Plant economics spectrum governs leaf nitrogen and phosphorus resorption in subtropical transitional forests.

BACKGROUND: Leaf nitrogen (N) and phosphorus (P) resorption is a fundamental adaptation strategy for plant nutrient conservation. However, the relative roles that environmental factors and plant functional traits play in regulating N and P resorption remain largely unclear, and little is known about the underlying mechanism of plant functional traits affecting nutrient resorption. Here, we measured leaf N and P resorption and 13 plant functional traits of leaf, petiole, and twig for 101 representative broad-leaved tree species in our target subtropical transitional forests. We integrated these multiple functional traits into the plant economics spectrum (PES). We further explored whether and how elevation-related environmental factors and these functional traits collectively control leaf N and P resorption. RESULTS: We found that deciduous and evergreen trees exhibited highly diversified PES strategies, tending to be acquisitive and conservative, respectively. The effects of PES, rather than of environmental factors, dominated leaf N and P resorption patterns along the elevational gradient. Specifically, the photosynthesis and nutrient recourse utilization axis positively affected N and P resorption for both deciduous and evergreen trees, whereas the structural and functional investment axis positively affected leaf N and P resorption for evergreen species only. Specific leaf area and green leaf nutrient concentrations were the most influential traits driving leaf N and P resorption. CONCLUSIONS: Our study simultaneously elucidated the relative contributions of environmental factors and plant functional traits to leaf N and P resorption by including more representative tree species than previous studies, expanding our understanding beyond the relatively well-studied tropical and temperate forests. We highlight that prioritizing the fundamental role of traits related to leaf resource capture and defense contributes to the monitoring and modeling of leaf nutrient resorption. Therefore, we need to integrate PES effects on leaf nutrient resorption into the current nutrient cycling model framework to better advance our general understanding of the consequences of shifting tree species composition for nutrient cycles across diverse forests.

Drought response strategies are coupled with leaf habit in 35 evergreen and deciduous oak (Quercus) species across a climatic gradient in the Americas.

Distinct survival strategies can result from trade-offs in plant function under contrasting environments. Investment in drought resistance mechanisms can enhance survivorship but result in conservative growth. We tested the hypothesis that the widespread oaks (Quercus spp.) of the Americas exhibit an interspecific trade-off between drought resistance and growth capacity. Using experimental water treatments, we isolated adaptive trait associations among species in relation to their broad climates of origin and tested for correlated evolution between plant functional responses to water availability and habitat. Across all lineages, oaks displayed plastic drought responses - typically acclimating through osmolyte accumulation in leaves and/or employing conservative growth. Oaks from xeric climates had higher osmolytes and reduced stomatal pore area index, which allows for moderated gas exchange and limits tissue loss. Patterns suggest drought resistance strategies are convergent and under strong adaptive pressure. Leaf habit, however, mediates the growth and drought resistance strategies of oaks. Deciduous species, and evergreen species from xeric climates, have increased drought tolerance through osmoregulation, which allows for continuous, conservative growth. Evergreen mesic species show limited drought resistance but could enhance growth under well-watered conditions. Consequently, evergreen species from mesic environments are especially vulnerable to chronic drought and climate change.

Ecologically driven selection of nonstructural carbohydrate storage in oak trees.

Leaf habit is a major axis of plant diversity that has consequences for carbon balance since the leaf is the primary site of photosynthesis. Nonstructural carbohydrates (NSCs) produced by photosynthesis can be allocated to storage and serve as a resiliency mechanism to future abiotic and biotic stress. However, how leaf habit affects NSC storage in an evolutionary context has not been shown. Using a comparative physiological framework and an analysis of evolutionary model fitting, we examined if variation in NSC storage is explained by leaf habit. We measured sugar and starch concentrations in 51 oak species (Quercus spp.) growing in a common garden and representing multiple evolutions of three different leaf habits (deciduous, brevideciduous and evergreen). The best fitting evolutionary models indicated that deciduous oak species are evolving towards higher NSC concentrations than their brevideciduous and evergreen relatives. Notably, this was observed for starch (the primary storage molecule) in the stem (a long-term C storage organ). Overall, our work provides insight into the evolutionary drivers of NSC storage and suggests that a deciduous strategy may confer an advantage against stress associated with a changing world. Future work should examine additional clades to further corroborate this idea.

