Acacia trees as keystone species in Negev desert ecosystems

Abstract. The only trees in most of the Negev desert are 3 native Acacia species. We tested the hypothesis that they act as keystone species as a result of the improved soil conditions under their canopies. Furthermore, because many Acacia populations suffer high levels of mortality due to water stress, we tested whether trees in high mortality populations had diminished effects on plant species and soil quality under their canopies. We show that plant species diversity beneath the tree canopies is higher than in the surrounding areas. There was also a clearly identifiable suite of species with higher occurrence under the trees. Plant species composition differed significantly between high and low mortality sites. However, there was higher species diversity in high mortality sites and under trees with higher water stress. Soil nutrient content was higher under the trees than in the open areas, especially under larger trees and trees with higher water status. The results indicate that there is a combination of positive and negative effects of Acacia trees on the under‐canopy environment, which may include positive effects of higher soil nutrients and a negative influence of higher soil salinity.     

Patterns of tree dieback in Queensland,Australia: the importance of drought stress and the role of resistance to cavitation

During the extreme 1992–1997 El Niño drought event, widespread stem mortality, or tree dieback, of both mature and juvenile eucalypts occurred within the tropical savannas of northeast Australia. Most of the dieback occurred in individuals of the ironbark species complex (Eucalyptus crebra – E. xanthoclada) while individuals of the bloodwood species Corymbia erythrophloia, exhibited significantly less stem mortality. Indicative of greater water stress, predawn and midday xylem water potentials of ironbark adults and saplings were significantly more negative than predawn values of bloodwoods. The very negative xylem water potentials in ironbarks suggest that stem mortality in both adult and juvenile ironbarks results from drought-induced embolism and that ironbarks perhaps have a shallower and less extensive root system than bloodwoods. Although predawn and midday water potentials for ironbark adults and saplings were similar, a census of mature and juvenile ironbark trees indicated that mortality was higher in adult trees. Cavitation vulnerability curves indicated that ironbark saplings may be better buffered against cavitation than adult trees. If they possess smaller root systems, saplings are more likely than adults to experience low xylem water potentials, even in non-drought years. Xylem conduits produced in adult trees during periods of normal rainfall, although perhaps more efficient in water conduction, may be more vulnerable to cavitation during infrequent severe droughts.     

Dieback and mortality of South African fynbos shrubs is likely driven by a novel pathogen and pathogen‐induced hydraulic failure

We examined whether extensive dry season dieback and mortality in a South African fynbos community were due to drought or pathogen attack. Plant dieback and mortality have been reported elsewhere in similar plant communities suggesting potential for a widespread climatic or biotic threat to this community. We collected tissue samples from Brunia noduliflora, the dominant plant in the community, and cultured them for potential plant pathogens. We also measured dry season predawn and midday water potentials of healthy and stressed plants and constructed pressure‐volume curves to assess turgor loss point. Plant stress and mortality were monitored over a 2‐year study period. Both healthy plants and plants that displayed moderate signs of stress had dry season predawn water potentials well above their turgor loss point suggesting plants were not water stressed. However, plants displaying >60% crown dieback had much lower water potentials (as low as ?12 MPa). A previously undescribed fungus (Pythium sp.) was isolated from the root vascular tissue of all stressed plants and was not present in healthy plants. The proximate cause of plant stress was likely pathogen‐induced, while the ultimate cause of plant death appears to be extreme water stress. The present study suggests that Brunia (Bruniceae), Leucadendron (Proteaceae) and Erica (Ericaceae), all emblematic and dominant genera within the diverse fynbos community, may be susceptible to Pythium infection. This may pose a serious threat to communities already threatened by climate change.     

Pine wilt disease as promising causal agent of the mass mortality of Pinus armandii Franch. var. amamiana (Koidz.) Hatusima in the field

The causal agent of the mass mortality of field populations of Pinus armandii Franch. var. amamiana (Koidz.) Hatusima (PAA) was investigated with special respect to the involvement of pine wilt disease. Wood chips, branches and/or increment cores for detecting the pinewood nematode, feeding marks of the vector insect and environmental stress in the past, respectively, were taken from live and dead PAA trees grown in three locations, Yaku-shima and Tanega-shima Islands and a plantation in Kagoshima City, from 1997 to 1998. Five trees died after the spring of 1996 and, of these, four were inhabited by the pinewood nematode. Feeding marks of the vector insects were found on the branches of all dead trees and most of the live trees investigated. These results suggest that the infection of pine wilt disease in PAA trees occurs in the field. Annual ring growth of the sample trees showed neither intervention nor growth reduction, which implies strong environmental stress that may cause mortality in PAA trees.     

