Lower Leaf Mass Per Area Research Articles

Intraspecific alternative phenotypes contribute to variation in species' strategies for growth.

Ecologists have historically sought to identify the mechanisms underlying the maintenance of local species diversity. High-dimensional trait-based relationships, such as alternative phenotypes, have been hypothesized as important for maintaining species diversity such that phenotypically dissimilar individuals compete less for resources but have similar performance in a given environment. The presence of alternative phenotypes has primarily been investigated at the community level, despite the importance of intraspecific variation to diversity maintenance. The aims of this research are to (1) determine the presence or absence of intraspecific alternative phenotypes in three species of tropical tree seedlings, (2) investigate if these different species use the same alternative phenotypes for growth success, and (3) evaluate how findings align with species co-occurrence patterns. We model species-specific relative growth rate with individual-level measurements of leaf mass per area (LMA) and root mass fraction (RMF), environmental data, and their interactions. We find that two of the three species have intraspecific alternative phenotypes, with individuals within species having different functional forms leading to similar growth. Interestingly, individuals within these species use the same trait combinations, high LMA low RMF and low LMA high RMF, in high soil nutrient environments to acquire resources for higher growth. This similarity among species in intraspecific alternative phenotypes and variables that contribute most to growth may lead to their negative spatial co-occurrence. Overall, we find that multiple traits or interactions between traits and the environment drive species-specific strategies for growth, but that individuals within species leverage this multi-dimensionality in different ways for growth success.

Large and thin leaves are compromised more by chewers

AbstractA major proportion of primary production is lost to insect herbivory. Leaf functional traits representing defense costs and palatability explain the intensity of herbivory at local scales. However, whether leaf traits explain herbivory at large scales needs clarification. By combining a leaf herbivory database and the TRY plant‐trait database, we compared 190 species from 65 families across 45 sites in nine countries. We hypothesized that defensive and/or nutrient‐poor leaves would compromise herbivory less globally. Both the absolute and percentage of maximum leaf‐area loss were higher in the tropical zone. Larger leaves lost more disproportionate areas to chewer herbivores whenever phylogenetic signals were considered. Thin leaves with low leaf mass per area (LMA) and short lifespan were compromised more by herbivory based on the leaf trait syndrome. Although thin leaves were rich in nitrogen, phosphorus, and calcium, only leaf nitrogen marginally explained the loss of leaf area. The contribution of leaf area dominated the herbivory intensity among all other morphological and chemical traits. Overall, the benefits of chewers on herbivory were based on leaf area, because the loss of leaf biomass and nitrogen increased proportionally with its loss. Leaf construction costs, but not nutrient content, determined the intensity of herbivory globally. However, the overall percentage of loss in leaf area was not high, demonstrating the importance of elucidating the relationship between leaf functional traits and induced defense in future studies.

Leaf structure and water relations of an allotetraploid Mediterranean fern and its diploid parents.

Allopolyploidy is a common speciation mechanism in plants; however, its physiological and ecological consequences in niche partitioning have been scarcely studied. In this sense, leaf traits are good proxies to study the adaptive capacity of allopolyploids and diploid parents to their respective environmental conditions. In the present work, leaf water relations (assessed through pressure-volume curves) and structural and anatomical traits of the allotetraploid fern Oeosporangium tinaei and its diploid parents, Oeosporangium hispanicum and Oeosporangium pteridioides, were studied under controlled conditions in response to a water stress (WS) cycle. O. hispanicum showed the lowest osmotic potential at turgor loss point (πtlp ) and leaf capacitance, together with higher leaf mass per area (LMA), leaf thickness (LT), leaf density (LD), and leaf dry matter content (LDMC), whereas O. pteridioides presented the opposite set of traits (high πtlp and capacitance, and low LMA, LT, LD, and LDMC). O. tinaei showed an intermediate position for most of the studied traits. The responsiveness (osmotic and elastic adjustments) to WS was low, although most of the traits explained the segregation of the three species across a range of drought tolerance according to the rank: O. hispanicum > O. tinaei > O. pteridioides. These trait differences may underlie the niche segregation among coexisting populations of the three species in the Mediterranean basin.

