Coordination of pinna, petiole, and root anatomical traits in 24 tropical-subtropical fern species.

Ferns are primitive vascular plants with diverse morphologies and structures. Plant anatomical traits and their linkages can reflect adaptation to the environment; however, these remain are still poorly understood in ferns. The main objective of this study was to explore whether there was structural coordination among and within organs in fern species. We measured 16 hydraulically related anatomical traits of pinnae, petioles, and roots of 24 representative fern species from the tropical and subtropical forest understory and analyzed trait correlation networks. In addition, we examined phylogenetic signals for the anatomical traits and analyzed co-evolutionary relationships. These results indicated that stomatal density and all petiole anatomical traits exhibited significant phylogenetic signals. Evolutionary correlations were observed between the tracheid diameter and wall thickness of the petiole and between the water transport capacity of the petiole and stomatal density. Conversely, anatomical traits of roots (e.g., root diameter) showed no phylogenetic signals and were not significantly correlated with those of the pinnae and petioles, indicating a lack of structural coordination between the below- and above-ground organs. Unlike angiosperms, vein density is unrelated to stomatal density or pinna thickness in ferns. As root diameter decreased, the cortex-to-stele diameter ratio decreased significantly (enhanced water absorption) in angiosperms but remained unchanged in ferns. These differences lead to different responses of ferns to climate change and improve our knowledge of the water adaptation strategies of ferns.

Correlations between leaf economics, mechanical resistance and drought tolerance across 41 cycad species.

BACKGROUND AND AIMS: We conducted a comprehensive analysis of the functional traits of leaves (leaflets) of cycads. The aim of this study was to clarify the functional divergence between the earlier origin Cycadaceae and the later differentiated Zamiaceae, and the differences in trait associations between cycads and angiosperms. METHODS: We selected 20 Cycadaceae species and 21 Zamiaceae species from the same cycad garden in South China, and measured their leaf structure, economic traits, mechanical resistance (Fp) and leaf water potential at the turgor loss point (πtlp). In addition, we compiled a dataset of geographical distribution along with climatic variables for these cycad species, and some leaf traits of tropical-sub-tropical angiosperm woody species from the literature for comparison. KEY RESULTS: The results showed significantly contrasting leaf trait syndromes between the two families, with Zamiaceae species exhibiting thicker leaves, higher carbon investments and greater Fp than Cycadaceae species. Leaf thickness (LT) and πtlp were correlated with mean climatic variables in their native distribution ranges, indicating their evolutionary adaptation to environmental conditions. Compared with the leaves of angiosperms, the cycad leaves were thicker and tougher, and more tolerant to desiccation. Greater Fp was associated with a higher structural investment in both angiosperms and cycads; however, cycads showed lower Fp at a given leaf mass per area or LT than angiosperms. Enhancement of Fp led to more negative πtlp in angiosperms, but the opposite trend was observed in cycads. CONCLUSIONS: Our results reveal that variations in leaf traits of cycads are mainly influenced by taxonomy and the environment of their native range. We also demonstrate similar leaf functional associations in terms of economics, but different relationships with regard to mechanics and drought tolerance between cycads and angiosperms. This study expands our understanding of the ecological strategies and likely responses of cycads to future climate change.

Co-ordination between leaf biomechanical resistance and hydraulic safety across 30 sub-tropical woody species.

