Leaf trait variations associated with habitat affinity of tropical karst tree species

Karst hills, that is, jagged topography created by dissolution of limestone and other soluble rocks, are distributed extensively in tropical forest regions, including southern parts of China. They are characterized by a sharp mosaic of water and nutrient availability, from exposed hilltops with poor soil development to valleys with occasional flooding, to which trees show species‐specific distributions. Here we report the relationship of leaf functional traits to habitat preference of tropical karst trees. We described leaf traits of 19 tropical tree species in a seasonal karst rainforest in Guangxi Province, China, 12 species in situ and 13 ex situ in a non‐karst arboretum, which served as a common garden, with six species sampled in both. We examined how the measured leaf traits differed in relation to species’ habitat affinity and evaluated trait consistency between natural habitats vs. the arboretum. Leaf mass per area (LMA) and optical traits (light absorption and reflectance characteristics between 400 and 1,050 nm) showed significant associations with each other and habitats, with hilltop species showing high values of LMA and low values of photochemical reflectance index (PRI). For the six species sampled in both the karst forest and the arboretum, LMA, leaf dry matter content, stomatal density, and vein length per area showed inconsistent within‐species variations, whereas some traits (stomatal pore index and lamina thickness) were similar between the two sites. In conclusion, trees specialized in exposed karst hilltops with little soils are characterized by thick leaves with high tissue density indicative of conservative resources use, and this trait syndrome could potentially be sensed remotely with PRI.     

Reflectance and transmittance spectra of leaves and shoots of 22 vascular plant species and reflectance spectra of trunks and branches of 12 tree species in Japan

This data paper reports spectral reflectance and transmittance data of leaves from 21 terrestrial vascular plant species (seven herbaceous, and 14 broadleaf and long-needle coniferous tree species) and of shoots from one short-needle coniferous tree species. The reflectance spectra of branches of one tree species, of the trunks of 12 tree species and ground surface of one deciduous broad-leaf forest are also reported. Optical measurements and leaf samplings were made at five sites on Honshu Island, Japan, which are typical vegetation types in East Asia, i.e., grassland, paddy field, and deciduous broad-leaf or coniferous forests. The collection and measurements were conducted for main species in each site. To include other common vegetation types in East Asia, such as evergreen broad-leaf or coniferous forests, the sample collection and the measurements were conducted at gardens and an experimental forest. Leaves of ten deciduous species were measured at different phenological stages from leaf expansion to senescence since those species shows significant seasonal changes in spectral reflectance and transmittance of leaves. Leaves at different position in a canopy (e.g., sunlit versus shaded leaves) were also measured for eight of 21 species. The spectral reflectance and transmittance from both adaxial and abaxial sides of the all leaves or needles, expect Picea abies needles. The measurements of the leaves were conducted with a spectroradiometer attached via an optical fiber to an integrating sphere. Two types of integrating spheres were used: a model LI-1800-12 (Li-Cor) and an RTS-3ZC integrating sphere (Analytical Spectral Devices). A leaf clip accessory was also used instead of an integrating sphere for measuring the leaves of two species. All data were measured within the 350–2,500-nm spectral range with 1-nm steps between measurements but the data obtained by LI-1800 is unavailable in 1,650–1,740, 1,890–1,950, and 2,050–2,500 nm because of a large amount of noise. These data are used as input parameters in a radiative transfer model designed to estimate the leaf area index from radiation reflected from a canopy surface.     

Ecological distribution of homobaric and heterobaric leaves in tree species of Malaysian lowland tropical rainforest