Implementation of trait-based ozone plant sensitivity in the Yale Interactive terrestrial Biosphere model v1.0 to assess global vegetation damage

Abstract. A major limitation in modeling global ozone (O3) vegetation damage has long been the reliance on empirical O3 sensitivity parameters derived from a limited number of species and applied at the level of plant functional types (PFTs), which ignore the large interspecific variations within the same PFT. Here, we present a major advance in large-scale assessments of O3 plant injury by linking the trait leaf mass per area (LMA) and plant O3 sensitivity in a broad and global perspective. Application of the new approach and a global LMA map in a dynamic global vegetation model reasonably represents the observed interspecific responses to O3 with a unified sensitivity parameter for all plant species. Simulations suggest a contemporary global mean reduction of 4.8 % in gross primary productivity by O3, with a range of 1.1 %–12.6 % for varied PFTs. Hotspots with damage >10 % are found in agricultural areas in the eastern US, western Europe, eastern China, and India, accompanied by moderate to high levels of surface O3. Furthermore, we simulate the distribution of plant sensitivity to O3, which is highly linked with the inherent leaf trait trade-off strategies of plants, revealing high risks for fast-growing species with low LMA, such as crops, grasses, and deciduous trees.

An integrated leaf trait analysis of two Paleogene leaf floras.

This study presents the Integrated Leaf Trait Analysis (ILTA), a workflow for the combined application of methodologies in leaf trait and insect herbivory analyses on fossil dicot leaf assemblages. The objectives were (1) to record the leaf morphological variability, (2) to describe the herbivory pattern on fossil leaves, (3) to explore relations between leaf morphological trait combination types (TCTs), quantitative leaf traits, and other plant characteristics (e.g., phenology), and (4) to explore relations of leaf traits and insect herbivory. The leaves of the early Oligocene floras Seifhennersdorf (Saxony, Germany) and Suletice-Berand (Ústí nad Labem Region, Czech Republic) were analyzed. The TCT approach was used to record the leaf morphological patterns. Metrics based on damage types on leaves were used to describe the kind and extent of insect herbivory. The leaf assemblages were characterized quantitatively (e.g., leaf area and leaf mass per area (LMA)) based on subsamples of 400 leaves per site. Multivariate analyses were performed to explore trait variations. In Seifhennersdorf, toothed leaves of TCT F from deciduous fossil-species are most frequent. The flora of Suletice-Berand is dominated by evergreen fossil-species, which is reflected by the occurrence of toothed and untoothed leaves with closed secondary venation types (TCTs A or E). Significant differences are observed for mean leaf area and LMA, with larger leaves tending to lower LMA in Seifhennersdorf and smaller leaves tending to higher LMA in Suletice-Berand. The frequency and richness of damage types are significantly higher in Suletice-Berand than in Seifhennersdorf. In Seifhennersdorf, the evidence of damage types is highest on deciduous fossil-species, whereas it is highest on evergreen fossil-species in Suletice-Berand. Overall, insect herbivory tends to be more frequently to occur on toothed leaves (TCTs E, F, and P) that are of low LMA. The frequency, richness, and occurrence of damage types vary among fossil-species with similar phenology and TCT. In general, they are highest on leaves of abundant fossil-species. TCTs reflect the diversity and abundance of leaf architectural types of fossil floras. Differences in TCT proportions and quantitative leaf traits may be consistent with local variations in the proportion of broad-leaved deciduous and evergreen elements in the ecotonal vegetation of the early Oligocene. A correlation between leaf size, LMA, and fossil-species indicates that trait variations are partly dependent on the taxonomic composition. Leaf morphology or TCTs itself cannot explain the difference in insect herbivory on leaves. It is a more complex relationship where leaf morphology, LMA, phenology, and taxonomic affiliation are crucial.

