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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.

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.

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.

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.

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.

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.

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.

Contents 1159 I. 1159 II. 1161 III. 1164 IV. 1166 1167 References 1167 SUMMARY: The search for a root economics spectrum (RES) has been sparked by recent interest in trait-based plant ecology. By analogy with the one-dimensional leaf economics spectrum (LES), fine-root traits are hypothesised to match leaf traits which are coordinated along one axis from resource acquisitive to conservative traits. However, our literature review and meta-level analysis reveal no consistent evidence of an RES mirroring an LES. Instead the RES appears to be multidimensional. We discuss three fundamental differences contributing to the discrepancy between these spectra. First, root traits are simultaneously constrained by various environmental drivers not necessarily related to resource uptake. Second, above- and belowground traits cannot be considered analogues, because they function differently and might not be related to resource uptake in a similar manner. Third, mycorrhizal interactions may offset selection for an RES. Understanding and explaining the belowground mechanisms and trade-offs that drive variation in root traits, resource acquisition and plant performance across species, thus requires a fundamentally different approach than applied aboveground. We therefore call for studies that can functionally incorporate the root traits involved in resource uptake, the complex soil environment and the various soil resource uptake mechanisms - particularly the mycorrhizal pathway - in a multidimensional root trait framework.

Dead wood provides a huge terrestrial carbon stock and a habitat to wide-ranging organisms during its decay. Our brief review highlights that, in order to understand environmental change impacts on these functions, we need to quantify the contributions of different interacting biotic and abiotic drivers to wood decomposition. LOGLIFE is a new long-term 'common-garden' experiment to disentangle the effects of species' wood traits and siterelated environmental drivers on wood decomposition dynamics and its associated diversity of microbial and invertebrate communities. This experiment is firmly rooted in pioneering experiments under the directorship of Terry Callaghan at Abisko Research Station, Sweden. LOGLIFE features two contrasting forest sites in the Netherlands, each hosting a similar set of coarse logs and branches of 10 tree species. LOGLIFE welcomes other researchers to test further questions concerning coarse wood decay that will also help to optimise forest management in view of carbon sequestration and biodiversity conservation. Copyright © Royal Swedish Academy of Sciences 2012.

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.

Under drought conditions, arbuscular mycorrhizal (AM) fungi alter water relationships of plants and improve their resistance to drought. In a factorial greenhouse experiment, we tested the effects of the AM symbiosis and precipitation regime on the performance (growth, gas exchange, nutrient status and mycorrhizal responsiveness) of Boswellia papyrifera seedlings. A continuous precipitation regime was imitated by continuous watering of plants to field capacity every other day during 4 months, and irregular precipitation by pulsed watering of plants where watering was switched every 15 days during these 4 months, with 15 days of watering followed by 15 days without watering. There were significantly higher levels of AM colonization under irregular precipitation regime than under continuous precipitation. Mycorrhizal seedlings had higher biomass than control seedlings. Stomatal conductance and phosphorus mass fraction in shoot and root were also significantly higher for mycorrhizal seedlings. Mycorrhizal seedlings under irregular watering had the highest biomass. Both a larger leaf area and higher assimilation rates contributed to higher biomass. Under irregular watering, the water use efficiency increased in non-mycorrhizal seedlings through a reduction in transpiration, while in mycorrhizal seedlings irregular watering increased transpiration. Because assimilation rates increased even more, mycorrhizal seedlings achieved an even higher water use efficiency. Boswellia seedlings allocated almost all carbon to the storage root. Boswellia seedlings had higher mass fractions of N, P, and K in roots than in shoots. Irregular precipitation conditions apparently benefit Boswellia seedlings when they are mycorrhizal. Electronic supplementary material The online version of this article (doi:10.1007/s00442-012-2258-3) contains supplementary material, which is available to authorized users.

