Mohammed Armani.

Recent empirical and theoretical approaches have called for an understanding of the processes underpinning ecosystem service provision. Environmental gradients have shown effects on key plant functional traits that subsequently explain ecosystem properties of several systems. However, little is known concerning how associations between plant functional traits, including both below- and aboveground plant components, predict ecosystem properties and independently measured final ecosystem services. Here, we modeled (1) the responses of the leaf and plant economics spectrum, Plant size axis, and root growth to environmental gradients and (2) how associations between plant functional traits explain trade-offs and synergies between multiple ecosystem properties and final services. Forty-four plots were studied in a coastal marsh landscape of the German North Sea Coast. We used a partial least square structural equation model approach to test the hypothesized model. We found (1) a negative covariation between plant traits pertaining to a size axis and traits explaining both plant growth (roots and stems) and the leaf economics spectrum; (2) this trade-off responded significantly to the land use gradient and nutrient availability, which were both strongly driven by the groundwater gradient; (3) this trade-off explained an initial major trade-off between carbon stocks, at one extreme of the axis, and both the habitat value to conserve endangered plants and forage production for meat and dairy products at the other extreme. However, a secondary trade-off between nature conservation value and forage production, explained by a trade-off between leaf economics spectrum and plant growth in response to the land use intensity gradient, was also found.

New PhytologistVolume 183, Issue 4 p. 1222-1222 Free Access Erratum This article corrects the following: Causes and consequences of variation in leaf mass per area (LMA): a meta-analysis Hendrik Poorter, Ülo Niinemets, Lourens Poorter, Ian J. Wright, Rafael Villar, Volume 182Issue 3New Phytologist pages: 565-588 First Published online: April 16, 2009 First published: 10 August 2009 https://doi.org/10.1111/j.1469-8137.2009.02972.xCitations: 2AboutSectionsPDF ToolsRequest permissionExport citationAdd to favoritesTrack citation ShareShare Give accessShare full text accessShare full-text accessPlease review our Terms and Conditions of Use and check box below to share full-text version of article.I have read and accept the Wiley Online Library Terms and Conditions of UseShareable LinkUse the link below to share a full-text version of this article with your friends and colleagues. Learn more.Copy URL Share a linkShare onFacebookTwitterLinked InRedditWechat New Phytologist 182 (2009), 565–588. Since its publication, the authors of Poorter et al. (2009) have brought to our attention that there is an error in Fig. 5(j), where all the data below the y= 1 line are absent. The correct panel (j), and the full legend to Fig. 5, are printed below. Fig. 5 Characterization of the response in leaf dry mass per unit area (LMA) of plants grown in a range of environmental conditions: (a) daily photon irradiance (DPI); (b) red : far-red ratio (R/FR); (c) UV-B; (d) CO2 concentration; (e) ozone; (f) nutrient availability; (g) water availability (drought stress); (h) waterlogging; (i) submergence; (j) temperature; (k) salinity; (l) soil compaction. Data are a compilation of the literature. For each environmental factor a reference condition was chosen (indicated by arrows), and all data from different species were normalized to that condition. For more information see Appendix A5. The shaded area indicates the interquartile range (between the 25th and the 75th percentile) of the observed ratios in that part of the response curve. The bold continuous line within the shaded area indicates the median value. Dashed lines indicate the median value for a specific subgroup of species and are labelled in the graph. Generally, species were classified with respect to their original habitat to grow at high, intermediate or low levels of the environmental factor studied. The total number of observations present in each part of the response curve is indicated at the top of each graph. The y-axis is logarithmic to correct for the fact that ratios are logarithmic by nature. We apologize to our readers for this mistake. Reference Poorter H, Niinemets U, Poorter L, Wright IJ, Villar R. 2009. Causes and consequences of variation in leaf mass per area (LMA): a meta-analysis. New Phytologist 182: 565– 588. Wiley Online LibraryCASPubMedWeb of Science®Google Scholar Citing Literature Volume183, Issue4September 2009Pages 1222-1222 ReferencesRelatedInformation

