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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.

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.

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.

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.

1. Plant organ biomass partitioning has been hypothesized to be driven by resources, such that species from drier environments allocate more biomass to roots than species from wetter environments to access water at greater soil depths. In savanna systems, fire may select for greater allocation to root biomass, especially in humid environments where fire is more frequent. Therefore, species from drier environments may have been under selection pressure to reach deeper soil water more effectively than species from humid environments, through faster root extension, more efficient depth penetration, and faster plant growth rates to respond rapidly to variable rainfall events. 2. We compared biomass partitioning, root morphology traits [root extension rate, RER; specific taproot length (STRL)] and relative growth rate (RGR) of seedlings of 51 savanna tree species, sampled from three continents (Africa, Australia and South America) in a greenhouse experiment. We used phylogenetically corrected and uncorrected analyses to compare the traits of the groups. We conducted a permanova on the combined traits to establish whether species could be distinguished on the basis of their combined traits. 3. On average, species from humid environments allocated more biomass to roots and less to stems than species from semi-arid environments, consistent with the expectation that fire pressure selects for greater allocation to roots in humid environments. However, some species from humid environments had fast growth rates instead of high allocation to roots. Both RER and STRL were greater among species of semi-arid environments than among species of humid environments, and also differed between continents. Differences between strategies under each climate type appear to be associated with leaf habit. 4. Synthesis. Plant biomass partitioning has been selected by defoliation pressure and the effects of this selection pressure can supersede any selection in response to local water constraints. Root morphological adaptations, but not plant growth rate, of tree seedlings, have been selected in response to water deficits

Summary 1. Plants exhibit a wide variety in traits at different organizational levels. Intraspecific and interspecific studies have potential to demonstrate functional relationships and trade-offs amongst traits, with potential consequences for growth. However, the distinction between the correlative and functional nature of trait covariation presents a challenge because traits interact in complex ways. 2. We present an intraspecific study on Scots pine branches and use functional multi-trait concepts to organize and understand trait interactions and their impacts on growth. Branch-level traits were assessed for 97 branches from 12 Scots pine sites across Europe. 3. To test alternative hypotheses on cause–effect relationships between anatomical traits, hydraulic traits and branch growth, we measured for each branch: the tracheid hydraulic diameter, double cell wall thickness, cell lumen span area, wood density, cavitation vulnerability, wood-specific hydraulic conductivity, the leaf area to sapwood area ratio and branch growth. We used mixed linear effect models and path models to show how anatomical traits determine hydraulic traits and, in turn, how those traits influence growth. 4. Tracheid hydraulic diameter was the best predictor of cavitation vulnerability (R 2 =0 AE09 explained by path model) and specific conductivity (R 2 =0 AE19) amongst anatomical traits. Leaf area to sapwood area ratio had the strongest direct effect on branch growth (R 2 =0 AE19) and was positively associated with the tracheid hydraulic diameter (R 2 =0 AE22). A number of bivariate correlations between traits could be explained by these functional relationships amongst traits. 5. The plasticity in tracheid hydraulic diameter (10.0–15.1 lm) and leaf area to sapwood area ratio (600–6051 cm 2 cm )2 ) and the maintenance of a minimum leaf water potential (between )2 and )2AE5 MPa) appear to drive the anatomical and hydraulic traits of Scots pine across Europe. These properties are major drivers of the functional trait network underlying the growth variation amongst pine branches and thus possibly contribute to the ecological success of pines at a local and continental scale.