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

In Ethiopia, forests near churches, are the last remnant forest patches. These forests are currently under threat, probably due to diminishing areas of the forest itself and repeated grazing for extended periods by cattle. We assessed the effect of livestock exclusion on the regeneration of four indigenous tree species in two church forests. The four species have a high abundance and socioeconomic value, but limited regeneration in the two forests. We investigated the effect of grazing and trampling on seed germination, seedling survival, and seedling growth. Livestock grazing had a strong negative effect on germination, seedling growth and mortality. In fenced plots, more seeds germinated, seedling survival was higher and seedlings grew faster. Seed germination was higher inside the forest than in the adjacent open area for all species. Seedling survival was not different between forest interior and open fields, except for unfenced plots in the open fields where survival was lower because of the higher grazing pressure. In unfenced plots, no seedlings survived until the end of the year, indicating that grazers destroyed the seedling bank in and around the forest. The significant interaction between fencing and species on seed germination and seedling survival revealed that the magnitude of damage due to grazing can vary with species. We conclude that for effective indigenous tree species regeneration in these church forests, the control of livestock pressure is necessary. Seeds dispersed outside the forest will not have a chance to establish seedlings, grow and colonize the surroundings. Livestock grazing thus has a paramount impact on the long-term sustainability of church forests and their role in restoring the degraded surroundings.

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

Many studies conclude that light is the most important resource that determines plant performance of tree saplings in tropical rain forests, and implicitly suggest that soil resources are less important. To provide a quantitative test for soil versus light effects on sapling performance, we studied how saplings of the shade-tolerant tree speciesBrosimum alicastrumresponded to contrasting levels of light availability and soil fertility in a Mexican tropical rain forest. Therefore saplings were selected from ten low-light exposure (crown position index

Most semiarid regions are facing an increasing scarcity of woody vegetation due mainly to anthropogenic deforestation aggravated by climate changes. However, there is insufficient information to reconstruct past changes in climate and to evaluate the implications of future climate changes on the vegetation. Tree-ring analysis is a powerful tool for studying tree age, population dynamics, growth behavior, and climate-growth relationships among tropical tree species and for gaining information about the environmental forces driving growth change as well as for developing proxies for climate reconstruction. Wood anatomical and dendrochronological methods were used on Juniperus procera trees from two Ethiopian highland forests to check (i) whether tree-ring series of juniper are cross-datable and hence suitable for building tree-ring chronologies, and if so, (ii) which climate factors mainly drive wood formation in juniper from this region. Visible growth layers of the juniper wood were shown to be annual rings. Tree-ring sequences could be cross-dated between trees growing at the same site and between trees growing at sites 350 km apart. Evidence was found that annual growth of junipers is mainly controlled by one climatic factor, precipitation. This strong precipitation influence proves the potential of African juniper chronologies for accurate climate reconstruction and points out the relevance of building a network of juniper chronologies across East Africa.

Summary Leaf specific conductivity (LSC; the ratio of stem conductivity (KP) to leaf area (AL)), a measure of the hydraulic capacity of the stem to supply leaves with water, varies with soil water content. Empirical evidence for LSC responses to drought is ambiguous, because previously published results were subject to many confounding factors.We tested howLSC of similar-sized trees of the same population, under similar climatic conditions, responds to persistently wet or dry soil. Scots pine (Pinus sylvestris L.) and pubescent oak (Quercus pubescensWilld.) trees were compared between a dry site and a wet site in theValais, an inner alpine valley in Switzerland. Soilwater strongly influenced AL and KP and the plant components affecting KP, such as conduit radius, conduit density and functional sapwood area. Trees at the dry site had lower LSC than trees with the same stem diameter at the wet site. Low LSC in trees at the dry site was associated with a smaller functional sapwood area and narrower conduits, resulting in a stronger reduction in KP than in AL. These observations support the hypothesis that trees maintain a homeostatic water pressure gradient. An alternative hypothesis is that relatively high investments in leaves compared with sapwood contribute to carbon gain over an entire season by enabling rapid whole-plant photosynthesis during periods of high water availability (e.g., in spring, after rain events and during morning hours when leafto- air vapor pressure deficit is small). Dynamic data and a hydraulic plant growth model are needed to test how investments in leaves versus sapwood and roots contribute to transpiration and to maximizing carbon gain throughout entire growth seasons. Keywords: Huber value, leaf area, soil water potential, stem conductivity.