Leaf mass per area (LMA), which is an important functional trait in leaf economic spectrum and plant growth analysis, is measured from leaf discs or whole leaves. Differences between the measurement methods may lead to large differences in the estimates of LMA values. We examined to what extent estimates of LMA based on whole leaves match those based on discs using 334 woody species from a wide range of biomes (tropics, subtropics, savanna, and temperate), whether the relationship varied by leaf morphology (tissue density, leaf area, leaf thickness), punch size (0.6- and 1.0-cm diameter), and whether the extent of intraspecifc variation for each species matches. Disc-based estimates of species mean LMA matched the whole-leaf estimates well, and whole-leaf LMA tended to be 9.69% higher than leaf-disc LMA. The ratio of whole-leaf LMA to leaf-disc LMA was higher for species with higher leaf tissue density and larger leaves, and variance in the ratio was greater for species with lower leaf tissue density and thinner leaves. Estimates based on small leaf discs also inflated the ratio. The extent of the intraspecific variation only weakly matched between whole-leaf and disc-based estimates (R2 = 0.08). Our results suggest that simple conversion between whole-leaf and leaf-disc LMA is difficult for species obtained with a small leaf punch, but it should be possible for species obtained with a large+ leaf punch. Accurately representing leaf traits will likely require careful selection between leaf-disc and whole-leaf traits depending on the objectives. Quantifying intraspecific variation using leaf discs should be also considered with caution.

Economic trade-offs of hydrophytes and neighbouring terrestrial herbaceous plants based on plant functional traits

Many studies focus on the relationships between plant functional traits and tree growth performances. However, little is known about the ontogenetic shifts of the relationships between functional traits and tree growth. This study examined associations between stem and leaf functional traits and growth rates and their ontogenetic shifts across 20 tropical tree species in a tropical seasonal rainforest in Xishuangbanna, south-west China. For each species, physiological active branches of individual trees belonged to three size classes (i.e. small, diameter at breast height (DBH) 5-10 cm; middle, DBH 10-20 cm; big, DBH >20 cm) were sampled respectively. We measured 18 morphological and structural traits, which characterised plant hydraulic properties or leaf economic spectrum. Associations between diameter growth rates and functional traits were analysed across three size classes. Our results revealed that diameter growth rates of big-sized trees were mainly related to traits related to plant hydraulic efficiency (i.e. theoretical hydraulic conductivity (Ktheo) and leaf vein density (Dvein)), which suggests that the growth of large trees is limited mainly by their xylem water transport capacity. For middle-sized trees, growth rates were significantly related to traits representing leaf economic spectrum (i.e. specific leaf area (SLA), individual leaf mass (ILM), palisade thickness (PT) and spongy thickness (SP)). Diameter growth rates of small-sized trees were not correlated with hydraulic or leaf economic traits. Thus, the associations between tree growth rates and functional traits are size dependent. Our results suggest ontogenetic shift of functional traits which could potential contribute to different growth response to climate change.

Plant functional traits are an important indicator for the comprehensive evaluation of community stability and resilience. Therefore, exploring the variations and relationships among leaf functional traits at different maturity levels during forest restoration can deepen the understanding of plant adaptation strategies and community assembly. In this study, we measured the leaf area (LA), specific leaf area (SLA), photosynthetic pigments, non-structural carbohydrates (NSCs), leaf nitrogen content (LNC), and leaf phosphorus content (LPC) of dominant tree species in three communities with different maturity levels (defined by species composition, biodiversity and spatial structure) in a monsoon forest located in the northern margin of the tropics in China, and explored the variation and relationships among different leaf traits at individual, species, and community scales. The results showed that maturity levels significantly affected leaf functional traits. With the increase in maturity levels, SLA increased, and leaf SS and NSCs decreased, while other leaf functional traits did not show a consistent pattern. In different communities, NSCs, Chl (a:b), SS:St or Car had a trade-off or synergistic relationship with leaf economic spectrum. Additionally, the LPC, LNC, and starch were the key traits in response to selection pressure at maturity levels, inter-specific and intra-specific scales, respectively, and the trait–trait relationships were stronger or more extreme as the scale was narrowed. Therefore, when evaluating the development and succession of tropical monsoon forest communities, the selection of leaf functional characteristics and the determination of the research scale should be comprehensively and systematically considered.