Physiological traits and response strategies of four subtropical tree species exposed to drought

World-wide, forests are experiencing drought-induced mortality. However, physiological adaptation to drought is highly variable, which may lead to differential mortality patterns and alter community structure. In this study, saplings of four common tree species ( Ormosia pinnata , Dalbergia odorifera , Castanopsis fissa and Michelia macclurei ) grown in subtropical plantations in southern China were subjected to average rainfall and three drought treatments, consisting of low (25% rainfall reduction), moderate (50% rainfall reduction) and high (75% rainfall reduction) drought. To investigate their drought response strategies and select tree species with higher drought tolerance, we examined plant traits, including tree growth, stomatal regulation, water relations, carbon dynamics and morphology, that might contribute to drought tolerance. We found that O. pinnata adapted to drought by decreasing turgor loss point (TLP) and increasing leaf dry matter content and leaf mass per area (LMA). D. odorifera exhibited low leaf minimum conductance and high LMA as the key drought traits to reduce water loss. C. fissa was slowest to initiate stomatal closure but completed stomatal closure more rapidly than the other species. Drought reduced light-saturated photosynthetic rates (A sat ) for O. pinnata , D. odorifera and C. fissa primarily due to declining stomatal conductance (g s ), with lesser limitations due to the biochemical capacity of Rubisco carboxylation (V cmax ) and electron transport (J max ). M. macclurei was the first species to close stomata. Drought did not affect A sat for M. macclurei due to stable photosynthetic biochemical processes; however, V cmax regulation of A sat increased with drought intensity. There was trait variation in the capacity of study trees to tolerate drought, with M. macclurei exhibiting the highest capacity to tolerate drought due to key stomatal (earlier stomatal closure) and hydraulic (lower TLP) traits reducing plant water loss with progressive drought stress. • Selecting species with morphological and physiological traits that confer high drought tolerance and resistance will be critical for designing and managing plantations. • M. macclurei might be the best adapted plantation tree for growth in future drier climates. • M. macclurei exhibited high drought avoidance and resistance, and maintained photosynthesis.

Intraspecific Leaf Trait Variation across and within Five Common Wine Grape Varieties.

Variability in traits forming the Leaf Economics Spectrum (LES) among and within crop species plays a key role in governing agroecosystem processes. However, studies evaluating the extent, causes, and consequences of within-species variation in LES traits for some of the world’s most common crops remain limited. This study quantified variations in nine leaf traits measured across 90 vines of five common wine grape (Vitis vinifera L.) varieties at two growth stages (post-flowering and veraison). Grape traits in these varieties covary along an intraspecific LES, in patterns similar to those documented in wild plants. Across the five varieties evaluated here, high rates of photosynthesis (A) and leaf nitrogen (N) concentrations were coupled with low leaf mass per area (LMA), whereas the opposite suite of traits defined the “resource-conserving end” of this intraspecific LES in grape. Variety identity was the strongest predictor of leaf physiological (A) and morphological traits (i.e., leaf area and leaf mass), whereas leaf chemical traits and LMA were best explained by growth stage. All five varieties expressed greater resource-conserving trait syndromes (i.e., higher LMA, lower N, and lower Amass) later in the growing season. Traits related to leaf hydraulics, including instantaneous water-use efficiency (WUE), were unrelated to LES and other resource capture traits, and were better explained by spatial location. These results highlight the relative contributions of genetic, developmental, and phenotypic factors in structuring trait variation in the five wine grape varieties evaluated here, and point to a key role of domestication in governing trait relationships in the world’s crops.

