Hiroko Kurokawa

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.

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.

Sustaining ecosystem functions in a changing world: a call for an integrated approach

With ever-increasing human pressure on ecosystems, it is critically important to predict how ecosystem functions will respond to such human-induced perturbations. We define perturbations as either changes to abiotic environment (e.g. eutrophication, climate change) that indirectly affects biota, or direct changes to biota (e.g. species introductions). While two lines of research in ecology, biodiversity-ecosystem function (BDEF) and ecological resilience (ER) research, have addressed this issue, both fields of research have nontrivial shortcomings in their abilities to address a wide range of realistic scenarios. We outline how an integrated research framework may foster a deeper understanding of the functional consequences of perturbations via simultaneous application of (i) process-based mechanistic predictions using trait-based approaches and (ii) detection of empirical patterns of functional changes along real perturbation gradients. In this context, the complexities of ecological interactions and evolutionary perspectives should be integrated into future research. Synthesis and applications. Management of human-impacted ecosystems can be guided most directly by understanding the response of ecosystem functions to controllable perturbations. In particular, we need to characterize the form of a wide range of perturbation-function relationships and to draw connections between those patterns and the underlying ecological processes. We anticipate that the integrated perspectives will also be helpful for managers to derive practical implications for management from academic literature. Management of human-impacted ecosystems can be guided most directly by understanding the response of ecosystem functions to controllable perturbations. In particular, we need to characterize the form of a wide range of perturbation-function relationships and to draw connections between those patterns and the underlying ecological processes. We anticipate that the integrated perspectives will also be helpful for managers to derive practical implications for management from academic literature.

Interspecific variation in the size-dependent resprouting ability of temperate woody species and its adaptive significance

Summary Resprouting of woody species after above-ground damage may help plants to persist longer at a given site and quickly reoccupy disturbed sites, thereby strongly influencing forest dynamics. Resprouting has been discussed from two adaptation perspectives: recovery from damage by catastrophic disturbance and survival in frequently disturbed shaded understorey. However, few studies have comprehensively dealt with both adaptation types to understand resprouting strategies. To understand the adaptive significance of resprouting, we assessed the size dependence of resprouting ability after stem clipping for 24 deciduous broad-leaved species, including shrubs, sub-canopy and canopy trees, in a cool-temperate forest in Japan. The community assembly includes species adapted to past catastrophic disturbances (e.g. fire, logging) and to stable forest with intermittent treefall (currently the dominant disturbance). We correlated resprouting ability with life-history strategies based on demographic parameters and plant functional traits, such as leaf mass per area (LMA), leaf toughness and wood density. All the studied species could resprout in juveniles, and resprouting ability increased as stump size increased. Most sub-canopy and canopy trees lost their ability to resprout after attaining a particular stump size, whereas shrub species retained the ability to resprout throughout their lifetimes. The relative growth rate, LMA and foliar nitrogen did not greatly influence the resprouting ability of a species. In contrast, species with smaller maximum size, lower leaf toughness and lower wood density had better juvenile resprouting ability. This better resprouting ability may have evolved because these characteristics make them more vulnerable to shaded understorey. However, species with larger maximum size and lower leaf toughness retained their ability to resprout to a larger size. Synthesis. A better resprouting ability is related to the ability to survive frequent disturbances, in juveniles, which are characteristics of both forest understorey and frequent fire or drought. To retain resprouting ability until grown seems to be an adaptation to survive infrequent large disturbances. Light-demanding species, which generally have better resprouting ability than shade-tolerants both in juveniles and adults, are adapted to disturbances of various scale and frequency; however, shade-tolerants could survive well in the understorey due to a combination of stronger physical defences and resprouting ability.

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.

Relationships between resprouting ability, species traits and resource allocation patterns in woody species in a temperate forest

Summary Many woody plants resprout to restore above-ground biomass after disturbances or to survive in stressful environments. Resprouting requires carbohydrate storage, but the general relationship between resource allocation patterns and resprouting ability remains unclear because it can be influenced by the disturbance regime to which species have adapted. We studied deciduous broadleaved trees that coexist in a Japanese cool-temperate forest to investigate the relationships among the biomass and total non-structural carbohydrate (TNC) allocation patterns of saplings, resprouting ability and functional traits. The study species comprised 16 single-stemmed species that only resprout when above-ground biomass loss occurs and eight multi-stemmed species that resprout regardless of whether above-ground damage occurs or not. Single-stemmed species with better juvenile resprouting ability had larger roots, whereas multi-stemmed species with better juvenile resprouting ability did not necessarily depend on below-ground reserves. Species that retain their ability to resprout until a larger size had a higher root TNC content as saplings, suggesting that they can survive major disturbances such as fire and coppicing by resprouting supported by TNC stored in their roots. Species with shade-tolerant traits (i.e. low foliar nitrogen indicating low photosynthetic capacity, high wood density indicating high defensive investment) had small below-ground TNC reserves irrespective of resprouting types. On the other hand, multi-stemmed species with high wood density and high LMA (indicating high photosynthetic capacity) had small above-ground TNC reserves. Contrary to our hypothesis, a species’ maximum size did not relate to the size of its below-ground reserves. By considering the differences in resprouting types, we suggest more complex control of resprouting than was formerly proposed. Variation in the resprouting ability of single-stemmed species was based on a trade-off between below-ground reserves for resprouting and shade-tolerant traits. However, multi-stemmed species can vigorously resprout irrespective of the size of its below-ground reserves. Their multi-stemmed architecture, well-defended wood, high photosynthetic capacity or large above-ground carbohydrate reserves seem to respectively contribute to their persistence. Such variation in the resprouting strategy based on a trade-off between shade tolerance and resource storage would promote species coexistence under a range of disturbance regimes and light environments.