Beyond leaf habit: generalities in plant function across 97 tropical dry forest tree species.

Leaf habit has been hypothesized to define a linkage between the slow-fast plant economic spectrum and the drought resistance-avoidance trade-off in tropical forests ('slow-safe vs fast-risky'). However, variation in hydraulic traits as a function of leaf habit has rarely been explored for a large number of species. We sampled leaf and branch functional traits of 97 tropical dry forest tree species from four sites to investigate whether patterns of trait variation varied consistently in relation to leaf habit along the 'slow-safe vs fast-risky' trade-off. Leaf habit explained from 0% to 43.69% of individual trait variation. We found that evergreen and semi-deciduous species differed in their location along the multivariate trait ordination when compared to deciduous species. While deciduous species showed consistent trait values, evergreen species trait values varied as a function of the site. Last, trait values varied in relation to the proportion of deciduous species in the plant community. We found that leaf habit describes the strategies that define drought avoidance and plant economics in tropical trees. However, leaf habit alone does not explain patterns of trait variation, which suggests quantifying site-specific or species-specific uncertainty in trait variation as the way forward.

Warming induces divergent stomatal dynamics in co-occurring boreal trees.

Climate warming will alter photosynthesis and respiration not only via direct temperature effects on leaf biochemistry but also by increasing atmospheric dryness, thereby reducing stomatal conductance and suppressing photosynthesis. Our knowledge on how climate warming affects these processes is mainly derived from seedlings grown under highly controlled conditions. However, little is known regarding temperature responses of trees growing under field settings. We exposed mature tamarack and black spruce trees growing in a peatland ecosystem to whole-ecosystem warming of up to +9°C above ambient air temperatures in an ongoing long-term experiment (SPRUCE: Spruce and Peatland Responses Under Changing Environments). Here, we report the responses of leaf gas exchange after the first two years of warming. We show that the two species exhibit divergent stomatal responses to warming and vapor pressure deficit. Warming of up to 9°C increased leaf N in both spruce and tamarack. However, higher leaf N in the warmer plots translate into higher photosynthesis in tamarack but not in spruce, with photosynthesis being more constrained by stomatal limitations in spruce than in tamarack under warm conditions. Surprisingly, dark respiration did not acclimate to warming in spruce, and thermal acclimation of respiration was only seen in tamarack once changes in leaf N were considered. Our results highlight how warming can lead to differing stomatal responses to warming in co-occurring species, with consequent effects on both vegetation carbon and water dynamics.

Loss of branches due to winter storms could favor deciduousness in oaks.

PREMISE: Ecologists have an incomplete understanding of the factors that select for deciduous, evergreen, and marcescent leaf habits. Evergreens have more opportunities for photosynthesis but may experience costs when abiotic conditions are unfavorable such as during ice and windstorms. METHODS: We documented branch loss for species of oaks (Quercus spp.) in a common garden in California during an unusual windstorm. RESULTS: Branches of marcescent trees were more likely to break during the storm, and this pattern had a negligible phylogenetic signature. Branches of evergreen and marcescent species were mostly alive before breaking, which likely accrued a fitness cost, while those of deciduous species were mostly already dead. One explanation for the overrepresentation of broken branches from marcescent species is that their petioles are inflexible and have greater wind loading compared to the flexible petioles of evergreen leaves and the leafless condition of deciduous branches. CONCLUSIONS: These results suggest that branch loss during unusual winter storms may be an important cost of a marcescent leaf habit.

