Responses of riparian trees to interannual variation in ground water depth in a semi-arid river basin

We investigated the physiological and growth responses of native (Populus fremontii S. Wats. and Salix gooddingii Ball) and exotic (Tamarix chinensis Lour.) riparian trees to ground water availability at the free‐flowing Hassayampa River, Arizona, during dry (1997) and wet (1998) years. In the drier year, all species experienced considerable water stress, as evidenced by low shoot water potentials, low leaf gas exchange rates and large amounts of canopy dieback. These parameters were significantly related to depth of ground water (DGW) in the native species, but not in T. chinensis, in 1997. Canopy dieback was greater in the native species than in T. chinensis when ground water was deep in 1997, and dieback increased rapidly at DGW > 2·5–3·0 m for the native species. Analysis of combined data from wet and dry years for T. chinensis tentatively suggests a similar physiological sensitivity to water availability and a similar DGW threshold for canopy dieback. In 1998, shoot water potential and leaf gas exchange rates were higher and canopy dieback was lower for all species because of increased water availability. However, T. chinensis showed a much larger increase in leaf gas exchange rates in the wet year than the native species. High leaf gas exchange rates, growth when water is abundant, drought tolerance and the maintenance of a viable canopy under dry conditions are characteristics that help explain the ability of T. chinensis to thrive in riparian ecosystems in the south‐western United States.     

Genetic variation in productivity of foundation riparian species at the edge of their distribution: implications for restoration and assisted migration in a warming climate

We examined the hypothesis that genotypic variation among populations of commonly co‐occurring phreatophytic trees (Populus fremontii, Salix gooddingii) and the shrub (Salix exigua) regulates aboveground net primary productivity (ANPP) at a hot site at the edge of the species’ distribution. We used a provenance trial in which replicated genotypes from populations varying in mean annual temperature were transplanted to a common garden adjacent to the Lower Colorado River in southeastern California. The garden environment represented an extreme maximum temperature for the study species. Four major findings emerged: (1) Genotypic variation in ANPP was significant for all species with broad‐sense heritability (H²) across populations of 0.11, 0.13, and 0.10 for P. fremontii, S. gooddingii, and S. exigua, respectively, and within‐population H² ranging from 0.00 to 0.25, 0.00 to 0.44, and 0.02 to 0.21, respectively. (2) Population ANPP decreased linearly as mean annual maximum temperature (MAMT) transfer distance increased for both P. fremontii (r² = 0.64) and S. gooddingii (r² = 0.37), whereas it did not change for S. exigua; (3) Populations with similar MAMT to that of the common garden were 1.5 and 1.2 times more productive than populations with 5.0 °C MAMT transfer distances for P. fremontii and S. gooddingii, respectively; and (4) Variation in regression slopes among species for the relationship between ANPP and MAMT indicate species‐specific responses to temperature. As these plant species characterize a threatened habitat type and support a diverse community that includes endangered species, ecosystem restoration programs should consider using both local genotypes and productive genotypes from warmer environments to maximize productivity of riparian ecosystems in the face of global climate change.     

Variation in structural and physiological leaf traits of eight species in karst desertification area of China

Leaf traits have long been recognized as influential factors in the acquisition and processing of resources by plants. However, there is less knowledge of between-species variations in seasonal changes in leaf traits and trait interrelationships. Therefore, we examined variations in leaf area (LA), dry biomass (DM), specific leaf area (SLA), and leaf gas-exchange parameters in one non-native and seven native tree species under field environmental conditions, in a karst area in China subjected to desertification. Measurements were taken three times during the growing season. The results show that the seven native trees had higher LA, DM, and water-use efficiency (WUE) than the non-native Cinnamomum camphora. In contrast, all the native tree species except Ligustrum lucidum had lower photosynthetic rates (P _N) than the non-native species. In all species, the relationship between LA and DM was less variable than the relationship between SLA and LA. However, leaves of the non-native C. camphora and native species Sterculia lanceolata, Cleidiocarpon cavalerei and Cyclobalanopsis glauca were highly sensitive to seasonal conditions, leaves of Sapindus mukorossi and Ligustrum lucidum were less sensitive to seasonal changes, and leaves of Syzygium cumini and Cephalomappa sinensis were insensitive. An understanding of leaf traits will aid the selection of suitable species for land restoration.     

