Demography and selection analysis of the incipient adaptive radiation of a Hawaiian woody species.

Ecological divergence in a species provides a valuable opportunity to study the early stages of speciation. We focused on Metrosideros polymorpha, a unique example of the incipient radiation of woody species, to examine how an ecological divergence continues in the face of gene flow. We analyzed the whole genomes of 70 plants collected throughout the island of Hawaii, which is the youngest island with the highest altitude in the archipelago and encompasses a wide range of environments. The continuous M. polymorpha forest stands on the island of Hawaii were differentiated into three genetic clusters, each of which grows in a distinctive environment and includes substantial genetic and phenotypic diversity. The three genetic clusters showed signatures of selection in genomic regions encompassing genes relevant to environmental adaptations, including genes associated with light utilization, oxidative stress, and leaf senescence, which are likely associated with the ecological differentiation of the species. Our demographic modeling suggested that the glaberrima cluster in wet environments maintained a relatively large population size and two clusters split: polymorpha in the subalpine zone and incana in dry and hot conditions. This ecological divergence possibly began before the species colonized the island of Hawaii. Interestingly, the three clusters recovered genetic connectivity coincidentally with a recent population bottleneck, in line with the weak reproductive isolation observed in the species. This study highlights that the degree of genetic differentiation between ecologically-diverged populations can vary depending on the strength of natural selection in the very early phases of speciation.

High exposure of global tree diversity to human pressure.

Safeguarding Earth's tree diversity is a conservation priority due to the importance of trees for biodiversity and ecosystem functions and services such as carbon sequestration. Here, we improve the foundation for effective conservation of global tree diversity by analyzing a recently developed database of tree species covering 46,752 species. We quantify range protection and anthropogenic pressures for each species and develop conservation priorities across taxonomic, phylogenetic, and functional diversity dimensions. We also assess the effectiveness of several influential proposed conservation prioritization frameworks to protect the top 17% and top 50% of tree priority areas. We find that an average of 50.2% of a tree species' range occurs in 110-km grid cells without any protected areas (PAs), with 6,377 small-range tree species fully unprotected, and that 83% of tree species experience nonnegligible human pressure across their range on average. Protecting high-priority areas for the top 17% and 50% priority thresholds would increase the average protected proportion of each tree species' range to 65.5% and 82.6%, respectively, leaving many fewer species (2,151 and 2,010) completely unprotected. The priority areas identified for trees match well to the Global 200 Ecoregions framework, revealing that priority areas for trees would in large part also optimize protection for terrestrial biodiversity overall. Based on range estimates for >46,000 tree species, our findings show that a large proportion of tree species receive limited protection by current PAs and are under substantial human pressure. Improved protection of biodiversity overall would also strongly benefit global tree diversity.

Light competition drives species replacement during secondary tropical forest succession.

Light competition is thought to drive successional shifts in species dominance in closed vegetations, but few studies have assessed this for species-rich and vertically structured tropical forests. We analyzed how light competition drives species replacement during succession, and how cross-species variation in light competition strategies is determined by underlying species traits. To do so, we used chronosequence approach in which we compared 14 Mexican tropical secondary rainforest stands that differ in age (8-32 year-old). For each tree, height and stem diameter were monitored for 2 years to calculate relative biomass growth rate (RGR, the aboveground biomass gain per unit aboveground tree biomass per year). For each stand, 3D light profiles were measured to estimate individuals' light interception to calculate light interception efficiency (LIE, intercepted light per unit biomass per year) and light use efficiency (LUE, biomass growth per intercepted light). Throughout succession, species with higher RGR attained higher changes in species dominance and thus increased their dominance over time. Both light competition strategies (LIE and LUE) increased RGR. In early succession, a high LIE and its associated traits (large crown leaf mass and low wood density) are more important for RGR. During succession, forest structure builds up, leading to lower understory light levels. In later succession, a high LUE and its associated traits (high wood density and leaf mass per area) become more important for RGR. Therefore, successional changes in relative importance of light competition strategies drive shifts in species dominance during tropical rainforest succession.