BACKGROUND AND AIMS: Leaf biomechanical resistance protects leaves from biotic and abiotic damage. Previous studies have revealed that enhancing leaf biomechanical resistance is costly for plant species and leads to an increase in leaf drought tolerance. We thus predicted that there is a functional correlation between leaf hydraulic safety and biomechanical characteristics. METHODS: We measured leaf morphological and anatomical traits, pressure-volume parameters, maximum leaf hydraulic conductance (Kleaf-max), leaf water potential at 50 % loss of hydraulic conductance (P50leaf), leaf hydraulic safety margin (SMleaf), and leaf force to tear (Ft) and punch (Fp) of 30 co-occurring woody species in a sub-tropical evergreen broadleaved forest. Linear regression analysis was performed to examine the relationships between biomechanical resistance and other leaf hydraulic traits. KEY RESULTS: We found that higher Ft and Fp values were significantly associated with a lower (more negative) P50leaf and a larger SMleaf, thereby confirming the correlation between leaf biomechanical resistance and hydraulic safety. However, leaf biomechanical resistance showed no correlation with Kleaf-max, although it was significantly and negatively correlated with leaf outside-xylem hydraulic conductance. In addition, we also found that there was a significant correlation between biomechanical resistance and the modulus of elasticity by excluding an outlier. CONCLUSIONS: The findings of this study reveal leaf biomechanical-hydraulic safety correlation in sub-tropical woody species.

Growing-season temperature and precipitation are independent drivers of global variation in xylem hydraulic conductivity.

Stem xylem-specific hydraulic conductivity (KS ) represents the potential for plant water transport normalized by xylem cross section, length, and driving force. Variation in KS has implications for plant transpiration and photosynthesis, growth and survival, and also the geographic distribution of species. Clarifying the global-scale patterns of KS and its major drivers is needed to achieve a better understanding of how plants adapt to different environmental conditions, particularly under climate change scenarios. Here, we compiled a xylem hydraulics dataset with 1,186 species-at-site combinations (975 woody species representing 146 families, from 199 sites worldwide), and investigated how KS varied with climatic variables, plant functional types, and biomes. Growing-season temperature and growing-season precipitation drove global variation in KS independently. Both the mean and the variation in KS were highest in the warm and wet tropical regions, and lower in cold and dry regions, such as tundra and desert biomes. Our results suggest that future warming and redistribution of seasonal precipitation may have a significant impact on species functional diversity, and is likely to be particularly important in regions becoming warmer or drier, such as high latitudes. This highlights an important role for KS in predicting shifts in community composition in the face of climate change.

Leaf hydraulic safety margin and safety-efficiency trade-off across angiosperm woody species.

Leaf hydraulic conductance and the vulnerability to water deficits have profound effects on plant distribution and mortality. In this study, we compiled a leaf hydraulic trait dataset with 311 species-at-site combinations from biomes worldwide. These traits included maximum leaf hydraulic conductance (Kleaf), water potential at 50% loss of Kleaf (P50leaf), and minimum leaf water potential (Ψmin). Leaf hydraulic safety margin (HSMleaf) was calculated as the difference between Ψmin and P50leaf. Our results indicated that 70% of the studied species had a narrow HSMleaf (less than 1 MPa), which was consistent with the global pattern of stem hydraulic safety margin. There was a positive relationship between HSMleaf and aridity index (the ratio of mean annual precipitation to potential evapotranspiration), as species from humid sites tended to have larger HSMleaf. We found a significant relationship between Kleaf and P50leaf across global angiosperm woody species and within each of the different plant groups. This global analysis of leaf hydraulic traits improves our understanding of plant hydraulic response to environmental change.

Leaf mechanical strength and photosynthetic capacity vary independently across 57 subtropical forest species with contrasting light requirements.

Leaf mechanical strength and photosynthetic capacity are critical plant life-history traits associated with tolerance and growth under various biotic and abiotic stresses. In principle, higher mechanical resistance achieved via higher relative allocation to cell walls should slow photosynthetic rates. However, interspecific relationships among these two leaf functions have not been reported. We measured leaf traits of 57 dominant woody species in a subtropical evergreen forest in China, focusing especially on photosynthetic rates, mechanical properties, and leaf lifespan (LLS). These species were assigned to two ecological strategy groups: shade-tolerant species and light-demanding species. On average, shade-tolerant species had longer LLS, higher leaf mechanical strength but lower photosynthetic rates, and exhibited longer LLS for a given leaf mass per area (LMA) or mechanical strength than light-demanding species. Depending on the traits and the basis of expression (per area or per mass), leaf mechanical resistance and photosynthetic capacity were either deemed unrelated, or only weakly negatively correlated. We found only weak support for the proposed trade-off between leaf biomechanics and photosynthesis among co-occurring woody species. This suggests there is considerable flexibility in these properties, and the observed relationships may result more so from trait coordination than any physically or physiologically enforced trade-off.