Leaf carbon isotope discrimination and nitrogen content for riparian trees along elevational transects

 Leaf carbon isotope discrimination (Δ), seasonal estimates of the leaf-to-air water vapor gradient on a molar basis (ω), and leaf nitrogen contents were examined in three riparian tree species (Populus fremontii, P. angustifolia, and Salix exigua) along elevational transects in northern and southern Utah USA (1500–2670 m and 600–1820 m elevational gradients, respectively). The ω values decreased with elevation for all species along transects. Plants growing at higher elevations exhibited lower Δ values than plants at lower elevations (P. fremontii, 22.9‰ and 19.5‰, respectively; P. angustifolia, 23.2‰ and 19.2‰, respectively; and S.␣exigua, 21.1‰ and 19.1‰, respectively). Leaf nitrogen content increased with elevation for all species, suggesting that photosynthetic capacity at a given intercellular carbon dioxide concentration was greater at higher elevations. Leaf Δ and nitrogen content values were highly correlated, implying that leaves with higher photosynthetic capacities also had lower intercellular carbon dioxide concentrations. No significant interannual differences were detected in carbon isotope discrimination. Received: 25 February 1996 / Accepted: 8 September 1996     

Leaf Longevity,Construction, and Maintenance Costs of Three Mangrove Species Under Field Conditions

This study assessed the effect of leaf age on construction cost (CC) in the mangrove species Avicennia germinans, Laguncularia racemosa, and Rhizophora mangle growing in their natural habitat. Leaf osmolality values were species-specific, the highest in A. germinans (1 693 mmol kg^–1) and the lowest in L. racemosa (1 270 mmol kg^–1). In the three species, contents of chlorophyll (a+b) (Chl_a+b) and nitrogen (N) per unit of leaf area were maximal in adult leaves and tended to decline with age. Leaf mass to leaf area ratio (LMA) and ash content increased during leaf ageing. Similarly, as leaves aged, a significant increase in leaf construction cost per leaf area (CC_a) was observed, while per leaf mass (CC_m) it remained almost constant, suggesting a sustained production of leaf compounds as leaves became older. CC was positively correlated with LMA and heat of combustion (Hc) per leaf area, suggesting differences among species in the quantity and composition of expensive compounds. Leaf half lifetime (t_0.5) showed contrasting values in the three mangrove species (60, 111, and 160 d in L. racemosa, R. mangle, and A. germinans, respectively). Overall, L. racemosa was the species with less expensive leaves to construct while leaves of A. germinans and R. mangle had the highest CC_m and CC_a, respectively. Leaf longevity was positively correlated with the ratio between CC and maximum photosynthetic rate (P _max), clearly showing the existence of a balance between leaf costs and benefits.     

Photosynthesis,Transpiration, and Water Use Efficiency of Two Puccinellia Species on the Songnen Grassland,Northeastern China

The intra- and inter-specific variations in net photosynthetic (P _N) and transpiration (E) rates and water use efficiency (WUE) of Puccinellia tenuiflora and Puccinellia chinampoensis leaves were compared. The two species experienced a similar habitat, but differed in leaf area, leaf colour, and nitrogen contents. Leaf P _N and E for both reproductive and vegetative shoots of the two species declined with leaf age. P _N for reproductive shoots was less than for vegetative shoots, but their E was greater than that of vegetative shoots in the dry season. The average P _N and E for reproductive shoots of P. tenuiflora were lower than those of P. chinampoensis, but higher for vegetative shoots.     

