Differences in leaf functional traits between red and green leaves of two evergreen shrubs Photinia × fraseri and Osmanthus fragrans

Leaf functional traits are adaptations that enable plants to live under different environmental conditions. This study aims to evaluate the differences in leaf functional traits between red and green leaves of two evergreen shrubs Photinia 9 fraseri and Osmanthus fragrans. Specific areas of red leaves are higher than that of green leaves in both species. Thus, the material investment per unit area and per lamina of red leaves is significantly lower than that of green leaves, implying an utmost effort of red leaves to increase light capture and use efficiency because of their low leafchlorophyll concentration. The higher petiole length of green leaves compared with that of red leaves indicates that adult green leaves may have large fractional biomass allocation to support the lamina structures in capturing light with maximum efficiency and obtaining a high growth rate. The high range of the phenotypic plasticity of leaf size, leaf thickness,single-leaf wet and dry weights, and leaf moisture of green leaves may be beneficial in achieving efficient control of water loss and nutrient deprivation. The high range of phenotypic plasticity of leaf chlorophyll concentration of red leaves may be advantageous in increasing resource(especially light) capture and use efficiency because this leaf type is juvenile in the growth stage and has low leaf-chlorophyll concentration.     

3个优势科树种叶功能性状及其关联特性

以小尺度范围内的木兰科、壳斗科和樟科各8种木本植物为研究对象,从分类单元科与种、功能群角度分析了叶功能性状特征及其关联性。结果表明:壳斗科树种的叶柄直径(1.08±0.22)和比叶面积(SLA)(90.87±25.35)显著低于樟科和木兰科,而樟科和木兰科之间差别不明显。叶含氮量在科间没有显著差异。从功能群比较,常绿树种比叶面积(90.10±23.15)显著低于落叶树种(134.65±27.77),但单位叶面积含氮量常绿树种(1.95±0.50)显著高于落叶树种(1.31±0.26)。在不同功能群中,叶面积与叶柄质量有很好的拟合直线关系。叶氮含量仅在常绿树种与其他叶性状有较好的拟合直线关系。叶性状相关性系数大于0.900的有:群落水平物种间叶面积与叶柄质量之间;功能群水平常绿树种叶面积与叶柄直径和叶柄质量间,落叶树种叶柄质量与叶面积和叶柄长间。其他叶性状间也表现不同程度的相关性,表明叶性状之间存在生长发育非同步也即异速生长现象。     

Differences in leaf functional traits between red and green leaves of two evergreen shrubs <Emphasis Type="Italic">Photinia</Emphasis> × <Emphasis Type="Italic">fraseri</Emphasis> and <Emphasis Type="Italic">Osmanthus fragrans</Emphasis>

Leaf functional traits are adaptations that enable plants to live under different environmental conditions. This study aims to evaluate the differences in leaf functional traits between red and green leaves of two evergreen shrubs Photinia × fraseri and Osmanthus fragrans. Specific areas of red leaves are higher than that of green leaves in both species. Thus, the material investment per unit area and per lamina of red leaves is significantly lower than that of green leaves, implying an utmost effort of red leaves to increase light capture and use efficiency because of their low leaf-chlorophyll concentration. The higher petiole length of green leaves compared with that of red leaves indicates that adult green leaves may have large fractional biomass allocation to support the lamina structures in capturing light with maximum efficiency and obtaining a high growth rate. The high range of the phenotypic plasticity of leaf size, leaf thickness, single-leaf wet and dry weights, and leaf moisture of green leaves may be beneficial in achieving efficient control of water loss and nutrient deprivation. The high range of phenotypic plasticity of leaf chlorophyll concentration of red leaves may be advantageous in increasing resource (especially light) capture and use efficiency because this leaf type is juvenile in the growth stage and has low leaf-chlorophyll concentration.     