Shifts in functional trait-species abundance relationships over secondary subalpine meadow succession in the Qinghai-Tibetan Plateau.

Although trait-based processes of community assembly during secondary succession invokes multiple factors that ultimately determine the presence or absence of a species, little is known regarding the impacts of functional traits on species abundance in successional plant communities. Here in species-rich subalpine secondary successional meadows of the Qinghai-Tibetan Plateau, we measured photosynthesis rate and leaf proline content that are related to plant growth and abiotic stress resistance, respectively, and seed germination rate that is closely correlated with plant germination strategy to test their influence on species abundance during succession. We used a linear mixed effects model framework to examine the shifts in trait-abundance relationships and the correlations among these three traits in successional communities. We observed significant shifts in trait-abundance relationships during succession, e.g., abundant species in early-successional meadows exhibited relatively high photosynthesis rates and leaf proline content, but showed low seed germination rates, whereas the converse were true in late successional communities. However, the correlations among the three traits were insignificant in most meadow communities. Our results show that functional traits associated with plant growth, stress resistance, and reproduction impose strong influence on species abundance during secondary subalpine meadow succession in the Qinghai-Tibetan Plateau.

Different leaf cost-benefit strategies of ferns distributed in contrasting light habitats of sub-tropical forests.

BACKGROUND AND AIMS: Ferns are abundant in sub-tropical forests in southern China, with some species being restricted to shaded understorey of natural forests, while others are widespread in disturbed, open habitats. To explain this distribution pattern, we hypothesize that ferns that occur in disturbed forests (FDF) have a different leaf cost-benefit strategy compared with ferns that occur in natural forests (FNF), with a quicker return on carbon investment in disturbed habitats compared with old-growth forests. METHODS: We chose 16 fern species from contrasting light habitats (eight FDF and eight FNF) and studied leaf functional traits, including leaf life span (LLS), specific leaf area (SLA), leaf nitrogen and phosphorus concentrations (N and P), maximum net photosynthetic rates (A), leaf construction cost (CC) and payback time (PBT), to conduct a leaf cost-benefit analysis for the two fern groups. KEY RESULTS: The two groups, FDF and FNF, did not differ significantly in SLA, leaf N and P, and CC, but FDF had significantly higher A, greater photosynthetic nitrogen- and phosphorus-use efficiencies (PNUE and PPUE), and shorter PBT and LLS compared with FNF. Further, across the 16 fern species, LLS was significantly correlated with A, PNUE, PPUE and PBT, but not with SLA and CC. CONCLUSIONS: Our results demonstrate that leaf cost-benefit analysis contributes to understanding the distribution pattern of ferns in contrasting light habitats of sub-tropical forests: FDF employing a quick-return strategy can pre-empt resources and rapidly grow in the high-resource environment of open habitats; while a slow-return strategy in FNF allows their persistence in the shaded understorey of old-growth forests.

Are functional traits a good predictor of global change impacts on tree species abundance dynamics in a subtropical forest?

Significant changes in the composition of tree species have been observed in various forests worldwide. We hypothesised that these changes might result from variable sensitivities of species to global change, and species sensitivities might be quantified, using functional traits. Employing long-term (1978-2010) species abundance data of 48 tree species from a permanent subtropical forest plot, where multiple global change factors have been observed, including soil drying, we examined the relationships between temporal trends in abundance and suits of functional traits. We found that species with high photosynthesis rates, leaf phosphorus and nitrogen concentrations, specific leaf area, hydraulic conductivity, turgor loss point and predawn leaf water potential had increased in abundance, while species with opposite trait patterns had decreased. Our results demonstrate that functional traits underlie tree species abundance dynamics in response to drought stress, thus linking traits to compositional shifts in this subtropical forest under global changes.