Incorporating pressure-volume traits into the leaf economics spectrum.

In recent years, attempts have been made in linking pressure-volume parameters and the leaf economics spectrum to expand our knowledge of the interrelationships among leaf traits. We provide theoretical and empirical evidence for the coordination of the turgor loss point and associated traits with net CO2 assimilation (An ) and leaf mass per area (LMA). We measured gas exchange, pressure-volume curves and leaf structure in 45 ferns and angiosperms, and explored the anatomical and chemical basis of the key traits. We propose that the coordination observed between mass-based An , capacitance and the turgor loss point (πtlp ) emerges from their shared link with leaf density (one of the components of LMA) and, specially, leaf saturated water content (LSWC), which in turn relates to cell size and nitrogen and carbon content. Thus, considering the components of LMA and LSWC in ecophysiological studies can provide a broader perspective on leaf structure and function.

Disentangling leaf structural and material properties in relationship to their anatomical and chemical compositional traits in oaks (Quercus L.).

BACKGROUND AND AIMS: The existence of sclerophyllous plants has been considered an adaptive strategy against different environmental stresses. Given that it literally means 'hard-leaved', it is essential to quantify the leaf mechanical properties to understand sclerophylly. However, the relative importance of each leaf trait for mechanical properties is not yet well established. METHODS: Genus Quercus is an excellent system to shed light on this because it minimizes phylogenetic variation while having a wide variation in sclerophylly. We measured leaf anatomical traits and cell wall composition, analysing their relationship with leaf mass per area and leaf mechanical properties in a set of 25 oak species. KEY RESULTS: The upper epidermis outer wall makes a strong and direct contribution to the leaf mechanical strength. Moreover, cellulose plays a crucial role in increasing leaf strength and toughness. The principal component analysis plot based on leaf trait values clearly separates Quercus species into two groups corresponding to evergreen and deciduous species. CONCLUSIONS: Sclerophyllous Quercus species are tougher and stronger owing to their thicker epidermis outer wall and/or higher cellulose concentration. Furthermore, section Ilex species share common traits, although they occupy different climates. In addition, evergreen species living in mediterranean-type climates share common leaf traits irrespective of their different phylogenetic origin.

A cost-benefit analysis of leaf carbon economy with consideration of seasonal changes in leaf traits for sympatric deciduous and evergreen congeners: implications for their coexistence.

Deciduous and evergreen species, which have evolved repeatedly across different clades, can coexist in a given environment despite substantial differences in their leaf traits. It remains unclear how these two groups differ in the development of leaf traits over their lifespans or how their carbon economy - the balance between lifetime carbon gain and leaf construction cost - is determined. We determined the photosynthetic rate (Aarea ), leaf mass per area (LMA), leaf mechanical strength and leaf water potentials and estimated the lifetime carbon gain and leaf construction cost of five closely related pairs of evergreen and deciduous species co-occurring in a temperate forest. Aarea of evergreen species was lower during their first spring, similar in summer and higher than the autumn until the following spring than their deciduous counterparts. Leaf mechanical strength, osmotic pressures and LMA increased continuously towards winter in evergreen species while remaining largely constant in deciduous species. The ratio of lifetime carbon gain to leaf construction cost was similar between the two groups. The additional cost associated with enduring winter is paid back by a longer revenue of photosynthesis in evergreen species, allowing evergreen and deciduous leaf habits to coexist in the seasonal environment.

Small and slow is safe: On the drought tolerance of tropical tree species.

