Sensitivity of leaf size and shape to climate within Acer rubrum and Quercus kelloggii

* Variation in the size and shape (physiognomy) of leaves has long been correlated to climate, and paleobotanists have used these correlations to reconstruct paleo-climate. Most studies focus on site-level means of largely nonoverlapping species sets. The sensitivity of leaf shape to climate within species is poorly known, which limits our general understanding of leaf-climate relationships and the value of intraspecific patterns for paleoclimate reconstructions. * The leaf physiognomy of two species whose native North American ranges span large climatic gradients (Acer rubrum and Quercus kelloggii) was quantified and correlated to mean annual temperature (MAT). Quercus kelloggii was sampled across a wide elevation range, but A. rubrum was sampled in strictly lowland areas. * Within A. rubrum, leaf shape correlates with MAT in a manner that is largely consistent with previous site-level studies; leaves from cold climates are toothier and more highly dissected. By contrast, Q. kelloggii is largely insensitive to MAT; instead, windy conditions with ample plant-available water may explain the preponderance of small teeth at high elevation sites, independent of MAT. * This study highlights the strong correspondence between leaf form and climate within some species, and demonstrates that intraspecific patterns may contribute useful information towards reconstructing paleoclimate.     

Paleotemperature proxies from leaf fossils reinterpreted in light of evolutionary history

Present-day correlations between leaf physiognomic traits (shape and size) and climate are widely used to estimate paleoclimate using fossil floras. For example, leaf-margin analysis estimates paleotemperature using the modern relation of mean annual temperature (MAT) and the site-proportion of untoothed-leaf species (NT). This uniformitarian approach should provide accurate paleoclimate reconstructions under the core assumption that leaf-trait variation principally results from adaptive environmental convergence, and because variation is thus largely independent of phylogeny it should be constant through geologic time. Although much research acknowledges and investigates possible pitfalls in paleoclimate estimation based on leaf physiognomy, the core assumption has never been explicitly tested in a phylogenetic comparative framework. Combining an extant dataset of 21 leaf traits and temperature with a phylogenetic hypothesis for 569 species-site pairs at 17 sites, we found varying amounts of non-random phylogenetic signal in all traits. Phylogenetic vs. standard regressions generally support prevailing ideas that leaf-traits are adaptively responding to temperature, but wider confidence intervals, and shifts in slope and intercept, indicate an overall reduced ability to predict climate precisely due to the non-random phylogenetic signal. Notably, the modern-day relation of proportion of untoothed taxa with mean annual temperature (NT-MAT), central in paleotemperature inference, was greatly modified and reduced, indicating that the modern correlation primarily results from biogeographic history. Importantly, some tooth traits, such as number of teeth, had similar or steeper slopes after taking phylogeny into account, suggesting that leaf teeth display a pattern of exaptive evolution in higher latitudes. This study shows that the assumption of convergence required for precise, quantitative temperature estimates using present-day leaf traits is not supported by empirical evidence, and thus we have very low confidence in previously published, numerical paleotemperature estimates. However, interpreting qualitative changes in paleotemperature remains warranted, given certain conditions such as stratigraphically closely-spaced samples with floristic continuity.     

Phenotypic Plasticity of Leaf Shape along a Temperature Gradient in Acer rubrum

Both phenotypic plasticity and genetic determination can be important for understanding how plants respond to environmental change. However, little is known about the plastic response of leaf teeth and leaf dissection to temperature. This gap is critical because these leaf traits are commonly used to reconstruct paleoclimate from fossils, and such studies tacitly assume that traits measured from fossils reflect the environment at the time of their deposition, even during periods of rapid climate change. We measured leaf size and shape in Acer rubrum derived from four seed sources with a broad temperature range and grown for two years in two gardens with contrasting climates (Rhode Island and Florida). Leaves in the Rhode Island garden have more teeth and are more highly dissected than leaves in Florida from the same seed source. Plasticity in these variables accounts for at least 6–19 % of the total variance, while genetic differences among ecotypes probably account for at most 69–87 %. This study highlights the role of phenotypic plasticity in leaf-climate relationships. We suggest that variables related to tooth count and leaf dissection in A. rubrum can respond quickly to climate change, which increases confidence in paleoclimate methods that use these variables.     