Linkages between stem vulnerability curves and tree demography and their implications for plant physiological modeling.

Vulnerability curves (VCs) have been measured extensively to describe the differences in plant vulnerability to cavitation. Although the roles of hydraulic conductivity (Ks,max) and hydraulic safety (P50, embolism resistance), both of which are parameters of VCs ('sigmoidal' type), in tree demography have been evaluated across different forests, the direct linkages between VCs and tree demography are rarely explored. In this study, we combined measured VCs and plot data of 16 tree species in Panamanian seasonal tropical forests to investigate the connections between VCs and tree mortality, recruitment and growth. We found that the mortality and recruitment rates of evergreen species were most significantly positively correlated with P50. However, the mortality and recruitment rates of deciduous species only exhibited significant positive correlations with parameter a, which describes the steepness of VCs and indicates the sensitivity of conductivity loss with water potential decline, but is often neglected. These differences among evergreen and deciduous species may contribute to the poor performance of existing quantitative relationships (such as the fitting relationships for all 16 species) in capturing tree mortality and recruitment dynamics. Additionally, evergreen species presented a significant positive relationship between relative growth rate (RGR) and Ks,max, while deciduous species did not display such relationship. The RGR of both evergreen and deciduous species also displayed no significant correlations with P50 and a. Further analysis demonstrated that species with steeper VCs tended to have high mortality and recruitment rates, while species with flatter VCs were usually those with low mortality and recruitment rates. Our results highlight the important role of parameter a in tree demography, especially for deciduous species. Given that VC is a key component of plant hydraulic models, integrating measured VC rather than optimizing its parameters will help improve the ability to simulate and predict forest response to water availability.

Vein hierarchy mediates the 2D relationship between leaf size and drought tolerance across subtropical forest tree species.

Previous studies have observed a 2D relationship (i.e. decoupled correlation) between leaf size (LS) and leaf economics as well as a tight correlation between leaf economics and drought tolerance. However, the underlying mechanism maintaining the relationship between LS and drought tolerance remains largely unknown. Here, we measured LS, water potential at 50% loss of hydraulic conductance, hydraulic safety margin and different orders of vein traits across 28 tree species in a subtropical forest in Southern China. We found that LS and drought tolerance were in two independent dimensions (R2 = 0.00, P > 0.05). Primary and secondary vein traits (i.e. vein diameter and density) explained the variation of LS, with R2 ranging from 0.37 to 0.70 (all Ps < 0.01), while minor vein traits accounted for the variation of leaf drought tolerance, with R2 ranging from 0.30 to 0.43 (all Ps < 0.01). Our results provide insight into the 2D relationship between LS and drought tolerance and highlight the importance of vein hierarchy in plant leaf functioning.

Water storage capacity is inversely associated with xylem embolism resistance in tropical karst tree species.

Tropical karst habitats are characterized by limited and patchy soil, large rocky outcrops and porous substrates, resulting in high habitat heterogeneity and soil moisture fluctuations. Xylem hydraulic efficiency and safety can determine the drought adaptation and spatial distribution of woody plants growing in karst environments. In this study, we measured sapwood-specific hydraulic conductivity (Ks), vulnerability to embolism, wood density, saturated water content, and vessel and pit anatomical characteristics in the branch stems of 12 evergreen tree species in a tropical karst seasonal rainforest in southwestern China. We aimed to characterize the effects of structural characteristics on hydraulic efficiency and safety. Our results showed that there was no significant correlation between Ks and hydraulic safety across the tropical karst woody species. Ks was correlated with hydraulic vessel diameter (r = 0.80, P < 0.05) and vessel density (r = -0.60, P < 0.05), while the stem water potential at 50 and 88% loss of hydraulic conductivity (P50 and P88) were both significantly correlated with wood density (P < 0.05) and saturated water content (P = 0.052 and P < 0.05, respectively). High stem water storage capacity was associated with low cavitation resistance possibly because of its buffering the moisture fluctuations in karst environments. However, both Ks and P50/P88 were decoupled from the anatomical traits of pit and pit membranes. This may explain the lack of tradeoff between hydraulic safety and efficiency in tropical karst evergreen tree species. Our results suggest that diverse hydraulic trait combination may facilitate species coexistence in karst environments with high spatial heterogeneity.