Understanding how evolutionary history and the coordination between trait trade-off axes shape the drought tolerance of trees is crucial to predict forest dynamics under climate change. Here, we compiled traits related to drought tolerance and the fast-slow and stature-recruitment trade-off axes in 601 tropical woody species to explore their covariations and phylogenetic signals. We found that xylem resistance to embolism (P50) determines the risk of hydraulic failure, while the functional significance of leaf turgor loss point (TLP) relies on its coordination with water use strategies. P50 and TLP exhibit weak phylogenetic signals and substantial variation within genera. TLP is closely associated with the fast-slow trait axis: slow species maintain leaf functioning under higher water stress. P50 is associated with both the fast-slow and stature-recruitment trait axes: slow and small species exhibit more resistant xylem. Lower leaf phosphorus concentration is associated with more resistant xylem, which suggests a (nutrient and drought) stress-tolerance syndrome in the tropics. Overall, our results imply that (1) drought tolerance is under strong selective pressure in tropical forests, and TLP and P50 result from the repeated evolutionary adaptation of closely related taxa, and (2) drought tolerance is coordinated with the ecological strategies governing tropical forest demography. These findings provide a physiological basis to interpret the drought-induced shift toward slow-growing, smaller, denser-wooded trees observed in the tropics, with implications for forest restoration programmes.

The acquisitive-conservative axis of leaf trait variation emerges even in homogeneous environments.

BACKGROUND AND AIMS: The acquisitive-conservative axis of plant ecological strategies results in a pattern of leaf trait covariation that captures the balance between leaf construction costs and plant growth potential. Studies evaluating trait covariation within species are scarcer, and have mostly dealt with variation in response to environmental gradients. Little work has been published on intraspecific patterns of leaf trait covariation in the absence of strong environmental variation. METHODS: We analysed covariation of four leaf functional traits [specific leaf area (SLA) leaf dry matter content (LDMC), force to tear (Ft) and leaf nitrogen content (Nm)] in six Poaceae and four Fabaceae species common in the dry Chaco forest of Central Argentina, growing in the field and in a common garden. We compared intraspecific covariation patterns (slopes, correlation and effect size) of leaf functional traits with global interspecific covariation patterns. Additionally, we checked for possible climatic and edaphic factors that could affect the intraspecific covariation pattern. KEY RESULTS: We found negative correlations for the LDMC-SLA, Ft-SLA, LDMC-Nm and Ft-Nm trait pairs. This intraspecific covariation pattern found both in the field and in the common garden and not explained by climatic or edaphic variation in the field follows the expected acquisitive-conservative axis. At the same time, we found quantitative differences in slopes among different species, and between these intraspecific patterns and the interspecific ones. Many of these differences seem to be idiosyncratic, but some appear consistent among species (e.g. all the intraspecific LDMC-SLA and LDMC-Nm slopes tend to be shallower than the global pattern). CONCLUSIONS: Our study indicates that the acquisitive-conservative leaf functional trait covariation pattern occurs at the intraspecific level even in the absence of relevant environmental variation in the field. This suggests a high degree of variation-covariation in leaf functional traits not driven by environmental variables.

Sources and consequences of mismatch between leaf disc and whole-leaf leaf mass per area (LMA).

PREMISE: Leaf mass per area (LMA), which is an important functional trait in leaf economic spectrum and plant growth analysis, is measured from leaf discs or whole leaves. Differences between the measurement methods may lead to large differences in the estimates of LMA values. METHODS: We examined to what extent estimates of LMA based on whole leaves match those based on discs using 334 woody species from a wide range of biomes (tropics, subtropics, savanna, and temperate), whether the relationship varied by leaf morphology (tissue density, leaf area, leaf thickness), punch size (0.6- and 1.0-cm diameter), and whether the extent of intraspecifc variation for each species matches. RESULTS: Disc-based estimates of species mean LMA matched the whole-leaf estimates well, and whole-leaf LMA tended to be 9.69% higher than leaf-disc LMA. The ratio of whole-leaf LMA to leaf-disc LMA was higher for species with higher leaf tissue density and larger leaves, and variance in the ratio was greater for species with lower leaf tissue density and thinner leaves. Estimates based on small leaf discs also inflated the ratio. The extent of the intraspecific variation only weakly matched between whole-leaf and disc-based estimates (R2 = 0.08). CONCLUSIONS: Our results suggest that simple conversion between whole-leaf and leaf-disc LMA is difficult for species obtained with a small leaf punch, but it should be possible for species obtained with a large+ leaf punch. Accurately representing leaf traits will likely require careful selection between leaf-disc and whole-leaf traits depending on the objectives. Quantifying intraspecific variation using leaf discs should be also considered with caution.