Leaf Shape Responds to Temperature but Not CO2 in Acer rubrum

The degree of leaf dissection and the presence of leaf teeth, along with tooth size and abundance, inversely correlate with mean annual temperature (MAT) across many plant communities. These relationships form the core of several methods for reconstructing MAT from fossils, yet the direct selection of temperature on tooth morphology has not been demonstrated experimentally. It is also not known if atmospheric CO₂ concentration affects leaf shape, limiting confidence in ancient climate reconstructions because CO₂ has varied widely on geologic timescales. Here I report the results of growing Acer rubrum (red maple) in growth cabinets at contrasting temperature and CO₂ conditions. The CO₂ treatment imparted no significant differences in leaf size and shape, while plants grown at cooler temperatures tended to have more teeth and more highly dissected leaves. These results provide direct evidence for the selection of temperature on leaf shape in one species, and support a key link in many leaf-climate methods. More broadly, these results increase confidence for using leaf shape in fossils to reconstruct paleoclimate.     

Correlations of climate and plant ecology to leaf size and shape: potential proxies for the fossil record

The sizes and shapes (physiognomy) of fossil leaves are widely applied as proxies for paleoclimatic and paleoecological variables. However, significant improvements to leaf-margin analysis, used for nearly a century to reconstruct mean annual temperature (MAT), have been elusive; also, relationships between physiognomy and many leaf ecological variables have not been quantified. Using the recently developed technique of digital leaf physiognomy, correlations of leaf physiognomy to MAT, leaf mass per area, and nitrogen content are quantified for a set of test sites from North and Central America. Many physiognomic variables correlate significantly with MAT, indicating a coordinated, convergent evolutionary response of fewer teeth, smaller tooth area, and lower degree of blade dissection in warmer environments. In addition, tooth area correlates negatively with leaf mass per area and positively with nitrogen content. Multiple linear regressions based on a subset of variables produce more accurate MAT estimates than leaf-margin analysis (standard errors of ±2 vs. ±3°C); improvements are greatest at sites with shallow water tables that are analogous to many fossil sites. The multivariate regressions remain robust even when based on one leaf per species, and the model most applicable to fossils shows no more signal degradation from leaf fragmentation than leaf-margin analysis.     

Estimation of temperature and precipitation from morphological characters of dicotyledonous leaves

The utility of regression and correspondence models for deducing climate from leaf physiognomy was evaluated by the comparative application of different predictive models to the same three leaf assemblages. Mean annual temperature (MAT), mean annual precipitation (MAP), and growing season precipitation (GSP) were estimated from the morphological characteristics of samples of living leaves from two extant forests and an assemblage of fossil leaves. The extant forests are located near Gainesville, Florida, and in the Florida Keys; the fossils were collected from the Eocene Clarno Nut Beds, Oregon. Simple linear regression (SLR), multiple linear regression (MLR), and canonical correspondence analysis (CCA) were used to estimate temperature and precipitation. The SLR models used only the percentage of species having entire leaf margins as a predictor for MAT and leaf size as a predictor for MAP. The MLR models used from two to six leaf characters as predictors, and the CCA used 31 characters. In comparisons between actual and predicted values for the extant forests, errors in prediction of MAT were 0.6°-5.7°C, and errors in prediction of precipitation were 6-89 cm (=6-66%). At the Gainesville site, seven models underestimated MAT and only one overestimated it, whereas at the Keys site, all eight models overestimated MAT. Precipitation was overestimated by all four models at Gainesville, and by three of them at the Keys. The MAT estimates from the Clarno leaf assemblage ranged from 14.3° to 18.8°C, and the precipitation estimates from 227 to 363 cm for MAP and from 195 to 295 cm for GSP.     

