Differential ecophysiological responses to seasonal drought of three co-existing oak species in northern Greece

Oaks (Quercus spp.) represent the most important broadleaf genus with respect to forest-shaping tree species in the Mediterranean. Considering future climate scenarios (increased drought conditions), the identification of drought tolerant oak species is of great importance for future forest management in this region. The objective of the study was the comparison of physiological status of three economically and ecologically valuable oak species (Quercus ilex, Quercus frainetto and Quercus pubescens) co-existing in natural coppice stands in NE Greece, in response to seasonal drought stress. Measurements were conducted between June and September 2016, every 15–20 days until leaf falling. The parameters studied were predawn leaf water potential and fast chlorophyll fluorescence induction curves (OJIP test), chlorophyll content, and relative water content. Meteorological data from the area were also collected. Photosynthetic parameters such as performance indices (PI_abs and PI_tot) reacted to summer drought conditions, with Q. frainetto showing the lowest values. The discrepancy between species increased with duration of drought period. Q. frainetto revealed the lowest predawn water potential values. The results indicate that Q. frainetto is less suitable for future forestry applications in the studied climate/elevation zone than Q. pubescens and Q. ilex.     

Wood anatomy and hydraulic architecture of stems and twigs of some Mediterranean trees and shrubs along a mesic-xeric gradient

Woody species populating the major Mediterranean ecosystems in the world are characterized by different levels of adaptation to the seasonal Mediterranean climate conditions. Many species of these ecosystems show wood features that allow high efficiency of transport when water is available, while maintaining hydraulic safety during drought periods. This study focuses on the anatomy of juvenile and mature wood of some species representative of continuous sequences of Mediterranean vegetation formations according to gradients of water availability, from xeric to relatively mesic: Cistus monspeliensis L., Rhamnus alaternus L., Myrtus communis L., Pistacia lentiscus L., Olea europaea L., Quercus ilex L., Fraxinus ornus L. and Ostrya carpinifolia L. Twigwood collected in Southern Italy was anatomically compared with the stemwood of the same species represented in the reference slide collection of the National Herbarium of the Netherlands (Lw). The “hydraulic distance” between the wood of main stems and twigs was estimated on the basis of suites of anatomical features related to water efficiency/safety. Although some attributes (i.e. porosity and type of imperforate tracheary elements) were similar in young twigs and older rings, other traits (i.e. vessel frequency and size) evidenced the different hydraulic properties of twig and stemwood. The difference between juvenile and mature structures was large in the species of the mesic end of the gradient while it was relatively small in those more xeric. This tendency is in agreement with the habit gradient from medium-sized trees to small evergreen/drought deciduous shrubs according to decreasing water availability in Mediterranean vegetation types.     

Wood anatomy of some trees with diffuse- and ring-porous wood: Some functional and ecological interpretations

Abstract

The xylem conduit dimensions (i.e. their width and length) have been measured in 1-year-old internodes, nodes and node-to-petiole (N-P) junctions of three species with diffuse-porous wood, namely Ceratonia siliqua L., Laurus nobilis L. and Olea europaea L. as well as of three species with ring-porous wood, namely Quercus ilex L., Q. suber L. and Q. pubescens Willd‥ The xylem conduit diameter and length distributions have been related to the drought resistance strategies adopted by the six species. C. siliqua and Q. ilex (drought avoiding water spenders) showed the widest xylem conduits (each species within its characteristic pattern of wood anatomy). This is consistent with their high demand of efficient water transport to leaves. L. nobilis (drought avoiding water saver) showed relatively narrow xylem conduits, efficient enough, however, to assure water supply to leaves at the reduced transpiration rate exhibited by the species. O. europaea, Q. suber and Q. pubescens (drought tolerants) showed the narrowest xylem conduits but also the longest ones. The xylem system of C. siliqua and Q. ilex represented a good compromise between efficiency and safety of the water transport, the former as due to wide xylem conduits, the latter to the reduced xylem conduit length as well as to the strong «hydraulic constrictions» at their nodes and N-P junctions. The ecological interpretation of such hydraulic architecture is discussed.     

Drought and shade deplete nonstructural carbohydrate reserves in seedlings of five temperate tree species

Plants that store nonstructural carbohydrates (NSC) may rely on carbon reserves to survive carbon‐limiting stress, assuming that reserves can be mobilized. We asked whether carbon reserves decrease in resource stressed seedlings, and if NSC allocation is related to species' relative stress tolerances. We tested the effects of stress (shade, drought, and defoliation) on NSC in seedlings of five temperate tree species (Acer rubrum Marsh., Betula papyrifera Marsh., Fraxinus americana L., Quercus rubra L., and Quercus velutina Lam.). In a greenhouse experiment, seedlings were subjected to combinations of shade, drought, and defoliation. We harvested seedlings over 32–97 days and measured biomass and NSC concentrations in stems and roots to estimate depletion rates. For all species and treatments, except for defoliation, seedling growth and NSC accumulation ceased. Shade and drought combined caused total NSC decreases in all species. For shade or drought alone, only some species experienced decreases. Starch followed similar patterns as total NSC, but soluble sugars increased under drought for drought‐tolerant species. These results provide evidence that species deplete stored carbon in response to carbon limiting stress and that species differences in NSC response may be important for understanding carbon depletion as a buffer against shade‐ and drought‐induced mortality.     

Vegetazione e ambiente delle Marche e relativi problemi di salvaguardia

Abstract

Vegetation and environment in the Marche and related problems of protection. – The « Marche » region is situated in central Italy and corresponds with the natural region east of the Appennine ridge and spreading as far as the coast which is washed by the Adriatic Sea.