Soil Compaction Drives an Intra-Genotype Leaf Economics Spectrum in Wine Grapes

Intraspecific trait variation is a critical determinant of ecosystem processes, especially in agroecosystems where single species or genotypes exist in very high abundance. Yet to date, only a small number of studies have evaluated if, how, or why traits forming the Leaf Economics Spectrum (LES) vary within crops, despite such studies informing our understanding of: (1) the environmental factors that drive crop LES trait variation and (2) how domestication has altered LES traits in crops vs. wild plants. We assess intragenotype variation in LES traits in ‘Chardonnay’ (Vitis vinifera)—one of the world’s most commercially important crops—across a soil compaction gradient: one of the most prominent characteristics of agricultural soils that may drive crop trait variation. Our early evidence indicates that ‘Chardonnay’ traits covary along an intragenotype LES in patterns that are qualitatively similar to those observed among wild plants: resource-acquiring vines expressed a combination of high mass-based photosynthesis (Amass), mass-based dark respiration (Rmass), and leaf nitrogen concentrations (N), coupled with low leaf mass per area (LMA); the opposite set of trait values defined the resource-conserving end of the ‘Chardonnay’ LES. Traits reflecting resource acquisition strategies (Amass, Rmass, and leaf N) declined with greater bulk density, while traits related to investment in leaf construction costs (LMA) increased with greater bulk density. Our findings contribute to an understanding of the domestication syndrome in grapevines and also provide information relevant for quantifying trait-based crop responses to environmental change and gradients.

Leaf habit determines the hydraulic and resource-use strategies in tree saplings from the Sonoran Desert.

The drought susceptibility of woody saplings may explain their low survival in arid environments. Therefore, it is critical to determine which morphological and physiological traits are more responsive to drought among young plants. This study tested whether plant responses to experimental drought differ between two plant functional groups: the deciduous and evergreen species. We predicted that deciduous species would present a tighter stomatal control under drought, coupled with fast carbon fixation under no stress, tending toward isohydry and faster growth rates than the evergreen species. Using 1-year-old saplings from three evergreen and four deciduous Sonoran Desert tree species, we evaluated their hydraulic and gas exchange traits under three experimental irrigation conditions: high, intermediate and low water availability. We measured CO2 assimilation rates (A), stomatal conductance (gs), the level of iso-anisohydry (as the plant's ability to maintain constant their water potential) and seven morphological and growth-related traits throughout 2 months. Under high water availability, saplings reached their maximum values of A and gs, which were significantly higher for deciduous than evergreen species. Correlations among hydroscape area (HA) and leaf traits positioned species along the iso/anisohydric continuum. Deciduous species presented isohydric characteristics, including low HA, high gs, A and Huber values (HVs), and traits indicative of a faster use of resources, such as low stem-specific density (SSD) and low leaf mass per area (LMA). By contrast, evergreen species showed traits that indicate slow resource use and anisohydric behavior, such as high HA, SSD and LMA, and low gs, A and HVs. Deciduous species drastically reduced gas exchange rates in response to drought, while evergreen maintained low rates independently of drought intensity. Overall, desert saplings showed strategies concordant with the iso-anisohydric continuum and the fast-slow use of resources.

The influence of biofertilizers on leaf economics spectrum traits in a herbaceous crop.

Microbial inoculations or 'biofertilizers' represent novel contributions to sustainable agriculture. While belowground mechanisms surrounding how biofertilizers enhance crop production are well described, their role in aboveground trait expression remains less well explored. We quantified infraspecific variation in leaf economics spectrum (LES) traits in response to 10 biofertilizer treatments in basil (Ocimum basiclicum) cultivated under hydroponic conditions. Multiple physiological (i.e. maximum photosynthesis rates (A), dark respiration (R), and leaf-level light compensation points) and morphological (i.e. leaf mass per area (LMA) and leaf thickness) traits varied significantly across microbial treatments. Following treatments, basil plants differentiated from one another along an infraspecific LES, with certain plants expressing more resource-acquiring LES trait values (i.e. high A, R, leaf N, and low LMA), versus others that expressed the opposite suite of resource-conserving LES trait values. Infraspecific trait covariation largely matched LES patterns observed among plants globally. Bivariate and multivariate trait analyses further revealed that certain treatments-namely those including closely related Bacillus and Brevibacillus species strains-increased leaf resource capture traits such as A and leaf N. Biofertilizers influence plant performance through a role in moderating infraspecific leaf trait variation, thereby suggesting aboveground leaf traits may be used to diagnose optimal biofertilizer formulations in basil and other crops.

