Drought-tolerance of an invasive alien tree, <Emphasis Type="Italic">Acacia mearnsii</Emphasis> and two native competitors in fynbos riparian ecotones

Invasive alien plants (IAPs) have successfully invaded many riparian zones in South Africa, especially Australian Acacia spp. which are prevalent along riverbanks in the south-western Cape of South Africa. This Mediterranean-type climate region is predicted to endure severe future water shortages under likely scenarios of increased population growth and climate change, and IAPs aggravate this problem due to their profligate water use. Acacia mearnsii competes aggressively with native species, however, it remains unclear what physiological advantage the species has over co-occurring native species under the predicted reduced streamflow scenarios. A mechanistic approach was used to investigate how key native fynbos riparian woody tree species compare in vulnerability to drought-induced cavitation against A. mearnsii by comparing findings from three Mediterranean-type fynbos river systems that differ in streamflow. A. mearnsii showed lower water potential at 50% hydraulic conductivity loss (P₅₀ values) compared to native species at certain sites, an indication of drought-tolerance. This suggests it is likely to persist under future drier conditions and it therefore remains a top priority for control. The native Brabejum stellatifolium had consistently higher water potentials across all sites than the other studied species, and is a potentially valuable species for restoration of south-western Cape riparian zones. Consistency in the shapes of species vulnerability curves across sites illustrated a species-specific hydraulic response to different water availability, strengthening the argument that this approach to distinguish site-level drought-tolerance between trees is a practical technique, with great application in understanding future geographic distribution under climate change, and potential for use in restoration research. Additionally, streamflow was an inaccurate predictor of species drought-tolerance along these riparian systems.     

Wood density and vessel traits as distinct correlates of ecological strategy in 51 California coast range angiosperms

Wood density and vessel characteristics are functionally interrelated, yet they may have distinct ecological associations. In a comparative study of 51 angiosperm species ranging from chaparral shrubs to riparian trees, we examined relationships among wood density and vessel traits and their ecological correlates. Mean vessel lumen area and vessel density (number mm(-2)) varied widely (7- to 10-fold). In multivariate analyses, both vessel traits were negatively correlated with wood density, which varied more narrowly (< 2-fold). Vessel density and lumen area were inversely related across species, allowing a broad range of vessel traits within a narrow range of wood density. Phylogenetic independent contrasts indicated correlated inverse evolutionary change in vessel traits. Each trait had a distinct pattern of ecological correlation -- wood density was most strongly associated with soil water, and vessel traits showed contrasting relationships with plant height. Within a narrow range of wood density, there was significant variation in vessel traits. Given their particular ecological associations, the results suggest that wood density and vessel traits describe two distinct ecological axes.     

Life-span growth dynamics and xylem anatomical patterns of diffuse-porous Afzelia africana Sm. (Fabaceae) in different ecological zones in Burkina Faso

Tropical West African savannas are exposed to high climatic variability with potential impacts on tree growth, forest dynamics and ecosystem productivity. In such context, understanding the long-term ecological responses of savanna trees to changing environmental conditions is of great relevance for taking appropriate conservation actions. We conducted the first study on tree-ring analysis and quantitative wood anatomy on Afzelia africana Sm. in Burkina Faso, to investigate the life-span growth trajectories and wood anatomical adjustment to site and to climate variations. A total of 24 stem discs was collected in four protected forests along the Sudano-sahelian and the Sudanian climatic zones. Wood samples were analyzed using standard dendrochronological methods and quantitative wood anatomy. The mean annual growth rates varied from 1.002 (± 0.249) mm. year⁻¹ in the Sudanian zone to 1.128 (± 0.436) mm. year⁻¹ in the Sudano-sahelian zone. Analysis of growth trajectories showed high variations within sites and between climatic zones. Wood anatomical traits significantly varied between sites. Principal Component Analysis revealed strong relationships between ring width, wood density and vessel traits, with 82.81 % of the total variance explained. Vessel size significantly increased from the pith to the bark, highlighting the ontogenetic effects on xylem anatomical variations. Inverse relationships were found between vessel size and vessel density across the driest site and the wettest site, suggesting that the higher the rainfall, the taller the tree, the larger vessel size, but the lower vessel density. By contrast, more arid conditions and high evapotranspiration lead to smaller vessel sizes and higher vessel density. Such anatomical adjustments highlight the trade-offs between water conductance efficiency and hydraulic safety, and emphasize physiological responses to climate variability. These variations on the long-term dynamics and xylem anatomical patterns underline complex interactions between ontogenetic effects and contrasting environmental factors that affect the eco-physiological functioning of A. africana throughout the Sudanian region.     

