Forest carbon storage and sink estimates under different management scenarios in China from 2020 to 2100.

Forests play a crucial role in mitigating climate change through carbon storage and sequestration, though environmental change drivers and management scenarios are likely to influence these contributions across multiple spatial and temporal scales. In this study, we employed three tree growth models-the Richard, Hossfeld, and Korf models-that account for the biological characteristics of trees, alongside national forest inventory (NFI) datasets from 1994 to 2018, to evaluate the carbon sink potential of existing forests and afforested regions in China from 2020 to 2100, assuming multiple afforestation and forest management scenarios. Our results indicate that the Richard, Hossfeld, and Korf models provided a good fit for 26 types of vegetation biomass in both natural and planted Chinese forests. These models estimate that in 2020, carbon stocks in existing Chinese forests are 7.62 ± 0.05 Pg C, equivalent to an average of 44.32 ± 0.32 Mg C/ ha. Our predictions then indicate this total forest carbon stock is expected to increase to 15.51 ± 0.99 Pg C (or 72.26 ± 4.6 Mg C/ha) in 2060, and further to 19.59 ± 1.36 Pg C (or 91.31 ± 6.33 Mg C/ha) in 2100. We also show that plantation management measures, namely tree species replacement, would increase carbon sinks to 0.09 Pg C/ year (contributing 38.9 %) in 2030 and 0.06 Pg C/ year (contributing 32.4 %) in 2060. Afforestation using tree species with strong carbon sink capacity in existing plantations would further significantly increase carbon sinks from 0.02 Pg C/year (contributing 10.3 %) in 2030 to 0.06 Pg C/year (contributing 28.2 %) in 2060. Our results quantify the role plantation management plays in providing a strong increase in forest carbon sequestration at national scales, pointing to afforestation with native tree species with high carbon sequestration as key in achieving China's 2060 carbon neutrality target.

Carbon fractions in wood for estimating embodied carbon in the built environment.

Determining wood carbon (C) fractions (CFs)-or the concentration of elemental C in wood on a per unit mass basis-in harvested wood products (HWP) is vital for accurately accounting embodied C in the built environment. Most estimates of embodied C assume that all wood-based building material is comprised of 50 % C on a per mass basis: an erroneous assumption that emerges from the literature on tree- and forest-scale C estimation, which has been shown to lead to substantial errors in C accounting. Here, we use published wood CF data from live trees, alongside laboratory analyses of sawn lumber, to quantify generalizable wood CFs for HWPs. Wood CFs in lumber average 51.7 %, deviating significantly from a 50 % default wood CF, as well as from CFs in live wood globally (which average 47.6 % across all species, and 47.1 % in tree species not typically employed in construction). Additionally, the volatile CF in lumber-i.e., the quantity of C lost upon heating of wood samples, but often overlooked in C accounting-is lower than the volatile CF in live wood, but significantly >0 % suggesting that industrial lumber drying processes remove some, but not all, of volatile C-based compounds. Our results demonstrate that empirically-supported wood CFs for construction material can correct meaningful systematic biases when estimating C storage in the built environment.

Leaf functional traits and pathogens: Linking coffee leaf rust with intraspecific trait variation in diversified agroecosystems.

Research has demonstrated that intraspecific functional trait variation underpins plant responses to environmental variability. However, few studies have evaluated how trait variation shifts in response to plant pathogens, even though pathogens are a major driver of plant demography and diversity, and despite evidence of plants expressing distinct strategies in response to pathogen pressures. Understanding trait-pathogen relationships can provide a more realistic understanding of global patterns of functional trait variation. We examined leaf intraspecific trait variability (ITV) in response to foliar disease severity, using Coffea arabica cv. Caturra as a model species. We quantified coffee leaf rust (CLR) severity-a fungal disease prominent in coffee systems-and measured key coffee leaf functional traits under contrasting, but widespread, management conditions in an agroforestry system. We found that coffee plants express significant ITV, which is largely related to shade tree treatment and leaf position within coffee canopy strata. Yet within a single plant canopy stratum, CLR severity increased with increasing resource conserving trait values. However, coffee leaves with visible signs of disease expressed overall greater resource acquiring trait values, as compared to plants without visible signs of disease. We provide among the first evidence that leaf traits are correlated with foliar disease severity in coffee, and that functional trait relationships and syndromes shift in response to increased disease prevalence in this plant-pathogen system. In doing so, we address a vital gap in our understanding of global patterns of functional trait variation and highlight the need to further explore the potential role of pathogens within established global trait relationships and spectra.