Over-winter decomposition and associated macroinvertebrate communities of three deciduous leaf species in forest streams on the Canadian Boreal Shield

The decomposition of deciduous leaf material provides a critical source of energy to aquatic food webs. Changes to riparian forests through harvesting practices may alter the species composition of deciduous leaf material entering streams. We compared over-winter decomposition of three different riparian leaf species (speckled alder (Alnus incana ssp. rugosa (Du Roi) J. Clausen), white birch (Betula papyrifera Marsh.), and trembling aspen (Populus tremuloides Michx.)) to determine their importance as a food resource for macroinvertebrate communities within Boreal Shield streams in northeastern Ontario, Canada. Leaf pack decomposition of the three leaf species formed a processing continuum throughout winter, where alder and birch leaf packs decomposed at a medium rate (k = 0.0065/day and 0.0053/day, respectively) and aspen leaf packs decomposed more slowly (k = 0.0035/day). Macroinvertebrate community colonization on leaf packs changed through time regardless of leaf species. Alder leaf packs supported higher abundances of macroinvertebrates in the fall while aspen leaf packs supported greater shredder abundances in the following spring. The study shows that leaf diversity may be important for providing a sustained food resource for aquatic macroinvertebrates throughout the relatively long over-winter period in Canadian Boreal Shield streams. Riparian forest management strategies should ensure that deciduous plant species richness is sustained in riparian areas.     

Photosynthetic Characteristics,Dark Respiration,and Leaf Mass Per Unit Area in Seedlings of Four Tropical Tree Species Grown Under Three Irradiances

We investigated the effect of growth irradiance (I) on photon-saturated photosynthetic rate (P _max), dark respiration rate (R _D), carboxylation efficiency (CE), and leaf mass per unit area (LMA) in seedlings of the following four tropical tree species with contrasting shade-tolerance. Anthocephalus chinensis (Rubiaceae) and Linociera insignis (Oleaceae) are light-demanding, Barringtonia macrostachya (Lecythidaceae) and Calophyllum polyanthum (Clusiaceae) are shade-tolerant. Their seedlings were pot-planted under shading nets with 8, 25, and 50 % daylight for five months. With increase of I, all species displayed the trends of increases of LMA, photosynthetic saturation irradiance, and chlorophyll-based P _max, and decreases of chlorophyll (Chl) content on both area and mass bases, and mass-based P _max, R _D, and CE. The area-based P _max and CE increased with I for the light-demanders only. Three of the four species significantly increased Chl-based CE with I. This indicated the increase of nitrogen (N) allocation to carboxylation enzyme relative to Chl with I. Compared to the two shade-tolerants, under the same I, the two light-demanders had greater area- and Chl-based P _max, photosynthetic saturation irradiance, lower Chl content per unit area, and greater plasticity in LMA and area- or Chl-based P _max. Our results support the hypothesis that light-demanding species is more plastic in leaf morphology and physiology than shade-tolerant species, and acclimation to I of tropical seedlings is more associated with leaf morphological adjustment relative to physiology. Leaf nitrogen partitioning between photosynthetic enzymes and Chl also play a role in the acclimation to I.     

Evidence for facultative deciduousness in Colophospermum mopane in semi‐arid African savannas

Leaf phenology dictates the time available for carbon assimilation, transpiration and nutrient uptake in plants. Understanding the environmental cues that control phenology is therefore vital for predicting climate‐related changes to plant and ecosystem function. In contrast to temperate systems, and to a lesser degree, tropical forest systems, the cues initiating leaf drop in tropical savannas are poorly studied. We investigated the cues for leaf fall in a tropical monodominant arid savanna species, Colophospermum mopane, using an irrigation experiment. We tracked soil moisture, solar radiation, air temperature, leaf water status, leaf health and leaf carbon balance through the dry season in both irrigated and control plants. Water was the primary cue driving leaf loss of C. mopane rather than temperature or light. Trees watered throughout the dry season retained their canopies. These leaves remained functional and continued photosynthesis throughout the dry season. Leaf carbon acquisition rates did not decline with leaf age but were affected by soil moisture availability and temperature. Leaf loss did not occur when leaf carbon gain was zero, or when a particular leaf carbon threshold was reached. Colophospermum mopane is facultatively deciduous as water availability determines leaf drop in this widespread arid savanna species. Obligate deciduosity is not the only successful strategy in climates with a long dry season.     