Flexible drought deciduousness in a neotropical understory herb.

PremiseAdaptive divergence across environmental gradients is a key driver of speciation. Precipitation seasonality gradients are common in the tropics, yet drought adaptation is nearly unexplored in neotropical understory herbs. Here, we examined two recently diverged neotropical spiral gingers, one adapted to seasonal drought and one reliant on perennial water, to uncover the basis of drought adaptation.MethodsWe combined ecophysiological trait measurements in the field and greenhouse with experimental and observational assessments of real‐time drought response to determine how Costus villosissimus (Costaceae) differs from C. allenii to achieve drought adaptation.ResultsWe found that drought‐adapted C. villosissimus has several characteristics indicating flexible dehydration avoidance via semi‐drought‐deciduousness and a fast economic strategy. Although the two species do not differ in water‐use efficiency, C. villosissimus has a more rapid growth rate, lower leaf mass per area, lower stem density, higher leaf nitrogen, and a strong trend of greater light‐saturated photosynthetic rates. These fast economic strategy traits align with both field‐based observations and experimental dry‐down results. During drought, C. villosissimus displays facultative drought‐deciduousness, losing lower leaves during the dry season and rapidly growing new leaves in the wet season.ConclusionsWe revealed a drought adaptation strategy that has not, to our knowledge, previously been documented in tropical herbs. This divergent drought adaptation evolved recently and is an important component of reproductive isolation between C. villosissimus and C. allenii, indicating that adaptive shifts to survive seasonal drought may be an underappreciated axis of neotropical understory plant diversification.

Intra-canopy leaf trait variation facilitates high leaf area index and compensatory growth in a clonal woody encroaching shrub.

Leaf trait variation enables plants to utilize large gradients of light availability that exist across canopies of high leaf area index (LAI), allowing for greater net carbon gain while reducing light availability for understory competitors. While these canopy dynamics are well understood in forest ecosystems, studies of canopy structure of woody shrubs in grasslands are lacking. To evaluate the investment strategy used by these shrubs, we investigated the vertical distribution of leaf traits and physiology across canopies of Cornus drummondii, the predominant woody encroaching shrub in the Kansas tallgrass prairie. We also examined the impact of disturbance by browsing and grazing on these factors. Our results reveal that leaf mass per area (LMA) and leaf nitrogen per area (Na) varied approximately threefold across canopies of C. drummondii, resulting in major differences in the physiological functioning of leaves. High LMA leaves had high photosynthetic capacity, while low LMA leaves had a novel strategy for maintaining light compensation points below ambient light levels. The vertical allocation of leaf traits in C. drummondii canopies was also modified in response to browsing, which increased light availability at deeper canopy depths. As a result, LMA and Na increased at lower canopy depths, leading to a greater photosynthetic capacity deeper in browsed canopies compared to control canopies. This response, along with increased light availability, facilitated greater photosynthesis and resource-use efficiency deeper in browsed canopies compared to control canopies. Our results illustrate how C. drummondii facilitates high LAI canopies and a compensatory growth response to browsing-both of which are key factors contributing to the success of C. drummondii and other species responsible for grassland woody encroachment.