Roles of climate and functional traits in controlling toothed vs. untoothed leaf margins

? Premise of the study: Leaf-margin state (toothed vs. untoothed) forms the basis of several popular methods for reconstructing temperature. Some potential confounding factors have not been investigated with large data sets, limiting our understanding of the adaptive significance of leaf teeth and their reliability to reconstruct paleoclimate. Here we test the strength of correlations between leaf-margin state and deciduousness, leaf thickness, wood type (ring-porous vs. diffuse-porous), height within community, and several leaf economic variables. ? Methods: We assembled a trait database for 3549 species from six continents based on published and original data. The strength of associations between traits was quantified using correlational and principal axes approaches. ? Key results: Toothed species, independent of temperature, are more likely to be deciduous and to have thin leaves, a high leaf nitrogen concentration, a low leaf mass per area, and ring-porous wood. Canopy trees display the highest sensitivity between leaf-margin state and temperature; subcanopy plants, especially herbs, are less sensitive. ? Conclusions: Our data support hypotheses linking the adaptive significance of teeth to leaf thickness and deciduousness (in addition to temperature). Toothed species associate with the "fast-return" end of the leaf economic spectrum, providing another functional link to thin leaves and the deciduous habit. Accounting for these confounding factors should improve climate estimates from tooth-based methods.     

Leaf size variations in a dominant desert shrub,Reaumuria soongarica,adapted to heterogeneous environments

The climate in arid Central Asia (ACA) has changed rapidly in recent decades, but the ecological consequences of this are far from clear. To predict the impacts of climate change on ecosystem functioning, greater attention should be given to the relationships between leaf functional traits and environmental heterogeneity. As a dominant constructive shrub widely distributed in ACA, Reaumuria soongarica provided us with an ideal model to understand how leaf functional traits of desert ecosystems responded to the heterogeneous environments of ACA. Here, to determine the influences of genetic and ecological factors, we characterized species‐wide variations in leaf traits among 30 wild populations of R. soongarica and 16 populations grown in a common garden. We found that the leaf length, width, and leaf length to width ratio (L/W) of the northern lineage were significantly larger than those of other genetic lineages, and principal component analysis based on the in situ environmental factors distinguished the northern lineage from the other lineages studied. With increasing latitude, leaf length, width, and L/W in the wild populations increased significantly. Leaf length and L/W were negatively correlated with altitude, and first increased and then decreased with increasing mean annual temperature (MAT) and mean annual precipitation (MAP). Stepwise regression analyses further indicated that leaf length variation was mainly affected by latitude. However, leaf width was uncorrelated with altitude, MAT, or MAP. The common garden trial showed that leaf width variation among the eastern populations was caused by both local adaptation and phenotypic plasticity. Our findings suggest that R. soongarica preferentially changes leaf length to adjust leaf size to cope with environmental change. We also reveal phenotypic evidence for ecological speciation of R. soongarica. These results will help us better understand and predict the consequences of climate change for desert ecosystem functioning.     

Historical and phylogenetic constraints on the incidence of entire leaf margins: insights from a new South American model

Aim The relationship between the proportion of species with an entire leaf margin (pE) and mean annual temperature (MAT) is one of the most powerful tools for estimating palaeotemperatures. However, phylogenetic and phytogeographic constraints on this relationship have remained unexplored. Here we investigate the pE–MAT relationship for modern floristic assemblages from southern South American forests, assess its conformity to other models and test for the existence of historical constraints on pE–MAT models. Location South America. Methods We used samples from 30 sites located in Chile between 32° and 44° S to test for a pE–MAT relationship and compared it with four regional models. We assessed the reliability of these models for predicting MAT from instrumental records in eight modern temperate‐forest localities in Chile. Additionally, palaeotemperatures for Cenozoic fossil floras were estimated. To assess historical constraints in pE, we measured the phylogenetic signal in leaf margin type and the association between leaf margin and phytogeographic affiliation, defined by the distribution of genera. Results We found a significant pE–MAT relationship for Chilean forest species that differed from Australia and Northern Hemisphere models, but not from tropical South America (TSA). Temperatures for southern South American localities predicted from the new regional model – combining Chilean and TSA datasets – were more accurate than those from previous models. We also showed that leaf margin type has a strong phylogenetic signal, which was further confirmed by the highly significant effect of phytogeographic element on leaf margin type. Main conclusions Differences between the Chilean and other regional models are explained by historical legacy, as Chilean leaf margin types are strongly affected by phylogenetic closeness and phytogeographic elements. We highlight that leaf margin analyses should be conducted within the context of a flora with a shared history. Thus, we propose a new model for South America to estimate palaeotemperatures for regional fossil floras.     