Tree species can generally be classified into two groups, heterobaric and homobaric leafed species, according to whether bundle-sheath extensions (BSEs) are found in the leaf (heterobaric leaf) or not (homobaric leaf). In this study, we study whether the leaf type is related to the growth environment and/or life form type, even in a tropical rain forest, where most trees have evergreen leaves that are generally homobaric. Accordingly, we investigated the distribution of leaf morphological differences across different life forms of 250 tree species in 45 families in a tropical rainforest. In total, 151 species (60%) in 36 families had homobaric leaves, and 99 species (40%) in 21 families had heterobaric leaves. We found that the proportion of heterobaric and homobaric leaf species differed clearly across taxonomic groups and life form types, which were divided into five life form types by their mature tree heights (understory, subcanopy, canopy, and emergent species) and as canopy gap species. Most understory (94%) and subcanopy (83%) species such as Annonaceae had homobaric leaves. In contrast, heterobaric leaf trees appeared more frequently in the canopy species (43%), the emergent species (96%) (such as Dipterocarpaceae), and the canopy gap species (62%). Our results suggest that tree species in the tropical rainforest adapt to spatial differences in the environmental conditions experienced at the mature height of each tree species, such as light intensity and vapor pressure difference, by having differing leaf types (heterobaric or homobaric) because these types potentially have different physiological and/or mechanical functions.     

A comparison of spectral reflectance indices in response to water: A case study of Quercus aliena var. acuteserrata

Aims Using leaf spectral reflectance to detect plant status in real time and non-destructively is a new method of forest drought assessment, but each spectral index possesses considerable moisture sensitivity. Therefore, determining moisture index applicable to tree leaf and its sensitive spectral index are both very important. Methods This study selected Quercus aliena var. acuteserrata leaves in different growth stages and canopy positions as the research object, and measured leaf moisture index and its synchronous reflectance spectral response curve during the dehydration process, explored the relationship between changes of leaf spectral reflectance and water status, compared and evaluated the advantages and disadvantages of correlation between the moisture indices of leaves in different growth stages and space positions and different spectral reflectance indices. Important findings Results indicated: (1) The variability of relative water content (RWC) and equivalent water thickness (EWT) in different growth stages and canopy positions was smaller than specific leaf water content (SWC) and leaf moisture percentage on fresh quality (LMP) as measured by the four different moisture indices. RWC and EWT could steadily characterize the holistic water status of trees, and they had greater spectral sensitivity. Therefore, they were suitable for application in remote sensing detection. (2) Spectral reflectance difference analysis and spectral reflectance sensitivity analysis showed that the leaf spectral sensitivity is strongly influenced by growth stage. In short wave infrared region, spectral reflectance of mature leaves changed slightly in the initial stage of dehydration stress, but new expended leaves showed obvious spectral differences during the dehydration process. (3) Through the correlation analysis between 15 different spectral indices and moisture indices, we found that water index (WI)-RWC and double difference index (DD_{n (1530,525)})-EWT has higher correlation. The fitted relations of WI-RWC are greatly influenced by leaf growth stage and canopy position, while those of DD_n(1530,525)- EWT are more stable.     

Leaf optical properties in higher plants: linking spectral characteristics to stress and chlorophyll concentration

A number of studies have linked responses in leaf spectral reflectance, transmittance, or absorptance to physiological stress. A variety of stressors including dehydration, flooding, freezing, ozone, herbicides, competition, disease, insects, and deficiencies in ectomycorrhizal development and N fertilization have been imposed on species ranging from grasses to conifers and deciduous trees. In all cases, the maximum difference in reflectance within the 400-850 nm wavelength range between control and stressed states occurred as a reflectance increase at wavelengths near 700 nm. In studies that included transmittance and absorptance as well as reflectance, maximum differences occurred as increases and decreases, respectively, near 700 nm. This common optical response to stress could be simulated closely by varying the chlorophyll concentration of model leaves (fiberglass filter pads) and by the natural variability in leaf chlorophyll concentrations in senescent leaves of five species. The optical response to stress near 700 nm, as well as corresponding changes in reflectance that occur in the green-yellow spectrum, can be explained by the general tendency of stress to reduce leaf chlorophyll concentration.     