Plant hydraulics and photosynthesis of 34 woody species from different successional stages of subtropical forests.

It is important to understand the ecophysiological characters of plants when exploring mechanisms underlying species substitution in the process of plant succession. In the present study, we selected 34 woody species from different stages of secondary succession in subtropical forests of southern China, and measured their hydraulic conductivity, gas exchange rates, leaf nutrients and drought-tolerance traits such as xylem resistance to cavitation, turgor loss point and carbon isotope ratio. Principal component analysis revealed that early-, mid- and late-successional species were significantly separated along axis 1, which was strongly associated with hydraulic-photosynthetic coordination. In contrast to species distributed in late-successional forest, early-successional species had the highest hydraulic conductivity, net photosynthetic rates, photosynthetic nitrogen and phosphorus use efficiencies, but had the lowest photosynthetic water-use efficiency. However, changes of the measured drought-tolerance traits of the 34 species along the succession did not demonstrate a clear trend - no significant correlations between these traits and plant successional stages were found. Moreover, the trade-off between hydraulic efficiency and safety was not identified. Taken together, our results suggested that hydraulic efficiency and photosynthetic function, rather than drought tolerance, play an important role in species distributions along plant succession in subtropical forests.

Tree growth is correlated with hydraulic efficiency and safety across 22 tree species in a subtropical karst forest.

Karst forests are habitats in which access to soil water can be challenging for plants. Therefore, safe and efficient xylem water transport and large internal water storage may benefit tree growth. In this study, we selected 22 tree species from a primary subtropical karst forest in southern China and measured their xylem anatomical traits, saturated water content (SWC), hydraulic conductivity (Ks) and embolism resistance (P50). Additionally, we monitored growth of diameter at breast height (DBH) in 440 individual trees of various sizes over three consecutive years. Our objective was to analyze the relationships between xylem structure, hydraulic efficiency, safety, water storage and growth of karst tree species. The results showed significant differences in structure but not in hydraulic traits between deciduous and evergreen species. Larger vessel diameter, paratracheal parenchyma and higher SWC were correlated with higher Ks. Embolism resistance was not correlated with the studied anatomical traits, and no tradeoff with Ks was observed. In small trees (5-15 cm DBH), diameter growth rate (DGR) was independent of hydraulic traits. In large trees (>15 cm DBH), higher Ks and more negative P50 accounted for higher DGR. From lower to greater embolism resistance, the size-growth relationship shifted from growth deceleration to acceleration with increasing tree size in eight of the 22 species. Our study highlights the vital contributions of xylem hydraulic efficiency and safety to growth rate and dynamics in karst tree species; therefore, we strongly recommend their integration into trait-based forest dynamic models.

Can leaf drought tolerance predict species abundance and its changes in tropical-subtropical forests?

Climate change has resulted in an increase in drought severity in the species-rich tropical and subtropical forests of southern China. Exploring the spatiotemporal relationship between drought-tolerance trait and tree abundance provides a means to elucidate the impact of droughts on community assembly and dynamics. In this study, we measured the leaf turgor loss point (πtlp) for 399 tree species from three tropical forest plots and three subtropical forest plots. The plot area was 1 ha and tree abundance was calculated as total basal area per hectare according to the nearest community census data. The first aim of this study was to explore πtlp abundance relationships in the six plots across a range of precipitation seasonality. Additionally, three of the six plots (two tropical forests and one subtropical forest) had consecutive community censuses data (12-22 years) and the mortality ratios and abundance year slope of tree species were analyzed. The second aim was to examine whether πtlp is a predictor of tree mortality and abundance changes. Our results showed that tree species with lower (more negative) πtlp were more abundant in the tropical forests with relative high seasonality. However, πtlp was not related to tree abundance in the subtropical forests with low seasonality. Moreover, πtlp was not a good predictor of tree mortality and abundance changes in both humid and dry forests. This study reveals the restricted role of πtlp in predicting the response of forests to increasing droughts under climate change.