Spatial and temporal variations in leaf area index, specific leaf area and leaf nitrogen of two co-occurring savanna tree species

Foliage growth, mass- and area-based leaf nitrogen concentrations (Nm and N a) and specific leaf area (SLA) were surveyed during a complete vegetation cycle for two co-occurring savanna tree species: Crossopteryx febrifuga (Afzel. ex G. Don) Benth. and Cussonia arborea A. Rich. The study was conducted in the natural reserve of Lamto, Ivory Coast, on isolated and clumped trees. Leaf flush occurred before the beginning of the rainy season. Maximum leaf area index (LAI), computed on a projected canopy basis for individual trees, was similar (mean of about 4) for both species. Seasonal courses of the ratio of actual to maximum LAI were similar for individuals of the same species, but differed between species. For C. febrifuga, clumped trees reached their maximum LAI before isolated trees. The LAI of C. arborea trees did not differ between clumped and isolated individuals, but maximum LAI was reached about 2 months later than for C. febrifuga. Leaf fall was associated with decreasing soil water content for C. arborea. For C. febrifuga, leaf fall started before the end of the rainy period and was independent of changes in soil water content. These features lead to a partial niche separation in time for light resource acquisition between the two species. Although Nm, N a and SLA decreased with time, SLA and N a decreased later in the vegetation cycle for C. arborea than for C. febrifuga. For both species, N a decreased and SLA increased with decreasing leaf irradiance within the canopy, although effects of light on leaf characteristics did not differ between isolated and clumped trees. Given relationships between N a and photosynthetic capacities previously reported for these species, our results show that C. arborea exhibits higher photosynthetic capacity than C. febrifuga during most of the vegetation cycle and at all irradiances.     

Leaf- and shoot-level plasticity in response to different nutrient and water availabilities

Phenotypic plasticity in response to environmental variation occurs at all levels of organization and across temporal scales within plants. However, the magnitude and functional significance of plasticity is largely unexplored in perennial species. We measured the plasticity of leaf- and shoot-level physiological, morphological and developmental traits in nursery-grown Populus deltoides Bartr. ex Marsh. individuals subjected to different nutrient and water availabilities. We also examined the extent to which nutrient and water availability influenced the relationships between these traits and productivity. Populus deltoides responded to changes in resource availability with high plasticity in shoot-level traits and moderate plasticity in leaf-level traits. Although shoot-level traits generally correlated strongly with productivity across fertilization and irrigation treatments, few leaf-level traits correlated with productivity, and the relationships depended on the resource examined. In fertilized plants, leaf nitrogen concentration was negatively correlated with productivity, suggesting that growth, rather than enhanced leaf quality, is an important response to fertilization in this species. With the exception of photosynthetic nitrogen-use efficiency, traits associated with resource conservation (leaf senescence rate, water-use efficiency and leaf mass per area) were uncorrelated with short-term productivity in nutrient- and water-stressed plants. Our results suggest that plasticity in shoot-level growth traits has a greater impact on plant productivity than does plasticity in leaf-level traits and that the relationships between traits and productivity are highly resource dependent.     

A 10-year evaluation of the functional basis for regeneration habitat preference of trees in an African evergreen forest