Habitat Use of Juvenile Fish in the Lower Danube and the Danube Delta: Implications for Ecotone Connectivity

Functional processes in freshwater ecosystems are highly influenced by acidic conditions. Foodwebs are affected and macroinvertebrate species diversity is decreased. This study aims to investigate leaf decomposition at very low pH in the acidic Banyupahit–Banyuputih river originating from the acidic crater lake Kawah Ijen in Indonesia. Leaf decomposition experiments were carried out for 200 days in the acidic river at pHs of approximately 0.7, 2.3 and 3.0 and in the neutral Kali Sengon river, using leaves from teak, Tectona grandis, and bamboo, Bambusa sp. Two different types of leaf packs were used: fine mesh size packs were used to exclude macroinvertebrates and coarse mesh size packs allowed macroinvertebrate colonization. Clear differences in decomposition rate were observed between the neutral Kali Sengon and the acidic Banyupahit–Banyuputih river with decomposition in the Kali Sengon river proceeding significantly faster for both leaf types. In the Kali Sengon k values (d⁻¹) over 46 days were 0.0202 for fine teak, 0.0236 for coarse teak, 0.0114 for fine bamboo and 0.0151 for coarse bamboo. No significant differences were observed between the three sites in the acidic Banyupahit–Banyuputih river with k values of 0.0034–0.0066 for fine teak, 0.0002–0.0057 for coarse teak, 0.0029–0.0054 for fine bamboo and 0.0000–0.0068 for coarse bamboo. Moreover, no clear adaptation of macroinvertebrates or microbes to low pH conditions could be detected. The coarse mesh leaf packs in the neutral Kali Sengon river revealed that macroinvertebrates are important in the breakdown process. Fine mesh packs revealed that microbial activity is depressed under acidic conditions. Based on this evidence, we conclude that the toxicity at low pH conditions, and probably also the precipitation of metals on the leaf material, seriously affects leaf decomposition.     

Influence of salinity on Na+ and K+ accumulation, and gas exchange in Avicennia germinans

We analysed plant growth, ion accumulation, leaf water relations, and gas exchange of Avicennia germinans (L.) L. subjected to a long-term, controlled salinity gradient from 0 to 55 ‰. Growth and leaf area were affected by salinity higher than 10 ‰. As salinity increased, the predawn leaf water potential (Ψ_w) and leaf osmotic potential (Ψ_s) decreased. Leaf Ψ_w was at least −0.32 MPa lower than the Ψ_w of solution. Na⁺ and K⁺ ions explained about 78 % of decrease in Ψ_s. K⁺ tissue water concentration decreased by more than 60 % in all salinity treatments as compared with those grown at 0 ‰. Inversely, Na⁺ concentration in tissue water increased with nutrient solution salinity. The maximum net photosynthetic rate (P _N) and stomatal conductance (g _s) decreased by 68 and 82 %, respectively, as salinity increased from 0 to 55 ‰; the intercellular CO₂ concentration (C _i) followed the same trend. The P _N as a function of C _i showed that both the initial linear slope and upper plateau of the P _N vs. C _i curve were markedly affected by high salinity (40 and 55 ‰).     

Leaf temperature effects on gas exchange in Quercus ilex L. growing under field conditions

ABSTRACT

Gas exchange temperature dependence in Quercus ilex shrubs growing in the Mediterranean maquis was analysed. The gas exchange trend was monitored during the year: photosynthetic activity (A _net) reached the highest average rates in early spring and autumn (12.5 µmol m^-2s^-1 was the absolute maximum A _net measured) and the lowest rates were monitored in the middle of June. There was a good correlation (r = 0.72) between A _net and g _s (A _net = 4.1246 ln g _s + 4316; P < 0.01), indicating that stomatal control of CO₂ diffusion plays an important role in controlling photosynthetic activity. Leaf temperature allowing the highest photosynthetic and stomatal conductance rates of Quercus ilex were in the range 17.5 – 29°C. A _net and g_s dropped below half its maximum value when leaf temperatures were below 11.5°C and above 35.7°C. Transpiration rates (E) were strongly related to leaf temperature; E increased as leaf temperature increased and the highest E rates were monitored in June, despite a 46% decrease in g _s. Leaf water loss from transpiration, during the drought period, could result in leaf water stress which would exacerbate heat effects on photosynthesis. During summer, the increase in leaf temperatures decreased g _s which in turn decreased A _net. Consequently, stomatal control in Quercus ilex may be considered as an adaptive strategy during drought.     