Summary 1. Resins are highly valued non-timber forest products (NTFP). One of the most widely traded resins is frankincense, tapped from several Boswellia tree species (Burseraceae). Exploited Boswellia populations often show poor regeneration, but the demographic consequences of these bottlenecks are unknown. Here we report on the first large-scale demographic study of frankincense-producing trees. 2. We studied 12 populations of Boswellia papyrifera in northern Ethiopia, varying in altitude and productivity. Six of these populations had been tapped before and were tapped during the study. Survival, growth and fecundity were determined for 4370 trees and 2228 seedlings, in 22.8 ha over a 2-year period. We also studied a remote population where no grazing and tapping took place. Matrix models were used to project population growth and frankincense production under four restoration scenarios. 3. Population structures of both tapped and untapped populations showed clear gaps. Small seedlings were abundant in all populations, but none developed into persistent saplings. Such saplings were only present in the remote population. Fire and grazing are the likely causes of this regeneration bottleneck. 4. Adult mortality was high (6–7% per year) in both tapped and untapped populations, probably caused by beetle attacks and fire. Unexpectedly, tapped populations presented higher diameter growth rates and fecundity compared to untapped populations. These differences are probably caused by non-random selection of exploited populations by tappers. 5. Under the ‘business as usual’ scenario, population models projected a 90% decline in the size of tapped and untapped populations within 50 years and a 50% decline in frankincense yield within 15 years. Model simulations for restoration scenarios revealed that populations and frankincense production could only be sustained with intensive management leading to full sapling recruitment and a 50–75% reduction in adult mortality. 6. Synthesis and applications. Regeneration bottlenecks and high adult mortality are causing rapid decline in frankincense-producing tree populations in Ethiopia. This decline is unlikely to be a consequence of harvesting and is probably driven by fire, grazing and beetle attacks. Fire prevention and the establishment of non-grazing areas are needed. Our results show that other factors than exploitation may seriously threaten populations yielding NTFP.

1. Because of its mechanical properties, wood density may affect the way that trees expand their stem and crown to exploit favourable light conditions in a mechanically stable way. From engineering theory and wood density properties, it is predicted that in terms of biomass investment, low-density wood is more efficient for vertical stem expansion, while high-density wood is more efficient for horizontal branch expansion. So far, these predictions have rarely been tested by empirical studies. 2. We tested these predictions for 145 co-occurring tree species in a Malaysian tropical rainforest. For each species, we selected trees across a broad size range and measured architectural dimensions (stem diameter, height of the lowest foliage and crown width). We used a hierarchical Bayesian model to estimate species-specific allometric relationships between architectural dimensions including estimated stem biomass. Then, we examined correlations between species wood density and estimated architectural variables at standardized heights. 3. When species were compared at standardized tree heights, wood density correlated negatively with stem diameter and positively with stem biomass at most reference heights. This indicates that species with low wood density produce thicker stems but at lower biomass costs. Wood density correlated positively with crown width and negatively with height of the lowest foliage, which indicates that high wood density species have wider and deeper crowns than low wood density species. These relationships were maintained at most reference heights. However, the relationship with crown width was nonsignificant above 18 m height. This may reflect large plastic response of lateral crown expansion to a local condition. 4. Wood density explains the trade-off between effective vertical stem expansion and horizontal crown expansion across co-occurring tropical tree species. Such mechanical constraints characterize the difference in tree architecture between low wood density species that show an efficient height expansion to attain better light conditions in the exposed canopy and high wood density species that show an efficient horizontal crown expansion to enhance current light interception and persistence in the shaded forest understorey. Our study thus suggests that the mechanical constraints set by wood density contribute to the co-existence of species differing in architecture and light capture strategy.

1. Tree architecture is thought to allow species to partition horizontal and vertical light gradients in the forest canopy. Tree architecture is closely related to light capture, carbon gain and the efficiency with which trees reach the canopy. Previous studies that investigated how light gradients drive differentiation in tree architecture have produced inconsistent results, partially because of the differences in which tree species and ontogenetic stages were studied. 2. We examined the relationship between stem diameter, tree height, foliage height, crown width and life-history strategy over a broad size range of 200 randomly selected, co-occurring tree species in a lowland rainforest in Peninsular Malaysia. We developed a hierarchical Bayesian model to account for both intra- and interspecific variation and describe the relationships among tree architectural variables. We analysed interspecific variation in tree architectural variables in relation to adult stature and light requirement for species regeneration as a function of tree size. 3. There was little interspecific variation in architectural variables, this is partly because of large intraspecific variation in response to canopy heterogeneity, but it also suggests architectural convergence within this community. However, interspecific analyses showed that, for large-statured species, small size classes had thinner stems with narrow and shallow crowns, whereas large-size classes had wider crowns. Light-demanding species (as indicated by high sapling mortality in shaded conditions) showed weak trends in tree architecture and were only characterized by wide crowns at intermediate sizes. 4. In summary, tree architectural traits overlapped across the species community. This suggests that architectural convergence and equalizing effects occur in this diverse tropical forest and that community-wide allometric equations can be used to describe forest height and carbon storage. Light resource partitioning also occurs, indicating stabilizing effects. Interspecific architectural variation in relation to adult stature supports the theory of the trade-off between early reproduction and vegetative growth. In closed rainforests, adult stature imposes a stronger force on architectural differentiation of species than regeneration light requirements.