Anthropogenic causes of high mortality and low recruitment in three Acacia tree taxa in the Negev desert, Israel

Large-scale mortality of native Acacia trees has been reported in the Negev desert, Israel. These trees may act as keystone species in this ecosystem due to their nitrogen-fixing abilities. Total mortality was variable and ranged from 0–61%. Contrary to widely-held opinion, aquifer depletion for agriculture and other human uses was not found to be a likely cause of this mortality. We found that the major probable cause of mortality was the absence of culverts under roads cross-cutting the ephemeral river beds where the Acacias are mostly found. Culverts allow water to pass from the upper to lower parts of the river during winter floods. In river beds without culverts, an average of 25% total mortality of trees was recorded downstream from the roads as compared with 12% upstream from the roads. We found that an average of five species of perennial plants disappeared downstream from the roads. An additional cause for concern is the rarity of recruitment of Acacia trees. This may be due to the absence of large mammalian herbivores (both wild and domestic) which eat seed pods and enhance the germination capacity of seeds through scarification of the hard seed coat during digestion.     

Ecophysiological responses of salt cedar (Tamarix spp. L.) to the northern tamarisk beetle (Diorhabdacarinulata Desbrochers) in a controlled environment

The northern tamarisk beetle (Diorhabda carinulata Desbrochers) was released in several western states as a biocontrol agent to suppress Tamarix spp. L. which has invaded riparian ecosystems; however, effects of beetle herbivory on Tamarix physiology are largely undocumented and may have ecosystem ramifications. Herbivory by this insect produces discoloration of leaves and premature leaf drop in these ecosystems, yet the cause of premature leaf drop and the effects of this leaf drop are still unknown. Insect herbivory may change leaf photosynthesis and respiration and may affect a plant’s ability to regulate water loss and increase water stress. Premature leaf drop may affect plant tissue chemistry and belowground carbon allocation. We conducted a greenhouse experiment to understand how Tamarix responds physiologically to adult beetle and larvae herbivory and to determine the proximate cause of premature leaf drop. We hypothesized that plants experiencing beetle herbivory would have greater leaf and root respiration rates, greater photosynthesis, increased water stress, inefficient leaf nitrogen retranslocation, lower root biomass and lower total non-structural carbohydrates in roots. Insect herbivory reduced photosynthesis rates, minimally affected respiration rates, but significantly increased water loss during daytime and nighttime hours and this produced increased water stress. The proximate cause for premature leaf drop appears to be desiccation. Plants exposed to herbivory were inefficient in their retranslocation of nitrogen before premature leaf drop. Root biomass showed a decreasing trend in plants subjected to herbivory. Stress induced by herbivory may render these trees less competitive in future growing seasons.     

Floodwater infiltration through root channels on a sodic clay floodplain and the influence on a local tree species Eucalyptus largiflorens

Dieback of riparian species on floodplains has been attributed to increased soil salinisation due to raised groundwater levels, resulting from irrigation and river regulation. This is exacerbated by a reduction in flooding frequency and duration of inundation. For the Chowilla floodplain on the River Murray raised water tables have increased the amount of salts mobilised in the soil profile, causing the trees to experience salt induced water stress. For the trees to survive in the long term, salts need to be leached from the root zone.This study investigated whether floodwater infiltrates through channels created by E. largiflorens (black box) roots, flushing salts away from roots, thereby allowing the trees to increase their water uptake. Trees at different sites on the floodplain were artificially flooded, by pumping 1.5 kL of creek water into impoundments constructed around the trees. Gas exchange parameters, and pre-dawn and midday water potential were measured the day before, the day after and one week after the artificial flood and compared against trees that were not flooded. Pre-dawn and midday water potentials were also measured one month after the flood. After flooding, the trees experienced less water stress, indicated by an increase in water potential of less than 0.2 MPa, in comparison to non-flooded control trees. However, this response was not evident one month after flooding. The response to flooding did not result in increased rates of transpiration, stomatal conductance or photosynthesis, even though flooding effectively doubled the trees yearly water supply.The infiltration of floodwater in the impoundments around E. largiflorens was also compared to that of impoundments on bare ground. Floodwater infiltrated 2 – 17 times faster around trees than on adjacent bare ground, for parts of the floodplain not grazed by livestock. Tracer dye experiments indicated that bulk flow of water through pores down the profile was the reason for the enhanced infiltration. Flooding leached salts in direct vicinity of tree roots, but only leached small amounts of salts from the bulk soil.     