Living on the edge: Legacy of water availability on Tetraclinis articulata secondary growth under semiarid conditions in Morocco

Tetraclinis articulata is a xerothermic Mediterranean conifer native from NW Africa that grows under semiarid conditions in the Atlas Mountains. This species is particularly well adapted to water scarcity and forms xeric woodlands subjected to recurring drought, aridification and overexploitation. Wood of T. articulata has been considered of limited dendrochronological value due to abundant anatomical anomalies in their growth rings. We studied tree-ring growth and tree-ring features to check its dendrochronological potential and evaluate their climatic signals and drought legacy effects at two semiarid sites with contrasting elevation in Morocco. Tree-ring boundaries were properly identified on wood cores from 60 to 68% of the sampled trees. Intra-annual wood density fluctuations (IADF), microrings, missing rings and rings with undefined limits were most abundant at the low-elevation coastal site than at the mountain site. Microring and missing ring abundance was inversely related to tree-ring growth, independent of cambial age, and mostly dependent on drought occurrence. IADF frequency was independent of tree-ring growth but inversely related to cambial age. A complex response of IADF formation to cool March to June conditions was found at the low-elevation coastal site, while IADF formation in the mountain site was related to May water availability. Undefined ring limits were observed only at the low-elevation site, showing positive relationships with elevated maximum temperatures in previous winter. Previous winter rainfall favored radial growth, but a significant legacy effect of water availability was evident up to three years prior to growth. Tree-ring growth was mostly related to water availability 33 and 26 months prior to growth at low-elevation and mountain sites, respectively. The obtained results reveal a very plastic cambial activity and a strong ability of T. articulata to withstand drought even for long periods.     

Correlation of drought traits and the predictability of osmotic potential at full leaf turgor in vegetation from New Zealand

Scientists do not know precisely how severe will be the impact of climate change on species. Evidence suggests that for some species, their future distributions might be jeopardized by local extinctions and drought‐induced tree mortality. Thus, we require models capable of estimating drought tolerance across many species. We can approach this goal by assessing functional traits. The trait osmotic potential at full turgor, π_O, is potentially a good drought indicator; however, few studies address its importance as a drought‐tolerance predictor and it is difficult to measure in the field with accuracy. In this work, we aim to answer the questions: which drought traits correlate with π_O?; do morpho‐anatomical traits correlate with π_O?; and which trees and shrubs are more (or less) vulnerable to drought? To achieve this aim, we assessed physiological and morpho‐anatomical traits for 14 native species from New Zealand forests. We included leaf‐ and wood‐related traits, π_O, water potential and stomatal conductance. We examined how these traits correlate with π_O and sought to generate models to predict π_O as a function of other traits. We tested 33 different models and evaluated them using Akaike's information criterion. Unfortunately, none of the morpho‐anatomical traits correlated well with π_O. Instead, water potential correlated most strongly with π_O. None of the models using only morpho‐anatomical traits produced plausible results. The model with the best predictive performance incorporated the effects of both morpho‐anatomical and physiological traits: water potential and wood saturated water content. Of the species analysed, and based on their π_O response, Lophozonia menziesii was considered the most vulnerable to drought stress, whereas Plagianthus regius was the least vulnerable. Our findings imply that it is potentially valuable to keep exploring the use of π_O as a drought indicator and that the effort required to measure some physiological traits, such as water potential, may be essential to consider plant drought responses and to predict π_O.     

Do tree-ring traits reflect different water deficit responses in young poplar clones (Populus × canadensis Mönch ‘I-214’ and P. deltoides ‘Dvina’)?

