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.

The contribution of photodegradation to litter decomposition in a temperate forest gap and understorey.

Litter decomposition determines carbon (C) backflow to the atmosphere and ecosystem nutrient cycling. Although sunlight provides the indispensable energy for terrestrial biogeochemical processes, the role of photodegradation in decomposition has been relatively neglected in productive mesic ecosystems. To quantify the effects of this variation, we conducted a factorial experiment in the understorey of a temperate deciduous forest and an adjacent gap, using spectral-attenuation-filter treatments. Exposure to the full spectrum of sunlight increased decay rates by nearly 120% and the effect of blue light contributed 75% of this increase. Scaled-up to the whole forest ecosystem, this translates to 13% loss of leaf-litter C through photodegradation over the year of our study for a scenario of 20% gap. Irrespective of the spectral composition, herbaceous and shrub litter lost mass faster than tree litter, with photodegradation contributing the most to surface litter decomposition in forest canopy gaps. Across species, the initial litter lignin and polyphenolic contents predicted photodegradation by blue light and ultraviolet B (UV-B) radiation, respectively. We concluded that photodegradation, modulated by litter quality, is an important driver of decomposition, not just in arid areas, but also in mesic ecosystems such as temperate deciduous forests following gap opening.

Leaf density and chemical composition explain variation in leaf mass area with spectral composition among 11 widespread forbs in a common garden.

Leaf mass per area (LMA) is a key leaf functional trait correlated with plant strategies dictating morphology, physiology, and biochemistry. Although sunlight is generally accepted as a dominant factor driving LMA, the contribution of each spectral region of sunlight in shaping LMA is poorly understood. In the present study, we grew 11 widespread forb species in a common garden and dissected the traits underpinning differences in LMA, such as its morphological components (leaf density [LD] and leaf thickness [LT]), macroelement, and metabolite composition under five spectral-attenuation treatments: (1) transmitting c. 95% of the whole solar spectrum (> 280 nm), (2) attenuating ultraviolet-B radiation (UV-B), (3) attenuating both UV-A and UV-B radiation, (4) attenuating UV radiation and blue light, (5) attenuating UV radiation, blue, and green light. We found that LMA, LD, and chemical traits varied significantly across species depending on spectral treatments. LMA was significantly increased by UV-B radiation and green light, while LD was increased by UV-A but decreased by blue light. LMA positively correlated with LD across treatments but was only weakly related to LT, suggesting that LD was a better determinate of LMA for this specific treatment. Regarding leaf elemental and metabolite composition, carbon, nitrogen, and total phenolics were all positively correlated with LMA, whereas lignin, non-structural carbohydrates, and soluble sugars had negative relationships with LMA. These trends imply a tradeoff between biomass allocation to structural and metabolically functional components. In conclusion, sunlight can spectrally drive LMA mainly through modifying functional and structural support.

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.

Soil fertility and disturbance interact to drive contrasting responses of co-occurring native and nonnative species.

Some plant functional groups such as nonnative invasive and nitrogen (N)-fixing plants are widely thought to have consistent, coordinated differences in their functional traits relative to other groups such as native and non -N-fixing plants. Recent evidence suggests that these trait differences between groups can be context dependent, varying with environmental factors such as resource availability and disturbance. However, many previous comparisons among plant groups differing in invasion status have not standardized growth form between groups or have compared species that do not co-occur, which could result in invasion status per se being confounded with other factors. We determined growth and leaf functional trait responses of 20 co-occurring woody species, that is, five species within each of four functional groups (native N-fixers, native non -N-fixers, nonnative [invasive] N-fixers and nonnative [invasive] non-N-fixers), to factorial combinations of soil fertility and defoliation treatments in a mesocosm experiment to test each of two hypotheses. First, we hypothesized that nonnative invasive and N-fixing species will have functional traits associated with rapid resource acquisition whereas natives and non -N-fixing species will have traits linked to resource conservation. Second, we hypothesized that plant growth and leaf traits of nonnative and N-fixing species will be more strongly influenced by environmental factors (i.e., soil fertility and disturbance) than will natives and non-N-fixers. Plant growth, foliar nutrients, and leaf structural traits varied among plant functional groups in a manner consistent with our first hypothesis. Support for our second hypothesis was mixed; origin (native vs. nonnative) and soil fertility rarely interacted to determine plant growth or variation in leaf traits whereas interactions involving N-fixing ability and soil fertility were common. Further, there were no consistent interactive effects between plant groupings and disturbance. Our results demonstrate that variation in growth and functional traits among plant species were driven primarily by the relatively large responses of nonnative N-fixers to soil fertility, rather than by consistent differences between other plant functional groups. These findings highlight the importance of resource availability in determining trait or performance differences among plant functional groups, and provide insights into the assembly of plant functional traits in novel communities of co-occurring native and nonnative species.

