Ian J. Wright

The worldwide leaf economics spectrum

Bringing together leaf trait data spanning 2,548 species and 175 sites we describe, for the first time at global scale, a universal spectrum of leaf economics consisting of key chemical, structural and physiological properties. The spectrum runs from quick to slow return on investments of nutrients and dry mass in leaves, and operates largely independently of growth form, plant functional type or biome. Categories along the spectrum would, in general, describe leaf economic variation at the global scale better than plant functional types, because functional types overlap substantially in their leaf traits. Overall, modulation of leaf traits and trait relationships by climate is surprisingly modest, although some striking and significant patterns can be seen. Reliable quantification of the leaf economics spectrum and its interaction with climate will prove valuable for modelling nutrient fluxes and vegetation boundaries under changing land-use and climate.

Bivariate line-fitting methods for allometry

Fitting a line to a bivariate dataset can be a deceptively complex problem, and there has been much debate on this issue in the literature. In this review, we describe for the practitioner the essential features of line‐fitting methods for estimating the relationship between two variables: what methods are commonly used, which method should be used when, and how to make inferences from these lines to answer common research questions.

Global convergence in the vulnerability of forests to drought.

Shifts in rainfall patterns and increasing temperatures associated with climate change are likely to cause widespread forest decline in regions where droughts are predicted to increase in duration and severity. One primary cause of productivity loss and plant mortality during drought is hydraulic failure. Drought stress creates trapped gas emboli in the water transport system, which reduces the ability of plants to supply water to leaves for photosynthetic gas exchange and can ultimately result in desiccation and mortality. At present we lack a clear picture of how thresholds to hydraulic failure vary across a broad range of species and environments, despite many individual experiments. Here we draw together published and unpublished data on the vulnerability of the transport system to drought-induced embolism for a large number of woody species, with a view to examining the likely consequences of climate change for forest biomes. We show that 70% of 226 forest species from 81 sites worldwide operate with narrow (<1 megapascal) hydraulic safety margins against injurious levels of drought stress and therefore potentially face long-term reductions in productivity and survival if temperature and aridity increase as predicted for many regions across the globe. Safety margins are largely independent of mean annual precipitation, showing that there is global convergence in the vulnerability of forests to drought, with all forest biomes equally vulnerable to hydraulic failure regardless of their current rainfall environment. These findings provide insight into why drought-induced forest decline is occurring not only in arid regions but also in wet forests not normally considered at drought risk.

New handbook for standardised measurement of plant functional traits worldwide

Plant functional traits are the features (morphological, physiological, phenological) that represent ecological strategies and determine how plants respond to environmental factors, affect other trophic levels and influence ecosystem properties. Variation in plant functional traits, and trait syndromes, has proven useful for tackling many important ecological questions at a range of scales, giving rise to a demand for standardised ways to measure ecologically meaningful plant traits. This line of research has been among the most fruitful avenues for understanding ecological and evolutionary patterns and processes. It also has the potential both to build a predictive set of local, regional and global relationships between plants and environment and to quantify a wide range of natural and human-driven processes, including changes in biodiversity, the impacts of species invasions, alterations in biogeochemical processes and vegetation–atmosphere interactions. The importance of these topics dictates the urgent need for more and better data, and increases the value of standardised protocols for quantifying trait variation of different species, in particular for traits with power to predict plant- and ecosystem-level processes, and for traits that can be measured relatively easily. Updated and expanded from the widely used previous version, this handbook retains the focus on clearly presented, widely applicable, step-by-step recipes, with a minimum of text on theory, and not only includes updated methods for the traits previously covered, but also introduces many new protocols for further traits. This new handbook has a better balance between whole-plant traits, leaf traits, root and stem traits and regenerative traits, and puts particular emphasis on traits important for predicting species’ effects on key ecosystem properties. We hope this new handbook becomes a standard companion in local and global efforts to learn about the responses and impacts of different plant species with respect to environmental changes in the present, past and future.

The Evolution of Plant Functional Variation: Traits, Spectra, and Strategies

Variation in plant functional traits results from evolutionary and environmental drivers that operate at a variety of different scales, which makes it a challenge to differentiate among them. In this article we describe patterns of functional trait variation and trait correlations within and among habitats in relation to several environmental and trade‐off axes. We then ask whether such patterns reflect natural selection and can be considered plant strategies. In so doing we highlight evidence that demonstrates that (1) patterns of trait variation across resource and environmental gradients (light, water, nutrients, and temperature) probably reflect adaptation, (2) plant trait variation typically involves multiple‐correlated traits that arise because of inevitable trade‐offs among traits and across levels of whole‐plant integration and that must be understood from a whole‐plant perspective, and (3) such adaptation may be globally generalizable for like conditions; i.e., the set of traits (collections of traits in syndromes) of taxa can be considered as “plant strategies.”