Plant functional traits have globally consistent effects on competition.

Phenotypic traits and their associated trade-offs have been shown to have globally consistent effects on individual plant physiological functions, but how these effects scale up to influence competition, a key driver of community assembly in terrestrial vegetation, has remained unclear. Here we use growth data from more than 3 million trees in over 140,000 plots across the world to show how three key functional traits--wood density, specific leaf area and maximum height--consistently influence competitive interactions. Fast maximum growth of a species was correlated negatively with its wood density in all biomes, and positively with its specific leaf area in most biomes. Low wood density was also correlated with a low ability to tolerate competition and a low competitive effect on neighbours, while high specific leaf area was correlated with a low competitive effect. Thus, traits generate trade-offs between performance with competition versus performance without competition, a fundamental ingredient in the classical hypothesis that the coexistence of plant species is enabled via differentiation in their successional strategies. Competition within species was stronger than between species, but an increase in trait dissimilarity between species had little influence in weakening competition. No benefit of dissimilarity was detected for specific leaf area or wood density, and only a weak benefit for maximum height. Our trait-based approach to modelling competition makes generalization possible across the forest ecosystems of the world and their highly diverse species composition.

Vertical and horizontal light heterogeneity along gradients of secondary succession in cool- and warm-temperate forests.

Aims: Light availability varies drastically in forests, both vertically and horizontally. Vertical light heterogeneity (i.e., patterns of light attenuation from the forest canopy to the floor) may be related to light competition among trees, while horizontal light heterogeneity (i.e., variations in light intensity at a given height within forests) may be associated with light-niche partitioning among tree species. However, light heterogeneity in vertical and horizontal directions and their associations with forest structure are rarely studied to date. Here we report the first comprehensive study to compare the vertical and horizontal light heterogeneity in differently aged forests in two forest types. Location: Twelve forest stands of different ages in cool-temperate forests (consisting of deciduous broad-leaved trees) and five of different ages in warm-temperate forests (evergreen conifer and deciduous broad-leaved trees) in Japan. Methods: We measured vertical light profiles at 1-m intervals from the understorey (1 m above the ground) to the top canopy (12-22 m depending on stands) at 16 locations for each stand (20 m × 20 m). We also measured structural parameters (diameter at breast height, height, and crown dimensions) for all major trees in these stands. Results: Along the secondary successional gradients, the vertical and horizontal light heterogeneity changed in a systematic manner in both forests. The vertical light attenuation rate was steeper in early succession and more gradual in late succession, and the horizontal light heterogeneity was relatively small in early succession and more pronounced in late succession. The vertical light attenuation rate was different between the two forest types; the light intensity dropped more sharply from the canopy surface in the cool-temperate forests due to the crown being vertically shorter and denser (i.e., higher leaf density per unit volume). Conclusion: In early succession, a steeper light attenuation rate is likely related to the strong light competition among co-occurring trees and thus a self-thinning process. In late succession, the high spatial light heterogeneity in forests (i.e., larger horizontal light heterogeneity and gradual light attenuation rate) may allow more species to partition light, and thus may enhance species coexistence and diversity.

How plants grow under gravity conditions besides 1 g: perspectives from hypergravity and space experiments that employ bryophytes as a model organism.