Global beta-diversity of angiosperm trees is shaped by Quaternary climate change.

As Earth's climate has varied strongly through geological time, studying the impacts of past climate change on biodiversity helps to understand the risks from future climate change. However, it remains unclear how paleoclimate shapes spatial variation in biodiversity. Here, we assessed the influence of Quaternary climate change on spatial dissimilarity in taxonomic, phylogenetic, and functional composition among neighboring 200-kilometer cells (beta-diversity) for angiosperm trees worldwide. We found that larger glacial-interglacial temperature change was strongly associated with lower spatial turnover (species replacements) and higher nestedness (richness changes) components of beta-diversity across all three biodiversity facets. Moreover, phylogenetic and functional turnover was lower and nestedness higher than random expectations based on taxonomic beta-diversity in regions that experienced large temperature change, reflecting phylogenetically and functionally selective processes in species replacement, extinction, and colonization during glacial-interglacial oscillations. Our results suggest that future human-driven climate change could cause local homogenization and reduction in taxonomic, phylogenetic, and functional diversity of angiosperm trees worldwide.

Intraspecific Leaf Trait Variation across and within Five Common Wine Grape Varieties.

Variability in traits forming the Leaf Economics Spectrum (LES) among and within crop species plays a key role in governing agroecosystem processes. However, studies evaluating the extent, causes, and consequences of within-species variation in LES traits for some of the world's most common crops remain limited. This study quantified variations in nine leaf traits measured across 90 vines of five common wine grape (Vitis vinifera L.) varieties at two growth stages (post-flowering and veraison). Grape traits in these varieties covary along an intraspecific LES, in patterns similar to those documented in wild plants. Across the five varieties evaluated here, high rates of photosynthesis (A) and leaf nitrogen (N) concentrations were coupled with low leaf mass per area (LMA), whereas the opposite suite of traits defined the "resource-conserving end" of this intraspecific LES in grape. Variety identity was the strongest predictor of leaf physiological (A) and morphological traits (i.e., leaf area and leaf mass), whereas leaf chemical traits and LMA were best explained by growth stage. All five varieties expressed greater resource-conserving trait syndromes (i.e., higher LMA, lower N, and lower Amass) later in the growing season. Traits related to leaf hydraulics, including instantaneous water-use efficiency (WUE), were unrelated to LES and other resource capture traits, and were better explained by spatial location. These results highlight the relative contributions of genetic, developmental, and phenotypic factors in structuring trait variation in the five wine grape varieties evaluated here, and point to a key role of domestication in governing trait relationships in the world's crops.

The influence of biofertilizers on leaf economics spectrum traits in a herbaceous crop.

Microbial inoculations or 'biofertilizers' represent novel contributions to sustainable agriculture. While belowground mechanisms surrounding how biofertilizers enhance crop production are well described, their role in aboveground trait expression remains less well explored. We quantified infraspecific variation in leaf economics spectrum (LES) traits in response to 10 biofertilizer treatments in basil (Ocimum basiclicum) cultivated under hydroponic conditions. Multiple physiological (i.e. maximum photosynthesis rates (A), dark respiration (R), and leaf-level light compensation points) and morphological (i.e. leaf mass per area (LMA) and leaf thickness) traits varied significantly across microbial treatments. Following treatments, basil plants differentiated from one another along an infraspecific LES, with certain plants expressing more resource-acquiring LES trait values (i.e. high A, R, leaf N, and low LMA), versus others that expressed the opposite suite of resource-conserving LES trait values. Infraspecific trait covariation largely matched LES patterns observed among plants globally. Bivariate and multivariate trait analyses further revealed that certain treatments-namely those including closely related Bacillus and Brevibacillus species strains-increased leaf resource capture traits such as A and leaf N. Biofertilizers influence plant performance through a role in moderating infraspecific leaf trait variation, thereby suggesting aboveground leaf traits may be used to diagnose optimal biofertilizer formulations in basil and other crops.

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.

National-scale changes in crop diversity through the Anthropocene.