Prediction of photosynthetic light-response curves using traits of the leaf economics spectrum for 75 woody species: effects of leaf habit and sun-shade dichotomy.

PREMISE: 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. METHODS: 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). RESULTS: 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. CONCLUSIONS: 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.

Phylogenomic insights into the temporal-spatial divergence history, evolution of leaf habit and hybridization in Stachyurus (Stachyuraceae).

Stachyuraceae, an East Asian endemic family of shrubs or small trees, comprises a single genus, Stachyurus (c. 11 spp.). Despite previous extensive studies based on both morphology and molecular data, species relationships within Stachyurus are still unresolved. Here, we employed transcriptome data aiming to elucidate the evolutionary history of Stachyurus and investigate possible causes of phylogenetic incongruence among individual gene trees in this genus. Our transcriptome phylogeny strongly supports four major clades of Stachyurus, with S. praecox from Japan being resolved as sister to the remainder of the genus on the Asian mainland. The deciduous S. praecox in Japan appears to have originated in the late Miocene, while the remainder diversified and expanded on the mainland over late Miocene to Pliocene/early Pleistocene times. These latter episodes of diversification and expansion were likely promoted by changes in paleoclimate and orogeny (e.g., late Miocene uplift of the Hengduan Mts. and/or enforcement of the East Asian summer monsoon). Species of this genus evolved from a deciduous ancestor, followed by multiple and independent transitions in leaf habit, possibly reflecting climate-related adaptations. Phylogenetic incongruence observed among individual gene trees may be attributable to incomplete lineage sorting following ancient rapid diversification and frequent interspecific gene flow caused by hybridization events. In sum, this study demonstrates the potential usefulness of genome-wide phylogenetic incongruence and network analyses for reconstructing complex evolutionary histories in rapidly diversifying and naturally hybridizing species groups.

Frost and leaf-size gradients in forests: global patterns and experimental evidence.

Explanations of leaf size variation commonly focus on water availability, yet leaf size also varies with latitude and elevation in environments where water is not strongly limiting. We provide the first conclusive test of a prediction of leaf energy balance theory that may explain this pattern: large leaves are more vulnerable to night-time chilling, because their thick boundary layers impede convective exchange with the surrounding air. Seedlings of 15 New Zealand evergreens spanning 12-fold variation in leaf width were exposed to clear night skies, and leaf temperatures were measured with thermocouples. We then used a global dataset to assess several climate variables as predictors of leaf size in forest assemblages. Leaf minus air temperature was strongly correlated with leaf width, ranging from -0.9 to -3.2°C in the smallest- and largest-leaved species, respectively. Mean annual temperature and frost-free period were good predictors of evergreen angiosperm leaf size in forest assemblages, but no climate variable predicted deciduous leaf size. Although winter deciduousness makes large leaves possible in strongly seasonal climates, large-leaved evergreens are largely confined to frost-free climates because of their susceptibility to radiative cooling. Evergreen leaf size data can therefore be used to enhance vegetation models, and to infer palaeotemperatures from fossil leaf assemblages.

Carbon dynamics in the deciduous broadleaf tree Erman's birch (Betula ermanii) at the subalpine treeline on Changbai Mountain, Northeast China.

PREMISE OF THE STUDY: The growth limitation hypothesis (GLH) and carbon limitation hypothesis (CLH) are two dominant explanations for treeline formation. The GLH proposes that low temperature drives the treeline through constraining C sinks more than C sources, and it predicts that non-structural carbohydrate (NSC) levels are static or increase with elevation. Although the GLH has received strong support globally for evergreen treelines, there is still no consensus for deciduous treelines, which experience great asynchrony between supply and demand throughout the year. METHODS: We investigated growth and the growing-season C dynamics in a common deciduous species, Erman's birch (Betula ermanii), along an elevational gradient from the closed forest to the treeline on Changbai Mountain, Northeast China. Samples were collected from developing organs (leaves and twigs) and main storage organs (stems and roots) for NSC analysis. KEY RESULTS: Tree growth decreased with increasing elevation, and NSC concentrations differed significantly among elevations, organs, and sampling times. In particular, NSC levels varied slightly during the growing season in leaves, peaked in the middle of the growing season in twigs and stems, and increased continuously throughout the growing season in roots. NSCs also tended to increase or vary slightly in developing organs but decreased significantly in mature organs with increasing elevation. CONCLUSIONS: The decrease in NSCs with elevation in main storage organs indicates support for the CLH, while the increasing or static trends in new developing organs indicate support for the GLH. Our results suggest that the growth limitation theory may be less applicable to deciduous species' growth than to that of evergreen species.

