to 2010. Several subplots had also been subjected to nitrogen fertilization from 2005 to 2010. Allometric equations were developed to predict tree height, stem volume, and aboveground biomass components for loblolly pine (Pinus taeda L.), the dominant tree species, and broad-leaved species. Using the same diameter-based allometric equations for biomass, it was estimated that plots with eCO2 contained 21% more aboveground biomass, consistent with previous studies. However, eCO2 significantly affected allometry, and these changes had an additional effect on biomass. In particular, P. taeda trees at a given diameter were observed to be taller under eCO2 than under ambient CO2 due to changes in both the allometric scaling exponent and intercept. Accounting for allometric change increased the treatment effect of eCO2 on aboveground biomass from a 21% to a 27% increase. No allometric changes for the nondominant broad-leaved species were identified, nor were allometric changes associated with nitrogen fertilization. For P. taeda, it is concluded that eCO2 affects allometries, and that knowledge of allometry changes is necessary to accurately compute biomass under eCO2 . Further observations are needed to determine whether this assessment holds for other taxa.

Tropical tree size-frequency distributions from airborne lidar.

In tropical rainforests, tree size and number density are influenced by disturbance history, soil, topography, climate, and biological factors that are difficult to predict without detailed and widespread forest inventory data. Here, we quantify tree size-frequency distributions over an old-growth wet tropical forest at the La Selva Biological Station in Costa Rica by using an individual tree crown (ITC) algorithm on airborne lidar measurements. The ITC provided tree height, crown area, the number of trees >10 m height and, predicted tree diameter, and aboveground biomass from field allometry. The number density showed strong agreement with field observations at the plot- (97.4%; 3% bias) and tree-height-classes level (97.4%; 3% bias). The lidar trees size spectra of tree diameter and height closely follow the distributions measured on the ground but showed less agreement with crown area observations. The model to convert lidar-derived tree height and crown area to tree diameter produced unbiased (0.8%) estimates of plot-level basal area and with low uncertainty (6%). Predictions on basal area for tree height classes were also unbiased (1.3%) but with larger uncertainties (22%). The biomass estimates had no significant bias at the plot- and tree-height-classes level (-5.2% and 2.1%). Our ITC method provides a powerful tool for tree- to landscape-level tropical forest inventory and biomass estimation by overcoming the limitations of lidar area-based approaches that require local calibration using a large number of inventory plots.

Forest age mapping based on multiple-resource remote sensing data.

Forest age is an important stand description factor and plays an important role in the carbon cycle of forest ecosystems. However, forest age data are typically lacking or are difficult to acquire at large spatial scale. Thus, it is important to develop a method of spatial forest age mapping. In this study, a method of forest age estimation based on multiple-resource remote sensing data was developed. Forest age was estimated by using average tree height estimated from the ICESat/GLAS and MODIS BRDF products. The results showed that forest age was significantly related to average tree height with a correlation coefficient of 0.752. Then, the average tree height was inversed using a waveform parameter extracted from ICESat/GLAS and was extended to continuous space with the help of the MODIS BRDF product. Thus, forest age mapping was realized. Lastly, the structure of forest age in the study area was evaluated. The results indicated that this method can be used to estimate forest age at the local scale and at large scale and can facilitate understandings of the real forest age structure features of a research area.

Within-crown plasticity in leaf traits among the tallest conifers.

PREMISE OF THE STUDY: Leaves are the sites of greatest water stress in trees and a key means of acclimation to the environment. We considered phenotypic plasticity of Pseudotsuga menziesii leaves in their ecological context, exploring responsiveness to natural gradients in water stress (indicated by sample height) and light availability (measured from hemispherical photos) to understand how leaf structure is controlled by abiotic factors in tall tree crowns. METHODS: After measuring anatomy, morphology, and carbon isotope composition (δ13 C) of leaves throughout crowns of P. menziesii >90 m tall, we compared structural plasticity of leaves among the three tallest conifer species using equivalent data from past work on Sequoia sempervirens and Picea sitchensis. KEY RESULTS: Leaf mass per projected area (LMA) and δ13 C increased and mesoporosity (airspace/area) decreased along the water-stress gradient, while light did not play a detectable role in leaf development. Overall, leaves of P. menziesii were far less phenotypically responsive to within-crown abiotic gradients than either P. sitchensis, whose leaves responded strongly to light availability, or S. sempervirens, whose leaves responded equally strongly to water stress. CONCLUSIONS: P. menziesii maintain remarkably consistent leaf structure despite pronounced vertical gradients in abiotic factors. Contrasting patterns of leaf structural plasticity underlie divergent ecological strategies of the three tallest conifer species, which coexist in Californian rainforests.

Factors influencing the accuracy of ground-based tree-height measurements for major European tree species.