The spatial distribution of tree juveniles in relation to light environments may reflect species differences in growth, survival, and functional traits and will shape the nature of forest regeneration. Long-term field experiments are important to evaluate this issue because of the potentially very long juvenile period in trees. Here, we combine a 10-year seedling survival–growth data with the results of community ordination and multivariate analyses of functional traits to ask how observed juvenile light guilds are related to species functional traits and seedling performance. We transplanted seedlings at a standardized height of 11 cm into the shaded understory and quantified their growth and survival for 10-years. Using the community-wide stem distribution data, we categorized 33 species including the focal 11 species to understory vs. gap/edge guilds. Then, we determined differences between the two guilds in seedling survival, growth, as well as seed size, adult height, and a series of leaf traits, including toughness and chemical traits (fiber, protein, phenolics, tannins, alkaloids, saponins). Among the 11 non-pioneer species whose seedlings were planted into the understory, there was no significant difference in 10-year survival between light guilds, but species in gap/edge guild tended to achieve greater height than species in the understory guild. The leaf chemical traits of 33 species did not differ between the two juvenile light guilds, but gap/edge species had smaller seeds, taller adults, and tougher leaves than understory species. We used logistic regression as a complementary approach to assess the extent to which plant traits varied between light guilds and the most parsimonious model based on AIC_c ranking included only leaf toughness and had an Akaike weight of 0.52. In addition, across the 11 species planted as seedlings, these traits were not significantly related to survivorship or growth over 10 years. A Principle Components Analysis illustrated associations among traits. We conclude that light guilds in terms of juvenile stem distribution could not be explained by long-term field performance of post-establishment seedlings alone. Earlier seedling stage or later sapling stage may be more important in differentiation of light guilds. For the species examined difference in growth rates could be linked to seed size and adult stature, but not to the adult leaf chemical traits considered. These results suggest the importance of examining ontogenetic shifts and relationships among functional traits for a better understanding of regeneration strategies of tropical trees.     

Ontogenetic changes in leaf phenology of Ulmus davidiana var. japonica, a deciduous broad-leaved tree

To determine how plants control leaf phenology to maximize annual carbon gain, I examined ontogenetic changes in leaf phenology of Japanese elm, Ulmus davidiana var. japonica Nakai plants of different ages growing in contrasting light environments. Leaf emergence occurred earlier in 1- and 2-year-old seedlings than in current-year seedlings. Although leaf emergence was not affected by light conditions at the sites, it was influenced by plant height. The delay in leaf emergence increased with increasing plant height. These traits indicate that seedlings that received the least light during the summer intercepted light for a long period during the spring; however, the advantage of earlier leaf emergence decreased with increasing plant height. At each site, 1-year-old seedlings had a longer duration of leaf emergence than adults, because of a longer period of favorable light conditions even in the forest understory. Duration of leaf emergence, leaf duration and leaf longevity were usually longer in sun than in shade for both seedlings and adults; however, flexibility in the response to light was greater in seedlings than in adults. The plastic response in leaf phenology during the juvenile stages may contribute to the optimization of light acquisition in habitats with differing light conditions, thereby enhancing seedling survival.     

Evaluation of leaf display of evergreen broadleaved tree species and deciduous tree species in warm temperate conifer plantations

We investigated the sapling leaf display in the shade among trees of various leaf lifespans co-occurring under the canopy of a warm-temperate conifer plantation. We measured leaf-area ratio (aLAR) and morphological traits of saplings of evergreen broadleaved tree species and a deciduous tree species. Although we found large interspecific and intraspecific differences in aLAR even among saplings of similar size in the homogeneous light environment, we did not find a consistent trend in aLAR with leaf lifespan among the species. While deciduous trees annually produced a large leaf area, some evergreen broadleaved trees retained their leaves across years and had aLAR values as high as those of deciduous trees. Among leaf-level, shoot-level, and individual-level morphological traits, aLAR was positively correlated with current-year shoots mass per aboveground biomass in deciduous trees, and with the area of old leaves per aboveground mass in evergreen broadleaved trees. Thus, tree-to-tree variation in the degrees of annual shoot production and the accumulation of old leaves were responsible for the interspecific and intraspecific variations in aLAR.     

Seasonal variations of leaf traits and drought adaptation strategies of four common woody species in South Texas,USA