Leaf morpho-physiological comparison between native and non-native plant species in a Mediterranean island

Islands tend to be more prone to plant invasions than mainland regions, with the Mediterranean ones not being an exception. So far, a large number of studies on comparing leaf morphological and physiological traits between native and non-native plants in Mediterranean environments have been performed, although none of them on Mediterranean islands. To fill this gap, this study focuses on 14 plant species grown in a controlled growth chamber in the absence of stress. The goal was (1) to differentiate leaf morpho-physiological traits between native and non-native plants on a Mediterranean island and (2) to deepen in the underlying causes of the differential photosynthetic traits displayed by non-native species. Results showed that in Mediterranean islands, non-native plant species show on average larger values of net CO2 assimilation, stomatal conductance (gm), photosynthetic nitrogen-use efficiency, among others, and lower leaf mass per area (LMA) and leaf thickness, compared to the native species. Among the assessed traits, this study reports for the first time larger gm, and lower mesophyll conductance limitation in non-native species, which seems to be linked to their lower LMA. These novel traits need to be added to the ‘leaf physiological trait invasive syndrome’. It was also found that on a Mediterranean island, native and non-native species are placed on opposite sides of the leaf economics spectrum, with non-native species being placed on the ‘‘fast-return’’ end. In conclusion, this study demonstrates that non-native species inhabiting a Mediterranean island possess distinct leaf morphological and physiological traits compared to co-occurring native species, at least during the favorable growth season, which increases the chances of a successful invasion.

Testing unified theories for ozone response in C4 species.

There is tremendous interspecific variability in O3 sensitivity among C3 species, but variation among C4 species has been less clearly documented. It is also unclear whether stomatal conductance and leaf structure such as leaf mass per area (LMA) determine the variation in sensitivity to O3 across species. In this study, we investigated leaf morphological, chemical, and photosynthetic responses of 22 genotypes of four C4 bioenergy species (switchgrass, sorghum, maize, and miscanthus) to elevated O3 in side‐by‐side field experiments using free‐air O3 concentration enrichment (FACE). The C4 species varied largely in leaf morphology, physiology, and nutrient composition. Elevated O3 did not alter leaf morphology, nutrient content, stomatal conductance, chlorophyll fluorescence, and respiration in most genotypes but reduced net CO2 assimilation in maize and photosynthetic capacity in sorghum and maize. Species with lower LMA and higher stomatal conductance tended to show greater losses in photosynthetic rate and capacity in elevated O3 compared with species with higher LMA and lower stomatal conductance. Stomatal conductance was the strongest determinant of leaf photosynthetic rate and capacity. The response of both area‐ and mass‐based leaf photosynthetic rate and capacity to elevated O3 were not affected by LMA directly but negatively influenced by LMA indirectly through stomatal conductance. These results demonstrate that there is significant variation in O3 sensitivity among C4 species with maize and sorghum showing greater sensitivity of photosynthesis to O3 than switchgrass and miscanthus. Interspecific variation in O3 sensitivity was determined by direct effects of stomatal conductance and indirect effects of LMA. This is the first study to provide a test of unifying theories explaining variation in O3 sensitivity in C4 bioenergy grasses. These findings advance understanding of O3 tolerance in C4 grasses and could aid in optimal placement of diverse C4 bioenergy feedstock across a polluted landscape.

Warming Responses of Leaf Morphology Are Highly Variable among Tropical Tree Species

Leaf morphological traits vary along climate gradients, but it is currently unclear to what extent this results from acclimation rather than adaptation. Knowing so is important for predicting the functioning of long-lived organisms, such as trees, in a rapidly changing climate. We investigated the leaf morphological warming responses of 18 tropical tree species with early (ES) abd late (LS) successional strategies, planted at three sites along an elevation gradient from 2400 m a.s.l. (15.2 °C mean temperature) to 1300 m a.s.l. (20.6 °C mean temperature) in Rwanda. Leaf size expressed as leaf area (LA) and leaf mass per area (LMA) decreased, while leaf width-to-length ratio (W/L) increased with warming, but only for one third to half of the species. While LA decreased in ES species, but mostly not in LS species, changes in LMA and leaf W/L were common in both successional groups. ES species had lower LMA and higher LA and leaf W/L compared to LS species. Values of LMA and LA of juvenile trees in this study were mostly similar to corresponding data on four mature tree species in another elevation-gradient study in Rwanda, indicating that our results are applicable also to mature forest trees. We conclude that leaf morphological responses to warming differ greatly between both successional groups and individual species, with potential consequences for species competitiveness and community composition in a warmer climate.