The evolutionary ecology of seed size.

Seed mass is a trait that occupies a pivotal position in the ecology of a species. It links the ecology of reproduction and seedling establishment with the ecology of vegetative growth, strategy sectors that are otherwise largely disconnected (Grime et al., 1988; Shipley et al., 1989; Leishman and Westoby, 1992). There is a startling diversity of shapes and sizes of seeds among the plant species of the world. Seeds range from the dust seeds of the Orchidaceae and some saprophytic and parasitic species (around 10 6 g), across ten orders of magnitude to the double coconut Lodoicea seychellarum (104 g) (Harper et al., 1970). Within species, seed size typically spans less than half an order of magnitude (about fourfold: Michaels et al., 1988). Most within-species variation occurs within plant rather than among plants or populations (Michaels et al., 1988; Obeso, 1993; Vaughton and Ramsey, 1998), indicating environmental effects during development rather than genetic differences between mothers. This chapter is concerned with the differences in seed size among species, and the consequences for vegetation dynamics and community composition. During the last 10–15 years, there has been considerable progress in the ecology of seed mass. Unlike many other areas of comparative plant ecology, we have substantial published information from several different scales and research styles. As well as field experiments and demographic studies with a few species at a time, we have simple experiments with larger numbers of species (ten to 50), quantification of seed mass and its correlates in whole-vegetation types (hundreds of species) and tests of consistency across different continents. The wide-scale quantification began as early as Salisbury (1942) and Baker (1972), but has been much added to and consolidated over the past 10 years (e.g. Mazer, 1989, 1990; Leishman and Westoby, 1994a; Leishman et al., 1995; Eriksson and Jakobsson, 1998). The work spanning large numbers of species is complementary to detailed experiments involving only a few species, giving a stronger sense of how widely the results from particular experiments can be generalized. Much of the literature examines how natural selection on seed size might be influenced by various environmental factors. In this context, it is at first glance surprising that seed size varies within communities across a remarkable five to six orders of magnitude (Leishman et al., 1995; Fig. 2.1). Further, there is strong overlap of seed-size distributions between quite different habitats. Within the temperate zone,

Global patterns of foliar nitrogen isotopes and their relationships with climate, mycorrhizal fungi, foliar nutrient concentrations, and nitrogen availability.

Ratios of nitrogen (N) isotopes in leaves could elucidate underlying patterns of N cycling across ecological gradients. To better understand global-scale patterns of N cycling, we compiled data on foliar N isotope ratios (delta(15)N), foliar N concentrations, mycorrhizal type and climate for over 11,000 plants worldwide. Arbuscular mycorrhizal, ectomycorrhizal, and ericoid mycorrhizal plants were depleted in foliar delta(15)N by 2 per thousand, 3.2 per thousand, 5.9 per thousand, respectively, relative to nonmycorrhizal plants. Foliar delta(15)N increased with decreasing mean annual precipitation and with increasing mean annual temperature (MAT) across sites with MAT >or= -0.5 degrees C, but was invariant with MAT across sites with MAT < -0.5 degrees C. In independent landscape-level to regional-level studies, foliar delta(15)N increased with increasing N availability; at the global scale, foliar delta(15)N increased with increasing foliar N concentrations and decreasing foliar phosphorus (P) concentrations. Together, these results suggest that warm, dry ecosystems have the highest N availability, while plants with high N concentrations, on average, occupy sites with higher N availability than plants with low N concentrations. Global-scale comparisons of other components of the N cycle are still required for better mechanistic understanding of the determinants of variation in foliar delta(15)N and ultimately global patterns in N cycling.