Efficacy of generic allometric equations for estimating biomass: a test in Japanese natural forests.

Accurate estimation of tree and forest biomass is key to evaluating forest ecosystem functions and the global carbon cycle. Allometric equations that estimate tree biomass from a set of predictors, such as stem diameter and tree height, are commonly used. Most allometric equations are site specific, usually developed from a small number of trees harvested in a small area, and are either species specific or ignore interspecific differences in allometry. Due to lack of site-specific allometries, local equations are often applied to sites for which they were not originally developed (foreign sites), sometimes leading to large errors in biomass estimates. In this study, we developed generic allometric equations for aboveground biomass and component (stem, branch, leaf, and root) biomass using large, compiled data sets of 1203 harvested trees belonging to 102 species (60 deciduous angiosperm, 32 evergreen angiosperm, and 10 evergreen gymnosperm species) from 70 boreal, temperate, and subtropical natural forests in Japan. The best generic equations provided better biomass estimates than did local equations that were applied to foreign sites. The best generic equations included explanatory variables that represent interspecific differences in allometry in addition to stem diameter, reducing error by 4-12% compared to the generic equations that did not include the interspecific difference. Different explanatory variables were selected for different components. For aboveground and stem biomass, the best generic equations had species-specific wood specific gravity as an explanatory variable. For branch, leaf, and root biomass, the best equations had functional types (deciduous angiosperm, evergreen angiosperm, and evergreen gymnosperm) instead of functional traits (wood specific gravity or leaf mass per area), suggesting importance of other traits in addition to these traits, such as canopy and root architecture. Inclusion of tree height in addition to stem diameter improved the performance of the generic equation only for stem biomass and had no apparent effect on aboveground, branch, leaf, and root biomass at the site level. The development of a generic allometric equation taking account of interspecific differences is an effective approach for accurately estimating aboveground and component biomass in boreal, temperate, and subtropical natural forests.

Masting is uncommon in trees that depend on mutualist dispersers in the context of global climate and fertility gradients.

The benefits of masting (volatile, quasi-synchronous seed production at lagged intervals) include satiation of seed predators, but these benefits come with a cost to mutualist pollen and seed dispersers. If the evolution of masting represents a balance between these benefits and costs, we expect mast avoidance in species that are heavily reliant on mutualist dispersers. These effects play out in the context of variable climate and site fertility among species that vary widely in nutrient demand. Meta-analyses of published data have focused on variation at the population scale, thus omitting periodicity within trees and synchronicity between trees. From raw data on 12 million tree-years worldwide, we quantified three components of masting that have not previously been analysed together: (i) volatility, defined as the frequency-weighted year-to-year variation; (ii) periodicity, representing the lag between high-seed years; and (iii) synchronicity, indicating the tree-to-tree correlation. Results show that mast avoidance (low volatility and low synchronicity) by species dependent on mutualist dispersers explains more variation than any other effect. Nutrient-demanding species have low volatility, and species that are most common on nutrient-rich and warm/wet sites exhibit short periods. The prevalence of masting in cold/dry sites coincides with climatic conditions where dependence on vertebrate dispersers is less common than in the wet tropics. Mutualist dispersers neutralize the benefits of masting for predator satiation, further balancing the effects of climate, site fertility and nutrient demands.