Leaf traits of natural populations of <Emphasis Type="Italic">Adiantum reniforme</Emphasis> var. <Emphasis Type="Italic">sinensis</Emphasis>, endemic to the Three Gorges region in China

Leaf mass per unit area (LMA), carbon and nitrogen contents, leaf construction cost, and photosynthetic capacity (P _max) of Adiantum reniforme var. sinensis, an endangered fern endemic to the Three Gorges region in southwest China, were compared in five populations differing in habitat such as soil moisture and irradiance. The low soil moisture and high irradiance habitat population exhibited significantly higher LMA, area-based leaf construction (CC_A), and carbon content (C_A), but lower leaf nitrogen content per unit dry mass (N_M) than the other habitat populations. The high soil moisture and low irradiance habitat populations had the lowest CC_A, but their cost/benefic ratios of CCA/P _max were similar to the medium soil moisture and irradiance habitat population due to their lower leaf P _max. Hence A. reniforme var. sinensis prefers partially shaded, moist but well-drained, slope habitats. Due to human activities, however, its main habitats now are cliffs or steeply sloped bare rocks with poor and thin soil. The relatively high energy requirements and low photosynthetic capacity in these habitats could limit the capability of the species in extending population or interspecific competition and hence increase its endangerment.     

Local biotic adaptation of trees and shrubs to plant neighbors

Natural selection as a result of plant–plant interactions can lead to local biotic adaptation. This may occur where species frequently interact and compete intensely for resources limiting growth, survival, and reproduction. Selection is demonstrated by comparing a genotype interacting with con‐ or hetero‐specific sympatric neighbor genotypes with a shared site‐level history (derived from the same source location), to the same genotype interacting with foreign neighbor genotypes (from different sources). Better genotype performance in sympatric than allopatric neighborhoods provides evidence of local biotic adaptation. This pattern might be explained by selection to avoid competition by shifting resource niches (differentiation) or by interactions benefitting one or more members (facilitation). We tested for local biotic adaptation among two riparian trees, Populus fremontii and Salix gooddingii, and the shrub Salix exigua by transplanting replicated genotypes from multiple source locations to a 17 000 tree common garden with sympatric and allopatric treatments along the Colorado River in California. Three major patterns were observed: 1) across species, 62 of 88 genotypes grew faster with sympatric neighbors than allopatric neighbors; 2) these growth rates, on an individual tree basis, were 44, 15 and 33% higher in sympatric than allopatric treatments for P. fremontii, S. exigua and S. gooddingii, respectively, and; 3) survivorship was higher in sympatric treatments for P. fremontii and S. exigua. These results support the view that fitness of foundation species supporting diverse communities and dominating ecosystem processes is determined by adaptive interactions among multiple plant species with the outcome that performance depends on the genetic identity of plant neighbors. The occurrence of evolution in a plant‐community context for trees and shrubs builds on ecological evolutionary research that has demonstrated co‐evolution among herbaceous taxa, and evolution of native species during exotic plants invasion, and taken together, refutes the concept that plant communities are always random associations.     