Influences of Environmental Factors on Leaf Morphology of Chinese Jujubes

Rainfall and temperature are the primary limiting factors for optimum quality and yield of cultivated jujube (Ziziphus jujuba Mill.). Adaptation to arid and cool environments has been and remains an important goal of many jujube improvement programs. This study summarized the survey results of 116 Chinese jujube varieties grown at 33 sites in China. The objective was to identify the environmental factors that influence leaf morphology, and the implications for breeding and introduction of new jujube varieties. Jujube leaf morphological traits were evaluated for their potential relationships with mean annual temperature (MAT) and mean annual precipitation (MAP). The results showed that many leaf morphological traits had a strong linear relationship with local precipitation and temperature. Longer veins per unit area (VLA) and reduced leaf area and leaf perimeter were typical of arid areas. VLA was inversely related to MAT and MAP at the centers of origin of jujube. There was a positive relationship between leaf shape (perimeter²/area) and both MAT and MAP. These results indicated that leaf vein traits of Chinese jujubes might have resulted from their adaptation to environmental factors in the course of long-term evolution. Principal component analysis allocated the 116 jujube varieties to three different groups, differentiated on the basis of morphological and physiological leaf characteristics. Jujube varieties from the Hebei, Shandong, Henan, southern Shanxi and central Shaanxi provinces were closely related, as were varieties from northwest Shanxi and northeast Shaanxi provinces, and varieties from the Gansu and Ningxia provinces. These close relationships were partially attributed to the frequent exchanges of varieties within each group. Leaf venation characteristics might be used as reference indices for jujube variety introduction between different locations.Influences of Environmental Factors on Leaf Morphology of Chinese Jujubes     

Leaf size of woody dicots predicts ecosystem primary productivity

A key challenge in ecology is to understand the relationships between organismal traits and ecosystem processes. Here, with a novel dataset of leaf length and width for 10 480 woody dicots in China and 2374 in North America, we show that the variation in community mean leaf size is highly correlated with the variation in climate and ecosystem primary productivity, independent of plant life form. These relationships likely reflect how natural selection modifies leaf size across varying climates in conjunction with how climate influences canopy total leaf area. We find that the leaf size?primary productivity functions based on the Chinese dataset can predict productivity in North America and vice‐versa. In addition to advancing understanding of the relationship between a climate‐driven trait and ecosystem functioning, our findings suggest that leaf size can also be a promising tool in palaeoecology for scaling from fossil leaves to palaeo‐primary productivity of woody ecosystems.     

Present-day testing of a paleoecological pattern: Is there really a latitudinal difference in leaf-feeding insect-damage diversity?

The fossil record suggests greater diversity of insect leaf feeding during warm climate intervals. Much published work in the paleobotanical literature has been based on the presumed validity of this pattern. However, the existence of this pattern in nature has never been tested from the present-day world. Here we ask, is it true that on average, in warmer climates, a leaf is being eaten in more ways?We compared forests at seven sites in northern Florida (30° N, MAT ca. 19.5 °C) to seven sites across the north-eastern USA (40-42° N, MAT 7-9 °C). Presence and absence of damage types were determined using a standard leaf damage guide; 93 damage types were found in the Florida samples and 80 in the north-eastern samples.In bulk floras, there was a consistent difference in damage diversity, on a per-leaf basis (as in the fossil studies), between Florida and north-eastern sites. Florida sites had a greater number of damage types. When northern and southern populations of individual tree species were compared, higher southern damage diversity was found in four species (Acer rubrum, Acer saccharum, Fagus grandifolia, and Quercus coccinea), though with no difference with latitude in a fifth species (Quercus alba).These results appear to validate that the trend seen in the fossil record is not a spurious effect of site differences. They also extend a pattern seen in deep time into the present day, adding greatly to its generality.     