Understanding physiological responses and drought adaptation strategies of woody plant leaf traits in sub-humid to semi-arid regions is of vital importance to understand the interplay between ecological processes and plant resource-allocation strategies of different tree species.Seasonal variations of leaf morphological traits,stoichiometric traits and their relationships of two drought tolerant woody species,live oak(Quercus virginiana)and honey mesquite(Prosopis glandulosa)and two less drought tolerant species,sugarberry(Celtis laevigata)and white ash(Fraxinus americana)were analyzed in a sub-humid to semi-arid area of south Texas,USA.Our findings demonstrate that for the two drought tolerant species,the leguminous P.glandulosa had the highest specific leaf area,leaf N,P,and lowest leaf area and dry mass,indicating that P.glandulosa adapts to an arid habitat by decreasing leaf area,thus reducing water loss,reflecting a resource acquisition strategy.While the evergreen species Q.virginiana exhibited higher leaf dry mass,leaf dry matter content,C content,C:N,C:P and N:P ratios,adapts to an arid habitat through increased leaf thickness and thus reduced water loss,reflecting a resource conservation strategy in south Texas.For the two less drought tolerant deciduous species,the variations of leaf traits in C.laevigata and F.americana varied between Q.virginiana and P.glandulosa,reflecting a trade-off between rapid plant growth and nutrient maintenance in a semi-arid environment.     

Light acclimation potential and carry-over effects vary among three evergreen tree species with contrasting patterns of leaf emergence and maturation

We compared light acclimation potential among three evergreen broadleaved species with contrasting patterns of shoot elongation, leaf emergence and leaf maturation. Understory saplings were transferred to a high-light environment before bud break, grown for 13 months, and then transferred back to the understory to observe subsequent carry-over effects. Acclimation potential was highest and sapling mortality was lowest for Cinnamomum japonicum Sieb. ex Nakai. Indeterminate growth and successive leaf emergence allowed this species to acclimate to both high and low light by adjusting leaf production as well as leaf properties. Sapling mortality occurred after both transfers for Camellia japonica L., which also has indeterminate growth and successive leaf emergence. In this species, carry-over effects were observed at the individual level, but leaf-level acclimation potential was high. Acclimation potential was lowest and sapling mortality occurred soon after the transfer to high light for Quercus glauca Thunb. ex Murray. Determinate growth and flush-type leaf emergence resulted in significant carry-over effects in this species. Indeterminate growth and successive leaf emergence increase whole-plant acclimation potential by extending the period of growth and architectural development during the growing season. Similarly, we inferred that delayed leaf maturation, observed in many evergreen species, increases the acclimation potential of current-year leaves by extending the period of leaf development. In evergreen species, the acclimation potential of preexisting leaves determines the role that leaf turnover plays in whole-plant light acclimation, resulting in diverse strategies for light acclimation among species, as observed in this study.     

Sex-related differences in leaf morphological and physiological responses in Hippophae rhamnoides along an altitudinal gradient

In most woody plants, leaf morphological and physiological characteristics are extremely variable across environmental gradients, particularly across altitudinal gradients. Hippophae rhamnoides L., a dioecious and deciduous shrub species, occupies a wide range of habitats in the Wolong Nature Reserve, southwest China. We measured growth, sex ratio and morphological and physiological characteristics of leaves in male and female H. rhamnoides individuals along an altitudinal gradient. Shoot height (HT), leaf N concentration per unit dry mass (N(mass)), leaf N concentration per unit area (N(area)) and leaf carbon isotope composition (delta(13)C) were higher in males than in females, whereas females had higher specific leaf area (SLA), stomatal length (SL) and stomatal index (SI) (i.e., total stomatal length per unit leaf area) than males along the altitudinal gradient. Females also had higher values of stomatal density (SD) at all altitudes except 2800 m. The male:female ratio (MFR) was biased toward males at all altitudes except at 2800 m. Changes in HT, MFR, SLA, SD, SL, SI, N(mass), N(area) and delta(13)C along the altitudinal gradient were nonlinear. Below 2800 m, HT, SLA, SD, SL and SI increased with increasing altitude, but above 2800 m they decreased with increasing altitude. In contrast, MFR, N(mass), N(area) and delta(13)C showed the opposite patterns with altitude. Consequently, we confirmed our hypotheses: (1) stressful environments have a more negative impact on females than on males in a variety of ways; (2) under optimal growth conditions the sex ratio is even, but becomes male-biased as resources become limited; and (3) there is an optimum altitudinal range at around 2800 m for the growth of H. rhamnoides in the Wolong Nature Reserve.     