The expression of light-related leaf functional traits depends on the location of individual leaves within the crown of isolated Olea europaea trees

Background The spatial arrangement and expression of foliar syndromes within tree crowns can reflect the coupling between crown form and function in a given environment. Isolated trees subjected to high irradiance and concomitant stress may adjust leaf phenotypes to cope with environmental gradients that are heterogeneous in space and time within the tree crown. The distinct expression of leaf phenotypes among crown positions could lead to complementary patterns in light interception at the crown scale.Methods We quantified eight light-related leaf traits across 12 crown positions of ten isolated Olea europaea trees in the field. Specifically, we investigated whether the phenotypic expression of foliar traits differed among crown sectors and layers and five periods of the day from sunrise to sunset. We investigated the consequences in terms of the exposed area of the leaves at the tree scale during a single day.Key Results All traits differed among crown positions except the length-to-width ratio of the leaves. We found a strong complementarity in the patterns of the potential exposed area of the leaves among day periods as a result of a non-random distribution of leaf angles across the crown. Leaf exposure at the outer layer was below 60 % of the displayed surface, reaching maximum interception during morning periods. Daily interception increased towards the inner layer, achieving consecutive maximization from east to west positions within the crown, matching the sun’s trajectory.Conclusions The expression of leaf traits within isolated trees of O. europaea varies continuously through the crown in a gradient of leaf morphotypes and leaf angles depending on the exposure and location of individual leaves. The distribution of light-related traits within the crown and the complementarity in the potential exposure patterns of the leaves during the day challenges the assumption of low trait variability within individuals.     

Leaf spectral signatures differ in plant species colonizing habitats along a hydrological gradient

Aims We aimed at determining differences in the leaf spectral signatures of plant species groups growing in habitats along the hydrological gradient of an intermittent wetland and to define leaf traits that explain their variability. We want to contribute to the understanding of the causes for plant spectrum variability at leaf and community levels.Methods We measured leaf reflectance spectra (300–887nm) of representative plant species from different habitats and analyzed spectral differences among species groups. To explain leaf spectra variability within a group, we performed detailed analyses of leaf morphological and biochemical traits in selected species.Important findings The reflectance spectra of the different species groups differed most in the green, yellow and red spectral ranges. The reflectance spectra of submerged leaves of hydrophytes with simple structures were explained by their biochemical traits (carotenoids), while for more complex aerial leaves, morphological traits were more important. In submerged and natant leaves of amphiphytes, total mesophyll and spongy tissue thickness were the most important traits, and these explained 44% and 47%, respectively, of the spectrum variability of each plant group. In general, the redundancy analysis biplots show that samples of different plant species colonizing the same habitat form separate clusters and are related to the explanatory variables in different ways. The redundancy analysis biplots of helophytes and wet meadow species show clustering of graminoids and dicots into two distinct groups. Leaf encrustation (prickle hair properties and epidermis thickness) is important for graminoids, while leaf thickness and specific leaf area have more important roles in dicots. Our results show that knowledge of the species composition and leaf traits is necessary to interpret the reflectance spectra of such plant communities.     

Is there a temporal niche separation in the leaf phenology of savanna trees and grasses?

Aim It has been proposed that, in tropical savannas, trees deploy their leaves earlier in the growing season and grasses deploy their leaves later. This hypothesis implies a mechanism that facilitates the coexistence of trees and grasses in savannas. If true, this hypothesis would also allow algorithms to use differences in the phenological timing of grass and tree leaves to partition the relative contribution of grasses and trees to net primary production. In this study we examine whether a temporal niche separation between grasses and trees exists in savanna. Location A semi‐arid, subtropical savanna, Kruger National Park, South Africa. Methods We use a multi‐spectral camera to track through an entire growing season the normalized difference vegetation index (NDVI) of individual canopies of grasses and trees at eight sites arranged along a precipitation and temperature gradient. Results Among trees, we identified two distinct phenological syndromes: an early flushing syndrome and a late‐flushing syndrome. Leaf flush in the tree strategies appears to pre‐empt rainfall, whereas grass leaf flush follows the rain. The growing season of trees is 20 (late‐flushing trees) to 27 (early flushing trees) days longer than that of the grasses. Main conclusions We show that grasses and trees have different leaf deployment strategies. Trees deployed leaves at lower temperatures than grasses and retained them for longer at the end of the growing season. The timing of the increase in NDVI is, however, similar between grasses and late‐flushing trees and this complicates the separation of grass and tree signals from multi‐spectral satellite imagery.     