Leaf traits are good predictors of woody species distribution after bamboo expansion in a mountain forest

Understanding the relationships between species' demography and functional traits is crucial for gaining a mechanistic understanding of community dynamics. While leaf morphology represents a key functional dimension for plants worldwide (i.e., the leaf economics spectrum), its ability to explain variation in trees' life history strategies remains limited. Plant growth is influenced by both leaf morphology and allocation; hence, incorporating both dimensions is essential but rarely done. Additionally, trait–performance relationships have mainly been studied in tropical communities, leaving gaps in our understanding of temperate forests where different seasonality patterns may alter these relationships. We examined whether species' leaf area index (leaf area per crown size, LAI), a measure of leaf allocation, explains the variation of juvenile tree species' potential growth rates in a winter‐deciduous broadleaf forest. LAI has not been characterized as a species‐level trait, but its ability to predict plant productivity at the ecosystem scale highlights its potential for explaining plant growth. We evaluated species' maximum LAI both individually and in conjunction with wood density (WD) and leaf mass per area (LMA). We expected that models would improve when both leaf morphology (LMA) and leaf allocation (LAI) were included and that species with denser crowns would have higher potential growth rates. LAI and LMA were significant predictors of growth but only when both were incorporated, and together explained a high proportion of species' growth variations (R2adj = 0.59). We found evidence of a trade‐off between LAI and LMA, with a negative relationship between them and each having a positive influence on species' growth, suggesting that there are multiple allocation strategies to achieve fast growth. A surprisingly positive LMA–growth relationship contrasts with observations from tropical forests. We did not find significant relationships with WD in this forest. Our results highlight that incorporating leaf allocation improves models of trait–performance relationships. They also suggest, in agreement with the limited literature, that temperate forests may exhibit different trait–performance relationships from those of tropical forests, where LMA is negatively related to growth and WD is often important. Clarifying the details and contexts of trait–performance relationships is crucial for applying the functional trait framework to understanding community structure and dynamics of forests globally.

Developmental phenotypic plasticity can allow plants to buffer the effects of abiotic and biotic environmental stressors. Therefore, it is vital to improve our understanding of how phenotypic plasticity in ecological functional traits is coordinated with variation in physiological performance in plants. To identify coordinated leaf responses to low-water (LW) versus low-light (LL) availability, we measured leaf mass per area (LMA), leaf anatomical characteristics and leaf gas exchange of juvenile Populus tremuloidesMichx. trees. Spongy mesophyll tissue surface area (Asmes/A) was correlated with intrinsic water-use efficiency (WUEi: photosynthesis, (Aarea)/stomatal conductance (gs)). Under LW availability, these changes occurred at the cost of greater leaf tissue density and reduced expansive growth, as leaves were denser but were only 20% the final area of control leaves, resulting in elevated LMA and elevated WUEi. Low light resulted in reduced palisade mesophyll surface area (Apmes/A) while spongy mesophyll surface area was maintained (Asmes/A), with no changes to WUEi. These leaf morphological changes may be a plastic strategy to increase laminar light capture while maintaining WUEi. With reduced density and thickness, however, leaves were 50% the area of control leaves, ultimately resulting in reduced LMA. Our results illustrate that P. tremuloides saplings partially maintain physiological function in response to water and light limitation by inducing developmental plasticity in LMA with underlying anatomical changes. We discuss additional implications of these results in the context of developmental plasticity, growth trade-offs and the ecological impacts of climate change.

AimsExtensive research on the leaf economics spectrum (LES) has improved insights into functional traits and their environmental interactions. Several studies explored trait correlations in angiosperms and ferns, expanding their ecological significance. However, knowledge gaps persist, especially concerning East‐Asian ferns and the differences between terrestrial and epiphytic ferns in LES.LocationEast Asia, Northern Taiwan.MethodsData were collected along an elevation gradient (870–2130 m a.s.l.) in northern Taiwan, where we measured nine leaf traits for 47 terrestrial and 34 epiphytic ferns across 59 vegetation plots within subtropical forests. We explored trait–trait and trait–environment relationships at the species and community levels for both terrestrial and epiphytic ferns while accounting for phylogenetic constraints.ResultsEpiphytes had lower specific leaf area (SLA) and leaf nitrogen, greater trait variance and trait space than terrestrial species, suggesting they are ecologically distinct. These differences observed at species and community levels appear to emerge because epiphytes undergo stronger water and nutritional stress along the elevation gradient. In addition, three groups were detected within epiphytes reflecting their strategies for water and nutrient stress adaptation. Trait correlations largely resembled angiosperm LES patterns but were less apparent in epiphytes. SLA and leaf dry matter content (LDMC) were consistently correlated to water stress traits, which might contribute to the restricted LES trait range of both life forms in our study. Traits exhibited a strong phylogenetic signal, with marked differences between SLA and LDMC correlations in phylogenetics‐corrected vs raw data. Trait–environment relationships were similar for several water stress‐related traits across both species groups, but stronger relationships were more evident in terrestrial ferns.ConclusionsTrait patterns are not entirely equivalent for epiphytic and terrestrial species or communities and should not be extrapolated across life forms. Distantly related species with various evolutionary backgrounds could dismantle LES patterns at the species level. Strong phylogenetic constraints may influence the trait–environment response of epiphytic species.