Foliar carbon isotope discrimination and related traits along light gradients in two different functional-type tree species

To understand how different plant functional types respond to light intensities, foliar carbon isotope discrimination (Δ¹³C) and related traits, i.e., specific leaf area (SLA), mass- and area-based nitrogen concentrations (N_mass and N_area), leaf dry mass content (LDMC) of two evergreen coniferous and three deciduous broad-leaved species, were measured under four light intensities. Foliar Δ¹³C and SLA increased significantly from full- to low-light conditions for all species. These indicate that species studied could increase their light capture capacity under low-light conditions, leading to lower water-use efficiency (higher ¹³C discrimination). There were significant differences in the responses of foliar N_mass or N_area to light variations in the two functional types, indicating that different functional-type tree species may have different N-use strategies to adapt to the light variations. It was found that there were large functional-type-dependent differences with regard to the relationships between foliar Δ¹³C and other leaf traits. Our findings suggest that all tree species could change foliar morphology to increase their light-harvesting ability under low-light conditions at the expense of decreasing their water-use efficiency. However, large differences in N-use strategy may exist between deciduous and evergreen species, which may be vital for the survival of these two functional-type tree species in a shaded understory. More important, our findings reveal that changes in Δ¹³C are not directly related to foliar N if N investment does not proportionally increase the photosynthetic capacity; this should be considered when exploring the relationships of nitrogen concentrations with Δ¹³C.     

Variability and grouping of leaf traits in multi-species reforestation (Leyte,Philippines)

Conventional reforestation in the tropics often results in stands with low tree species and functional diversities. A different approach to reforestation, the so-called rainforestation, has been developed in the Philippines. It emphasizes mixed stands and the preferential use of native species supplemented by fruit trees. In such stands, we studied several functional leaf traits (stomatal conductance for water vapour, leaf morphology and chemistry) with the objectives (1) of assessing the species-specific variation of leaf traits and in particular that of maximal leaf stomatal conductance (g_smax), (2) of determining relationships between g_smax and other tree variables, and (3) of assessing whether leaf traits group the species studied. Sixteen broad-leaved species were studied, using five individual trees per species and ten fully expanded sunlit leaves per individual tree. Species-specific g_smax differed fivefold (165–772 mmol m⁻² s⁻¹). Among studied leaf traits, only the carbon isotope ratio δ¹³C exhibited a simple linear correlation with g_smax. A separate analysis for dipterocarp species indicated a strong negative relationship between g_smax and specific leaf area (SLA) (r² = 0.96, P < 0.001, n = 5). For all 16 species, a multiple linear regression with the combinations leaf size/tree height and leaf size/canopy projection area also resulted in significant relationships, which partly explained the variability in g_smax. A multivariate approach (principal component analysis) combining the leaf traits provided an explanation of 75% of the variability along the first two axes. All native dipterocarps species, a native Guttiferae and the durian tree (Durio zibethinus) were associated with more depleted δ¹³C, small leaves and a low leaf width to length ratio. Two exotic species frequently used for reforestation (Gmelina arborea and Swietenia macrophylla) and the native early successional Terminalia microcarpa were differentiated by their high SLA and high leaf nitrogen content per leaf area (N_area). Both species of Artocarpus (A. blancoi and A. odoratissima) were also differentiated and had large leaves with low SLA and low N_area. These associations of species with leaf traits as variables indicate that species have different leaf investment strategies, which may imply that there are differences in whole plant performance. We conclude that rainforestation creates substantial variation in leaf traits, which is based on the combination of species with different leaf trait groupings. This can be seen as an important step towards – partly – restoring the functional diversity which characterizes many natural tropical rainforests.     