Tree height is one of the most important forest characteristics and is one of the crucial measurements taken for either practical or scientific reasons. However, the accuracy of a tree-height measurement may vary in relation to many factors. The work described here thus sought to evaluate the accuracy of ground-based tree-height measurements for major forest-forming tree species of the temperate and boreal zones. The focus was on the importance of factors affecting accuracy of the measurements in question at larger geographical scales. In line with the above research goals, data were gathered from 299 stands throughout Poland and heights of 2388 sample trees of eight species, growing in different stands and site conditions, were measured; heights were then compared with measured lengths of felled trees as a reference. In total, 10 variables to determine factors that may influence ground-based tree-height measurement accuracy were used. We merged them into 4 groups: measurements, topography, stand and biometric-related factors. Results showed that biometric and topographic factors had the greatest relative influence on the accuracy of measurements of tree height. Tree length and species, followed by the slope of the terrain, tree age, and height above sea level were the most important factors found to affect accuracy. In most of the cases studied the terrestrial tree-height measurements were underestimated when set against definitive measurements of length. This was true for all species studied except oak, for which height measurements were typically overestimated. Notwithstanding the broad geographical scope of the work, the particular device used and the team factor were only found to have a marginal influence on measurement accuracy.

Tree distribution, morphology and modelled air pollution in urban parks of Hong Kong.

Trees offer a range of ecosystem services and remain important in providing human benefits. However, emerging literature questions the long-accepted view of trees being able to improve air quality in urban parks. The aerodynamic effect of trees was identified as a major reason for the change of pollutant distribution in near-road parks, where trees can act as porous barriers and cause localised concentration increase. Although not yet fully developed, planting strategies aiming to mitigate the negative effect of vegetation on air quality should be encouraged in future park design. In this study, we explored the effect of tree planting design on pollutant diffusion by integrating field surveys in urban parks in Hong Kong with computational fluid dynamic (CFD) modelling. A series of indicators associated with tree morphology and landscape were derived from the surveys and their influence on air pollutant distribution in parks was examined using ENVI-MET. Dense trees with low crown base were found effective in improving air quality within parks when planted as barriers with a width of ~15 m at borders. However, more extensive planting led to a decrease in wind velocity and an increase in pollutant concentrations, which should be avoided. Tall trees tended to have little influence on airflow at the pedestrian level, which means they seem appropriate for small urban parks where wide barriers are not applicable and rapid ventilation should be encouraged. The tree distribution also altered the airflow and pollutant dispersion in parks. Our study provides clues for thoughtful planting strategies which can optimise air quality in urban parks.

Water transport through tall trees: A vertically explicit, analytical model of xylem hydraulic conductance in stems.

Trees grow by vertically extending their stems, so accurate stem hydraulic models are fundamental to understanding the hydraulic challenges faced by tall trees. Using a literature survey, we showed that many tree species exhibit continuous vertical variation in hydraulic traits. To examine the effects of this variation on hydraulic function, we developed a spatially explicit, analytical water transport model for stems. Our model allows Huber ratio, stem-saturated conductivity, pressure at 50% loss of conductivity, leaf area, and transpiration rate to vary continuously along the hydraulic path. Predictions from our model differ from a matric flux potential model parameterized with uniform traits. Analyses show that cavitation is a whole-stem emergent property resulting from non-linear pressure-conductivity feedbacks that, with gravity, cause impaired water transport to accumulate along the path. Because of the compounding effects of vertical trait variation on hydraulic function, growing proportionally more sapwood and building tapered xylem with height, as well as reducing xylem vulnerability only at branch tips while maintaining transport capacity at the stem base, can compensate for these effects. We therefore conclude that the adaptive significance of vertical variation in stem hydraulic traits is to allow trees to grow tall and tolerate operating near their hydraulic limits.

Traits and trade-offs in whole-tree hydraulic architecture along the vertical axis of Eucalyptus grandis.

Background and Aims: Sapwood traits like vessel diameter and intervessel pit characteristics play key roles in maintaining hydraulic integrity of trees. Surprisingly little is known about how sapwood traits covary with tree height and how such trait-based variation could affect the efficiency of water transport in tall trees. This study presents a detailed analysis of structural and functional traits along the vertical axes of tall Eucalyptus grandis trees. Methods: To assess a wide range of anatomical and physiological traits, light and electron microscopy was used, as well as field measurements of tree architecture, water use, stem water potential and leaf area distribution. Key Results: Strong apical dominance of water transport resulted in increased volumetric water supply per unit leaf area with tree height. This was realized by continued narrowing (from 250 to 20 µm) and an exponential increase in frequency (from 600 to 13 000 cm-2) of vessels towards the apex. The widest vessels were detected at least 4 m above the stem base, where they were associated with the thickest intervessel pit membranes. In addition, this study established the lower limit of pit membrane thickness in tall E. grandis at ~375 nm. This minimum thickness was maintained over a large distance in the upper stem, where vessel diameters continued to narrow. Conclusions: The analyses of xylem ultrastructure revealed complex, synchronized trait covariation and trade-offs with increasing height in E. grandis. Anatomical traits related to xylem vessels and those related to architecture of pit membranes were found to increase efficiency and apical dominance of water transport. This study underlines the importance of studying tree hydraulic functioning at organismal scale. Results presented here will improve understanding height-dependent structure-function patterns in tall trees.