Plants have evolved and grown under the selection pressure of gravitational force at 1 g on Earth. In response to this selection pressure, plants have acquired gravitropism to sense gravity and change their growth direction. In addition, plants also adjust their morphogenesis in response to different gravitational forces in a phenomenon known as gravity resistance. However, the gravity resistance phenomenon in plants is poorly understood due to the prevalence of 1 g gravitational force on Earth: not only it is difficult to culture plants at gravity > 1 g(hypergravity) for a long period of time but it is also impossible to create a < 1 genvironment (µg, micro g) on Earth without specialized facilities. Despite these technical challenges, it is important to understand how plants grow in different gravity conditions in order to understand land plant adaptation to the 1 g environment or for outer space exploration. To address this, we have developed a centrifugal device for a prolonged duration of plant culture in hypergravity conditions, and a project to grow plants under the µg environment in the International Space Station is also underway. Our plant material of choice is Physcomitrium (Physcomitrella) patens, one of the pioneer plants on land and a model bryophyte often used in plant biology. In this review, we summarize our latest findings regarding P. patens growth response to hypergravity, with reference to our on-going "Space moss" project. In our ground-based hypergravity experiments, we analyzed the morphological and physiological changes and found unexpected increments of chloroplast size and photosynthesis rate, which might underlie the enhancement of growth and increase in the number of gametophores and rhizoids. We further discussed our approaches at the cellular level and compare the gravity resistance in mosses and that in angiosperms. Finally, we highlight the advantages and perspectives from the space experiments and conclude that research with bryophytes is beneficial to comprehensively and precisely understand gravitational responses in plants.

Global patterns of leaf construction traits and their covariation along climate and soil environmental gradients.

Leaf functional traits and their covariation underlie plant ecological adaptations along environmental gradients, but there is limited information on the global covariation patterns of key leaf construction traits. To explore how leaf construction traits co-vary across diverse climate and soil environmental conditions, we compiled a global dataset including cell wall mass per unit leaf mass (CWmass ), leaf carbon (C) and calcium (Ca) concentrations, and specific leaf area (SLA) for 2348 angiosperm species from 340 sites world-wide. Our results demonstrated negative correlations between leaf C and Ca concentrations and between leaf C and SLA across diverse nongraminoid angiosperms. Leaf C concentration increased with increasing mean annual temperature (MAT) and mean annual precipitation (MAP) and with decreasing soil pH and calcium carbonate (CaCO3 ) concentration, whereas leaf Ca concentration and SLA exhibited the opposite responses to these environmental variables. The covariations of leaf Ca-C and of leaf SLA-C were stronger in habitats with lower MAT and MAP, and/or higher soil CaCO3 content. This global-scale analysis demonstrates that the leaf C and Ca concentrations and SLA together govern the C and biomass investment strategies in leaves of nongraminoids. We conclude that environmental conditions strongly shape leaf construction traits and their covariation patterns.

Leaf trichomes in Metrosideros polymorpha can contribute to avoiding extra water stress by impeding gall formation.

BACKGROUND AND AIMS: Plants inhabiting arid environments tend to have leaf trichomes, but their adaptive significance remains unclear. Leaf trichomes are known to play a role in plant defence against herbivores, including gall makers. Because gall formation can increase water loss partly through increased surface area, we tested the novel hypothesis that leaf trichomes could contribute to avoiding extra water stress by impeding gall formation, which would have adaptive advantages in arid environments. METHODS: We focused on Metrosideros polymorpha, an endemic tree species in the Hawaiian Islands, whose leaves often suffer from galls formed by specialist insects, Hawaiian psyllids (Pariaconus spp.). There is large variation in the amount of leaf trichomes (0-40 % of leaf mass) in M. polymorpha. Three gall types are found on the island of Hawaii: the largest is the 'cone' type, followed by 'flat' and 'pit' types. We conducted laboratory experiments to quantify the extent to which gall formation is associated with leaf water relations. We also conducted a field census of 1779 individuals from 48 populations across the entire range of habitats of M. polymorpha on the island of Hawaii to evaluate associations between gall formation (presence and abundance) and the amount of leaf trichomes. KEY RESULTS: Our laboratory experiment showed that leaf minimum conductance was significantly higher in leaves with a greater number of cone- or flat-type galls but not pit-type galls. Our field census suggested that the amount of trichomes was negatively associated with probabilities of the presence of cone- or flat-type galls but not pit-type galls, irrespective of environmental factors. CONCLUSION: Our results suggest that leaf trichomes in M. polymorpha can contribute to the avoidance of extra water stress through interactions with some gall-making species, and potentially increase the fitness of plants under arid conditions.

Does the leaf economic spectrum hold within plant functional types? A Bayesian multivariate trait meta-analysis.