Seedling biomass allocation and vital rates of cloud forest tree species: Responses to light in shade house conditions

Patterns of above- and below-ground biomass allocation in seedlings of nine common cloud forest (CF) tree species of western Mexico were examined under varying controlled light conditions using artificial shade houses. We analysed the relationships between vital rates (growth and survival) and four morphological traits (SLA, biomass allocation to stems, leaves and roots). We hypothesised that these traits represent differentiation axes in the way seedlings face the heterogeneous light regime typical of the CF understorey. For all species, traits between the different light levels, i.e. allocation to leaves, roots and stems differed among light levels. Five species had the largest SLA in the lowest light levels at the end of the experiment (Citharexylum, Dendropanax, Fraxinus, Quercus and Magnolia). Juglans was the only species with a large SLA at the highest light level (377.47 cm² g⁻¹). In contrast, light levels did not cause any significant variation in SLA of Persea and Simplococarpon at the end of the experiment. The relative height growth rates (RHGR) of the seedlings of five species were significantly different between light levels (P < 0.05). Overall, all species grew better in the highest light levels. The RHGR of three species were correlated positively with SLA. In turn, allocation to stem, leaves and root biomass were strongly correlated with the RHGR of five species (e.g. Citharexylum, Dendropanax and Fraxinus). Survival did not vary significantly between treatments in any species, only in the case of Simplococarpon (P < 0.05) and was correlated with all morphological variables. For this species, Peto and Peto's test showed a significantly larger survival of seedlings in the highest light level. The mean responses of these species based on all traits to the controlled light variation did not differed significantly. Our results show that these species display a wide range of resource allocation patterns when exposed to the varying light conditions that may be found in the forest understorey and highlight the role of morphological traits in this variation.     

Tree allometry, leaf size and adult tree size in old-growth forests of western Oregon

Relationships between tree height and crown dimensions and trunk diameter were determined for shade-tolerant species of old-growth forests of western Oregon. The study included both understory and overstory species, deciduous and evergreen angiosperms and evergreen conifers. A comparison of adult understory species with sapling overstory species of similar height showed greater crown width and trunk diameter in the former, whether the comparison is made among conifers or deciduous trees. Conifer saplings had wider crowns than deciduous saplings, but the crown widths of the two groups converged with increase in tree height. Conifer saplings had thicker trunks than deciduous saplings of similar crown width, possibly because of selection for resistance to stem bending under snow loads. The results suggest that understory species have morphologies that increase light interception and persistence in the understory, whereas overstory species allocate their biomass for efficient height growth, thereby attaining the high-light environment of the canopy. The greater crown widths and the additional strength requirements imposed by snow loads on conifer saplings result in less height growth per biomass increment in conifer saplings than in deciduous saplings. However, the convergence in crown width of the two groups at heights greater than 20 m, and the proportionately smaller effect of snow loads on large trees, may result in older conifers equalling or surpassing deciduous trees in biomass allocation to height growth.     

三种园林植物叶形态生理指标及其环境指示作用

通过测定3种南方园林植物的叶面积、叶片含水量、比叶面积和叶绿素指数4项形态和生理指标,并对两种不同环境条件下各指标的差异进行分析,验证3种园林植物利用资源及对环境适应的能力。结果表明,高山榕(Ficus altissima)生长较快但不能较好地适应大气污染,白千层(Melaleuca quinquenervia)能抗大气污染,而尖叶杜英(Elaeocarpus apiculatus)更能适应资源丰富的环境条件。     

Production physiology of three fast-growing hardwood species along a soil resource gradient