Leaf nitrogen and phosphorus concentration and the empirical regulations in dominant woody plants of shrublands across southern China

Aims Understanding the changes in N and P concentration in plant organs along the environmental gradients can provide meaningful information to reveal the underline mechanisms for the geochemical cycles and adaptation strategies of plants to the changing environment. In this paper, we aimed to answer: (1) How did the N and P concentration in leaves of evergreen and deciduous woody plants change along the environmental gradients? (2) What were the main factors regulating the N and P concentration in leaves of woody plants in the shrublands across southern China?
Methods Using a stratified random sampling method, we sampled 193 dominant woody plants in 462 sites of 12 provinces in southern China. Leaf samples of dominant woody plants, including 91 evergreen and 102 deciduous shrubs, and soil samples at each site were collected. N and P concentration of the leaves and soils were measured after lapping and sieving. Kruskal-Wallis and Nemenyi tests were applied to quantify the difference among the organs and life-forms. For each life-form, the binary linear regression was used to estimate the relationships between leaf log [N] and log [P] concentration and mean annual air temperature (MAT), mean annual precipitation (MAP) and log soil total [N], [P]. The effects of climate, soil and plant life-form on leaf chemical traits were modeled through the general linear models (GLMs) and F-tests.
Important findings 1) The geometric means of leaf N and P concentrations of the dominant woody plants were 16.57 mg·g^-1 and 1.02 mg·g^-1, respectively. The N and P concentration in leaves (17.91 mg·g^-1, 1.14 mg·g^-1) of deciduous woody plants was higher than those of evergreen woody plants (15.19 mg·g^-1, 0.89 mg·g^-1). The dependent of leaf P concentration on environmental (climate and soil) appeared more variable than N concentration. 2) Leaf N and P in evergreen woody plants decreased with MAT and but increased with MAP, whereas those in deciduous woody plants showed opposite trends. With increase in MAP, leaf P concentration decreased for both evergreen and deciduous woody plants. 3) Soil N concentration had no significant effect on both evergreen and deciduous woody plants. However, leaf P concentration of the tow increased significantly with soil P concentration. (4) GLMs showed that plant growth form explained 7.6% and 14.4% of variation in leaf N and P, respectively. MAP and soil P concentration contributed 0.8% and 16.4% of the variation in leaf P, respectively. These results suggested that leaf N was mainly influenced by plant growth form, while leaf P concentration was driven by soil, plant life-form, and climate at our study sites.     

Evidence of a Cooler Continental Climate in East China during the Warm Early Cenozoic

The early Cenozoic was characterized by a very warm climate especially during the Early Eocene. To understand climatic changes in eastern Asia, we reconstructed the Early Eocene vegetation and climate based on palynological data of a borehole from Wutu coal mine, East China and evaluated the climatic differences between eastern Asia and Central Europe. The Wutu palynological assemblages indicated a warm temperate vegetation succession comprising mixed needle- and broad-leaved forests. Three periods of vegetation succession over time were recognized. The changes of palynomorph relative abundance indicated that period 1 was warm and humid, period 2 was relatively warmer and wetter, and period 3 was cooler and drier again. The climatic parameters estimated by the coexistence approach (CA) suggested that the Early Eocene climate in Wutu was warmer and wetter. Mean annual temperature (MAT) was approximately 16°C and mean annual precipitation (MAP) was 800–1400 mm. Comparison of the Early Eocene climatic parameters of Wutu with those of 39 other fossil floras of different age in East China, reveals that 1) the climate became gradually cooler during the last 65 million years, with MAT dropping by 9.3°C. This cooling trend coincided with the ocean temperature changes but with weaker amplitude; 2) the Early Eocene climate was cooler in East China than in Central Europe; 3) the cooling trend in East China (MAT dropped by 6.9°C) was gentler than in Central Europe (MAT dropped by 13°C) during the last 45 million years.     

Leaf litter carbon,nitrogen, and phosphorus stoichiometric patterns as related to climatic factors and leaf habits across Chinese broad-leaved tree species