Poplar clones are known to display a wide range of tolerance to drought and water-use efficiency, but the effects of water deficit on stem growth and tree-ring characteristics are rarely taken into account. This study was conducted in order to investigate whether the main tree-ring traits correlate with irrigation regimes during the growing season in ‘I-214’ and ‘Dvina’ 4-year-old poplar clone saplings grown in concrete tanks, during three consecutive years. Total carbon, stable carbon isotope, Klason lignin and α-cellulose contents were analyzed to characterize wood biochemistry; ring width, wood density, mean vessel density and mean vessel lumen area were analyzed to characterize wood anatomy to assess the influence of irrigation regime. In both clones, wood formed in 2005 was more enriched in ¹³C, suggesting drought-induced stomatal closure. Wood formed in 2006 was less variable in δ¹³C in relation to irrigation regimes. ‘Dvina’ showed higher Klason lignin content and wood density than ‘I-214’, whatever the irrigation regime, despite the larger ring widths. ‘Dvina’ has the potential to recover promptly after drought stress, but at the expense of poor wood technological properties, while ‘I-214’ could continue to grow more uniformly under limited water availability, though at a lower rate.     

Phenotypic variation and water selection potential in the stem structure of invasive alligator weed

The morphological and anatomical characteristics of stems have been found to be related to drought resistance in plants. Testing the phenotypic selection of water availability on stem anatomical traits would be useful for exploring the evolutionary potential of the stem in response to water availability. To test the phenotypic variation of the stem anatomical traits of an invasive plant in response to water availability, we collected a total of 320 individuals of Alternanthera philoxeroides from 16 populations from terrestrial and aquatic habitats in 8 plots in China and then analyzed the variation, differentiation, plasticity and selection potential of water availability on the stem anatomical traits. We found that except for the thickness of the cortex, all of the examined phenotypic parameters of the A. philoxeroides stem were significantly and positively correlated with soil water availability. The phenotypic differentiation coefficient for all of the anatomical structural parameters indicated that most of the variation existed between habitats within the same plot, whereas there was little variation among plots or among individuals within the same habitat except for variation in the thickness of the cortex. A significant phenotypic plasticity response to water availability was found for all of the anatomical traits of A. philoxeroides stem except for the thickness of the cortex. The associations between fitness and some of the anatomical traits, such as the stem diameter, the cortex area-to-stem area ratio, the pith cavity area-to-stem area ratio and the density of vascular bundles, differed with heterogeneous water availability. In both the aquatic and terrestrial habitats, no significant directional selection gradient was found for the stem diameter, the cortex area-to-stem area ratio or the density of vascular bundles. These results indicated that the anatomical structure of the A. philoxeroides stem may play an important role in the adaptation to changes in water availability.     

Wood anatomy and wood density in shrubs: Responses to varying aridity along transcontinental transects

Wood density plays a key role in ecological strategies and life history variation in woody plants, but little is known about its anatomical basis in shrubs. We quantified the relationships between wood density, anatomy, and climate in 61 shrub species from eight field sites along latitudinal belts between 31° and 35° in North and South America. Measurements included cell dimensions, transverse areas of each xylem cell type and percentage contact between different cell types and vessels. Wood density was more significantly correlated with precipitation and aridity than with temperature. High wood density was achieved through reductions in cell size and increases in the proportion of wall relative to lumen. Wood density was independent of vessel traits, suggesting that this trait does not impose conduction limitations in shrubs. The proportion of fibers in direct contact with vessels decreased with and was independent of wood density, indicating that the number of fiber-vessel contacts does not explain the previously observed correlation between wood density and implosion resistance. Axial and radial parenchyma each had a significant but opposite association with wood density. Fiber size and wall thickness link wood density, life history, and ecological strategies by controlling the proportion of carbon invested per unit stem volume.     