Question: Thousands of small isolated forest fragments remain around churches (“church forests”) in the almost completely deforested Ethiopian Highlands. We questioned how the forest structure and composition varied with altitude, forest area and human influence. Location: South Gondar, Amhara National Regional State, Northern Ethiopia. Methods: The structure and species composition was assessed for 810 plots in 28 church forests. All woody plants were inventoried, identified and measured (stem diameter) in seven to 56 10 m x 10-m plots per forest. Results: In total, 168 woody species were recorded, of which 160 were indigeneous. The basal area decreased with tree harvest intensity; understorey and middle-storey density (5 cm DBH trees) increased with altitude. The dominance of a small set of species increased with altitude and grazing intensity. Species richness decreased with altitude, mainly due to variation in the richness of the overstorey community. Moreover, species richness in the understorey decreased with grazing intensity. Conclusions: We show how tree harvesting intensity, grazing intensity and altitude contribute to observed variations in forest structure, composition and species richness. Species richness was, however, not related to forest area. Our study emphasizes the significant role played by the remaining church forests for conservation of woody plant species in North Ethiopian Highlands, and the need to protect these forests for plant species conservation purposes

The strength of competitive and facilitative interactions in plant communities is expected to change along resource gradients. Contrasting theoretical models predict that with increasing abiotic stress, facilitative effects are higher, lower, or similar than those found under more productive conditions. While these predictions have been tested in stressful environments such as arid and alpine ecosystems, they have hardly been tested for more productive African woodlands. We experimentally assessed the strength of tree seedling facilitation by nurse trees in mesic and dry woodlands in Benin, West Africa. We planted seedlings of the drought-sensitive Afzelia africana and the drought-tolerant Khaya senegalensis under three microsite conditions (closed woodland, woodland gap, and open fields). Seedling survival was greater within woodlands compared with open fields in both the mesic and dry woodlands. The relative benefits in seedling survival were larger at the dry site, especially for the drought-sensitive species. Nevertheless, plant interactions became neutral or negative during the dry season in the drier woodland, indicating that the net positive effects may be lost under very stressful abiotic conditions. We conclude that facilitation also occurs in the relatively more productive conditions of African woodlands. Our results underscore the role of environmental variation in space and time, and the stress tolerance of species, in explaining competitive and facilitative interactions within plant communities

Basic knowledge of the relationships between tree growth and environmental variables is crucial for understanding forest dynamics and predicting vegetation responses to climate variations. Trees growing in tropical areas with a clear seasonality in rainfall often form annual growth rings. In the understory, however, tree growth is supposed to be mainly affected by interference for access to light and other resources. In the semi-deciduous Mayombe forest of the Democratic Republic of Congo, the evergreen species Aidia ochroleuca, Corynanthe paniculata and Xylopia wilwerthii dominate the understory. We studied their wood to determine whether they form annual growth rings in response to changing climate conditions. Distinct growth rings were proved to be annual and triggered by a common external factor for the three species. Species-specific site chronologies were thus constructed from the cross-dated individual growth-ring series. Correlation analysis with climatic variables revealed that annual radial stem growth is positively related to precipitation during the rainy season but at different months. The growth was found to associate with precipitation during the early rainy season for Aidia but at the end of the rainy season for Corynanthe and Xylopia. Our results suggest that a dendrochronological approach allows the understanding of climate–growth relationships in tropical forests, not only for canopy trees but also for evergreen understory species and thus arguably for the whole tree community. Global climate change influences climatic seasonality in tropical forest areas, which is likely to result in differential responses across species with a possible effect on forest composition over time