The leaf economic spectrum is a widely studied axis of plant trait variability that defines a trade-off between leaf longevity and productivity. While this has been investigated at the global scale, where it is robust, and at local scales, where deviations from it are common, it has received less attention at the intermediate scale of plant functional types (PFTs). We investigated whether global leaf economic relationships are also present within the scale of plant functional types (PFTs) commonly used by Earth System models, and the extent to which this global-PFT hierarchy can be used to constrain trait estimates. We developed a hierarchical multivariate Bayesian model that assumes separate means and covariance structures within and across PFTs and fit this model to seven leaf traits from the TRY database related to leaf longevity, morphology, biochemistry, and photosynthetic metabolism. Although patterns of trait covariation were generally consistent with the leaf economic spectrum, we found three approximate tiers to this consistency. Relationships among morphological and biochemical traits (specific leaf area [SLA], N, P) were the most robust within and across PFTs, suggesting that covariation in these traits is driven by universal leaf construction trade-offs and stoichiometry. Relationships among metabolic traits (dark respiration [Rd ], maximum RuBisCo carboxylation rate [Vc,max ], maximum electron transport rate [Jmax ]) were slightly less consistent, reflecting in part their much sparser sampling (especially for high-latitude PFTs), but also pointing to more flexible plasticity in plant metabolistm. Finally, relationships involving leaf lifespan were the least consistent, indicating that leaf economic relationships related to leaf lifespan are dominated by across-PFT differences and that within-PFT variation in leaf lifespan is more complex and idiosyncratic. Across all traits, this covariance was an important source of information, as evidenced by the improved imputation accuracy and reduced predictive uncertainty in multivariate models compared to univariate models. Ultimately, our study reaffirms the value of studying not just individual traits but the multivariate trait space and the utility of hierarchical modeling for studying the scale dependence of trait relationships.

Mapping local and global variability in plant trait distributions.

Our ability to understand and predict the response of ecosystems to a changing environment depends on quantifying vegetation functional diversity. However, representing this diversity at the global scale is challenging. Typically, in Earth system models, characterization of plant diversity has been limited to grouping related species into plant functional types (PFTs), with all trait variation in a PFT collapsed into a single mean value that is applied globally. Using the largest global plant trait database and state of the art Bayesian modeling, we created fine-grained global maps of plant trait distributions that can be applied to Earth system models. Focusing on a set of plant traits closely coupled to photosynthesis and foliar respiration-specific leaf area (SLA) and dry mass-based concentrations of leaf nitrogen ([Formula: see text]) and phosphorus ([Formula: see text]), we characterize how traits vary within and among over 50,000 [Formula: see text]-km cells across the entire vegetated land surface. We do this in several ways-without defining the PFT of each grid cell and using 4 or 14 PFTs; each model's predictions are evaluated against out-of-sample data. This endeavor advances prior trait mapping by generating global maps that preserve variability across scales by using modern Bayesian spatial statistical modeling in combination with a database over three times larger than that in previous analyses. Our maps reveal that the most diverse grid cells possess trait variability close to the range of global PFT means.

Leaf mechanical strength and photosynthetic capacity vary independently across 57 subtropical forest species with contrasting light requirements.

Leaf mechanical strength and photosynthetic capacity are critical plant life-history traits associated with tolerance and growth under various biotic and abiotic stresses. In principle, higher mechanical resistance achieved via higher relative allocation to cell walls should slow photosynthetic rates. However, interspecific relationships among these two leaf functions have not been reported. We measured leaf traits of 57 dominant woody species in a subtropical evergreen forest in China, focusing especially on photosynthetic rates, mechanical properties, and leaf lifespan (LLS). These species were assigned to two ecological strategy groups: shade-tolerant species and light-demanding species. On average, shade-tolerant species had longer LLS, higher leaf mechanical strength but lower photosynthetic rates, and exhibited longer LLS for a given leaf mass per area (LMA) or mechanical strength than light-demanding species. Depending on the traits and the basis of expression (per area or per mass), leaf mechanical resistance and photosynthetic capacity were either deemed unrelated, or only weakly negatively correlated. We found only weak support for the proposed trade-off between leaf biomechanics and photosynthesis among co-occurring woody species. This suggests there is considerable flexibility in these properties, and the observed relationships may result more so from trait coordination than any physically or physiologically enforced trade-off.