Variation in crown light utilization characteristics among tropical canopy trees

BACKGROUND AND AIMS: Light extinction through crowns of canopy trees determines light availability at lower levels within forests. The goal of this paper is the exploration of foliage distribution and light extinction in crowns of five canopy tree species in relation to their shoot architecture, leaf traits (mean leaf angle, life span, photosynthetic characteristics) and successional status (from pioneers to persistent). METHODS: Light extinction was examined at three hierarchical levels of foliage organization, the whole crown, the outermost canopy and the individual shoots, in a tropical moist forest with direct canopy access with a tower crane. Photon flux density and cumulative leaf area index (LAI) were measured at intervals of 0.25-1 m along multiple vertical transects through three to five mature tree crowns of each species to estimate light extinction coefficients (K). RESULTS: Cecropia longipes, a pioneer species with the shortest leaf life span, had crown LAI <0.5. Among the remaining four species, crown LAI ranged from 2 to 8, and species with orthotropic terminal shoots exhibited lower light extinction coefficients (0.35) than those with plagiotropic shoots (0.53-0.80). Within each type, later successional species exhibited greater maximum LAI and total light extinction. A dense layer of leaves at the outermost crown of a late successional species resulted in an average light extinction of 61% within 0.5 m from the surface. In late successional species, leaf position within individual shoots does not predict the light availability at the individual leaf surface, which may explain their slow decline of photosynthetic capacity with leaf age and weak differentiation of sun and shade leaves. CONCLUSION: Later-successional tree crowns, especially those with orthotropic branches, exhibit lower light extinction coefficients, but greater total LAI and total light extinction, which contribute to their efficient use of light and competitive dominance.     

Relationships between endophyte diversity and leaf optical properties

A single tropical plant species can harbour hundreds of endophyte species within its tissues. Beyond this, little is known about the relationship between endophyte colonization, leaf traits and spectral properties of leaves. We explore these relationships in Coccoloba cereifera, a plant well known for its symbiotic properties. Endophyte richness in C. cereifera was statistically correlated with leaf traits such as water content, the ratio of fresh weight/dry weight and polyphenol/leaf specific weight. Endophyte diversity was also related to spectral vegetation indices of chlorophyll content. The associations among endophyte diversity, leaf traits and spectral reflectance pose new questions and present new opportunities to better understand plant–fungal symbioses and related leaf optical properties.     

Increased leaf reflectance in tropical trees under elevated CO2

Globally increasing atmospheric CO₂ concentrations are known to affect many aspects of plant physiology and development; however, little attention has been given to leaf and canopy optical properties. Three tropical trees in the Leguminosae, an important canopy tree family in many tropical forests, responded similarly to an experimental doubling of CO₂ partial pressure with a 9–23% increase in spectral leaf reflectance to light in the visible (400–700 nm) waveband. Decreased leaf chlorophyll content under elevated CO₂ may explain part of the observed increase in reflectance. However, analyses that statistically corrected for chlorophyll content effects on reflectance still indicated a significant CO₂ effect. This results, in conjunction with the spectral pattern of the response, suggests that the primary mechanism is increased optical masking of chlorophyll under elevated CO₂. The magnitude of the increase in leaf reflectance is sufficient to suggest that increased canopy reflectance of tropical forests (and possibly other terrestrial ecosystems) may be an important negative feedback in the response of global net radiative climate forcing to increasing atmospheric CO₂.     