Convergence, Consilience, and the Evolution of Temperate Deciduous Forests.

The deciduous habit of northern temperate trees and shrubs provides one of the most obvious examples of convergent evolution, but how did it evolve? Hypotheses based on the fossil record posit that deciduousness evolved first in response to drought or darkness and preadapted certain lineages as cold climates spread. An alternative is that evergreens first established in freezing environments and later evolved the deciduous habit. We monitored phenological patterns of 20 species of Viburnum spanning tropical, lucidophyllous (subtropical montane and warm temperate), and cool temperate Asian forests. In lucidophyllous forests, all viburnums were evergreen plants that exhibited coordinated leaf flushes with the onset of the rainy season but varied greatly in the timing of leaf senescence. In contrast, deciduous species exhibited tight coordination of both flushing and senescence, and we found a perfect correlation between the deciduous habit and prolonged annual freezing. In contrast to previous stepwise hypotheses, a consilience of independent lines of evidence supports a lockstep model in which deciduousness evolved in situ, in parallel, and concurrent with a gradual cooling climate. A pervasive selective force combined with the elevated evolutionary accessibility of a particular response may explain the massive convergence of adaptive strategies that characterizes the world's biomes.

Relationships between the timing of budburst, plant traits, and distribution of 24 coexisting woody species in a warm-temperate forest in Japan.

PREMISE OF THE STUDY: Timing of budburst (DBB) may be related to the functional traits and distributions of woody species in temperate regions. Although many previous studies have investigated DBB in a number of temperate species, it has seldom been linked to multiple plant trait relationships. METHODS: DBB and plant traits were investigated for 24 woody species for 2 years in a warm-temperate secondary forest in Japan. Particular attention was paid to differences in trait relationships between coexisting deciduous and evergreen broad-leaved species. KEY RESULTS: DBB was correlated with plant traits in deciduous but not evergreen broad-leaved species; DBB was later for deciduous species with greater leaf mass, leaf area, vessel diameter, and leaf nitrogen content per unit mass. In addition, DBB was later for species with more northern distributions in deciduous and evergreen species. CONCLUSIONS: Clear differences in the trait relationships between deciduous and evergreen broad-leaved species might be caused by different selection pressures on DBB; selection is expected to be more severe in deciduous species. Overall, the continuous variable of vessel diameter might be used as a simple and effective trait to predict DBB of deciduous species regardless of wood anatomy; however, no such traits were detected as effective predictors of DBB in evergreen species at this study site. In addition, DBB was earlier for the species of more southern distributions, suggesting that such species benefit more from warming.

Is the relationship between mast-seeding and weather in oaks related to their life-history or phylogeny?