Functional traits and the growth?mortality trade-off in tropical trees

A trade-off between growth and mortality rates characterizes tree species in closed canopy forests. This trade-off is maintained by inherent differences among species and spatial variation in light availability caused by canopy-opening disturbances. We evaluated conditions under which the trade-off is expressed and relationships with four key functional traits for 103 tree species from Barro Colorado Island, Panama. The trade-off is strongest for saplings for growth rates of the fastest growing individuals and mortality rates of the slowest growing individuals (r2 = 0.69), intermediate for saplings for average growth rates and overall mortality rates (r2 = 0.46), and much weaker for large trees (r2 < or = 0.10). This parallels likely levels of spatial variation in light availability, which is greatest for fast- vs. slow-growing saplings and least for large trees with foliage in the forest canopy. Inherent attributes of species contributing to the trade-off include abilities to disperse, acquire resources, grow rapidly, and tolerate shade and other stresses. There is growing interest in the possibility that functional traits might provide insight into such ecological differences and a growing consensus that seed mass (SM), leaf mass per area (LMA), wood density (WD), and maximum height (H(max)) are key traits among forest trees. Seed mass, LMA, WD, and H(max) are predicted to be small for light-demanding species with rapid growth and mortality and large for shade-tolerant species with slow growth and mortality. Six of these trait-demographic rate predictions were realized for saplings; however, with the exception of WD, the relationships were weak (r2 < 0.1 for three and r2 < 0.2 for five of the six remaining relationships). The four traits together explained 43-44% of interspecific variation in species positions on the growth-mortality trade-off; however, WD alone accounted for > 80% of the explained variation and, after WD was included, LMA and H(max) made insignificant contributions. Virtually the full range of values of SM, LMA, and H(max) occurred at all positions on the growth-mortality trade-off. Although WD provides a promising start, a successful trait-based ecology of tropical forest trees will require consideration of additional traits.

Three keys to the radiation of angiosperms into freezing environments

Early flowering plants are thought to have been woody species restricted to warm habitats. This lineage has since radiated into almost every climate, with manifold growth forms. As angiosperms spread and climate changed, they evolved mechanisms to cope with episodic freezing. To explore the evolution of traits underpinning the ability to persist in freezing conditions, we assembled a large species-level database of growth habit (woody or herbaceous; 49,064 species), as well as leaf phenology (evergreen or deciduous), diameter of hydraulic conduits (that is, xylem vessels and tracheids) and climate occupancies (exposure to freezing). To model the evolution of species’ traits and climate occupancies, we combined these data with an unparalleled dated molecular phylogeny (32,223 species) for land plants. Here we show that woody clades successfully moved into freezing-prone environments by either possessing transport networks of small safe conduits and/or shutting down hydraulic function by dropping leaves during freezing. Herbaceous species largely avoided freezing periods by senescing cheaply constructed aboveground tissue. Growth habit has long been considered labile, but we find that growth habit was less labile than climate occupancy. Additionally, freezing environments were largely filled by lineages that had already become herbs or, when remaining woody, already had small conduits (that is, the trait evolved before the climate occupancy). By contrast, most deciduous woody lineages had an evolutionary shift to seasonally shedding their leaves only after exposure to freezing (that is, the climate occupancy evolved before the trait). For angiosperms to inhabit novel cold environments they had to gain new structural and functional trait solutions; our results suggest that many of these solutions were probably acquired before their foray into the cold.

Are functional traits good predictors of demographic rates? Evidence from five neotropical forests

A central goal of comparative plant ecology is to understand how functional traits vary among species and to what extent this variation has adaptive value. Here we evaluate relationships between four functional traits (seed volume, specific leaf area, wood density, and adult stature) and two demographic attributes (diameter growth and tree mortality) for large trees of 240 tree species from five Neotropical forests. We evaluate how these key functional traits are related to survival and growth and whether similar relationships between traits and demography hold across different tropical forests. There was a tendency for a trade-off between growth and survival across rain forest tree species. Wood density, seed volume, and adult stature were significant predictors of growth and/or mortality. Both growth and mortality rates declined with an increase in wood density. This is consistent with greater construction costs and greater resistance to stem damage for denser wood. Growth and mortality rates also declined as seed volume increased. This is consistent with an adaptive syndrome in which species tolerant of low resource availability (in this case shade-tolerant species) have large seeds to establish successfully and low inherent growth and mortality rates. Growth increased and mortality decreased with an increase in adult stature, because taller species have a greater access to light and longer life spans. Specific leaf area was, surprisingly, only modestly informative for the performance of large trees and had ambiguous relationships with growth and survival. Single traits accounted for 9-55% of the interspecific variation in growth and mortality rates at individual sites. Significant correlations with demographic rates tended to be similar across forests and for phylogenetically independent contrasts as well as for cross-species analyses that treated each species as an independent observation. In combination, the morphological traits explained 41% of the variation in growth rate and 54% of the variation in mortality rate, with wood density being the best predictor of growth and mortality. Relationships between functional traits and demographic rates were statistically similar across a wide range of Neotropical forests. The consistency of these results strongly suggests that tropical rain forest species face similar trade-offs in different sites and converge on similar sets of solutions.