We determined how specific leaf area (SLA), specific leaf nitrogen (SLN), leaf area index (LAI), light-saturated photosynthesis (Amax) and aboveground net primary productivity (ANPP) of three commercially important hardwood species, eastern cottonwood (Populus deltoides Bartr.), American sycamore (Platanus occidentalis L.) and cherrybark oak (Quercus falcata var.pagodafolia Ell.), vary across a soil resource gradient. Five treatments were applied in a randomized block design (control, irrigation only (IRR), and irrigation plus fertilization with 56, 112 or 224 kg N ha-1 year-1 (N56, N112 and N224)) with four replications per species. When trees were 6 years old, Amax, SLA, SLN, LAI and ANPP were quantified during peak leaf production within a single growing season. In all species, Amax for sun leaves was significantly higher than for shade leaves (34, 32 and 29 micromol m2 s-1 versus 27, 23 and 23 micromol m2 s-1 for cottonwood, cherrybark oak and sycamore sun and shade leaves, respectively) and tended to plateau in the N112 treatment. The SLA was significantly lower in sun than in shade leaves and reached a plateau in IRR-treated cottonwood and sycamore, and in N56-treated oak. Values of SLN peaked in the N122 treatment for cottonwood sun leaves (1.73 g N m2) and in the N56 treatment for sycamore and oak (1.54 and 1.90 g N m2, respectively). In sun and shade leaves of all species, Amax increased with increasing SLN. Cherrybark oak LAI reached a plateau across the resource gradient in the N56 treatment, whereas cottonwood and sycamore LAI reached a plateau in the IRR treatment. All species exhibited significant curvilinear relationships between canopy Amax and ANPP. These findings indicate that nutrients and water regulate leaf-level traits such as SLA and SLN, which in turn influence LAI and canopy photosynthesis, thereby affecting ANPP at the tree and stand levels.     

Leaf and wood carbon isotope ratios, specific leaf areas and wood growth of Eucalyptus species across a rainfall gradient in Australia

Leaves and samples of recent wood of Eucalyptus species were collected along a rainfall gradient parallel to the coast of Western Australia between Perth in the north and Walpole in the south and along a southwest to northeast transect from Walpole in southwestern Australia, to near Mount Olga in central Australia. The collection included 65 species of Eucalyptus sampled at 73 sites and many of the species were collected at several sites along the rainfall gradient. Specific leaf area (SLA) and isotopic ratio of 13C to 12C (delta 13C) of leaves that grew in 2002, and tree ring growth and delta 13C of individual cell layers of the wood were measured. Rainfall data were obtained from the Australian Bureau of Meteorology for 29 locations that represented one or a few closely located collection sites. Site-averaged data and species-specific values of delta 13C decreased with decreasing annual rainfall between 1200 and 300 mm at a rate of 1.63 per thousand per 1000 mm decrease in rainfall. Responses became variable in the low rainfall region (< 300 mm), with some species showing decreasing delta 13C with rainfall, whereas delta 13C increased or remained constant in other species. The range of delta 13C values in the low rainfall region was as large as the range observed at sites receiving > 300 mm of annual rainfall. Specific leaf area varied between 2 and 6 m2 kg(-1) and tended to increase with decreasing annual rainfall in some species, but not all, whereas delta 13C decreased with SLA. The relationship between delta 13C and SLA was highly species and soil-type specific. Leaf-area-based nitrogen (N) content varied between 2 and almost 6 g m(-2) and decreased with rainfall. Thus, thicker leaves were associated with higher N content and this compensated for the effect of drought on delta 13C. Nitrogen content was also related to soil type and species identity. Based on a linear mixed model, statistical analysis of the whole data set showed that 27% of the variation in delta 13C was associated with changes in SLA, 16% with soil type and only 1% with rainfall. Additionally, 21% was associated with species identity. For a subset of sites with > 300 mm rainfall, 43% of the variation was explained by SLA, 13% by soil type and only 3% by rainfall. The species effect decreased to 9% because there were fewer species in the subset of sites. The small effect of rainfall on delta 13C was further supported by a path analysis that yielded a standardized path coefficient of 0.38 for the effect of rainfall on SLA and -0.50 for the effect of SLA on delta 13C, but an insignificantly low standardized path coefficient of -0.05 for the direct effect of rainfall on delta 13C. Thus, in contrast to our hypothesis that delta 13C decreases with rainfall independent of soil type and species, we detected no statistically significant relationship between rainfall and delta 13C in leaves of trees growing at sites receiving < 300 mm of rainfall annually. Rainfall affected delta 13C indirectly through soil type (a surrogate for water-holding capacity) across the rainfall gradient. Annual tree rings are not clearly visible in evergreen Eucalyptus species, even in the seasonally cool climate of SW Australia. Generally, visible density transitions in the wood are related not to a strict annual cycle but to periods of growth associated mainly with rainfall. The relationship between delta 13C of leaves and the width of these stem increments was not statistically significant. Analysis of stem growth periods showed that delta 13C in wood responded to rainfall events, but carbohydrate storage and reallocation also affected the isotopic signature. Although delta 13C in wood of any one species varied over a range of 2 to 4 per thousand, there was a general relationship between delta 13C of the leaves and the annual range of delta 13C in wood. We conclude that species-specific traits are important in understanding the response of Eucalyptus to rainfall and that the diversity of the genus may reflect its response to the large climatic gradient in Australia and to the large annual and interannual variations in rainfall at any one location.     