Mapping local and global variability in plant trait distributions

Significance Currently, Earth system models (ESMs) represent variation in plant life through the presence of a small set of plant functional types (PFTs), each of which accounts for hundreds or thousands of species across thousands of vegetated grid cells on land. By expanding plant traits from a single mean value per PFT to a full distribution per PFT that varies among grid cells, the trait variation present in nature is restored and may be propagated to estimates of ecosystem processes. Indeed, critical ecosystem processes tend to depend on the full trait distribution, which therefore needs to be represented accurately. These maps reintroduce substantial local variation and will allow for a more accurate representation of the land surface in ESMs. 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 (Nm) and phosphorus (Pm), we characterize how traits vary within and among over 50,000 ∼50×50-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.

Context‐dependent changes in the functional composition of tree communities along successional gradients after land‐use change

Summary Land-use change can modify the functional composition of tree communities, which is an essential determinant of the ecosystem functions. The lack of consensus about the functional responses of tree communities to land-use change is a major uncertainty in the assessments of human impacts on terrestrial ecosystem functions. In this study, we applied a machine-learning method to a large data set consisting of 2574 tree communities across Japan to examine changes in the functional composition of tree communities after land-use change while considering contexts including successional trajectories, forest types and the presence of gymnosperms. Specifically, we hypothesized that functional changes along successional gradients after land-use change can be different in different contexts. Effects of two successional variables (stand age and basal area) on functional composition were highly significant throughout the study region. Changes in functional composition with changes in the two successional variables differed greatly, and the effects of basal area often outweighed those of stand age. Tree communities with small basal area were generally characterized by functional traits related to the resource-acquisitive strategy, that is short adult stature, low leaf mass per unit area, small seeds, low wood density and large leaves, especially when gymnosperms were excluded from the analysis. Decreasing basal area but not decreasing stand age often led to a considerable loss in functional diversity. Despite these general trends, functional changes along successional gradients after land-use change were not necessarily parallel and the opposite patterns were sometimes observed among forest types, traits and taxonomic groups. Synthesis. As a whole, our analyses demonstrate that the functional changes in tree communities after land-use change are highly evident in a given context but can be different under different contexts. These changes in functional composition can trigger variable changes in ecosystem functions such as carbon and nutrient cycling that depend on the context.

Ecological consequences through responses of plant and soil communities to changing winter climate

Community processes are now undergoing substantial reconfiguration because of climate change. Although the effects of climate change on ecosystems are currently a major concern, the issues tightly associated with winter climate change have been underrepresented. Given the importance of winter climate variables and events for determining the spatial distribution of communities and their phenological and physiological responses, and the functional roles of each species, all of which are expected to substantially influence community dis/re-assemble in the future, this review focuses on the ecological responses and consequences of terrestrial communities to changing winter climate. In particular, the effects on processes supported by biological interactions are largely undetermined. In this context, focusing on plant–soil feedback as a major interactive multi-system is worthwhile; these interactions can be disentangled through careful evaluation of the functional roles of organisms involved in the feedback (i.e., plants and soil organisms). The underlying mechanisms are indeed complex because direct (i.e., changes in physical conditions) and indirect pathways (i.e., plant-mediated influences on soil-organisms and vice versa) from winter climate change influence the functionality of future ecosystems. To face these issues, the framework of response–effect-traits deserves research priority since this can define community re-organization as the accumulated responses of individual species, which determines the stability and performance of ecosystem functioning. Thus, research that quantifies functional responses and roles of organisms under a changing climate will continue to be essential for the issues of winter climate change, which may become more serious and significant in the near future.

Simulation of natural capital and ecosystem services in a watershed in Northern Japan focusing on the future underuse of nature: by linking forest landscape model and social scenarios

A quantitative scenario approach to compare the future state of natural capital and ecosystem services (ESs) plays a key role in facilitating decision-making for the sustainable management of landscapes. In Japan, the shrinking and aging population will likely lead to a situation of underuse of natural resources, resulting in rewilding of terrestrial ecosystems. This study conducted a quantitative scenario analysis of natural capital and ESs by linking model and social scenarios on a local scale. The case study area was the Bekanbeushi River Watershed in Northern Japan. LANDIS-II model (a forest landscape model) was used to simulate the vegetation dynamics in species composition, age structure, and biomass considering impacts of forest and pasture land management. Four “population distribution” and “capital preference” scenarios were translated into forest and pasture land management. The population distribution and capital preference assumptions resulted in different consequences for natural capital and ESs. The population distribution affected the spatial allocation of abandoned pasture land and level of isolation of managed pasture land. The capital preference assumptions largely affected the consequences for ESs. Finally, these simulation results demonstrated the capacity to feed quantitative information to the narrative scenarios. Our process-based approach provides insight into the relationships among social drivers, ecological processes, and the consequences that will affect natural capital and ESs, which can contribute to decision-making and sustainability design of regions, which may face issues associated with underuse in the future.