Functional relationship between chlorophyll content and leaf reflectance, and light-capturing efficiency of Japanese forest species

We examined the functional relationship between chlorophyll concentrations and light spectral absorption in 16 species of woody, vine and herbaceous plants in northern Japan. Leaves of each species from under forest shade and in more open sites were measured for chlorophyll, specific leaf area (SLA) and spectral absorption. In all species, SLA increased and the Chl a : b ratio declined in shade- vs open-grown leaves indicating an adaptive adjustment to forest shade in these leaf characters. However, the expected increase in the ratio of 680 to 700 nm absorption in shade leaves did not occur in all species. Light absorption at 680 relative to 700 nm was lower in the shade leaves of Acer japonicum. Kalopanax pictus, Panax japonicus and Petasites japonicus even with a reduced Chl a : b , a commonly accepted indicator of shade adaptation. Therefore, spectral measurements in these species failed to support Chl concentrations that were expected to confer an improvement in the absorption of red light (<680nm) deficient relative to far-red light (>700 nm) in the forest shade. Compared with other species, the absorption pattern of these four 'non-conforming' species is associated with a higher ratio of shade:open leaves in reflectance spectra in the 600–750 nm range. This suggests an increased reflectance in shade leaves caused by changes in leaf surface properties which are not immediately apparent. We conclude that adaptive spectral absorption cannot always be inferred from changes in specific leaf area and chlorophyll a and b concentrations.     

Canopy cover and leaf age affect colonization by tropical fungal endophytes: Ecological pattern and process in Theobroma cacao (Malvaceae)

Fungal endophytes inhabit healthy tissues of all terrestrial plant taxa studied to date and are diverse and abundant in leaves of tropical woody angiosperms. Studies have demonstrated that plant location and leaf age influence density of endophyte infection in leaves of tropical forest trees. However, ecological factors underlying these observations have not been explored in detail. Here, we establish that foliar endophytes of a tropical tree (Theobroma cacao, Malvaceae) are transmitted horizontally and that endophyte-free seedlings can be produced for experimental manipulation by protecting aerial tissues from surface wetting. At Barro Colorado Island, Panama, we used transects of endophyte-free seedlings to determine the importance of several factors (canopy cover, abundance of aerial and epiphytic propagules, leaf age, leaf chemistry, leaf toughness and duration of exposure to viable air spora) in shaping colonization by endophytic fungi. Endophytes colonized leaves of T. cacao more rapidly beneath the forest canopy than in cleared sites, reflecting local abundance of aerial and epiphytic propagules. The duration of exposure, rather than absolute leaf age, influenced endophyte infection, whereas leaf toughness and chemistry had no observed effect. Endophytes isolated from mature T. cacao grew more rapidly on media containing leaf extracts of T. cacao than on media containing extracts from other co-occurring tree species, suggesting that interspecific differences in leaf chemistry influence endophyte assemblages. Together, these data allow us to identify factors underlying patterns of endophyte colonization within healthy leaves of this tropical tree.     

Variability in leaf optical properties among 26 species from a broad range of habitats

Leaves from 26 species with growth forms from annual herbs to trees were collected from open, intermediate, and shaded understory habitats in Mississippi and Kansas, USA. Leaf optical properties including reflectance, transmittance, and absorptance in visible and near infrared (NIR) wavelengths were measured along with leaf thickness and specific leaf mass (SLM). These leaf properties and internal light scattering have been reported to vary with light availability in studies that have focused on a limited number of species. Our objective was to determine whether these patterns in leaf optics and light availability were consistent when a greater number of species were evaluated. Leaf thickness and SLM varied by tenfold among species sampled, but within-habitat variance was high. Although there was a strong trend toward thicker leaves in open habitats, only SLM was significantly greater in open vs. understory habitats. In contrast, leaf optical properties were strikingly similar among habitats. Reflectance and reflectance/transmittance in the NIR were used to estimate internal light scattering and there were strong relationships (r1 > 0.65) between these optical properties and leaf thickness. We concluded that leaf thickness, which did not vary consistently among habitats, was the best predictor of NIR reflectance and internal light scattering. However, because carbon allocation to leaves was lower in understory species (low SLM) yet gross optical properties were similar among all habitats, the energy investment by shade leaves required to achieve optical equivalence with sun leaves was lower. Differences in leaf longevity and growth form within a habitat may help explain the lack of consistent patterns in leaf optics as the number of species sampled increases.     