Changes in shoot allometry with increasing tree height in a tropical canopy species,Elateriospermum tapos

Allometry of shoot extension units (hereafter termed "current shoots") was analyzed in a Malaysian canopy species, Elateriospermum tapos Bl. (Euphorbiaceae). Changes in current shoot allometry with increasing tree height were related to growth and maintenance of tree crowns. Total biomass, biomass allocation ratio of non-photosynthetic to photosynthetic organs, and wood density of current shoots were unrelated to tree height. However, shoot structure changed with tree height. Compared with short trees, tall trees produced current shoots of the same mass but with thicker and shorter stems. Current shoots with thin and long stems enhanced height growth in short trees, whereas in tall trees, thick and short current shoots may reduce mechanical and hydraulic stresses. Furthermore, compared with short trees, tall trees produced current shoots with more leaves of lower dry mass, smaller area, and smaller specific leaf area (SLA). Short trees adapted to low light flux density by reducing mutual shading with large leaves having a large SLA. In contrast, tall trees reduced mutual shading within a shoot by producing more small leaves in distal than in proximal parts of the shoot stem. The production of a large number of small leaves promoted light penetration into the dense crowns of tall trees. All of these characteristics suggest that the change in current shoot structure with increasing tree height is adaptive in E. tapos, enabling short trees to maximize height growth and tall trees to maximize light capture.     

Seasonal patterns of leaf gas exchange and water relations in dry rain forest trees of contrasting leaf phenology

Diurnal and seasonal patterns of leaf gas exchange and water relations were examined in tree species of contrasting leaf phenology growing in a seasonally dry tropical rain forest in north-eastern Australia. Two drought-deciduous species, Brachychiton australis (Schott and Endl.) A. Terracc. and Cochlospermum gillivraei Benth., and two evergreen species, Alphitonia excelsa (Fenzal) Benth. and Austromyrtus bidwillii (Benth.) Burret. were studied. The deciduous species had higher specific leaf areas and maximum photosynthetic rates per leaf dry mass in the wet season than the evergreens. During the transition from wet season to dry season, total canopy area was reduced by 70-90% in the deciduous species and stomatal conductance (g(s)) and assimilation rate (A) were markedly lower in the remaining leaves. Deciduous species maintained daytime leaf water potentials (Psi(L)) at close to or above wet season values by a combination of stomatal regulation and reduction in leaf area. Thus, the timing of leaf drop in deciduous species was not associated with large negative values of daytime Psi(L) (greater than -1.6 MPa) or predawn Psi(L) (greater than -1.0 MPa). The deciduous species appeared sensitive to small perturbations in soil and leaf water status that signalled the onset of drought. The evergreen species were less sensitive to the onset of drought and g(s) values were not significantly lower during the transitional period. In the dry season, the evergreen species maintained their canopies despite increasing water-stress; however, unlike Eucalyptus species from northern Australian savannas, A and g(s) were significantly lower than wet season values.