Expansion of crops beyond their centres of domestication is a defining feature of the Anthropocene Epoch. This process has fundamentally altered the diversity of croplands, with likely consequences for the ecological functioning and socio-economic stability of agriculture under environmental change. While changes in crop diversity through the Anthropocene have been quantified at large spatial scales, the patterns, drivers, and consequences of change in crop diversity and biogeography at national-scales remains less explored. We use production data on 339 crops, grown in over 150 countries from 1961 to 2017, to quantify changes in country-level crop richness and evenness. Virtually all countries globally have experienced significant increases in crop richness since 1961, with the early 1980s marking a clear onset of a ~ 9-year period of increase in crop richness in countries worldwide. While these changes have increased the similarity of diversity of croplands among countries, only half of countries experienced increases in crop evenness through time. Ubiquitous increases in crop richness within nearly all countries between 1980 and 2000 are a unique biogeographical feature of the Anthropocene. At the same time, we detected opposing changes in crop evenness, and only modest signatures of increased homogenization of croplands among countries. Therefore context-dependent and, at least, national-scale assessments are needed to understand and predict how changes in crop diversity influence agricultural resistance and resilience to environmental change.

Carbon concentration in the world's trees across climatic gradients.

Wood carbon (C) concentration is a key wood trait that varies widely among tree species, but our understanding of the factors governing this trait is limited, despite reason to hypothesize that wood C varies systematically across environmental gradients. We compiled a novel database of 1145 geo-referenced wood C observations from 415 species, to elucidate climate correlates of wood C concentrations, and test if these relationships differ across tissue types and major taxonomic divisions (i.e. angiosperms vs gymnosperms). Climate variables, including mean annual temperature (MAT) and precipitation and temperature seasonality, are significantly correlated with wood C concentrations. Relationships between wood C and these variables differ across tissue types and taxonomic divisions, yet there is a negative relationship between wood C and MAT that exists across all tissues and species groups. Wood C concentrations in trees are influenced by climate, with experimental evidence (albeit scant) indicating that climate-driven changes in lignin concentrations likely govern these relationships. Our study presents among the first lines of evidence indicating that wood C concentrations are correlated with environmental conditions, thereby enhancing our understanding of the potential adaptive significance of wood C variation in trees.

Dimensions of invasiveness: Links between local abundance, geographic range size, and habitat breadth in Europe's alien and native floras.

Understanding drivers of success for alien species can inform on potential future invasions. Recent conceptual advances highlight that species may achieve invasiveness via performance along at least three distinct dimensions: 1) local abundance, 2) geographic range size, and 3) habitat breadth in naturalized distributions. Associations among these dimensions and the factors that determine success in each have yet to be assessed at large geographic scales. Here, we combine data from over one million vegetation plots covering the extent of Europe and its habitat diversity with databases on species' distributions, traits, and historical origins to provide a comprehensive assessment of invasiveness dimensions for the European alien seed plant flora. Invasiveness dimensions are linked in alien distributions, leading to a continuum from overall poor invaders to super invaders-abundant, widespread aliens that invade diverse habitats. This pattern echoes relationships among analogous dimensions measured for native European species. Success along invasiveness dimensions was associated with details of alien species' introduction histories: earlier introduction dates were positively associated with all three dimensions, and consistent with theory-based expectations, species originating from other continents, particularly acquisitive growth strategists, were among the most successful invaders in Europe. Despite general correlations among invasiveness dimensions, we identified habitats and traits associated with atypical patterns of success in only one or two dimensions-for example, the role of disturbed habitats in facilitating widespread specialists. We conclude that considering invasiveness within a multidimensional framework can provide insights into invasion processes while also informing general understanding of the dynamics of species distributions.

Carbon fractions in the world's dead wood.

A key uncertainty in quantifying dead wood carbon (C) stocks-which comprise ~8% of total forest C pools globally-is a lack of accurate dead wood C fractions (CFs) that are employed to convert dead woody biomass into C. Most C estimation protocols utilize a default dead wood CF of 50%, but live tree studies suggest this value is an over-estimate. Here, we compile and analyze a global database of dead wood CFs in trees, showing that dead wood CFs average 48.5% across forests, deviating significantly from 50%, and varying systematically among biomes, taxonomic divisions, tissue types, and decay classes. Utilizing data-driven dead wood CFs in tropical forests alone may correct systematic overestimates in dead wood C stocks of ~3.0 Pg C: an estimate approaching nearly the entire dead wood C pool in the temperate forest biome. We provide for the first time, robust empirical dead wood CFs to inform global forest C estimation.

The leaf economics spectrum's morning coffee: plant size-dependent changes in leaf traits and reproductive onset in a perennial tree crop.

