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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.\"

Forests cover 30% of the terrestrial Earth surface and are a major component of the global carbon (C) cycle. Humans have doubled the amount of global reactive nitrogen (N), increasing deposition of N onto forests worldwide. However, other global changes—especially climate change and elevated atmospheric carbon dioxide concentrations—are increasing demand for N, the element limiting primary productivity in temperate forests, which could be reducing N availability. To determine the long-term, integrated effects of global changes on forest N cycling, we measured stable N isotopes in wood, a proxy for N supply relative to demand, on large spatial and temporal scales across the continental U.S.A. Here, we show that forest N availability has generally declined across much of the U.S. since at least 1850 C.E. with cool, wet forests demonstrating the greatest declines. Across sites, recent trajectories of N availability were independent of recent atmospheric N deposition rates, implying a minor role for modern N deposition on the trajectory of N status of North American forests. Our results demonstrate that current trends of global changes are likely to be consistent with forest oligotrophication into the foreseeable future, further constraining forest C fixation and potentially storage.

1. Our goal was to determine the relationships among ecophysiological, whole-plant and ecosystem traits of a wide variety of grassland species grown under field conditions in the long term. We measured 87 traits for 33 species (32 perennial, one annual) grown in monoculture for 5 years on sandy soils, and determined the relationship among traits and their correspondence with current functional classifications. 2. Among non-legumes, species that produced and maintained large amounts of biomass had tough, low-activity leaves and roots, high root: shoot ratios, and low extractable inorganic nitrogen and N mineralization in their soils. The set of correlations among the functional traits of fine roots for non-legumes parallels the set of correlations for leaf functional traits. Low-N species maintained greater biomass than high-N species, more by producing tissues with low N concentrations and greater longevity than by acquiring more N. Greater relative production below ground, and the production of long-lived below-ground structures, were both important in determining the high root: shoot ratio of species. 3. For legumes, N2fixation not only led to greater above-ground biomass production, but also was associated with low fine root production; greater relative production of stem biomass; and accelerated ecosystem N cycling compared to non-legumes. 4. The measured traits, as condensed via principal components analysis, differentiated the 32 species into groups that corresponded with a common grassland functional classification scheme (C3grasses, C4grasses, forbs, legumes, woody species) as well as an alternative, continuous approach. For all traits, species can be arrayed well along two continuous axes. The first axis separates cool-season and warm-season legumes; the second low-N and high-N non-legumes. 5. These continuous classifications show the generality of the two strategies for dealing with low nitrogen availability (N2fixation and the low-N suite of traits) and extends the strategies to span organ-level traits to ecosystem processes including roots, whole-plant patterns of productivity, and nutrient cycling. The correlations of traits among species will also be useful in predicting a large number of important parameters associated with plant growth from the measurement of a few, key traits.

$\\bullet$ Here, we tested hypothesized relationships among leaf and fine root traits of grass, forb, legume, and woody plant species of a savannah community. $\\bullet$ CO2 exchange rates, structural traits, chemistry, and longevity were measured in tissues of 39 species grown in long-term monocultures. $\\bullet$ Across species, respiration rates of leaves and fine roots exhibited a common regression relationship with tissue nitrogen (N) concentration, although legumes had lower rates at comparable N concentrations. Respiration rates and N concentration declined with increasing longevity of leaves and roots. Species rankings of leaf and fine-root N and longevity were correlated, but not specific leaf area and specific root length. The C3 and C4 grasses had lower N concentrations than forbs and legumes, but higher photosynthesis rates across a similar range of leaf N. $\\bullet$ Despite contrasting photosynthetic pathways and N2-fixing ability among these species, concordance in above- and below-ground traits was evident in comparable rankings in leaf and root longevity, N and respiration rates, which is evidence of a common leaf and root trait syndrome linking traits to effects on plant and ecosystem processes.

For 76 annual, biennial, and perennial species common in the grasslands of central Minnesota, USA, we determined the patterns of correlations among seven organ-level traits (specific leaf area, leaf thickness, leaf tissue density, leaf angle, specific root length, average fine root diameter, and fine root tissue density) and their relationships with two traits relating to growth form (whether species existed for part of the growing season in basal, non-caulescent form and whether species were rhizomatous or not). The first correlation of traits showed that grasses had thin, dense leaves and thin roots while forbs had thick, low-density leaves and thick roots without any significant differences in growth form or life history. The second correlation of traits showed a gradient of species from those with high-density roots and high-density erect leaves to species with low-density roots and low-density leaves that were held parallel to the ground. High tissue density species were more likely to exist as a basal rosette for part of the season, were less likely to be rhizomatous, and less likely to be annuals. We examined the relationships between the two axes that represent the correlations of traits and previously collected data on the relative abundance of species across gradients of nitrogen addition and disturbance. Grasses were generally more abundant than forbs and the relative abundance of grasses and forbs did not change with increasing nitrogen addition or soil disturbance. High tissue density species became less common as fertility and disturbance increased.