Although the functional basis of variable and synchronous seed production (masting behavior) has been extensively investigated, only recently has attention been focused on the proximate mechanisms driving this phenomenon. We analyzed the relationship between weather and acorn production in 15 species of oaks (genus Quercus) from three geographic regions on two continents, with the goals of determining the extent to which similar sets of weather factors affect masting behavior across species and to explore the ecological basis for the similarities detected. Lag-1 temporal autocorrelations were predominantly negative, supporting the hypothesis that stored resources play a role in masting behavior across this genus, and we were able to determine environmental variables correlating with acorn production in all but one of the species. Standard weather variables outperformed "differential-cue" variables based on the difference between successive years in a majority of species, which is consistent with the hypothesis that weather is linked directly to the proximate mechanism driving seed production and that masting in these species is likely to be sensitive to climate change. Based on the correlations between weather variables and acorn production, cluster analysis failed to generate any obvious groups of species corresponding to phylogeny or life-history. Discriminant function analyses, however, were able to identify the phylogenetic section to which the species belonged and, controlling for phylogeny, the length of time species required to mature acorns, whether they were evergreen or deciduous, and, to a lesser extent, the geographic region to which they are endemic. These results indicate that similar proximate mechanisms are driving acorn production in these species of oaks, that the environmental factors driving seed production in oaks are to some extent phylogenetically conserved, and that the shared mechanisms driving acorn production result in some degree of synchrony among coexisting species in a way that potentially enhances predator satiation, at least when they have acorns requiring the same length of time to mature.

An experimental approach to explain the southern Andes elevational treeline.

UNLABELLED: • PREMISE OF THE STUDY: The growth limitation hypothesis (GLH) is the most accepted mechanistic explanation for treeline formation, although it is still uncertain whether it applies across taxa. The successful establishment of Pinus contorta--an exotic conifer species in the southern hemisphere--above the Nothofagus treeline in New Zealand may suggest a different mechanism. We tested the GLH in Nothofagus pumilio and Pinus contorta by comparing seedling performance and carbon (C) balance in response to low temperatures.• METHODS: At a southern Chilean treeline, we grew seedlings of both species 2 m above ground level, to simulate coupling between temperatures at the meristem and in the air (colder), and at ground level, i.e., decoupling air temperature (relatively milder). We recorded soil and air temperatures as well. After 3 yr, we measured seedling survival and biomass (as a surrogate of growth) and determined nonstructural carbohydrates (NSC).• KEY RESULTS: Nothofagus and Pinus did not differ in survival, which, as a whole, was higher at ground level than at the 2-m height. The root-zone temperature for the growing season was 6.6°C. While biomass and NSC decreased significantly for Nothofagus at the 2-m height compared with ground level (C limitation), these trends were not significant for Pinus• CONCLUSIONS: The treeline for Nothofagus pumilio is located at an isotherm that fully matches global patterns; however, its physiological responses to low temperatures differed from those of other treeline species. Support for C limitation in N. pumilio but not in P. contorta indicates that the physiological mechanism explaining their survival and growth at treeline may be taxon-dependent.

Non-structural carbohydrate concentrations in tree organs vary across biomes and leaf habits, but are independent of the fast-slow plant economic spectrum.

Carbohydrate reserves play a vital role in plant survival during periods of negative carbon balance. Under a carbon-limited scenario, we expect a trade-offs between carbon allocation to growth, reserves, and defense. A resulting hypothesis is that carbon allocation to reserves exhibits a coordinated variation with functional traits associated with the 'fast-slow' plant economics spectrum. We tested the relationship between non-structural carbohydrates (NSC) of tree organs and functional traits using 61 angiosperm tree species from temperate and tropical forests with phylogenetic hierarchical Bayesian models. Our results provide evidence that NSC concentrations in stems and branches are decoupled from plant functional traits. while those in roots are weakly coupled with plant functional traits. In contrast, we found that variation between NSC concentrations in leaves and the fast-slow trait spectrum was coordinated, as species with higher leaf NSC had trait values associated with resource conservative species, such as lower SLA, leaf N, and leaf P. We also detected a small effect of leaf habit on the variation of NSC concentrations in branches and roots. Efforts to predict the response of ecosystems to global change will need to integrate a suite of plant traits, such as NSC concentrations in woody organs, that are independent of the 'fast-slow' plant economics spectrum and that capture how species respond to a broad range of global change drivers.