Acclimation and adaptation components of the temperature dependence of plant photosynthesis at the global scale.

The temperature response of photosynthesis is one of the key factors determining predicted responses to warming in global vegetation models (GVMs). The response may vary geographically, owing to genetic adaptation to climate, and temporally, as a result of acclimation to changes in ambient temperature. Our goal was to develop a robust quantitative global model representing acclimation and adaptation of photosynthetic temperature responses. We quantified and modelled key mechanisms responsible for photosynthetic temperature acclimation and adaptation using a global dataset of photosynthetic CO2 response curves, including data from 141 C3 species from tropical rainforest to Arctic tundra. We separated temperature acclimation and adaptation processes by considering seasonal and common-garden datasets, respectively. The observed global variation in the temperature optimum of photosynthesis was primarily explained by biochemical limitations to photosynthesis, rather than stomatal conductance or respiration. We found acclimation to growth temperature to be a stronger driver of this variation than adaptation to temperature at climate of origin. We developed a summary model to represent photosynthetic temperature responses and showed that it predicted the observed global variation in optimal temperatures with high accuracy. This novel algorithm should enable improved prediction of the function of global ecosystems in a warming climate.

Physiological and structural tradeoffs underlying the leaf economics spectrum.

The leaf economics spectrum (LES) represents a suite of intercorrelated leaf traits concerning construction costs per unit leaf area, nutrient concentrations, and rates of carbon fixation and tissue turnover. Although broad trade-offs among leaf structural and physiological traits have been demonstrated, we still do not have a comprehensive view of the fundamental constraints underlying the LES trade-offs. Here, we investigated physiological and structural mechanisms underpinning the LES by analysing a novel data compilation incorporating rarely considered traits such as the dry mass fraction in cell walls, nitrogen allocation, mesophyll CO2 diffusion and associated anatomical traits for hundreds of species covering major growth forms. The analysis demonstrates that cell wall constituents are major components of leaf dry mass (18-70%), especially in leaves with high leaf mass per unit area (LMA) and long lifespan. A greater fraction of leaf mass in cell walls is typically associated with a lower fraction of leaf nitrogen (N) invested in photosynthetic proteins; and lower within-leaf CO2 diffusion rates, as a result of thicker mesophyll cell walls. The costs associated with greater investments in cell walls underpin the LES: long leaf lifespans are achieved via higher LMA and in turn by higher cell wall mass fraction, but this inevitably reduces the efficiency of photosynthesis.

The population genomic signature of environmental association and gene flow in an ecologically divergent tree species Metrosideros polymorpha (Myrtaceae).

Genomewide markers enable us to study genetic differentiation within a species and the factors underlying it at a much higher resolution than before, which advances our understanding of adaptation in organisms. We investigated genomic divergence in Metrosideros polymorpha, a woody species that occupies a wide range of ecological habitats across the Hawaiian Islands and shows remarkable phenotypic variation. Using 1659 single nucleotide polymorphism (SNP) markers annotated with the genome assembly, we examined the population genetic structure and demographic history of nine populations across five elevations and two ages of substrates on Mauna Loa, the island of Hawaii. The nine populations were differentiated into two genetic clusters distributed on the lower and higher elevations and were largely admixed on the middle elevation. Demographic modelling revealed that the two genetic clusters have been maintained in the face of gene flow, and the effective population size of the high-altitude cluster was much smaller. A FST -based outlier search among the 1659 SNPs revealed that 34 SNPs (2.05%) were likely to be under divergent selection and the allele frequencies of 21 of them were associated with environmental changes along elevations, such as temperature and precipitation. This study shows a genomic mosaic of M. polymorpha, in which contrasting divergence patterns were found. While most genomic polymorphisms were shared among populations, a small fraction of the genome was significantly differentiated between populations in diverse environments and could be responsible for the dramatic adaptation to a wide range of environments.