Micro-evolutionary patterns of juvenile wood density in a pine species

Wood density can be considered an adaptive trait, because it ensures the safe and efficient transport of water from the roots to the leaves, mechanical support for the body of the plant and the storage of biological chemicals. Its variability has been extensively described in narrow genetic backgrounds and in wide ranges of forest tree species, but little is known about the extent of natural genetic and phenotypic variability within species. This information is essential to our understanding of the evolutionary forces that have shaped this trait, and for the evaluation of its inclusion in breeding programs. We assessed juvenile wood density, leaf area, total aboveground biomass, and growth in six Pinus pinaster populations of different geographic origins (France, Spain, and Morocco) growing in a provenance-progeny trial. No genetic differentiation was found for wood density, whereas all other traits significantly differed between populations. Heritability of this trait was moderate, with a low additive genetic variance. For retrospective identification of the evolutionary forces acting on juvenile wood density, we compared the distribution of neutral markers (F _ST) and quantitative genetic differentiation (Q _ST). We found that Q _ST was significantly lower than F _ST, suggesting evolutionary stasis. Furthermore, we did not detect any relationship between juvenile wood density and drought tolerance (resistance to cavitation), suggesting that this trait could not be used as a proxy for drought tolerance at the intraspecific level.     

Variation in wood density and anatomy in a widespread mangrove species

Wood density is an important plant trait that influences a range of ecological processes, including resistance to damage and growth rates. Wood density is highly dependent on anatomical characteristics associated with the conductive tissue of trees (xylem and phloem) and the fibre matrix in which they occur. Here, we investigated variation in the wood density of the widespread mangrove species Avicennia marina in the Exmouth Gulf in Western Australia and in the Firth of Thames in New Zealand. We assessed how variation in xylem vessel size, fibre wall thickness and proportion of phloem within the wood contributed to variation in wood density and how these characteristics were linked to growth rates. We found the wood density of A. marina to be higher in Western Australia than in New Zealand and to be higher in taller seaward fringing trees than in scrub trees growing high in the intertidal. At the cellular level, high wood density was associated with large xylem vessels and thick fibre walls. Additionally, wood density increased with decreasing proportions of phloem per growth layer of wood. Tree growth rates were positively correlated with xylem vessel size and wood density. We conclude that A. marina can have large xylem vessel sizes and high growth rates while still maintaining high wood density because of the abundance and thickness of fibres in which vessels are found.     

Impact of extreme drought on tree-ring width and vessel anatomical features of Chukrasia tabularis

Multiple sources of evidence suggest an increasing frequency of extreme climatic events during the past century. In Bangladesh, a country strongly influenced by the South Asian monsoon climate, the years 1999 and 2006 were the most severe droughts among the ten drought events identified over the last four decades. We investigated the impact of these two drought events on radial growth and xylem anatomical features of the brevi-deciduous tree species Chukrasia tabularis in a moist tropical forest in Bangladesh. Tree radial growth declined by 54% during the 1999 and 48.7% during the 2006 droughts, respectively. Among the wood anatomical features, the number of vessels (NV) showed the highest sensitivity to drought, with a 45% decrease in the 1999 drought year, followed by total vessel area (TVA) and mean vessel area (MVA). On the other hand, Vessel density (VD) increased by 13% during the 1999 drought but the increase in VD was very low in the drought year 2006. The decreasing vessel area and increasing vessel density indicate xylem hydraulic adaptation of C. tabularis to minimize drought induced cavitation risk and to avoid hydraulic failure. The significant correlations between the Standardized Precipitation Evapotranspiration Index (SPEI) and time series of tree-ring width and vessel variables imply that decline in radial growth and changes in vessel features in C. tabularis are likely to be caused by drought induced water stress. Our analyses suggest that radial growth and wood anatomical features of C. tabularis are highly sensitive to extreme drought events in South Asian moist tropical forests and can be used to reconstruct past droughts and to model tree response to drought stress under future climate conditions.     