Deciduous and evergreen trees differ in juvenile biomass allometries because of differences in allocation to root storage.

BACKGROUND AND AIMS: Biomass partitioning for resource conservation might affect plant allometry, accounting for a substantial amount of unexplained variation in existing plant allometry models. One means of resource conservation is through direct allocation to storage in particular organs. In this study, storage allocation and biomass allometry of deciduous and evergreen tree species from seasonal environments were considered. It was expected that deciduous species would have greater allocation to storage in roots to support leaf regrowth in subsequent growing seasons, and consequently have lower scaling exponents for leaf to root and stem to root partitioning, than evergreen species. It was further expected that changes to root carbohydrate storage and biomass allometry under different soil nutrient supply conditions would be greater for deciduous species than for evergreen species. METHODS: Root carbohydrate storage and organ biomass allometries were compared for juveniles of 20 savanna tree species of different leaf habit (nine evergreen, 11 deciduous) grown in two nutrient treatments for periods of 5 and 20 weeks (total dry mass of individual plants ranged from 0·003 to 258·724 g). KEY RESULTS: Deciduous species had greater root non-structural carbohydrate than evergreen species, and lower scaling exponents for leaf to root and stem to root partitioning than evergreen species. Across species, leaf to stem scaling was positively related, and stem to root scaling was negatively related to root carbohydrate concentration. Under lower nutrient supply, trees displayed increased partitioning to non-structural carbohydrate, and to roots and leaves over stems with increasing plant size, but this change did not differ between leaf habits. CONCLUSIONS: Substantial unexplained variation in biomass allometry of woody species may be related to selection for resource conservation against environmental stresses, such as resource seasonality. Further differences in plant allometry could arise due to selection for different types of biomass allocation in response to different environmental stressors (e.g. fire vs. herbivory).

Carbon use strategies in shoot and acorn growth of two evergreen broadleaf trees unraveled by seasonal carbohydrate measurements and carbon isotope analysis.

Woody species have evolved carbon (C) storage processes that meet needs for reserves associated with asynchronies between C supply and demand. However, our understanding of storage dynamics is still elusive in mature trees, especially when reproduction is involved. Integrated analyses of isotope ratios, concentrations, and biomass may enhance understanding of stored C fractions' dynamics and roles. Thus, we monitored starch and soluble sugars (SSs), C isotope ratios, and biomass, in leaves, twigs and reproductive organs of two mature evergreen broadleaf trees, Quercus glauca and Lithocarpus edulis, for two years. During the growing season, no starch was observed in twigs, while constant starch levels were observed in leaves. Increase in SSs for winter hardening was earlier in L. edulis than in Q. glauca, in line with L. edulis acorns' earlier ripening. Decrease in SSs and increase in starch occurred simultaneously in the next spring. In addition, sucrose accounted for less than 10% of total SSs in leaves of both species, whereas mannose accounted for up to 75% in Q. glauca and myo-inositol up to 23% in L. edulis, indicating species specific sugar composition. These results indicate that seasonal variation of SSs fraction was more reflective to climatic change and NSC storage was less influenced by reproduction. No starch was detected in acorn organs of either Q. glauca or L. edulis except in ripening seeds. The biomass of ripe acorns was 1.7- and 6.4-fold greater than that of current-year twigs in Q. glauca and L. edulis, respectively. Bulk twigs and reproductive organs were ca. 1.0‰ 13C enriched relative to bulk leaves, which was lower than in deciduous trees. These results indicate that new photo-assimilate is the predominant C source for reproductive growth. These findings provide new insights into the dynamics of C storage in relation to reproduction in evergreen broadleaf trees.

Leaf Venation Architecture in Relation to Leaf Size Across Leaf Habits and Vein Types in Subtropical Woody Plants.