Anthropogenic effects on population structure of Acacia tortilis subsp. raddiana along a gradient of water availability in South Sinai,Egypt

Acacia trees in Sinai desert are suffering population decline. This study aims to explore this assumption and to determine the effects of grazing and urbanization on size structure of Acacia tortilis subsp. raddiana along a gradient of water availability in Wadi Feiran basin, South Sinai, Egypt. Size structure of 289 Acacia trees in thirteen isolated populations was analysed. Catchment area and lineament density as water availability indicators were calculated. Effects of grazing and urbanization on Acacia populations were evaluated. A strong positive correlation was recognized between tree height, crown diameter and trunk circumference. Acacia tree density and crown diameter are positively correlated with water availability indicators. Population structure of A. tortilis is characterized by absence of juveniles, clear reduction in numbers of small and large categories and increasing in medium categories. This pattern indicates very low recruitment and high mortality of small and large trees. The variation in water availability may have considerable effects on the structure of A. tortilis. Overgrazing, cutting and urbanization are the main causes of population decline of Acacia in Sinai desert. In addition, drought conditions and bruchid seed beetles aggravate the anthropogenic effect on recruitment of Acacia in South Sinai.     

Mechanisms of plant survival and mortality during drought: why do some plants survive while others succumb to drought?

Severe droughts have been associated with regional-scale forest mortality worldwide. Climate change is expected to exacerbate regional mortality events; however, prediction remains difficult because the physiological mechanisms underlying drought survival and mortality are poorly understood. We developed a hydraulically based theory considering carbon balance and insect resistance that allowed development and examination of hypotheses regarding survival and mortality. Multiple mechanisms may cause mortality during drought. A common mechanism for plants with isohydric regulation of water status results from avoidance of drought-induced hydraulic failure via stomatal closure, resulting in carbon starvation and a cascade of downstream effects such as reduced resistance to biotic agents. Mortality by hydraulic failure per se may occur for isohydric seedlings or trees near their maximum height. Although anisohydric plants are relatively drought-tolerant, they are predisposed to hydraulic failure because they operate with narrower hydraulic safety margins during drought. Elevated temperatures should exacerbate carbon starvation and hydraulic failure. Biotic agents may amplify and be amplified by drought-induced plant stress. Wet multidecadal climate oscillations may increase plant susceptibility to drought-induced mortality by stimulating shifts in hydraulic architecture, effectively predisposing plants to water stress. Climate warming and increased frequency of extreme events will probably cause increased regional mortality episodes. Isohydric and anisohydric water potential regulation may partition species between survival and mortality, and, as such, incorporating this hydraulic framework may be effective for modeling plant survival and mortality under future climate conditions.     

Effect of liana cutting on water potential and growth of adult<Emphasis Type="Italic"> Senna multijuga</Emphasis> (Caesalpinioideae) trees in a Bolivian tropical forest