Testing the divergent adaptation of two congeneric tree species on a rainfall gradient using eco‐physio‐morphological traits

In tropical Africa, evidence of widely distributed genera transcending biomes or habitat boundaries has been reported. The evolutionary processes that allowed these lineages to disperse and adapt into new environments are far from being resolved. To better understand these processes, we propose an integrated approach, based on the eco‐physio‐morphological traits of two sister species with adjacent distributions along a rainfall gradient. We used wood anatomical traits, plant hydraulics (vulnerability to cavitation, wood volumetric water content, and hydraulic capacitance), and growth data from the natural habitat, in a common garden, to compare species with known phylogeny, very similar morphologically, but occupying contrasting habitats: Erythrophleum ivorense (wet forest) and Erythrophleum suaveolens (moist forest and forest gallery). We identified some slight differences in wood anatomical traits between the two species associated with strong differences in hydraulics, growth, and overall species distribution. The moist forest species, E. suaveolens, had narrower vessels and intervessel pits, and higher vessel cell‐wall reinforcement than E. ivorense. These traits allow a high resistance to cavitation and a continuous internal water supply of the xylem during water shortage, allowing a higher fitness during drought periods, but limiting growth. Our results confirm a trade‐off between drought tolerance and growth, controlled by subtle adaptations in wood traits, as a key mechanism leading to the niche partitioning between the two Erythrophleum species. The generality of this trade‐off and its importance in the diversification of the African tree flora remains to be tested. Our integrated eco‐physio‐morpho approach could be the way forward.     

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 (π₀) and higher bulk modulus of elasticity. There were evolutionary correlations between leaf life span and stem hydraulic efficiency, A_m, and dry season π₀. Xylem-cavitation resistance was evolutionarily correlated with stem hydraulic efficiency, D_h, as well as dry season π₀. 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.Key words: Tropical dry forest, karst, leaf habit, hydraulic conductivity, cavitation resistance, leaf water-stress tolerance, wood density, leaf density, phylogenetic independent contrasts     

Wood day capacitance is related to water content,wood density,and anatomy across 30 temperate tree species

Water released from wood during transpiration (capacitance) can meaningfully affect daily water use and drought response. To provide context for better understanding of capacitance mechanisms, we investigated links between capacitance and wood anatomy. On twigs of 30 temperate angiosperm tree species, we measured day capacitance (between predawn and midday), water content, wood density, and anatomical traits, that is, vessel dimensions, tissue fractions, and vessel–tissue contact fractions (fraction of vessel circumference in contact with other tissues). Across all species, wood density (WD) and predawn lumen volumetric water content (VWC_L-pd) together were the strongest predictors of day capacitance (r²_adj = .44). Vessel–tissue contact fractions explained an additional ~10% of the variation in day capacitance. Regression models were not improved by including tissue lumen fractions. Among diffuse-porous species, VWC_L-pd and vessel–ray contact fraction together were the best predictors of day capacitance, whereas among semi/ring-porous species, VWC_L-pd, WD and vessel–fibre contact fraction were the best predictors. At predawn, wood was less than fully saturated for all species (lumen relative water content = 0.52 ± 0.17). Our findings imply that day capacitance depends on the amount of stored water, tissue connectivity and the bulk wood properties arising from WD (e.g., elasticity), rather than the fraction of any particular tissue.     

The Structural Adaptation of Aerial Parts of Invasive <Emphasis Type="Italic">Alternanthera philoxeroides</Emphasis> to Water Regime

Alternanthera philoxeroides has successfully invaded diverse habitats with considerably various water availability, threatening biological diversity in many parts of the world. Because its genetic variation is very low, phenotypic plasticity is believed to be the primary strategy for adapting to the diverse habitats. In the present paper, we investigated the plastic changes of anatomical traits of the aerial parts of A. philoxeroides from flooding to wet then to drought habitat; the results are as follows: A. philoxeroides could change anatomical structures sensitively to adapt to water regime. As a whole, effects of water regime on structures in stem were greater than those in leaf. Except for principal vein diameter and stoma density on leaf surfaces, all other structural traits were significantly affected by water regime. Among which, cuticular wax layer, collenchyma cell wall, phloem fiber cell wall, and hair density on both leaf surfaces thickened significantly with decrease of water availability, whereas, pith cavity and vessel lumen in stem lessened significantly; wet habitat is vital for the spread of A. philoxeroides from flooding to drought habitat and vice versa, because in this habitat, it had the greatest structural variations; when switching from flooding to wet then to drought habitat, the variations of cuticular wax layer, collenchyma cell wall, phloem fiber cell wall, pith cavity area ratio, diameter of vessel lumen, and hair density on both leaf surfaces, played the most important role. These responsive variables contribute most to the adaptation of A. philoxeroides to diverse habitats with considerably various water availability.     