Our understanding of how climate and leaf habit (evergreen vs. deciduous) drive leaf litter carbon (C), nitrogen (N), and phosphorus (P) stoichiometric patterns is largely limited but is particularly important for broad-leaved forests, since the forest is sensitive to climate change. Here, we investigated leaf litter C, N, and P stoichiometric ratios of broad-leaved tree species in relation to climate and leaf habit using previous publications and our additional samplings across China. We found that mean leaf litter C:N across Chinese broad-leaved tree species was within the range of the global flora, whereas C:P was lower and N:P higher. Evergreen species displayed higher leaf litter C:N, C:P, and N:P than their deciduous counterparts. Both leaf litter C:P and N:P for all species pooled were negatively correlated with latitude, driven by mean annual precipitation (MAP) and mean annual temperature, respectively, while leaf litter C:N displayed no clear latitudinal trend. The direction and magnitude of leaf litter C, N, and P stoichiometric ratios in response to climate diverged between leaf habits. For example, evergreen leaf litter C:N was negatively correlated with MAP, while deciduous counterparts did not respond significantly to MAP. We conclude that leaf litter C, N, and P stoichiometric ratios shifted along the climatic gradient, and the strength of such shifts differed between leaf habits. Therefore, leaf litter stoichiometric patterns across leaf habits suggest that any climate change-driven shift in species distribution may potentially alter the ecosystem’s nutrient cycling processes of evergreen- and deciduous-dominated broad-leaved forests differentially.     

A climatic and taxonomic comparison between leaf litter and standing vegetation from a Florida swamp woodland

One method to determine past climate has been the use of leaf morphological characteristics of fossil leaves quantified using modern climate and canopy leaf characteristics. Fossil assemblages are composed of abscised leaves, and climate may be more accurately determined by using leaves from leaf litter instead of the canopy. To better understand whether taphonomic processes make a difference in this relationship, a north-central Florida woodland was sampled to determine the morphologically based climate estimates from these leaves. Leaves from woody, dicotyledonous plants were collected and identified, then compared using presence/absence data and analyzed using several linear regression equations and the CLAMP data set. Although the majority of standing vegetation was reflected in leaf litter, some inconsistencies were observed, which may reflect plant community structure or sampling technique. Mean annual temperature (MAT) and growing season precipitation (GSP) were estimated from leaf litter morphological characters and living leaves. Overall, values for MAT estimated from litter and living leaves were cooler than actual MATs, although several accurate and high estimates were obtained depending on the predictive method used. Estimated GSP values were higher than actual GSPs. Statistically, no difference was observed between MAT and GSP estimates derived from leaf litter vs. estimates derived from living leaves, with one exception.     

Evolution of leaf form correlates with tropical-temperate transitions in Viburnum (Adoxaceae)

Strong latitudinal patterns in leaf form are well documented in floristic comparisons and palaeobotanical studies. However, there is little agreement about their functional significance; in fact, it is still unknown to what degree these patterns were generated by repeated evolutionary adaptation. We analysed leaf form in the woody angiosperm clade Viburnum (Adoxaceae) and document evolutionarily correlated shifts in leafing habit, leaf margin morphology, leaf shape and climate. Multiple independent shifts between tropical and temperate forest habitats have repeatedly been accompanied by a change between evergreen, elliptical leaves with entire margins and deciduous, more rounded leaves with toothed or lobed margins. These consistent shifts in Viburnum support repeated evolutionary adaptation as a major determinant of the global correlation between leaf form and mean annual temperature. Our results provide a new theoretical grounding for the inference of past climates using fossil leaf assemblages.     

Do leaf margins of the temperate forest flora of southern South America reflect a warmer past?

Aim   Our aims were: (1) to characterize the linear relationship between the proportion of woody dicotyledonous species with entire-margined leaves ( E ) and mean annual temperature (MAT) from a southern temperate flora that still harbours many lineages that originated under warmer climates; (2) to compare this relationship with those developed from floras of different regions of the world; and (3) to contrast temperature predictions based on leaf margins of the native southern flora versus the naturalized alien flora, mostly of boreal origin.
Location   The temperate forest of southern South America (TFSA).
Methods   At each 1° latitudinal band, we estimated E based on species latitudinal ranges and MAT from both an isotherm map and a global temperature grid. We also calculated E from five local floras located between 40 and 43° S, and from the naturalized alien flora of Nahuel Huapi National Park in southern Argentina.
Results   We found a close relationship between E and MAT for the TFSA. Equations developed from floras of the Northern Hemisphere overestimated extant temperatures of this biome by 6–10 °C at both geographical and local spatial scales. On the other hand, MAT predictions from leaf margins of the alien flora were similar to the actual MAT. A published regression between E and MAT from tropical South America was remarkably similar to the one we estimated from the TFSA. This tropical equation predicted accurately the temperatures observed for this temperate biome based on leaf margins of the native flora.
Main conclusions   Despite massive plant extinction due to environmental cooling and biogeographical isolation during the Tertiary, leaf-margin analysis reveals that the flora of the TFSA still reflects its original development under the warmer conditions of western Gondwana and its past connections with low-latitude forest floras of tropical South America.     