Lianas, or woody climbing plants, are a major constituent of seasonally dry tropical forests, and are thought to impact negatively their host trees. In this study we evaluated whether liana presence was associated with reduced leaf water potentials and growth in adult Senna multijuga trees during the dry season in a lowland Bolivian forest. We used leaf water potentials in trees as a first approach to assess trees’ water status, under the assumption that leaf water potentials become more negative when water losses (via transpiration) exceed gains (by uptake). We measured relative growth in girth at 1.5 m height (gbh) to quantify tree growth. At the beginning of the 1996 dry season (early June), we selected 20 S. multijuga trees 10–20 cm dbh, and measured their gbh. We also recorded pre-dawn and mid-day leaf water potentials in these trees. In ten experimental trees all lianas were then cut, while the remaining trees were used as controls. Pre-dawn and mid-day water potentials were re-measured 1 day after liana-cutting, and then every week in all trees for 1 month and then at 3 and 5 months, until the beginning of the next rainy season (November); gbh was measured again in July 1997 to estimate relative growth rate. Liana removal was associated with less negative pre-dawn (–0.3 vs –0.4 MPa) and mid-day (–0.5 vs –0.7 MPa) water potentials in trees during the dry season. This difference appeared as early as 1 day after cutting, and disappeared once the rainy season began. Liana-cut trees grew more (0.4 mm/mm year) than liana-uncut trees (0.2 mm/mm year). These findings suggest that lianas may interfere with water availability to these trees during the dry season, and may also hinder tree growth. Received: 16 November 1999 / Accepted: 23 March 2000     

Variations in stream water uptake by Eucalyptus camaldulensis with differing access to stream water

The stable isotopes ²H and ¹⁸O were used to determine the water sources of Eucalyptus camaldulensis at three sites with varying exposure to stream water, all underlain by moderately saline groundwater. Water uptake patterns were a function of the long-term availability of surface water. Trees with permanent access to a stream used some stream water at all times. However, water from soils or the water table commonly made up 50% of these trees' water. Trees beside an ephemeral stream had access to the stream 40–50% of the time (depending on the level of the stream). No more than 30% of the water they used was stream water when it was available. However, stream water use did not vary greatly whether the trees had access to the stream for 2 weeks or 10 months prior to sampling. Trees at the third site only had access to surface water during a flood. These trees did not change their uptake patterns during 2 months inundation compared with dry times, so were not utilising the low-salinity flood water. Pre-dawn leaf water potentials and leaf ¹³C measurements showed that the trees with permanent access to the stream experienced lower water stress and had lower water use efficiencies than trees at the least frequently flooded site. The trees beside the ephemeral stream appeared to change their water use efficiency in response to the availability of surface water; it was similar to the perennial-stream trees when stream water was available and higher at other times. Despite causing water stress, uptake of soil water and groundwater would be advantageous to E. camaldulensis in this semi-arid area, as it would provide the trees with a supply of nutrients and a reliable source of water. E. camaldulensis at the study site may not be as vulnerable to changes in stream flow and water quality as previously thought.     

Tree distribution on a steep environmental gradient in an arid savanna

Aim The structure of savannas ranges from scrub to woodland over broad geographical gradients. Here we examine the hypothesis that water availability is a major determinant of these structural differences by ascertaining the relationship between water availability and tree growth across a steep moisture gradient. Location The study site is a sub‐tropical savanna, with a mean annual precipitation of 400 mm, located in the Phugwane river basin, Kruger National Park, Limpopo Province, South Africa. Methods We determined plant moisture stress using xylem pressure potentials, stem growth using dendrometer bands, and estimated the water sources available to plants using stable hydrogen and oxygen isotope ratios. The primary objective was to understand tree growth relative to available water along an environmental gradient. Results We found that Philenoptera violacea trees growing close to the stream have lower water stress (least negative xylem pressure potentials) and higher cumulative growth than those growing away from the stream. The stem growth of P. violacea was characterized by steady incremental growth and could not be related to antecedent rainfall. Colophospermum mopane trees experienced higher water stress, yet trees growing adjacent to the stream achieved the highest cumulative growth rates over the study period. The growth of C. mopane could be clearly linked to antecedent rainfall, and most growth was achieved during short growth pulses that followed rainfall events. δ¹⁸O values become progressively more enriched in the heavier isotope with distance from the stream, suggesting that access to groundwater decreases with distance from the stream side. The isotopic data suggest that P. violacea has access to groundwater, but that C. mopane does not. Main conclusions Our results show that water stress increases with distance from the stream side as a result of reduced access to groundwater. Trees without access to deep water adopt an opportunistic growth strategy.     