Wood anatomy and vessel characteristics of spiny monkey orange (Strychnos spinosa) in Benin (West Africa)

The response of plant species to varying climate conditions in tropical Africa remains poorly understood but can be assessed using wood anatomical traits. These traits play an important role for the adaptive capacity of a species to environmental stress, since environmental conditions can modify the proportion, size, and morphology of wood anatomical elements. This study reports quantitative data on vessel characteristics of the diffuse porous angiosperm Spiny monkey orange (Strychnos spinosa Lam.) in Benin in tropical West Africa. The vessel-related anatomical traits varied with high amplitude (coefficient of variation CV ˃ 25%) between different sites located in different climate zones. The variability of the traits is higher within one climatic zone than between climatic zones, and even more pronounced within trees. Consequently, the climatic zones have less influence on the studied features than local site conditions. However, the study showed that S. spinosa individuals that have numerous vessels also have a high lumen fraction and total ring area. On the other hand, individuals presenting a high vessel density also display vessels of smaller size. The correlation between vessel number and total ring area on the one hand, and between vessel size and lumen fraction on the other hand are highly significant and positive. In Benin, S. spinosa wood anatomical traits are likely linked to local site factors rather than to regional climatic factors.     

Drought-induced adaptive changes in the seedling anatomy of Acacia ehrenbergiana and Acacia tortilis subsp. raddiana

This study, conducted on Acacia species (Acacia ehrenbergiana Hayne and Acacia tortilis subsp. raddiana (Savi) Brenan) to determine their adaptive capacity to tolerate drought and suitability for reforestation, revealed that leaf water potential (ψ) decreased in both the species with increase in drought intensity. With increase in the intensity of drought, vessel diameter increased in A. ehrenbergiana, causing a significant decline in vessel frequency mm^?2 of the transverse wood surface, while it decreased in A. tortilis, leading to a crowded vessel population. Vessel-wall thickness, in conjunction with inter-vessel pit membrane thickness, showed a positive correlation with drought stress in both the species. Ray dimensions generally decreased in A. ehrenbergiana but increased in A. tortilis under increasing degree of drought. The transverse fiber-wall area decreased in A. ehrenbergiana, thus lowering the density (r = 0.996) and enhancing the vulnerability of wood (r = ?0.979) under the drought stress, but increased in A. tortilis, causing a high density (r = ?0.979) and low vulnerability of wood (r = 0.869), under the same set of conditions. Correlation of wood density with vulnerability index was stronger in A. ehrenbergiana (r = ?0.993) than in A. tortilis (r = ?0.753). Diameters and thickness of inter-vessel pit membrane were linearly correlated with increasing intensity of drought in both the species, but its area fraction per vessel segment increased due to water stress in A. ehrenbergiana and decreased in A. tortilis. This study indicated that, on the whole, A. tortilis has a greater capacity to tolerate the harshness of drought than A. ehrenbergiana.     

Broad Anatomical Variation within a Narrow Wood Density Range—A Study of Twig Wood across 69 Australian Angiosperms