Leaf element concentrations of terrestrial plants across China are influenced by taxonomy and the environment

Aim The productivity, functioning and biogeochemical cycles of terrestrial ecosystems are strongly affected by leaf element concentrations. Understanding the biological and ecological factors affecting leaf element concentrations is therefore important for modelling the productivity and nutrient fluxes of ecosystems and their responses to global change. The present study aimed to determine how leaf element concentrations are linked to taxonomy and the environment. Location China. Methods The concentrations of 10 leaf elements of 702 terrestrial plant species from different biomes were extracted from publications. The links between environmental variables, taxonomy and leaf elements were analyzed using phylogenetically comparative methods and partial Mantel tests. Results Taxonomy had stronger effects on leaf S and SiO₂ than latitude, explaining 40.2–43.9% of total variation, whereas latitude had stronger effects on leaf N, P, K, Fe, Al, Mn, Na and Ca concentrations, explaining 19.5–52.1% of total variation. Leaf N, S, Al, Fe and Na concentrations were correlated with mean annual precipitation (MAP), while leaf N, P and Fe concentrations were correlated with mean annual temperature (MAT). Latitude, MAP and MAT were significantly correlated with the first axis of a principal components analysis (PCA). This first axis was associated with leaf elements involved in protein synthesis and photosynthesis. The other PCA axes, which were not correlated with MAT, latitude and MAP, were associated with leaf elements responsible for cell structure and enzymes. Main conclusions Leaf element concentrations of terrestrial plants in China were correlated with climate, latitude and taxonomy. With the exception of S and SiO₂, the environmental factors were more important in explaining leaf element variation than taxonomy. Therefore, changes in temperature and precipitation will directly affect the spatial patterns of leaf elements and thus the associated nutrient fluxes and ecosystem functioning.     

Global trends in senesced-leaf nitrogen and phosphorus

Aim Senesced‐leaf litter plays an important role in the functioning of terrestrial ecosystems. While green‐leaf nutrients have been reported to be affected by climatic factors at the global scale, the global patterns of senesced‐leaf nutrients are not well understood. Location Global. Methods Here, bringing together a global dataset of senesced‐leaf N and P spanning 1253 observations and 638 plant species at 365 sites and of associated mean climatic indices, we describe the world‐wide trends in senesced‐leaf N and P and their stoichiometric ratios. Results Concentration of senesced‐leaf N was highest in tropical forests, intermediate in boreal, temperate, and mediterranean forests and grasslands, and lowest in tundra, whereas P concentration was highest in grasslands, lowest in tropical forests and intermediate in other ecosystems. Tropical forests had the highest N : P and C : P ratios in senesced leaves. When all data were pooled, N concentration significantly increased, but senesced‐leaf P concentration decreased with increasing mean annual temperature (MAT) and mean annual precipitation (MAP). The N : P and C : P ratios also increased with MAT and MAP, but C : N ratios decreased. Plant functional type (PFT), i.e. life‐form (grass, herb, shrub or tree), phylogeny (angiosperm versus gymnosperm) and leaf habit (deciduous versus evergreen), affected senesced‐leaf N, P, N : P, C : N and C : P with a ranking of senesced‐leaf N from high to low: forbs ≈ shrubs ≈ trees > grasses, while the ranking of P was forbs ≈ shrubs ≈ trees < grasses. The climatic trends of senesced‐leaf N and P and their stoichiometric ratios were similar between PFTs. Main conclusions Globally, senesced‐leaf N and P concentrations differed among ecosystem types, from tropical forest to tundra. Differences were significantly related to global climate variables such as MAT and MAP and also related to plant functional types. These results at the global scale suggest that nutrient feedback to soil through leaf senescence depends on both the climatic conditions and the plant composition of an ecosystem.