Leaf surface properties of Norway spruce needles exposed to sulphur dioxide and ozone in an open-air fumigation system at Liphook

Samples of current-year and 1-year-old foliage were taken from Norway spruce (Picea abies (L.) Karst.) trees in April 1991, 4 months after a 3–4 year controlled fumigation with O₃ and SO₂ in the open at Liphook, south-east England. Trees were grown in seven plots, and treated in a factorial design with three levels of SO₂ and two levels of O₃ (ambient and c. 1.3 × ambient), with an extra ambient air plot. All statistical analyses were made on plot means. Leaf wettability, as measured by the contact angle of water droplets, was significantly affected by needle age and by SO₂ treatment (P≤0–05. in older needles, decreasing with increasing SO₂ concentration. There was no effect of O₃ on wettability, and no effect of any treatment on amounts of surface wax extracted by immersion of needles in chloroform. Electrolyte leakage rates from detached current-year needles were not affected by prior exposure to O₃, but decreased significantly (P= 0.034) with increasing exposure to SO₂. There was no detectable effect of fumigation on the rate of water loss from detached needles. Similarly, there was no effect of fumigation on the dry weight/fresh weight ratio of needles. The total sulphur content of needles increased significantly (P≤0.0001) with exposure to SO₂ and with needle age. Amounts of water-extractable sulphate, however, varied greatly among plots, but with no pattern with respect to fumigation treatment. It is concluded that leaf wettability and electrolyte leakage rates may be good indicators of the persistent effects of SO₂ on Norway spruce growing in the open air, and that the observed changes in leaf surface properties in response to SO₂ fumigation have implications for the processes, both biotic and abiotic, that occur on leaf surfaces.     

Feeding habits of two sympatric loliginid squids,Uroteuthis (Photololigo) chinensis (Gray, 1849) and Uroteuthis (Photololigo) duvaucelii (d’Orbigny, 1835), in the lower part of the South China Sea

The feeding habits of two sympatric squid species, Uroteuthis (Photololigo) chinensis and Uroteuthis (Photololigo) duvaucelii from the southwestern Gulf of Thailand were studied. They fed on low numbers of food types (AF) and had a low diet breadth; 1.18 and 0.01 for U. (P.) chinensis and 1.49 and 0.05 for U. (P.) duvaucelii, respectively. Three major prey types (fishes, crustaceans and molluscs) were always detected and cannibalism was observed. Fish was the greatest contributor to the diet of both species, contributing 89.5% for U. (P.) chinensis and 69.9% for U. (P.) duvaucelii. Fish size significantly affected fullness index (FL) and AF for U. (P.) chinensis (P?<?0.001) and U. (P.) duvaucelii (P?<?0.001). Depth affected the FL of U. (P.) chinensis (P?<?0.001) but not of U. (P.) duvaucelii (P?>?0.05). Maturity stages of both male and female U. (P.) chinensis influenced FL (male: P?<?0.001; female: P?<?0.05) and AF (male: P?<?0.05; female: P?<?0.01). The FL of squid from cast nets was higher than those from trawls. The multivariate results showed dietary grouping between size classes of both species.     

Photosynthesis and water-relation traits of the summer annual C4 grasses, Eleusine indica and Digitaria adscendens, with contrasting trampling tolerance

Two summer annual C₄ grasses with different trampling susceptibilities were grown as potted plants, and diurnal leaf gas exchange and leaf water potential in each grass were compared. The maximum net photosynthetic rate, leaf conductance and transpiration rate were higher in the trampling-tolerant Eleusine indica (L.) Gaertn. than in trampling sensitive Digitaria adscendens (H. B. K.) Henr. Leaf water potential was much lower in E. indica than in D. adscendens. There were no differences in soil-to-leaf hydraulic conductance and leaf osmotic potential at full turgor as obtained by pressure–volume analysis. However, the bulk modulus of elasticity in cell walls was higher in E. indica leaves than in D. adscendens leaves. This shows that the leaves of E. indica are less elastic. Therefore, the rigid cell walls of E. indica leaves reduced leaf water potential rapidly by decreasing the leaf water content, supporting a high transpiration rate with high leaf conductance. In trampled habitats, such lowering of leaf water potential in E. indica might play a role in water absorption from the compacted soil. In contrast, the ability of D. adscendens to colonize dry habitats such as coastal sand dunes appears to be due to its lower transpiration rate and its higher leaf water potential which is not strongly affected by decreasing leaf water content.     