Natural regeneration in gaps in Douglas fir forest stands in the Netherlands mainly consists of Betula pendula (Roth.), Pinus sylvestris (L.), Larix kaempferi (Carr.), and Pseudotsuga menziesii (Mirb. Franco). Even though these species are well known, the autogenic development of these species in an unmanaged plant community falls outside of traditional silvicultural experience. We therefore present an explorative study, aimed to map out differences between these species under conditions of co-occurrence, and develop hypotheses on the further development of these sapling communities. We focused on differences in height, and particularly the architectural contribution to height. We analyzed interspecific differences in the height–mass and height–DBH relationship, biomass partitioning, and slenderness for a total of 177 saplings up to 13.7 cm DBH and 11.8 m high. Average annual height and diameter growth rates of an additional 48 saplings are presented for comparison with the static architectural data. Betula had a high stem biomass, high stem slenderness, and low leaf mass. The reverse was true for Pseudotsuga . Larix behaved as an intermediate species. Pinus however deviated from the classical pioneer–climax dichotomy in having a low slenderness and low height per unit biomass while being a highly light-demanding species. The static height-mass relationship shows a similar species ranking when compared to average annual height growth rates, but not when compared to diameter growth rates. This underlines the importance of a species’ architecture for its height development and suggesting a predictable stratification between species. Based on the results regarding species’ stratification in heights and a priori information on light demand, it is hypothesized that in the studied gap-phase regeneration Betula , Larix and Pseudotsuga will partition the available light. Pinus however will be overtopped and given its need for high-light environments, is expected to be competitively excluded.

Crown development was monitored for juvenile trees (4–18 m) of the canopy tree species Dicorynia guianensis and Vouacapoua americana in a tropical rain forest in French Guiana. A comparison was made between crown development in closed understorey forest sites (low light) and in canopy gaps (high light); development was expressed by plant traits at multiple levels of organisation. Dicorynia and Vouacapoua responded to gap creation at all organisational levels, but not for all traits. Both species increased their light interception efficiency in the understorey. Firstly, understorey trees produced relatively wide crowns because they did not favour the growth of shoots in the summit of the crown, as did gap trees. Secondly, they reduced self-shading (expressed by leaf area index) by lower sympodial unit production rates, lower leaf production rates per growth unit (only Vouacapoua), and smaller leaf size (only Dicorynia). The reduction in self-shading was weakened by other traits that had the opposite effect on self shading (e.g. longer leaf life span in understorey), and cannot be considered an adaptive response in itself. (3) Understorey trees of Vouacapoua reduced leaf display costs by producing a higher specific leaf area, a shorter space among leaves, a smaller leaf spacing to leaf size ratio, and a longer leaf life span. Thus, all traits contributed to the more economical use of carbon in the understorey. Dicorynia showed the same trends, but not significantly. These results suggest that light availability plays a major role in the development and morphology of trees through its influences at multiple levels of organisation within the crown hierarchy. The two species studied were rather similar in their response at crown level, while they sometimes differed in their responses at the underlying lower organisational levels. The response directions of some individual plant traits were similar for the tall trees studied here and small saplings studied elsewhere. Some of traits investigated here may not be important in the crown level responses of smaller seedlings and saplings, while they are important mechanisms for crown level responses in taller trees.

Light fluctuations and crown traits were studied for saplings of four tree species in a Costa Rican rain forest. Light fluctuations (1988-1994) were assessed by annual light estimations above saplings, using a visual crown position index. Crown traits in 1994 and growth between 1994 and 1995 were measured. Crown position values varied between 1.5 and 2.5. Of the 70 saplings only four were found at higher light levels (crown position of 3 or 3.5), but for 1-2 y only. Over the 6 y of investigation, 55-75 % of the saplings experienced no or only one light fluctuation, and 25-45 % two or three fluctuations. Crown traits in 1994 were either most strongly correlated with light levels in 1993 (Lecythis) and with light levels in 1991 and 1992 (Dipteryx and Simarouba), or they were not significantly correlated with light levels (Minquartia). It is hypothesised that: (1) the saplings require 1-3 y to establish a crown trait in response to light levels in the forest; and (2) and species that can economically produce a leaf can adjust crown traits more quickly in an environment dominated by low light levels than species that are less economic. Crown trait responses may track environmental changes in three of the four species, in particular in Lecythis. In this latter species, leaf area had no significant effect on growth and survival, whereas light level had a positive effect. Conversely, in the other three species, light levels had no discernible effect on growth and survival (due in part to low variation in crown position in two of these species), whereas leaf area had a positive effect on both.