Leaves are enormously diverse in their size and venation architecture, both of which are core determinants of plant adaptation to environments. Leaf size is an important determinant of leaf function and ecological strategy, while leaf venation, the main structure for support and transport, determines the growth, development, and performance of a leaf. The scaling relationship between venation architecture and leaf size has been explored, but the relationship within a community and its potential variations among species with different vein types and leaf habits have not been investigated. Here, we measured vein traits and leaf size across 39 broad-leaved woody species within a subtropical forest community in China and analyzed the scaling relationship using ordinary least squares and standard major axis method. Then, we compared our results with the global dataset. The major vein density, and the ratio of major (1° and 2°) to minor (3° and higher) vein density both geometrically declined with leaf size across different vein types and leaf habits. Further, palmate-veined species have higher major vein density and a higher ratio of major to minor vein density at the given leaf size than pinnate-veined species, while evergreen and deciduous species showed no difference. These robust trends were confirmed by reanalyzing the global dataset using the same major vein classification as ours. We also found a tradeoff between the cell wall mass per vein length of the major vein and the major vein density. These vein scaling relationships have important implications on the optimization of leaf size, niche differentiation of coexisting species, plant drought tolerance, and species distribution.

Differentiation in stem and leaf traits among sympatric lianas, scandent shrubs and trees in a subalpine cold temperate forest.

The scandent shrub plant form is a variant of liana that has upright and self-supporting stems when young but later becomes a climber. We aimed to explore the associations of stem and leaf traits among sympatric lianas, scandent shrubs and trees, and the effects of growth form and leaf habit on variation in stem or leaf traits. We measured 16 functional traits related to stem xylem anatomy, leaf morphology and nutrient stoichiometry in eight liana, eight scandent shrub and 21 tree species co-occurring in a subalpine cold temperate forest at an elevation of 2600-3200 m in Southwest China. Overall, lianas, scandent shrubs and trees were ordered along a fast-slow continuum of stem and leaf functional traits, with some traits overlapping. We found a consistent pattern of lianas > scandent shrubs > trees for hydraulically weighted vessel diameter, maximum vessel diameter and theoretical hydraulic conductivity. Vessel density and sapwood density showed a pattern of lianas = scandent shrubs < trees, and lianas < scandent shrubs = trees, respectively. Lianas had significantly higher specific leaf area and lower carbon concentration than co-occurring trees, with scandent shrubs showing intermediate values that overlapped with lianas and trees. The differentiation among lianas, scandent shrubs and trees was mainly explained by variation in stem traits. Additionally, deciduous lianas were positioned at the fast end of the trait spectrum, and evergreen trees at the slow end of the spectrum. Our results showed for the first time clear differentiation in stem and leaf traits among sympatric liana, scandent shrub and tree species in a subalpine cold temperate forest. This work will contribute to understanding the mechanisms responsible for variation in ecological strategies of different growth forms of woody plants.

Highly Species-Specific Foliar Metabolomes of Diverse Woody Species and Relationships with the Leaf Economics Spectrum.

Plants show an extraordinary diversity in chemical composition and are characterized by different functional traits. However, relationships between the foliar primary and specialized metabolism in terms of metabolite numbers and composition as well as links with the leaf economics spectrum have rarely been explored. We investigated these relationships in leaves of 20 woody species from the Mediterranean region grown as saplings in a common garden, using a comparative ecometabolomics approach that included (semi-)polar primary and specialized metabolites. Our analyses revealed significant positive correlations between both the numbers and relative composition of primary and specialized metabolites. The leaf metabolomes were highly species-specific but in addition showed some phylogenetic imprints. Moreover, metabolomes of deciduous species were distinct from those of evergreens. Significant relationships were found between the primary metabolome and nitrogen content and carbon/nitrogen ratio, important traits of the leaf economics spectrum, ranging from acquisitive (mostly deciduous) to conservative (evergreen) leaves. A comprehensive understanding of various leaf traits and their coordination in different plant species may facilitate our understanding of plant functioning in ecosystems. Chemodiversity is thereby an important component of biodiversity.