Investigating the effects of ontogenetic stage and leaf age on leaf traits is important for understanding the utilization and distribution of resources in the process of plant growth. However, few studies have been conducted to show how traits and trait-trait relationships change across a range of ontogenetic stage and leaf age for evergreen coniferous species. We divided 67 Pinus koraiensis Sieb. et Zucc. of various sizes (0.3–100 cm diameter at breast height, DBH) into four ontogenetic stages, i.e., young trees, middle-aged trees, mature trees and over-mature trees, and measured the leaf mass per area (LMA), leaf dry matter content (LDMC), and mass-based leaf nitrogen content (N) and phosphorus content (P) of each leaf age group for each sampled tree. One-way analysis of variance (ANOVA) was used to describe the variation in leaf traits by ontogenetic stage and leaf age. The standardized major axis method was used to explore the effects of ontogenetic stage and leaf age on trait-trait relationships. We found that LMA and LDMC increased significantly and N and P decreased significantly with increases in the ontogenetic stage and leaf age. Most trait-trait relationships were consistent with the leaf economic spectrum (LES) at a global scale. Among them, leaf N content and LDMC showed a significant negative correlation, leaf N and P contents showed a significant positive correlation, and the absolute value of the slopes of the trait-trait relationships showed a gradually increasing trend with an increasing ontogenetic stage. LMA and LDMC showed a significant positive correlation, and the slopes of the trait-trait relationships showed a gradually decreasing trend with leaf age. Additionally, there were no significant relationships between leaf N content and LMA in most groups, which is contrary to the expectation of the LES. Overall, in the early ontogenetic stages and leaf ages, the leaf traits tend to be related to a "low investment-quick returns" resource strategy. In contrast, in the late ontogenetic stages and leaf ages, they tend to be related to a "high investment-slow returns" resource strategy. Our results reflect the optimal allocation of resources in Pinus koraiensis according to its functional needs during tree and leaf ontogeny.

Phenotypic plasticity is a key mechanism by which plants respond to changing or heterogeneous conditions. Efforts to predict phenotypic plasticity across plant species have mainly focused on environmental variability or abiotic conditions, i.e. site characteristics. However, the considerable variation in phenotypic plasticity within sites calls for alternative approaches. Different functional groups are thought to differ in their plasticity levels. Further, traits such as leaf specific area (SLA), leaf area (LA) and maximum photosynthetic rate ( A max ) reflect central aspects of plant strategies. Lower values of SLA, LA and A max are indicative of a resource‐conservative strategy, which is thought to be associated with lower phenotypic plasticity. We used meta‐analytical data to test whether plant functional group (herbs, woody deciduous and woody evergreens) and SLA, LA and A max are associated with phenotypic plasticity in four trait types: biomass allocation, plant size, leaf morphology and physiology. We obtained data from 168 plant species and accounted for phylogenetic relationships in all analyses. We found a positive relationship between SLA and phenotypic plasticity in biomass allocation, leaf morphology and physiology, with differences across functional groups. In contrast, there was no evidence of greater plasticity in plant size in species with higher SLA; rather the opposite was true for woody evergreens. A max and LA showed similar, but less consistent associations with phenotypic plasticity. Our results show the potential of building predictive frameworks for phenotypic plasticity based on easily measured plant functional characteristics. Results also provide insights into plant strategies and suggest the existence of potential compromises: resource‐conservative, low‐SLA species tend to be more stress‐tolerant but may be less able to cope with variable conditions due to their generally lower phenotypic plasticity. Further studies are needed to explore the mechanisms and the potential implications of this association.