Summary •In a comparative study of 42 rainforest tree species we examined relationships amongst wood traits, diameter growth and survival of large trees in the field, and shade tolerance and adult stature of the species. •The species show two orthogonal axes of trait variation: a primary axis related to the vessel size–number trade-off (reflecting investment in hydraulic conductance vs hydraulic safety) and a secondary axis related to investment in parenchyma vs fibres (storage vs strength). Across species, growth rate was positively related to vessel diameter and potential specific hydraulic conductance (Kp), and negatively related to wood density. Survival rate was only positively related to wood density. •Light-demanding species were characterized by low wood and vessel density and wide vessels. Tall species were characterized by wide vessels with low density and large Kp. Hydraulic traits were more closely associated with adult stature than with light demand, possibly because tall canopy species experience more drought stress and face a higher cavitation risk. •Vessel traits affect growth and wood density affects growth and survival of large trees in the field. Vessel traits and wood density are therefore important components of the performance and life history strategies of tropical tree species.

Almost all dry Afromontane forests of Northern Ethiopia have been converted to agricultural, grazing or scrub lands except for small fragments left around churches ('Church forests'). Species regeneration in these forests is limited. We investigated (i) how intense postdispersal seed predation was in church forest, and if this seed predation varied with species and/or habitat, and (ii) for how long tree seeds maintained their viability while buried in forest soil. In the seed predation experiment, we monitored seeds of six tree species in four habitats for a period of 14 weeks (the peak seeding season). In the seed viability experiment, we assessed seed viability of five species in four habitats after being buried 6, 12, or 18 months. Ninety-two percent of the tree seeds were predated within 3.5 months. Predation was mainly dependent on species whereas habitat had a weaker effect. Seed viability decreased sharply with burial time in soil for all species except for Juniperus. To minimize seed availability limitation for regeneration of such species in the forest, the standing vegetation needs to be persistently managed and conserved for a continuous seed rain supply. Additional seed sowing, and seed and seedling protection (by e.g. animal exclosures) may increase successful regeneration of important species in these forests

Big leaf mahogany Swietenia macrophylla is the most valuable timber species in the tropics but its future as a commercial timber species is at risk. This study evaluates whether recovery of overexploited mahogany populations is enhanced by actively managing the species and its surrounding forest. We assessed the effect of four different management interventions that varied in their intensities of harvesting and silvicultural treatments. We tested the hypothesis that intensive forest management stimulates population growth rates. Data were gathered over a 4-year period in the plots (326 ha) of the Long Term Silvicultural Research Program in Bolivia. Plants > 1·3 m tall were identified and monitored in the plots, while seedlings and saplings ( 70 cm diameter at 1·3 m height) that produce large numbers of seedlings. Harvesting simulations indicate that mahogany populations can only be sustainably harvested by increasing the cutting cycle length, reducing harvesting intensity and by maintaining optimal growing conditions. Synthesis and applications. Mahogany is the most valuable timber species in the tropics, and its range has dramatically decreased mostly due to commercial harvesting. The results of simulation modelling based on field and experimental data suggest that overexploited populations are recovering and that sustainable harvesting will be possible in the future when cutting cycle length is increased, harvesting intensity is reduced and silvicultural treatments are applied regularly throughout the cutting cycle

Size, allometry, and mechanical design were measured for trees of three canopy species in a tropical rain forest in French Guiana. Mechanical design was expressed as the safety factor, using the elastic-stability model, and the wind resistance factor, using the constant-stress model. Changes with ontogeny were described as regressions using stem diameter as the independent variable, and they were compared between species. Height, crown size, and the wind resistance factor increased with ontogeny. The safety factor decreased to a minimum and then increased continuously in thicker trees. The crown width/height ratio did not change with ontogeny. Interspecific differences in allometry and mechanical design were related to the adult stature of the species, and not to shade tolerance. The short stature species (Vouacapoua americana) was less slender (height:DBH [stem diameter at 1.3 m] ratio) and had a higher crown width/height ratio than the tall stature species (Goupia glabra and Dicorynia guianensis). Vouacapoua had a higher safety factor, but a similar wind resistance factor. The safety factors of our study species were lower than those of two temperate tree species because of a higher slenderness. Differences in safety factors between tropical and temperate trees may result from unrealistic assumptions of the elastic-stability model, and may also be related to lower light levels and-or wind rates in the tropics.