ObjectivesJust as people with the same weight can have different body builds, woods with the same wood density can have different anatomies. Here, our aim was to assess the magnitude of anatomical variation within a restricted range of wood density and explore its potential ecological implications.MethodsTwig wood of 69 angiosperm tree and shrub species was analyzed. Species were selected so that wood density varied within a relatively narrow range (0.38–0.62 g cm^-3). Anatomical traits quantified included wood tissue fractions (fibres, axial parenchyma, ray parenchyma, vessels, and conduits with maximum lumen diameter below 15 μm), vessel properties, and pith area. To search for potential ecological correlates of anatomical variation the species were sampled across rainfall and temperature contrasts, and several other ecologically-relevant traits were measured (plant height, leaf area to sapwood area ratio, and modulus of elasticity).ResultsDespite the limited range in wood density, substantial anatomical variation was observed. Total parenchyma fraction varied from 0.12 to 0.66 and fibre fraction from 0.20 to 0.74, and these two traits were strongly inversely correlated (r = -0.86, P < 0.001). Parenchyma was weakly (0.24 ≤|r|≤ 0.35, P < 0.05) or not associated with vessel properties nor with height, leaf area to sapwood area ratio, and modulus of elasticity (0.24 ≤|r|≤ 0.41, P < 0.05). However, vessel traits were fairly well correlated with height and leaf area to sapwood area ratio (0.47 ≤|r|≤ 0.65, all P < 0.001). Modulus of elasticity was mainly driven by fibre wall plus vessel wall fraction rather than by the parenchyma component.ConclusionsOverall, there seem to be at least three axes of variation in xylem, substantially independent of each other: a wood density spectrum, a fibre-parenchyma spectrum, and a vessel area spectrum. The fibre-parenchyma spectrum does not yet have any clear or convincing ecological interpretation.     

An exotic Australian <Emphasis Type="Italic">Acacia</Emphasis> fixes more N than a coexisting indigenous <Emphasis Type="Italic">Acacia</Emphasis> in a South African riparian zone

Acacia mearnsii is an introduced Australian acacia in South Africa and has invaded more than 2.5 million ha, primarily establishing in rangeland and riparian areas. Because acacias have the capability to fix N, A. mearnsii invasions may fundamentally change N dynamics in invaded systems. This study compares biological N₂-fixation in the alien invasive A. mearnsii and the native A. caffra growing in a grassland riparian zone in the Komati Gorge Reserve, Mpumalanga, South Africa. A ¹⁵N natural abundance field survey suggested that both mature alien and native acacias fix N under current conditions in the riparian zone. Significantly depleted δ¹⁵N was observed in both acacias relative to reference species, although variation in δ¹⁵N was not correlated with N concentrations. Calculated contributions of N₂-fixation (%Ndfa) suggest that alien acacias fix significantly more of their N than native acacias (~75 ± 5% SE and 53 ± 9% SE, respectively). There was a larger variation in δ¹⁵N and %Ndfa in the native acacia, suggesting relatively high plasticity in its N₂-fixation contributions. This plasticity was interpreted as a facultative N₂-fixation strategy for the native acacia, while the N₂-fixation strategy of the alien acacia remained unclear. Our results emphasize the importance of potentially elevated N inputs through N₂-fixation by invasive legumes in invaded landscapes. Furthermore, they suggest that N₂-fixation by invasive acacias may not respond to fine-scale patchiness in soil N in the same manner as native acacias, making them potential contributors to N excess in Southern Africa.     

Hydraulics and life history of tropical dry forest tree species: coordination of species' drought and shade tolerance

Plant hydraulic architecture has been studied extensively, yet we know little about how hydraulic properties relate to species' life history strategies, such as drought and shade tolerance. The prevailing theories seem contradictory. We measured the sapwood (K(s) ) and leaf (K(l) ) hydraulic conductivities of 40 coexisting tree species in a Bolivian dry forest, and examined associations with functional stem and leaf traits and indices of species' drought (dry-season leaf water potential) and shade (juvenile crown exposure) tolerance. Hydraulic properties varied across species and between life-history groups (pioneers vs shade-tolerant, and deciduous vs evergreen species). In addition to the expected negative correlation of K(l) with drought tolerance, we found a strong, negative correlation between K(l) and species' shade tolerance. Across species, K(s) and K(l) were negatively correlated with wood density and positively with maximum vessel length. Consequently, drought and shade tolerance scaled similarly with hydraulic properties, wood density and leaf dry matter content. We found that deciduous species also had traits conferring efficient water transport relative to evergreen species. Hydraulic properties varied across species, corresponding to the classical trade-off between hydraulic efficiency and safety, which for these dry forest trees resulted in coordinated drought and shade tolerance across species rather than the frequently hypothesized trade-off.