Photosynthetic light-response (PLR) curves for leaves are important components of models related to carbon fixation in forest ecosystems, linking the Mitscherlich equation and Michaelis-Menten equation to traits of the leaf economics spectrum (LES). However, models do not consider changes in leaf habits (i.e., evergreen and deciduous) and within-canopy shading variation in these PLR curves. Here, we measured the PLR curves in sun and shade leaves of 44 evergreen and 31 deciduous species to examine the relationships between variables of the Mitscherlich equation and Michaelis-Menten equation, leaf nitrogen (N) and phosphorus (P) content, and leaf mass per area (LMA). Small changes were caused by different leaf habits and shade variations in relationships linking variables of the two equations to leaf N and P content and LMA. Values of the scaling exponents for PLR curve parameters did not differ regardless of canopy position and leaf habit (P > 0.05). The PLR curves in species with different leaf habits (i.e., evergreen and deciduous) at different canopy positions could be predicted using the general allometric relations between leaf traits and PLR parameters in the two equations. For photosynthetic photon flux densities from 0 to 2000 μmol m-2 s-1 , approximately 71% (Mitscherlich equation) and 70% (Michaelis-Menten equation) of the net assimilation rates could be predicted. These findings indicate that leaf net assimilation rates can be predicted through the large available data for LES traits. Incorporation of values for these traits available in the LES databases into ecosystem models of forest productivity and carbon fixation warrants further investigation.

Physiological ecology and plant functional traits are often used to explain plant invasion. To gain a better understanding of how traits influence invasion, studies usually compare the invasive plant to a native congener, but there are few conspecific examples in the literature. In North America, the presence of native and introduced genetic lineages of the common reed, Phragmites australis, presents a unique example to evaluate how traits influence plant invasion. We reviewed the literature on functional traits of P. australis lineages in North America, specifically contrasting lineages present on the Atlantic Coast. We focused on differences in physiology between the lineage introduced from Eurasia and the lineage native to North America, specifically seeking to identify the causes underlying the recent expansion of the introduced lineage. Our goals were to better understand which traits may confer invasiveness, provide predictions of how these lineages may respond to interspecific competition or imminent global change, and provide guidance for future research. We reviewed published studies and articles in press, and conducted personal communications with appropriate researchers and managers to develop a comparative dataset. We compared the native and introduced lineages and focused on plant physiological ecology and functional traits. Under both stressful and favourable conditions, our review showed that introduced P. australis consistently exhibited greater ramet density, height and biomass, higher and more plastic relative growth rate, nitrogen productivity and specific leaf area, higher mass specific nitrogen uptake rates, as well as greater phenotypic plasticity compared with the native lineage. We suggest that ecophysiological and other plant functional traits elucidate potential mechanisms for the introduced lineage's invasiveness under current and predicted global change conditions. However, our review identified a disconnect between field surveys, experiments, natural competition and plant ecophysiology that must be addressed in future field studies. Given the likelihood of hybridization between lineages, a better understanding of plant traits in native, non-native and hybrid lineages is needed to manage current invasions and to predict the outcome of interactions among novel genotypes. Comparative physiology and other plant functional traits may provide additional tools to predict the trajectory of current and potential future invasions.

The Leaf Economic Spectrum (LES) describes the different strategies of leaves in capturing and using resources, ranging from fast-growing species with short-lived leaves to those that adopt a more conservative approach, investing in durable and efficient leaves in the use of resources. The objective of this systematic review is to quantify and investigate studies on the Leaf Economic Spectrum (LES) in tropical vegetation, analyzing the association with functional traits, their geographic distribution, vegetation types, plant life habits and the main biotic and abiotic factors that shape these characteristics. The current study performed a systematic review of the indexed databases (Web of Science, Scopus) in the last 20 years (2004-2024). We identified 160 publications. The discovery of articles investigating leaf functional properties and the leaf economic spectrum in the tropics, the topic of our systematic review, has grown considerably over the past eight years. We identified a dominant association of studies published with the Neotropical region. The most common functional leaf attributes studied in the framework of LES (e.g. specific leaf area, leaf nitrogen concentration and photosynthetic capacity) were all found to be associated with the plant growth rate in our research. The distribution of LES research shows that LES studies of tropical vegetation cover a relatively higher amount of sampling for tropical and subtropical dry and moist broadleaf forests than others. Despite the advances, significant knowledge gaps remain, particularly regarding physiological traits and understudied plant groups such as lianas, epiphytes, and bryophytes. Future research efforts must prioritize a broader range of life forms and functional traits, integrating eco-physiological and biogeochemical approaches to better predict the responses of tropical vegetation to environmental changes and to support more effective conservation and restoration strategies.