BACKGROUND AND AIMS: Size-dependent changes in plant traits are an important source of intraspecific trait variation. However, there are few studies that have tested if leaf trait co-variation and/or trade-offs follow a within-genotype leaf economics spectrum (LES) related to plant size and reproductive onset. To our knowledge, there are no studies on any plant species that have tested whether or not the shape of a within-genotype LES that describes how traits covary across whole plant sizes, is the same as the shape of a within-genotype LES that represents environmentally driven trait plasticity. METHODS: We quantified size-dependent variation in eight leaf traits in a single coffee genotype (Coffea arabica var. Caturra) in managed agroecosystems with different environmental conditions (light and fertilization treatments), and evaluated these patterns with respect to reproductive onset. We also evaluated if trait covariation along a within-genotype plant-size LES differed from a within-genotype environmental LES defined with trait data from coffee growing in different environmental conditions. KEY RESULTS: Leaf economics traits related to resource acquisition - maximum photosynthetic rates (A) and mass-based leaf nitrogen (N) concentrations - declined linearly with plant size. Structural traits - leaf mass, leaf thickness, and leaf mass per unit area (LMA) - and leaf area increased with plant size beyond reproductive onset, then declined in larger plants. Three primary LES traits (mass-based A, leaf N and LMA) covaried across a within-genotype plant-size LES, with plants moving towards the 'resource-conserving' end of the LES as they grow larger; in coffee these patterns were nearly identical to a within-genotype environmental LES. CONCLUSIONS: Our results demonstrate that a plant-size LES exists within a single genotype. Our findings indicate that in managed agroecosystems where resource availability is high the role of reproductive onset in driving within-genotype trait variability, and the strength of covariation and trade-offs among LES traits, are less pronounced compared with plants in natural systems. The consistency in trait covariation in coffee along both plant-size and environmental LES axes indicates strong constraints on leaf form and function that exist within plant genotypes.

Accumulating crop functional trait data with citizen science.

Trait-based ecology is greatly informed by large datasets for the analyses of inter- and intraspecific trait variation (ITV) in plants. This is especially true in trait-based agricultural research where crop ITV is high, yet crop trait data remains limited. Based on farmer-led collections, we developed and evaluated the first citizen science plant trait initiative. Here we generated a dataset of eight leaf traits for a commercially important crop species (Daucus carota), sampled from two distinct regions in Canada, which is 25-fold larger than datasets available in existing trait databases. Citizen-collected trait data supported analyses addressing theoretical and applied questions related to (i) intraspecific trait dimensionality, (ii) the extent and drivers of ITV, and (iii) the sampling intensity needed to derive accurate trait values. Citizen science is a viable means to enhance functional trait data coverage across terrestrial ecosystems, and in doing so, can directly support theoretical and applied trait-based analyses of plants.

Regional and global shifts in crop diversity through the Anthropocene.

The Anthropocene epoch is partly defined by anthropogenic spread of crops beyond their centres of origin. At global scales, evidence indicates that species-level taxonomic diversity of crops being cultivated on large-scale agricultural lands has increased linearly over the past 50 years. Yet environmental and socio-economic differences support expectations that temporal changes in crop diversity vary across regions. Ecological theory also suggests that changes in crop taxonomic diversity may not necessarily reflect changes in the evolutionary diversity of crops. We used data from the Food and Agricultural Organization (FAO) of the United Nations to assess changes in crop taxonomic- and phylogenetic diversity across 22 subcontinental-scale regions from 1961-2014. We document certain broad consistencies across nearly all regions: i) little change in crop diversity from 1961 through to the late 1970s; followed by ii) a 10-year period of sharp diversification through the early 1980s; followed by iii) a "levelling-off" of crop diversification beginning in the early 1990s. However, the specific onset and duration of these distinct periods differs significantly across regions and are unrelated to agricultural expansion, indicating that unique policy or environmental conditions influence the crops being grown within a given region. Additionally, while the 1970s and 1980s are defined by region-scale increases in crop diversity this period marks the increasing dominance of a small number of crop species and lineages; a trend resulting in detectable increases in the similarity of crops being grown across regions. Broad similarities in the species-level taxonomic and phylogenetic diversity of crops being grown across regions, primarily at large industrial scales captured by FAO data, represent a unique feature of the Anthropocene epoch. Yet nuanced asymmetries in regional-scale trends suggest that environmental and socio-economic factors play a key role in shaping observed macro-ecological changes in the plant diversity on agricultural lands.

Intraspecific variation in soy across the leaf economics spectrum.