Leaf Economics of Early- and Late-Successional Plants.

AbstractThe leaf economics spectrum ranges from cheap, short-lived leaves to expensive, long-lived leaves. Species with low leaf mass per area (LMA) and short leaf life span tend to be fast growing and shade intolerant (early successional), whereas species with high LMA and long leaf life span tend to be slow growing and shade tolerant (late successional). However, we have limited understanding of how different leaf mass components (e.g., metabolically active photosynthetic components vs. structural toughness components) contribute to variation in LMA and other leaf economics spectrum traits. Here, we develop a model of plant community dynamics in which species differ in just two traits, photosynthetic and structural LMA components, and we identify optimal values of these traits for early- and late-successional species. Most of the predicted increase in LMA from early- to late-successional species was due to structural LMA. Photosynthetic LMA did not differ consistently between early- and late-successional species, but the photosynthetic LMA to structural LMA ratio declined from early- to late-successional species. Early-successional species had high rates of instantaneous return on leaf mass investment, whereas late-successional species had high lifetime return. Our results provide theoretical support for the primary role of structural (rather than photosynthetic) LMA variation in driving relationships among leaf economics spectrum traits.

Assessing the relationship between trait-based and horticultural classifications of plant responses to drought

Water availability can be a major abiotic constraint for plant success in urban areas. To ensure resilient green spaces, understanding which species are able to cope with low water availability is paramount for plant selection. However, in the horticultural industry, descriptions of species’ responses to low water availability are oftentimes poorly defined, inconsistent across sources, and informed from the natural habitat of species, personal experience, or both. Therefore, a method to objectively assess tolerances of species to low water availability is needed. Using a multivariate approach, we characterised the drought strategies of 113 horticultural plant species spanning five different growth forms (graminoids, herbs, climbers, shrubs and trees) by using five leaf-level traits (leaf mass per area (LMA), leaf thickness, unit leaf area, leaf dry matter content (LDMC) and leaf water potential at turgor loss point (TLP)). Based on this approach, we found that species clustered into three distinct groups, which correspond to fundamental drought strategies based on previous ecological literature (i.e., ‘dehydration avoiders’ (species with high LMA and LDMC, thick leaves and low (more negative) TLP), ‘dehydration tolerators’ (species with low LMA and LDMC and high (less negative) TLP) and an ‘intermediate’ mixture of the two strategies). We found that the majority of species that were considered as ‘drought tolerant’ by the horticultural industry were classified as either ‘dehydration tolerators’ (31.9 %) or ‘dehydration avoiders’ (42.5 %) using our trait-based approach. We did not find any strong connections between the climate of origin of our species and our trait-based classifications. Distinguishing between dehydration avoidance and dehydration tolerance may help to enhance horticultural guidelines by better informing which species are suitable for particular biophysical and landscape contexts. This trait-based approach allows for evidence-based evaluation of drought strategies of new and emerging species and cultivars that may be suitable for urban landscapes.

Deciduous and evergreen oaks show contrasting adaptive responses in leaf mass per area across environments.