小秦岭自然保护区秦岭冷杉死亡原因

小秦岭国家级自然保护区濒危植物秦岭冷杉(Abies chensiensis)近几年出现了不明原因的衰败死亡现象,并有不断扩展的趋势,分析研究就地保护这一珍稀植物资源迫在眉睫。在对小秦岭全部秦岭冷杉定位监测的基础上,通过对活树、死树分布和结构的对比分析,并采用相关分析、偏相关分析和结构等式模型(Structural Equation Models,SEM)结合环境因子,探讨了秦岭冷杉的生存状况和死亡原因。结果如下:(1)枯死树与活树分布基本相一致,活树个体中小树和大树相对较少,死亡个体主要为小树和中龄树;(2)秦岭冷杉的生长与海拔、坡度、土壤含水量和土壤温度呈显著的相关性;(3)SEM分析结果显示,秦岭冷杉活树与土壤含水量和温度呈明显的正相关系,而死树却表现出负相关关系,枯死树与空间位置和地形表现出的负相关关系要明显大于活树。上述结果说明,秦岭冷杉幼苗和小树较少,更新能力差,秦岭冷杉的死亡不是个别现象和局部环境变化所造成的;秦岭冷杉的生长与海拔、坡度、土壤含水量和土壤温度关系密切;土壤含水量和温度对秦岭冷杉生长具有促进作用,是影响秦岭冷杉死亡和衰败的重要环境因素。在今后秦岭冷杉经营保护过程中要注意微环境的变化,减少人为因素对环境的干扰。     

Relationships between individual‐tree mortality and water‐balance variables indicate positive trends in water stress‐induced tree mortality across North America

Accounting for water stress‐induced tree mortality in forest productivity models remains a challenge due to uncertainty in stress tolerance of tree populations. In this study, logistic regression models were developed to assess species‐specific relationships between probability of mortality (P_m) and drought, drawing on 8.1 million observations of change in vital status (m) of individual trees across North America. Drought was defined by standardized (relative) values of soil water content (W_s,z) and reference evapotranspiration (ET_r,z) at each field plot. The models additionally tested for interactions between the water‐balance variables, aridity class of the site (AC), and estimated tree height (h). Considering drought improved model performance in 95 (80) per cent of the 64 tested species during calibration (cross‐validation). On average, sensitivity to relative drought increased with site AC (i.e. aridity). Interaction between water‐balance variables and estimated tree height indicated that drought sensitivity commonly decreased during early height development and increased during late height development, which may reflect expansion of the root system and decreasing whole‐plant, leaf‐specific hydraulic conductance, respectively. Across North America, predictions suggested that changes in the water balance caused mortality to increase from 1.1% yr^?1 in 1951 to 2.0% yr^?1 in 2014 (a net change of 0.9 ± 0.3% yr^?1). Interannual variation in mortality also increased, driven by increasingly severe droughts in 1988, 1998, 2006, 2007 and 2012. With strong confidence, this study indicates that water stress is a common cause of tree mortality. With weak‐to‐moderate confidence, this study strengthens previous claims attributing positive trends in mortality to increasing levels of water stress. This ‘learn‐as‐we‐go’ approach – defined by sampling rare drought events as they continue to intensify – will help to constrain the hydraulic limits of dominant tree species and the viability of boreal and temperate forest biomes under continued climate change.     

Effect of soil temperature on stem sap flow,shoot gas exchange and water potential of Picea engelmannii (Parry) during snowmelt

Summary The effect of cold soils on stem sap flow, shoot gas exchange and water potential of Picea engelmannii (Parry) was investigated during the snowmelt period in the Medicine Bow Mountains, Wyoming, USA. Shoot net photosynthetic rates were higher in young trees (1.5–1.8 m in height) growing in cold soils (<3.5° C) associated with snowpack, than trees in warm soils until about 1500 h. Higher shoot photosynthetic rates of trees in cold soils continued after snow was removed and could not be completely explained by higher visible irradiance over highly reflective snow. Following soil warming higher photosynthetic rates were evident in these trees for five days. High nutrient availability associated with snowmelt may improve shoot nutrient status leading to higher gas-exchange rates during snowmelt. Shoot conductance to water vapor was higher in trees in cold soil until midday, when declining shoot conductance led to lower intercellular CO₂ concentrations. Midday through afternoon shoot water potentials of trees in cold soils were similar or higher than those of trees in warm soils and the lower afternoon shoot conductances in cold soils were not the result of lower bulk shoot water potentials. Decline in net photosynthesis of trees in cold soils at 1500 h paralleled increases in intercellular CO₂ concentrations, implying a nonstomatal limitation of photosynthesis. This scenario occurred consistently in mid-afternoon following higher morning and midday photosynthesis in cold soils, suggesting a carbohydrate feedback inhibition of photosynthesis. Diurnal patterns in stem sap flow of all trees (cold and warm soils) reflected patterns of shoot conductance, although changes in stem sap flow lagged 1–3 h behind shoot conductance apparently due to stem water storage. Total daily stem sap flow was similar in trees in cold and warm soils, although diel patterns differed. The morning surge and night-time drop in sap flow commenced 1–2 h earlier in trees in cold soils. Overnight stem sap flow was lower in trees in cold soils, possibly due to higher resistance to root water uptake in cold soils, which may explain lower predawn shoot water potentials. However, midday shoot water potentials of trees in cold soils equalled or exceeded those of trees in warm soils. Higher resistance to root water uptake in P. engelmannii in cold soils was apparently overshadowed by transpirational forces and significant shoot water deficits did not develop.     