Stem hydraulic traits and leaf water-stress tolerance are co-ordinated with the leaf phenology of angiosperm trees in an Asian tropical dry karst forest.

BACKGROUND AND AIMS: The co-occurring of evergreen and deciduous angiosperm trees in Asian tropical dry forests on karst substrates suggests the existence of different water-use strategies among species. In this study it is hypothesized that the co-occurring evergreen and deciduous trees differ in stem hydraulic traits and leaf water relationships, and there will be correlated evolution in drought tolerance between leaves and stems. METHODS: A comparison was made of stem hydraulic conductivity, vulnerability curves, wood anatomy, leaf life span, leaf pressure-volume characteristics and photosynthetic capacity of six evergreen and six deciduous tree species co-occurring in a tropical dry karst forest in south-west China. The correlated evolution of leaf and stem traits was examined using both traditional and phylogenetic independent contrasts correlations. KEY RESULTS: It was found that the deciduous trees had higher stem hydraulic efficiency, greater hydraulically weighted vessel diameter (D(h)) and higher mass-based photosynthetic rate (A(m)); while the evergreen species had greater xylem-cavitation resistance, lower leaf turgor-loss point water potential (π(0)) and higher bulk modulus of elasticity. There were evolutionary correlations between leaf life span and stem hydraulic efficiency, A(m), and dry season π(0). Xylem-cavitation resistance was evolutionarily correlated with stem hydraulic efficiency, D(h), as well as dry season π(0). Both wood density and leaf density were closely correlated with leaf water-stress tolerance and A(m). CONCLUSIONS: The results reveal the clear distinctions in stem hydraulic traits and leaf water-stress tolerance between the co-occurring evergreen and deciduous angiosperm trees in an Asian dry karst forest. A novel pattern was demonstrated linking leaf longevity with stem hydraulic efficiency and leaf water-stress tolerance. The results show the correlated evolution in drought tolerance between stems and leaves.

Functional Traits Are Good Predictors of Tree Species Abundance Across 101 Subtropical Forest Species in China.

What causes variation in species abundance for a given site remains a central question in community ecology. Foundational to trait-based ecology is the expectation that functional traits determine species abundance. However, the relative success of using functional traits to predict relative abundance is questionable. One reason is that the diversity in plant function is greater than that characterized by the few most commonly and easily measurable traits. Here, we measured 10 functional traits and the stem density of 101 woody plant species in a 200,000 m2 permanent, mature, subtropical forest plot (high precipitation and high nitrogen, but generally light- and phosphorus-limited) in southern China to determine how well relative species abundance could be predicted by functional traits. We found that: (1) leaf phosphorus content, specific leaf area, maximum CO2 assimilation rate, maximum stomata conductance, and stem hydraulic conductivity were significantly and negatively associated with species abundance, (2) the ratio of leaf nitrogen content to leaf phosphorus content (N:P) and wood density were significantly positively correlated with species abundance; (3) neither leaf nitrogen content nor leaf turgor loss point were related to species abundance; (4) a combination of N:P and maximum stomata conductance accounted for 44% of the variation in species' abundances. Taken together, our findings suggested that the combination of these functional traits are powerful predictors of species abundance. Species with a resource-conservative strategy that invest more in their tissues are dominant in the mature, subtropical, evergreen forest.

Coordinated variation in stem and leaf functional traits of temperate broadleaf tree species in the isohydric-anisohydric spectrum.