Canopy structure and phenology modulate the impacts of solar radiation on C and N dynamics during litter decomposition in a temperate forest.

Decomposition of plant organic matter plays a key role in the terrestrial biogeochemical cycles. Sunlight has recently been identified as an important contributor to carbon [C] turnover through photodegradation, accelerating decomposition even in forest ecosystems where understorey solar irradiance remains relatively low. However, it is uncertain how C and nutrients dynamics respond to fluctuations in solar spectral irradiance caused by canopy structure (understorey vs. gaps) and season (open vs. closed canopy phenology). Spectral-attenuation treatments were used to compare litter decomposition over eight months, covering canopy phenology, in a temperate deciduous forest and an adjacent gap. Exposure to the full spectrum of sunlight increased the loss of litter C and lignin by 75% and 64% in the forest gap, and blue light was responsible for respectively 27% and 42% of that loss. Whereas in the understorey, C and lignin loss were similar among spectral-attenuation treatments over the experimental period, except prior to and during spring canopy flush when exposure to the full spectrum of sunlight promoted C loss by 15% overall, 80% of which was attributable to ultraviolet-B (UV-B) radiation. Nitrogen [N] was immobilized in the understorey during canopy flush before the canopy completely closed but N was swiftly released during canopy leaf-fall. Our study suggests that blue-driven photodegradation plays an important role in lignin decomposition and N dynamics in canopy gaps, whereas seasonal canopy phenology affecting sunlight reaching the forest floor drastically changes patterns of C and N in litter during decomposition. Hence, including sunlight dynamics driven by canopy structure and phenology would improve estimates of biogeochemical cycling in forests responding to changes in climate and land-use.

Global patterns of leaf mechanical properties.

Leaf mechanical properties strongly influence leaf lifespan, plant-herbivore interactions, litter decomposition and nutrient cycling, but global patterns in their interspecific variation and underlying mechanisms remain poorly understood. We synthesize data across the three major measurement methods, permitting the first global analyses of leaf mechanics and associated traits, for 2819 species from 90 sites worldwide. Key measures of leaf mechanical resistance varied c. 500-800-fold among species. Contrary to a long-standing hypothesis, tropical leaves were not mechanically more resistant than temperate leaves. Leaf mechanical resistance was modestly related to rainfall and local light environment. By partitioning leaf mechanical resistance into three different components we discovered that toughness per density contributed a surprisingly large fraction to variation in mechanical resistance, larger than the fractions contributed by lamina thickness and tissue density. Higher toughness per density was associated with long leaf lifespan especially in forest understory. Seldom appreciated in the past, toughness per density is a key factor in leaf mechanical resistance, which itself influences plant-animal interactions and ecosystem functions across the globe.

Factors Influencing Leaf- and Root-Associated Communities of Bacteria and Fungi Across 33 Plant Orders in a Grassland.

In terrestrial ecosystems, plants interact with diverse taxonomic groups of bacteria and fungi in the phyllosphere and rhizosphere. Although recent studies based on high-throughput DNA sequencing have drastically increased our understanding of plant-associated microbiomes, we still have limited knowledge of how plant species in a species-rich community differ in their leaf and root microbiome compositions. In a cool-temperate semi-natural grassland in Japan, we compared leaf- and root-associated microbiomes across 137 plant species belonging to 33 plant orders. Based on the whole-microbiome inventory data, we analyzed how sampling season as well as the taxonomy, nativeness (native or alien), lifeform (herbaceous or woody), and mycorrhizal type of host plants could contribute to variation in microbiome compositions among co-occurring plant species. The data also allowed us to explore prokaryote and fungal lineages showing preferences for specific host characteristics. The list of microbial taxa showing significant host preferences involved those potentially having some impacts on survival, growth, or environmental resistance of host plants. Overall, this study provides a platform for understanding how plant and microbial communities are linked with each other at the ecosystem level.

Plant species traits are the predominant control on litter decomposition rates within biomes worldwide.