Background and Aims: Intraspecific trait variation (ITV) is an important dimension of plant ecological diversity, particularly in agroecosystems, where phenotypic ITV (within crop genotypes) is an important correlate of key agroecosystem processes including yield. There are few studies that have evaluated whether plants of the same genotype vary along well-defined axes of biological variation, such as the leaf economics spectrum (LES). There is even less information disentangling environmental and ontogenetic determinants of crop ITV along an intraspecific LES, and whether or not a plant's position along an intraspecific LES is correlated with reproductive output. Methods: We sought to capture the extent of phenotypic ITV within a single cultivar of soy (Glycine max) - the world's most commonly cultivated legume - using a data set of nine leaf traits measured on 402 leaves, sampled from 134 plants in both agroforestry and monoculture management systems, across three distinct whole-plant ontogenetic stages (while holding leaf age and canopy position stable). Key Results: Leaf traits covaried strongly along an intraspecific LES, in patterns that were largely statistically indistinguishable from the 'universal LES' observed across non-domesticated plants. Whole-plant ontogenetic stage explained the highest proportion of phenotypic ITV in LES traits, with plants progressively expressing more 'resource-conservative' LES syndromes throughout development. Within ontogenetic stages, leaf traits differed systematically across management systems, with plants growing in monoculture expressing more 'resource-conservative' trait syndromes: trends largely owing to an approximately ≥50% increases in leaf mass per area (LMA) in high-light monoculture vs. shaded agroforestry systems. Certain traits, particularly LMA, leaf area and maximum photosynthetic rates, correlated closely with plant-level reproductive output. Conclusions: Phenotypic ITV in soy is governed by constraints in trait trade-offs along an intraspecific LES, which in turn (1) underpins plant responses to managed environmental gradients, and (2) reflects shifts in plant functional biology and resource allocation that occur throughout whole-plant ontogeny.

Inter- and intraspecific variation in leaf economic traits in wheat and maize.

Leaf Economics Spectrum (LES) trait variation underpins multiple agroecological processes and many prominent crop yield models. While there are numerous independent studies assessing trait variation in crops, to date there have been no comprehensive assessments of intraspecific trait variation (ITV) in LES traits for wheat and maize: the world's most widespread crops. Using trait databases and peer-reviewed literature, we compiled over 700 records of specific leaf area (SLA), maximum photosynthetic rates (Amax) and leaf nitrogen (N) concentrations, for wheat and maize. We evaluated intraspecific LES trait variation, and intraspecific trait-environment relationships. While wheat and maize occupy the upper 90th percentile of LES trait values observed across a global species pool, ITV ranged widely across the LES in wheat and maize. Fertilization treatments had strong impacts on leaf N, while plant developmental stage (here standardized as the number of days since planting) had strong impacts on Amax; days since planting, N fertilization and irrigation all influenced SLA. When controlling for these factors, intraspecific responses to temperature and precipitation explained 39.4 and 43.7 % of the variation in Amax and SLA, respectively, but only 5.4 % of the variation in leaf N. Despite a long history of domestication in these species, ITV in wheat and maize among and within cultivars remains large. Intraspecific trait variation is a critical consideration to refine regional to global models of agroecosystem structure, function and food security. Considerable opportunities and benefits exist for consolidating a crop trait database for a wider range of domesticated plant species.

Intraspecific Trait Variation and Coordination: Root and Leaf Economics Spectra in Coffee across Environmental Gradients.

Hypotheses on the existence of a universal "Root Economics Spectrum" (RES) have received arguably the least attention of all trait spectra, despite the key role root trait variation plays in resource acquisition potential. There is growing interest in quantifying intraspecific trait variation (ITV) in plants, but there are few studies evaluating (i) the existence of an intraspecific RES within a plant species, or (ii) how a RES may be coordinated with other trait spectra within species, such as a leaf economics spectrum (LES). Using Coffea arabica (Rubiaceae) as a model species, we measured seven morphological and chemical traits of intact lateral roots, which were paired with information on four key LES traits. Field collections were completed across four nested levels of biological organization. The intraspecific trait coefficient of variation (cv) ranged from 25 to 87% with root diameter and specific root tip density showing the lowest and highest cv, respectively. Between 27 and 68% of root ITV was explained by site identity alone for five of the seven traits measured. A single principal component explained 56.2% of root trait covariation, with plants falling along a RES from resource acquiring to conserving traits. Multiple factor analysis revealed significant orthogonal relationships between root and leaf spectra. RES traits were strongly orthogonal with respect to LES traits, suggesting these traits vary independently from one another in response to environmental cues. This study provides among the first evidence that plants from the same species differentiate from one another along an intraspecific RES. We find that in one of the world's most widely cultivated crops, an intraspecific RES is orthogonal to an intraspecific LES, indicating that above and belowground responses of plants to managed (or natural) environmental gradients are likely to occur independently from one another.