Stomatal regulation serves as an important strategy for plants to adapt to drought. However, the understanding of how complexes of plant-functional traits vary along the continuum from isohydry to anisohydry remains insufficient. In this study, we investigated a proxy of the degree of iso/anisohydry-the water potential at stomatal closure-and a series of functional traits of leaves and branches in 20 temperate broadleaf species planted in an arid limestone habitat in northern China. The results showed that the water potential at stomatal closure was significantly correlated with many functional traits. At the anisohydric end of the spectrum, species had a higher leaf carbon content and vein density, a greater stomatal length, a thicker lower leaf epidermis, higher embolism resistance, higher wood density, a greater Huber value, a greater ratio of fiber wall thickness to xylem lumen diameter, a larger proportion of total fiber wall area to xylem cross-sectional area, a lower water potential at the turgor loss point (TLP), a smaller relative water content at the TLP, a lower osmotic potential at full turgor and a smaller specific leaf area. It is concluded that a continuum of coordination and trade-offs among co-evolved anatomical and physiological traits gives rise to the spectrum from isohydry to anisohydry spanned by the 20 tree species, and the anisohydric species showed stronger stress resistance, with greater investment in stems and leaves than the isohydric species to maintain stomatal opening under drought conditions.

Contrasting cost-benefit strategy between lianas and trees in a tropical seasonal rain forest in southwestern China.

Lianas are an important component of tropical forests and often abundant in open habitats, such as tree-fall gaps, forest edges, and disturbed forests. The abundance of lianas in tropical forests has been increasing as a result of global environmental change and increasing forest fragmentation. In order to understand this phenomenon in terms of leaf functional traits and to evaluate their competitive potential, we conducted a cost-benefit analysis of leaves from 18 liana species and 19 tree species in a tropical seasonal rain forest. The results revealed that lianas were scattered in a group distinct from trees along the first axis of a principal component analysis using 15 leaf ecophysiological traits, being located at the quick-return end of the leaf economics spectrum, with higher specific leaf area and photosynthetic rates (A), higher photosynthetic nitrogen (N) and phosphorus (P) use efficiencies, a lower leaf construction cost per unit leaf area (CC) and cost-benefit ratio (CC/A), and a shorter leaf life span (LLS). Trees showed the opposite trends. The results indicate that lianas can grow faster and capture resources more efficiently than trees in disturbed, open habitats. The positive relationship between LLS and CC/A revealed a trade-off between leaf construction cost and benefit over time. The 37 species analyzed had a mean foliar N/P ratio of 20, indicating that the forest was characterized by a P deficit. With an increasing atmospheric CO(2) concentration, the higher nutrient use efficiency could benefit lianas more than trees in terms of productivity, possibly also contributing to the increasing abundance of lianas in nutrient-limited tropical forests.

Tradeoff between storage capacity and embolism resistance in the xylem of temperate broadleaf tree species.

Xylem traits are critical plant functional traits associated with water transport, mechanical support, and carbohydrate and water storage. Studies on the xylem hydraulic efficiency-safety tradeoff are numerous; however, the storage function of xylem parenchyma is rarely considered. The effects of a substantial number of xylem traits on water transport, embolism resistance, mechanical support, storage capacity and nonstructural carbohydrate (NSC) content were investigated in 19 temperate broadleaf species planted in an arid limestone habitat in northern China. There was no xylem hydraulic efficiency-safety tradeoff in the 19 broadleaf species. The total parenchyma fraction was negatively correlated with the fiber fraction. Embolism resistance was positively correlated with indicators of xylem mechanical strength such as vessel wall reinforcement, vessel wall thickness and fiber wall thickness, and was negatively related to the axial parenchyma fraction, especially the paratracheal parenchyma fraction. The paratracheal parenchyma fraction was positively correlated with the ratio of the paratracheal parenchyma fraction to the vessel fraction. In addition, the xylem NSC concentration was positively related to the total parenchyma fraction and axial parenchyma fraction. There was a storage capacity-embolism resistance tradeoff in the xylem of 19 broadleaf species in arid limestone habitats. We speculate that the temperate broadleaf species may show a spectrum of xylem hydraulic strategies, from the embolism resistance strategy related to a more negative P50 (the water potential corresponding to 50% loss of xylem conductivity) to the embolization repair strategy based on more paratracheal parenchyma.