Plant growth-form alters the relationship between foliar morphology and species shade-tolerance ranking in temperate woody taxa

Variation in leaf size (area per leaf) and leaf dry weight per area (LWA) in relation to species shade- and drought-tolerance, characterised by Ellenberg's light (ELD) and water demand (EWD) values, respectively, were examined in 60 temperate woody taxa at constant relative irradiance. LWA was independent of plant size, but leaf size increased with total plant height at constant ELD. Canopy position also affected leaf morphology: leaves from the upper crown third had higher LWA and were larger than leaves from the lower third. Leaf size and LWA were negatively correlated, and leaf size decreased and LWA increased with decreasing species shade-tolerance. Mean LWA was similar for trees and shrubs, but trees had larger leaves than shrubs. Furthermore, all relationships were altered by plant growth-form: none of the qualitative tendencies was significant for trees. This implies the considerably lower plasticity of foliar parameters in trees than those in shrubs. Accordingly, shade-tolerance of trees, having relatively constant leaf structure, may be most affected by the variability in biomass partitioning and crown geometry which influence foliage distribution and spacing and finally determine canopy light absorptance. Alteration of leaf form and investment pattern for construction of unit foliar surface area which change the efficiency of light interception per unit biomass investment in leaves, is a competitive strategy inherent to shrubs. EWD as well as wood anatomy did not control LWA and leaf size, though there was a trend of ring-porous tree species to be more shade-tolerant than diffuse-porous trees. Since ring-porous species are more vulnerable to cavitation than diffuse-porous species, they may be constrained to environments where irradiances and consequently evaporative demand is lower.     

Leaf litter inputs reinforce islands of nitrogen fertility in a lowland tropical forest

The role of lowland tropical forest tree communities in shaping soil nutrient cycling has been challenging to elucidate in the face of high species diversity. Previously, we showed that differences in tree species composition and canopy foliar nitrogen (N) concentrations correlated with differences in soil N availability in a mature Costa Rican rainforest. Here, we investigate potential mechanisms explaining this correlation. We used imaging spectroscopy to identify study plots containing 10–20 canopy trees with either high or low mean canopy N relative to the landscape mean. Plots were restricted to an uplifted terrace with relatively uniform parent material and climate. In order to assess whether canopy and soil N could be linked by litterfall inputs, we tracked litter production in the plots and measured rates of litter decay and the carbon and N content of leaf litter and leaf litter leachate. We also compared the abundance of putative N fixing trees and rates of free-living N fixation as well as soil pH, texture, cation exchange capacity, and topographic curvature to assess whether biological N fixation and/or soil properties could account for differences in soil N that were, in turn, imprinted on the canopy. We found no evidence of differences in legume communities, free-living N fixation, or abiotic properties. However, soils beneath high canopy N assemblages received ~ 60% more N via leaf litterfall due to variability in litter N content between plot types. The correlation of N in canopy leaves, leaf litter, and soil suggests that, under similar abiotic conditions, litterfall-mediated feedbacks can help maintain soil N differences among tropical tree assemblages in this diverse tropical forest.

     

Distinctive Phyllosphere Bacterial Communities in Tropical Trees

Recent work has suggested that in temperate and subtropical trees, leaf surface bacterial communities are distinctive to each individual tree species and dominated by Alpha- and Gammaproteobacteria. In order to understand how general this pattern is, we studied the phyllosphere bacterial community on leaves of six species of tropical trees at a rainforest arboretum in Malaysia. This represents the first detailed study of ‘true’ tropical lowland tree phyllosphere communities. Leaf surface DNA was extracted and pyrosequenced targeting the V1–V3 region of 16S rRNA gene. As was previously found in temperate and subtropical trees, each tree species had a distinctive bacterial community on its leaves, clustering separately from other tree species in an ordination analysis. Bacterial communities in the phyllosphere were unique to plant leaves in that very few operational taxonomic units (0.5%) co-occurred in the surrounding soil environment. A novel and distinctive aspect of tropical phyllosphere communities is that Acidobacteria were one of the most abundant phyla across all samples (on average, 17%), a pattern not previously recognized. Sequences belonging to Acidobacteria were classified into subgroups 1–6 among known 24 subdivisions, and subgroup 1 (84%) was the most abundant group, followed by subgroup 3 (15%). The high abundance of Acidobacteria on leaves of tropical trees indicates that there is a strong relationship between host plants and Acidobacteria in tropical rain forest, which needs to be investigated further. The similarity of phyllosphere bacterial communities amongst the tree species sampled shows a significant tendency to follow host plant phylogeny, with more similar communities on more closely related hosts.     