Resistance and resilience of root fungal communities to water limitation in a temperate agroecosystem.

Understanding crop resilience to environmental stress is critical in predicting the consequences of global climate change for agricultural systems worldwide, but to date studies addressing crop resiliency have focused primarily on plant physiological and molecular responses. Arbuscular mycorrhizal fungi (AMF) form mutualisms with many crop species, and these relationships are key in mitigating the effects of abiotic stress in many agricultural systems. However, to date there is little research examining whether (1) fungal community structure in agroecosystems is resistant to changing environmental conditions, specifically water limitation and (2) resilience of fungal community structure is moderated by agricultural management systems, namely the integration of trees into cropping systems. Here, we address these uncertainties through a rainfall reduction field experiment that manipulated short-term water availability in a soybean-based (Glycine max L. Merr.) agroforest in Southern Ontario, Canada. We employed terminal restriction fragment length polymorphism analysis to determine the molecular diversity of both general fungal and AMF communities in soybean roots under no stress, stress (rainfall shelters added), and poststress (rainfall shelters removed). We found that general fungal and AMF communities sampled from soybean roots were resistant to rainfall reduction in a monoculture, but not in an agroforest. While AMF communities were unchanged after stress removal, general fungal communities were significantly different poststress in the agroforest, indicating a capacity for resiliency. Our study indicates that generalist fungi and AMF are responsive to changes in environmental conditions and that agroecosystem management plays a key role in the resistance and resilience of fungal communities to water limitation.

Wood nitrogen concentrations in tropical trees: phylogenetic patterns and ecological correlates.

In tropical and temperate trees, wood chemical traits are hypothesized to covary with species' life-history strategy along a 'wood economics spectrum' (WES), but evidence supporting these expected patterns remains scarce. Due to its role in nutrient storage, we hypothesize that wood nitrogen (N) concentration will covary along the WES, being higher in slow-growing species with high wood density (WD), and lower in fast-growing species with low WD. In order to test this hypothesis we quantified wood N concentrations in 59 Panamanian hardwood species, and used this dataset to examine ecological correlates and phylogenetic patterns of wood N. Wood N varied > 14-fold among species between 0.04 and 0.59%; closely related species were more similar in wood N than expected by chance. Wood N was positively correlated with WD, and negatively correlated with log-transformed relative growth rates, although these relationships were relatively weak. We found evidence for co-evolution between wood N and both WD and log-transformed mortality rates. Our study provides evidence that wood N covaries with tree life-history parameters, and that these patterns consistently co-evolve in tropical hardwoods. These results provide some support for the hypothesized WES, and suggest that wood is an increasingly important N pool through tropical forest succession.

Size-dependent changes in leaf and wood chemical traits in two Caribbean rainforest trees.

Tree functional traits and their link to patterns of growth and demography are central to informing trait-based analyses of forest communities, and mechanistic models of forest dynamics. However, few data are available on how functional traits in trees vary through ontogeny, particularly in tropical species; and less is known about how patterns of size-dependent changes in traits may differ across species of contrasting life-history strategies. Here we describe size-dependent variation in seven leaf functional traits and four wood chemical traits, in two Dominican rainforest tree species (Dacryodes excelsa Vahl. and Miconia mirabilis (Aubl.) L.O. Williams), ranging from small saplings to the largest canopy trees. With one exception, all traits showed pronounced variation with tree size (diameter at breast height, DBH). Leaf mass per area (LMA), thickness and tissue density increased monotonically with DBH in both species. Leaf area, leaf nitrogen (N) and carbon (C) : nitrogen (N) ratios also varied significantly with DBH; however, these patterns were unimodal, with peak trait values preceding the DBH at reproductive onset in both species. Size-dependent changes in leaf structural traits (LMA and leaf thickness) were generally similar in both species, while traits associated with leaf-level investment in C gain (leaf area, leaf C : N ratio) showed contrasting ontogenetic trends between species. Wood starch concentration varied with DBH in both species, also showing unimodal patterns with peaks preceding size at reproductive onset. Wood C concentration increased linearly with DBH in both species, though significantly only in M. mirabilis. Size-dependent patterns in wood chemical traits were similar between both species. Our data demonstrate pronounced variation in functional traits through tree ontogeny, probably due to a combination of environmental factors and shifts in resource allocation. Such ontogenetic variation is comparable in magnitude with interspecific variation, and so should be accounted for in trait-based studies of forest dynamics, structure and function.