Humans have drastically altered the global nitrogen (N) cycle, and these alterations have begun to affect a variety of ecosystems. In North America, N deposition rates are highest in the central US, yet there are few studies that examine whether N availability has been increasing to different tree species in the forests of the region. To determine the species-specific trajectories of N availability in secondary temperate forests experiencing high N deposition, we measured the N concentrations and composition of stable N isotopes in wood of four tree species from six hardwood forest remnants in northern Indiana, USA. Annual nitrogen deposition rates averaged 5.8 kg ha−1 from 2000 to 2008 in this region. On average, wood δ15N values in Quercus alba have been increasing steadily over the past 100 years. In contrast, wood δ15N values have been declining in three other hardwood species – Acer saccharum, Carya ovata, and Fagus grandifolia – over the same time period. The species-specific trends suggest a change in the partitioning of ammonium and nitrate among species, due to an increase in nitrification rates over time. With no apparent net change in wood δ15N over the past century at the stand level, there is currently little evidence for consistent trends in stand-level N availability over time in the Indiana forests.

A universal challenge in methodology used to study the ecology, conservation and evolutionary biology of migratory species is the quantification of connectivity among breeding, wintering and stopover sites. For the avian Eurasian-Afrotropical migratory system, knowledge of geographical wintering areas used by migrants that breed in Europe remains deficient, despite the advent of satellite transmitters and geolocators. Here we explored the use of theoretical plant δ 13 C and δ 15 N landscape distributions coupled with δ 2 H hydrologic models to construct multi-isotopic avian foodweb clusters for Africa. The cluster analysis identified four distinct regions of Africa based on all three isotopes ( 13 C, 2 H, 15 N), and five regions based only on 13 C and 15 N. We applied known isotopic diet-tissue discrimination factors to map equivalent feather isotopic clusters for Africa. The validity of these feather isotopic clusters was tested by examining how well known- and unknown-origin species were placed in regions of Africa using previously published feather isotope data. The success of this multi-isotopic cluster model depended upon the species of interest and additionally on how well potential winter molt origins in Africa were constrained by prior information. Ground-truthing data suggested this approach will be useful for first-order approximation of overwintering regions for Afrotropical migrants and will be improved as our understanding of the nature of isoscapes for Africa is refined.

The majority of variation in six traits critical to the growth, survival and reproduction of plant species is thought to be organised along just two dimensions, corresponding to strategies of plant size and resource acquisition. However, it is unknown whether global plant trait relationships extend to climatic extremes, and if these interspecific relationships are confounded by trait variation within species. We test whether trait relationships extend to the cold extremes of life on Earth using the largest database of tundra plant traits yet compiled. We show that tundra plants demonstrate remarkably similar resource economic traits, but not size traits, compared to global distributions, and exhibit the same two dimensions of trait variation. Three quarters of trait variation occurs among species, mirroring global estimates of interspecific trait variation. Plant trait relationships are thus generalizable to the edge of global trait-space, informing prediction of plant community change in a warming world.

Across 30 grassland sites in New Zealand that ranged from native alpine grasslands to low elevation improved pastures, there were consistent patterns of leaf and root traits and significant differences between native and non-native grasses. Plants of high altitude sites have low N concentrations in both their leaves and roots, have thick leaves and roots, yet no differences in tissue density or photosynthetic water use efficiency when compared to plants of low altitude sites. Both the leaves and roots of the low altitude plants were enriched in15N relative to the plants of higher altitude, indicating that the low-N set of traits is associated with a more closed N cycle at high altitude. A second independent set of correlations shows that plants of wetter habitats have lower photosynthetic water use efficiency (more negative$\\partial {}^{13}\\text{C}$) and lower leaf and root tissue density than the plants of drier sites. For both leaves and roots, plants of native species consistently had traits associated with lower resource availability: lower N concentrations, denser tissues, more negative$\\partial {}^{15}\\text{N}$, and more positive$\\partial {}^{13}\\text{C}$than non-native species. If root %N is correlated with root longevity as has been shown in other systems, root longevity may be able to be predicted from simple measurements of leaf %N, though a hysteresis in the relationship between leaf and root N concentrations may make prediction of high longevity roots difficult.