Increases in leaf mass per area (LMA) are commonly observed in response to environmental stresses and are achieved through increases in leaf thickness and/or leaf density. Here, we investigated how the two underlying components of LMA differ in relation to species native climates and phylogeny, across deciduous and evergreen species. Using a phylogenetic approach, we quantified anatomical, compositional and climatic variables from 40 deciduous and 45 evergreen Quercus species from across the Northern Hemisphere growing in a common garden. Deciduous species from shorter growing seasons tended to have leaves with lower LMA and leaf thickness than those from longer growing seasons, while the opposite pattern was found for evergreens. For both habits, LMA and thickness increased in arid environments. However, this shift was associated with increased leaf density in evergreens but reduced density in deciduous species. Deciduous and evergreen oaks showed fundamental leaf morphological differences that revealed a diverse adaptive response. While LMA in deciduous species may have diversified in tight coordination with thickness mainly modulated by aridity, diversification of LMA within evergreens appears to be dependent on the infrageneric group, with diversification in leaf thickness modulated by both aridity and cold, while diversification in leaf density is only modulated by aridity.

Photosynthetic and Morphological Acclimation to High and Low Light Environments in Petasites japonicus subsp. giganteus

Within each species, leaf traits such as light-saturated photosynthetic rate or dark respiration rate acclimate to local light environment. Comparing only static physiological traits, however, may not be sufficient to evaluate the effects of such acclimation in the shade because the light environment changes diurnally. We investigated leaf photosynthetic and morphological acclimation for a perennial herb, butterbur (Petasites japonicus (Siebold et Zucc.) Maxim. subsp. giganteus (G.Nicholson) Kitam.) (Asteraceae), in both a well-lit clearing and a shaded understory of a temperate forest. Diurnal changes in light intensity incident on the leaves were also measured on a sunny day and an overcast day. Leaves in the clearing were more folded and upright, whereas leaves in the understory were flatter. Leaf mass per area (LMA) was approximately twofold higher in the clearing than in the understory, while light-saturated photosynthetic rate and dark respiration rate per unit mass of leaf were similar between the sites. Consequently, both light-saturated photosynthetic rate and dark respiration rate per unit area of leaf were approximately twofold higher in the clearing than in the understory, consistent with previous studies on different species. Using this experimental dataset, we performed a simulation in which sun and shade leaves were hypothetically exchanged to investigate whether such plasticity increased carbon gain at each local environment. As expected, in the clearing, the locally acclimated sun leaves gained more carbon than the hypothetically transferred shade leaves. By contrast, in the understory, the daily net carbon gain was similar between the simulated sun and shade leaves on the sunny day due to the frequent sunflecks. Lower LMA and lower photosynthetic capacity in the understory reduced leaf construction cost per area rather than maximizing net daily carbon gain. These results indicate that information on static photosynthetic parameters may not be sufficient to evaluate shade acclimation in forest understories.

The worldwide leaf economic spectrum traits are closely linked with mycorrhizal traits

Abstract Leaf economic spectrum (LES) traits are considered to be the most important plant functional traits, greatly influencing element cycling and ecosystem processes. Mycorrhizal features are considered as part of the root economic spectrum constituting the whole plant economic spectrum, and thus probably linked to LES. Arbuscular mycorrhizas (AMs) are one of the most abundant symbiotic associations in nature, however, the relationship between the global LES traits and AMs remains unknown. Here, based on the global leaf traits database, 1037 plant species with ascertained mycorrhizal types are classified into arbuscular mycorrhizal (AM) and nonAM groups in order to explore the differences in LES traits between AM and nonAM plants. Results show the worldwide LES traits are greatly linked with mycorrhizal traits. For all plants, LES traits vary greatly, but AM plants display larger variations in the six LES traits than nonAM species. AM woody plants, especially trees, have shorter leaf lifespan (LL), lower leaf mass per area (LMA), and higher leaf nitrogen concentration (Nmass), photosynthetic capacity (Amass) and dark respiration rate (Rmass) than nonAM woody ones, but similar leaf phosphorus concentration (Pmass). However, the LES traits of herbaceous plants showed no significant correlation with AMs. Compared to nonAM plants, more AM plants tend to adopt a investment-return strategy. In most cases, LES traits correlate significantly, but their interrelationships vary with mycorrhizal traits. In conclusion, AM plants possess highly varied LES traits and preference for a quick strategy, leading them to a high responsiveness to environmental challenges.