Inter‐specific Variation in Foliar Nutrients and Resorption of Nine Canopy‐tree Species in a Secondary Neotropical Rain Forest

The influence of environmental gradients on the foliar nutrient economy of forests has been well documented; however, we have little understanding of what drives variability among individuals within a single forest stand, especially tropical forests. We evaluated inter‐ and intra‐specific variation in nutrient resorption, foliar nutrient concentrations and physical leaf traits of nine canopy tree species within a 1‐ha secondary tropical rain forest in northeastern Costa Rica. Both nitrogen (N) and phosphorus (P) resorption efficiency (RE) and proficiency of the nine tree species varied significantly among species, but not within. Both N and P RE were significantly negatively related to leaf specific strength. Green leaf N and P concentrations were strongly negatively related to leaf mass per area, and senesced leaf nutrient concentrations were significantly positively related to green leaf nutrient concentrations. This study reveals a strong influence of physical leaf traits on foliar nutrient and resorption traits of co‐occurring species in a secondary wet tropical forest stand.     

Light-dependent leaf trait variation in 43 tropical dry forest tree species

Our understanding of leaf acclimation in relation to irradiance of fully grown or juvenile trees is mainly based on research involving tropical wet forest species. We studied sun-shade plasticity of 24 leaf traits of 43 tree species in a Bolivian dry deciduous forest. Sampling was confined to small trees. For each species, leaves were taken from five of the most and five of the least illuminated crowns. Trees were selected based on the percentage of the hemisphere uncovered by other crowns. We examined leaf trait variation and the relation between trait plasticity and light demand, maximum adult stature, and ontogenetic changes in crown exposure of the species. Leaf trait variation was mainly related to differences among species and to a minor extent to differences in light availability. Traits related to the palisade layer, thickness of the outer cell wall, and N(area) and P(area) had the greatest plasticity, suggesting their importance for leaf function in different light environments. Short-lived pioneers had the highest trait plasticity. Overall plasticity was modest and rarely associated with juvenile light requirements, adult stature, or ontogenetic changes in crown exposure. Dry forest tree species had a lower light-related plasticity than wet forest species, probably because wet forests cast deeper shade. In dry forests light availability may be less limiting, and low water availability may constrain leaf trait plasticity in response to irradiance.     

Influence of Seasonal Variations in Soil Water Availability on Gas Exchange of Tropical Canopy Trees

Seasonal variations in environmental conditions influence the functioning of the whole ecosystem of tropical rain forests, but as yet little is known about how such variations directly influence the leaf gas exchange and transpiration of individual canopy tree species. We examined the influence of seasonal variations in relative extractable water in the upper soil layers on predawn leaf water potential, saturated net photosynthesis, leaf dark respiration, stomatal conductance, and tree transpiration of 13 tropical rain forest canopy trees (eight species) over 2 yr in French Guiana. The canopies were accessed by climbing ropes attached to the trees and to a tower. Our results indicate that a small proportion of the studied trees were unaffected by soil water depletion during seasonal dry periods, probably thanks to efficient deep root systems. The trees showing decreased tree water status (i.e., predawn leaf water potential) displayed a wide range of leaf gas exchange responses. Some trees strongly regulated photosynthesis and transpiration when relative extractable water decreased drastically. In contrast, other trees showed little variation, thus indicating good adaptation to soil drought conditions. These results have important applications to modeling approaches: indeed, precise evaluation and grouping of these response patterns are required before any tree‐based functional models can efficiently describe the response of tropical rain forest ecosystems to future changes in environmental conditions.