Drought avoidance and the effect of local topography on trees in the understorey of Bornean lowland rain forest

The water relations of two tree species in the Euphorbiaceae werecompared to test in part a hypothesis that the forest understorey plays anintegral role in drought response. At Danum, Sabah, the relatively commonspecies Dimorphocalyx muricatus is associated with ridgeswhilst another species, Mallotus wrayi, occurs widely bothon ridges and lower slopes. Sets of subplots within two 4 -hapermanent plots in this lowland dipterocarp rain forest, were positioned onridges and lower slopes. Soil water potentials were recorded in1995–1997,and leaf water potentials were measured on six occasions. Soil water potentialson the ridges (–0.047 MPa) were significantly lower than onthe lower slopes (–0.012 MPa), but during the driest periodin May 1997 they fell to similarly low levels on both sites (–0.53MPa). A weighted 40-day accumulated rainfall index was developedtomodel the soil water potentials. At dry times, D.muricatus(ridge) had significantly higher pre-dawn (–0.21 v.–0.57 MPa) and mid-day (–0.59 v.–1.77 MPa) leaf water potentials than M.wrayi (mean of ridge and lower slope). Leaf osmotic potentials ofM. wrayi on the ridges were lower (–1.63MPa) than on lower slopes (–1.09 MPa), withthose for D. muricatus being intermediate (–1.29MPa): both species adjusted osmotically between wet and dry times.D. muricatus trees were more deeply rooted thanM. wrayi trees (97 v. 70cm). M. wrayi trees had greaterlateral root cross-sectional areas than D. muricatus treesalthough a greater proportion of this sectional area for D.muricatus was further down the soil profile. D.muricatus appeared to maintain relatively high water potentialsduring dry periods because of its access to deeper water supplies and thus itlargely avoided drought effects, but M. wrayi seemed to bemore affected yet tolerant of drought and was more plastic in its response. Theinteraction between water availability and topography determines these species'distributions and provides insights into how rain forests can withstandoccasional strong droughts.     

Comparative hydraulic and anatomic properties in palm trees (Washingtonia robusta) of varying heights: implications for hydraulic limitation to increased height growth

As trees grow taller, the energetic cost of moving water to the leaves becomes higher and could begin to limit carbon gain and subsequent growth. The hydraulic limitation hypothesis states that as trees grow taller, the path length and therefore frictional resistance of water flow increases, leading to stomatal closure, reduced photosynthesis and decreased height growth in tall trees. Although this hypothesis is supported by the physical laws governing water movement in trees, its validation has been complicated by the complex structure of most tree species. Therefore, this study tested the hydraulic limitation hypothesis in Washingtonia robusta (H. Wendl.), a palm that, while growing to tall heights, is still structurally simple enough to act as a model organism for testing. There were no discernable relationships between tree height and stomatal conductance, stomatal densities, guard cell lengths, leaf dry mass per unit area (LMA) or sap flux, suggesting that these key aspects of hydraulic limitation are not reduced in taller palms. Taller palms did, however, have higher maximum daily photosynthetic assimilation rates, lower minimum leaf water potentials that occurred earlier in the day and fewer, smaller leaves than did shorter palms. Leaf epidermal cells were also smaller in taller palms compared with shorter ones. These findings are consistent with hydraulic compensation in that tall palms may be overcoming the increased path length resistance through smaller, more efficient leaves and lower leaf water potentials than shorter palms.     