Physiological divergences between two rhizomatous grasses from a desertification steppe,North China

The inter-and intra-specific physiological differences, e.g. rates of net photosynthesis (P _N) and transpiration (E), stomatal conductance (g _s), and water use efficiency (WUE), were compared between two grasses, Calamagrostis epigeios (L.) Roth. and Psammochloa villosa (Trin.) Bor., and between their leaf types in a desertification steppe in North China. The two species had a similar habitat, but differed in leaf area and rhizome depth. Leaf P _N, E, and g _s for P. villosa were significantly greater than those for C. epigeios in the growing season, but WUE for the former species was only 50 and 80 % of that for the latter one in dry and rainy seasons, respectively. In general, leaf P _N, E, g _s, and WUE for both vegetative and reproductive shoots of the two species exhibited little variations between leaf types or with leaf age, even though there were some remarkable differences between dry and rainy seasons. The mean leaf P _N and E in reproductive shoots of P. villosa were significantly lower than those in its vegetative shoots in rainy season, while these differences were much smaller for those of C. epigeios. P. villosa with deeper rhizome roots has relative higher leaf P _N, E, and g _s, but a smaller WUE in the arid desertification steppe region.     

Functional relationships between hydraulic traits and the timing of diurnal depression of photosynthesis

The hydraulic coordination along the water transport pathway helps trees provide adequate water supply to the canopy, ensuring that water deficits are minimized and that stomata remain open for CO₂ uptake. We evaluated the stem and leaf hydraulic coordination and the linkages between hydraulic traits and the timing of diurnal depression of photosynthesis across seven evergreen tree species in the southern Andes. There was a positive correlation between stem hydraulic conductivity (k_s) and leaf hydraulic conductance (K_Leaf) across species. All species had similar maximum photosynthetic rates (A_max). The species with higher k_s and K_Leaf attained A_max in the morning, whereas the species with lower k_s and K_Leaf exhibited their A_max in the early afternoon concurrently with turgor loss. These latter species had very negative leaf water potentials, but far from the pressure at which the 88% of leaf hydraulic conductance is lost. Our results suggest that diurnal gas exchange dynamics may be determined by leaf hydraulic vulnerability such that a species more vulnerable to drought restrict water loss and carbon assimilation earlier than species less vulnerable. However, under stronger drought, species with earlier CO₂ uptake depression may increase the risk of hydraulic failure, as their safety margins are relatively narrow.     

Contrasting foliar responses to drought in Quercus ilex and Phillyrea latifolia

Leaf morphology, longevity, and demography were examined in Quercus ilex and Phillyrea latifolia growing in a holm oak forest in Prades mountains (northeast Spain). Four plots (10 × 15 m) of this forest were submitted to an experimental drought during three years (soil moisture was reduced about 15 %). Leaf area, thickness and leaf mass per area ratio (LMA) were measured in sun and shade leaves of both species. Leaf longevity, the mean number of current-year shoots produced per previous-year shoot (Sn/Sn-1), the mean number of current-year leaves per previous-year shoot (Ln/Sn-1), and the percentage of previous-year shoots that developed new ones were measured once a year, just after leaf flushing. LMA and leaf thickness increased since leaf unfolding except in summer periods, when stomatal closure imposed low photosynthetic rates and leaves consumed their reserves. LMA, leaf area, and leaf thickness were higher in Q. ilex than in P. latifolia, but leaf density was higher in the latter species. Drought reduced the leaf thickness and the LMA of both species ca. 2.5 %. Drought also increased leaf shedding up to ca. 20 % in Phillyrea latifolia and decreased it up to ca. 20 % in Q. ilex. In the later species, Sn/Sn-1 decreased by 32 %, Ln/Sn-1 by 41 %, percentage of shoots developed new ones by 26 %, and leaf area by 17 %. Thus the decrease of leaf number and area was stronger in the less drought-resistant Q. ilex, which, under increasingly drier conditions, might lose its current competitive advantage in these Mediterranean holm oak forests.