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.

Most palm species occur in the shaded lower strata of tropical rain forests, but how their traits relate to shade adaptation is poorly understood. We hypothesized that palms are adapted to the shade of their native habitats by convergent evolution towards high net carbon gain efficiency (CGEn), which is given by the maximum photosynthetic rate to dark respiration rate ratio. Leaf mass per area, maximum photosynthetic rate, dark respiration and N and P concentrations were measured in 80 palm species grown in a common garden, and combined with data of 30 palm species growing in their native habitats. Compared to other species from the global leaf economics data, dicotyledonous broad-leaved trees in tropical rainforest or other monocots in the global leaf economics data, palms possessed consistently higher CGEn, achieved by lowered dark respiration and fairly high foliar P concentration. Combined phylogenetic analyses of evolutionary signal and trait evolution revealed convergent evolution towards high CGEn in palms. We conclude that high CGEn is an evolutionary strategy that enables palms to better adapt to shady environments than coexisting dicot tree species, and may convey advantages in competing with them in the tropical forest understory. These findings provide important insights for understanding the evolution and ecology of palms, and for understanding plant shade adaptations of lower rainforest strata. Moreover, given the dominant role of palms in tropical forests, these findings are important for modelling carbon and nutrient cycling in tropical forest ecosystems.

Carbon allocation to sapwood in tropical canopy trees is a key process determining forest carbon sequestration, and is at the heart of tree growth and dynamic global vegetation models (DGVM). Several allocation hypotheses exist including those applying assumptions on fixed allocation, pipe model, and hierarchical allocation between plant organs. We use a tree growth model (IBTREE) to evaluate these hypotheses by comparing simulated sapwood growth with 30 year tree ring records of the tropical long-lived tree Toona ciliata M. Roem. in Thailand. Simulated annual variation in wood production varied among hypotheses. Observed and simulated growth patterns matched most closely (r2 = 0.70) when hierarchical allocation was implemented, with low priority for sapwood. This allocation method showed realistic results with respect to reserve dynamics, partitioning and productivity and was the only one able to capture the large annual variation in tree ring width. Consequently, this method might also explain the large temporal variation in diameter growth and the occurrence of missing rings often encountered in other tropical tree species. Overall, our results show that sapwood growth is highly sensitive to allocation principles, and that allocation assumptions may greatly influence estimated carbon sequestration of tropical forests under climatic change.

Intraspecific and interspecific architectural patterns were studied for eight tree species of a Bornean rain forest. Trees 5–19 m tall in two 4-ha permanent sample plots in primary forest were selected, and three light descriptors and seven architectural traits for each tree were measured. Two general predictions were made: (1) Slow growing individuals (or short ones) encounter lower light, and have flatter crowns, fewer leaf layers, and thinner stems, than do fast growing individuals (or tall ones). (2) Species with higher shade-tolerance receive less light and have flatter crowns, fewer leaf layers, and thinner stems, than do species with lower shade-tolerance. Shade-tolerance is assumed to decrease with maximum growth rate, mortality rate, and adult stature of a species.

Seasonal rainfall affects tropical forest dynamics and behavior of species that are part of these ecosystems. The positive correlation between amphibian activity patterns and rainfall has been demonstrated repeatedly. Members of Dendrobatidae, a clade of Neotropical dartpoison frogs, are well known for their habitat use and behavior during the rainy season, but their behavior during the dry season has received little attention. We studied habitat use and diet of the dendrobatid frog Dendrobates tinctorius in French Guiana during the rainy and dry seasons. Unlike many other dendrobatid frogs, D. tinctorius does not maintain territories for the entire rainy season. Both sexes colonize recently formed canopygaps and stay in these forest patches for only a few weeks. The frogs in these patches consume a great diversity of prey, consisting of ants, beetles, wasps, insect larvae, and mites. During the dry season, frogs move to retreat sites in mature forest, such as palm bracts and tree holes. The frogs are less active and consume fewer prey items in the dry season, and they consume fewer wasps and insect larvae, but more termites. Ants are the most common prey items during both the wet and dry seasons. We discuss the effects of shifts in seasonal habitat use on the territorial behavior of dendrobatid frogs.

Journal of Experimental Botany, 58 (8), ISSN:1460-2431, ISSN:0022-0957