Responses of stomata to environmental factors-experiments with isolated epidermal strips of Polypodium vulgare

Summary Stomatal apertures of isolated and suitably conditioned epidermal strips of Polypodium vulgare are described as the stomata respond to the influences of temperature, air humidity, and water potential at the epidermal inner walls. Water stress as a result of reduced water potential in the substomatal airspace leads to narrower stomatal pores when water potential falls below -8 bar. Water potentials above this threshold value show minor influence. Stomatal responses to such water stress strongly interact with the responses to humidity changes in ambient air and to temperature. The linear dependence of stomatal apertures on the vapor saturation deficit of the air (closing) is shifted to lower values (more closed) by lower leaf bulk water potentials.Stomatal behavior depending on the temperature factor seems to be reversed by higher water stress. Without water stress, rising temperatures between 20 and 28° C are accompanied by further opening of the pores, whereas an increase of temperature within this range leads to narrowing of the stomata under the influence of lower water potentials within the substomatal airspace. It can be demonstrated that stomatal aperture values of Polypodium vulgare depending on temperature always describe optimum curves. With no water stress, closing does not occur before rather high temperatures are reached and above a broad range of maximal opening. Water stress, on the other hand, results in more pronounced narrowing of stomatal pores and shifts the onset to considerably lower temperatures.     

Population-specific Traits and their Implication for the Evolution of a Drought-adapted Ecotype in Armeria maritima

Traits contributing to drought resistance of Armeria maritima were investigated by comparing six populations of this species from sandy grasslands, heavy metal mines and salt marsh sites. The sandy soil ecotype that is influenced by periodic drought was found to allocate constitutively a higher proportion of biomass to the root system, especially to the fine roots, than the other two ecotypes. The selective advantage of this lower shoot-root ratio is presumably the delayed onset of the critical water saturation deficit because of the ability to take up additional water from deeper soil layers. Under water stress, all populations of A. maritima showed a decrease in the shoot-root ratio. Additionally, under moderate long-term water stress a decrease in (= more negative) osmotic potential was found in leaves of plants from all populations. Lowest osmotic potentials were shown by the salt marsh ecotype and the highest by one of the heavy metal populations. Osmotic adjustment was achieved passively by a reduction in osmotic volume. Accumulation of osmotically active compounds was seen only under severe water stress, when the considerable betaine contents found in all populations of A. maritima even under control conditions were further increased by severe drought. The evolution of traits related to drought adaptation in the sandy soil ecotype of A. maritima is discussed.     

Causes of reduced leaf‐level photosynthesis during strong El Niño drought in a Central Amazon forest

Sustained drought and concomitant high temperature may reduce photosynthesis and cause tree mortality. Possible causes of reduced photosynthesis include stomatal closure and biochemical inhibition, but their relative roles are unknown in Amazon trees during strong drought events. We assessed the effects of the recent (2015) strong El Niño drought on leaf‐level photosynthesis of Central Amazon trees via these two mechanisms. Through four seasons of 2015, we measured leaf gas exchange, chlorophyll a fluorescence parameters, chlorophyll concentration, and nutrient content in leaves of 57 upper canopy and understory trees of a lowland terra firme forest on well‐drained infertile oxisol. Photosynthesis decreased 28% in the upper canopy and 17% in understory trees during the extreme dry season of 2015, relative to other 2015 seasons and was also lower than the climatically normal dry season of the following non‐El Niño year. Photosynthesis reduction under extreme drought and high temperature in the 2015 dry season was related only to stomatal closure in both upper canopy and understory trees, and not to chlorophyll a fluorescence parameters, chlorophyll, or leaf nutrient concentration. The distinction is important because stomatal closure is a transient regulatory response that can reverse when water becomes available, whereas the other responses reflect more permanent changes or damage to the photosynthetic apparatus. Photosynthesis decrease due to stomatal closure during the 2015 extreme dry season was followed 2 months later by an increase in photosynthesis as rains returned, indicating a margin of resilience to one‐off extreme climatic events in Amazonian forests.