Worldwide decomposition rates depend both on climate and the legacy of plant functional traits as litter quality. To quantify the degree to which functional differentiation among species affects their litter decomposition rates, we brought together leaf trait and litter mass loss data for 818 species from 66 decomposition experiments on six continents. We show that: (i) the magnitude of species-driven differences is much larger than previously thought and greater than climate-driven variation; (ii) the decomposability of a species' litter is consistently correlated with that species' ecological strategy within different ecosystems globally, representing a new connection between whole plant carbon strategy and biogeochemical cycling. This connection between plant strategies and decomposability is crucial for both understanding vegetation-soil feedbacks, and for improving forecasts of the global carbon cycle.

Leaf herbivory and decomposability in a Malaysian tropical rain forest.

There is accumulating evidence that similar suites of plant traits may affect leaf palatability and leaf litter decomposability. However, the possible association between leaf herbivory and litter decomposition rates across species in species-diverse natural ecosystems such as tropical rain forests remains unexplored, despite its importance in estimating the herbivory effects on carbon and nutrient cycling of ecosystems. We found no strong association between leaf herbivory and litter decomposition rates across 40 tree species in a Malaysian tropical rain forest, even though the leaf and litter traits were tightly correlated. This is because the leaf and litter traits related to herbivory and decomposition rates in the field were inconsistent. Leaf toughness accounted for only a small part of the variation in the herbivory rate, whereas a number of litter traits (the leaf mass per area, lignin to nitrogen ratio, and condensed tannin concentration) accurately predicted the decomposition rate across species. These results suggest that herbivory rate across species may not be strongly related to single leaf traits, probably because plant-herbivore interactions in tropical rain forests are highly diverse; on the other hand, plant-decomposer interactions are less specific and can be governed by litter chemicals. We also investigated two factors, phylogeny and tree functional types, that could affect the relationship between herbivory and decomposition across species. Phylogenetic relatedness among the species did not affect the relationship between herbivory and decomposition. In contrast, when the plants were segregated according to their leaf emergence pattern, we found a significant positive relationship between herbivory and decomposition rates for continuous-leafing species. In these species, the condensed tannin to N ratios in leaves and litter were related to herbivory and decomposition rates, respectively. However, we did not observe a similar trend for synchronous-leafing species. These results suggest that the relationship between herbivory and decomposition may be more greatly affected by functional types than by phylogenetic relatedness among species. In conclusion, our results suggest that well-defended leaves are not necessarily less decomposable litter in a tropical rain forest community, implying that herbivory may not generate positive feedback for carbon and nutrient cycling in this type of ecosystem.

Allocation to defense or growth in dipterocarp forest seedlings in Borneo.

We quantified the allocation of net production to plant secondary metabolites (especially condensed tannins and lignins) to evaluate the investment into defense mechanisms of three tropical forest canopy species in Borneo vis-à-vis the resource availability hypothesis. In particular, we focused on Borneo ironwood (Eusideroxylon zwageri, Lauraceae), which seemed to employ an extreme defensive strategy. The wood of this species is extremely durable and has a high specific gravity with a very slow growth rate. The allocation to defense by Borneo ironwood was compared to two emergent species of Dipterocarpaceae, the dominant family in this forest community. We conducted shade-house experiments on seedlings under four controlled conditions (two light levels x two nutrient levels) and showed that the growth rate of E. zwageri was much lower than those of the other two species, and it allocated more of its net production to leaves and roots than to stems. The concentrations of condensed tannins and lignins were very high in the leaves and stems of this species, at about 20 and 30%, respectively. In total, E. zwageri allocated a maximum of about 35% of its net production to defensive substances (i.e., condensed tannins + lignins). In contrast, the two dipterocarp species allocated about 20-25% of their net production to defensive substances. The condensed tannins in E. zwageri help to prolong the lifespan of the leaves, and the lignins in the stems enhance the durability of the wood. Thus, although E. zwageri grows very slowly, the allocation to defensive substances seems to be an advantageous strategy for survival under dark conditions.