As to the climate, we may distinguish a coastal strip with Mesomediterranean climate with dry season, corresponding to the distribution area of Quercus ilex; a hilly strip with Submediterranean climate, corresponding to the distribution area of Quercus pubescens; a mountain strip with temperate axeric climate with sub-dry period, corresponding to the distribution area of Fagus silvatica and finally a high mountain strip with cold axeric climate, corresponding to altitude pastures. The Marche vegetation is shown in the vegetation map of Fabriano sheet (scale 1:50,000) plotted according Ozenda's vegetational series.

The following vegetation series have been plotted: submediterranean series of Quercus ilex; series of thermophilic oak wood with priority of Quercus pubescens or Quercus cerris; series of Ostrya carpinifolia; series of Quercus petraea and series of Fagus silvatica.

As far as the problems to safegard environment are concerned, a plan is given providing for the institution of 4 complete nature reserves, 34 nature reserves, and 9 regional nature parks among which those of Monte Conero, of Monti Sibillini and Monti della Laga.     

Stem hydraulic capacitance decreases with drought stress: implications for modelling tree hydraulics in the Mediterranean oak Quercus ilex

Hydraulic modelling is a primary tool to predict plant performance in future drier scenarios. However, as most tree models are validated under non‐stress conditions, they may fail when water becomes limiting. To simulate tree hydraulic functioning under moist and dry conditions, the current version of a water flow and storage mechanistic model was further developed by implementing equations that describe variation in xylem hydraulic resistance (R_X) and stem hydraulic capacitance (C_S) with predawn water potential (Ψ_PD). The model was applied in a Mediterranean forest experiencing intense summer drought, where six Quercus ilex trees were instrumented to monitor stem diameter variations and sap flow, concurrently with measurements of predawn and midday leaf water potential. Best model performance was observed when C_S was allowed to decrease with decreasing Ψ_PD. Hydraulic capacitance decreased from 62 to 25 kg m^?3 MPa^?1 across the growing season. In parallel, tree transpiration decreased to a greater extent than the capacitive water release and the contribution of stored water to transpiration increased from 2.0 to 5.1%. Our results demonstrate the importance of stored water and seasonality in C_S for tree hydraulic functioning, and they suggest that C_S should be considered to predict the drought response of trees with models.     

Elevated growth temperatures alter hydraulic characteristics in trembling aspen (Populus tremuloides) seedlings: implications for tree drought tolerance

Although climate change will alter both soil water availability and evaporative demand, our understanding of how future climate conditions will alter tree hydraulic architecture is limited. Here, we demonstrate that growth at elevated temperatures (ambient +5 °C) affects hydraulic traits in seedlings of the deciduous boreal tree species Populus tremuloides, with the strength of the effect varying with the plant organ studied. Temperature altered the partitioning of hydraulic resistance, with greater resistance attributed to stems and less to roots in warm‐grown seedlings (P < 0.02), and a 46% (but marginally significant, P = 0.08) increase in whole plant conductance at elevated temperature. Vulnerability to cavitation was greater in leaves grown at high than at ambient temperatures, but vulnerability in stems was similar between treatments. A soil–plant–atmosphere (SPA) model suggests that these coordinated changes in hydraulic physiology would lead to more frequent drought stress and reduced water‐use efficiency in aspen that develop at warmer temperatures. Tissue‐specific trade‐offs in hydraulic traits in response to high growth temperatures would be difficult to detect when relying solely on whole plant measurements, but may have large‐scale ecological implications for plant water use, carbon cycling and, possibly, plant drought survival.     

Leaf and stem physiological responses to summer and winter extremes of woody species across temperate ecosystems

Winter cold limits temperate plant performance, as does summer water stress in drought‐prone ecosystems. The relative impact of seasonal extremes on plant performance has received considerable attention for individual systems. An integrated study compiling the existing literature was needed to identify overall trends. First, we conducted a meta‐analysis of the impacts of summer and winter on ecophysiology for three woody plant functional types (winter deciduous angiosperms, evergreen angiosperms and conifers), including data for 210 records from 75 studies of ecosystems with and without summer drought across the temperate zone. Second, we tested predictions by conducting a case study in a drought‐prone Mediterranean ecosystem subject to winter freezing. As indicators of physiological response of leaves and xylem to seasonal stress, we focused on stomatal conductance (g_s), percent loss of stem xylem hydraulic conductivity (PLC) and photochemical efficiency of photosystem II (F_v/F_m). Our meta‐analysis showed that in ecosystems without summer drought, g_s was higher during summer than winter. By contrast, in drought‐prone ecosystems many species maintained open stomata during winter, with potential strong consequences for plant carbon gain over the year. Further, PLC tended to increase and F_v/F_m to decrease from summer to winter for most functional types and ecosystems due to low temperatures. Overall, deciduous angiosperms were most sensitive to climatic stress. Leaf gas exchange and stem xylem hydraulics showed a coordinated seasonal response at ecosystems without summer drought. In our Mediterranean site subjected to winter freezing the species showed similar responses to those typically found for ecosystems without summer drought. We conclude that winter stress is most extreme for systems without summer drought and systems with summer drought and winter freezing, and less extreme for drought‐prone systems without freezing. In all cases the evergreen species show less pronounced seasonal responses in both leaves and stems than deciduous species.     

Radial growth responses to drought of Pinus sylvestris and Quercus pubescens in an inner‐Alpine dry valley

Question: Lower montane treeline ecotones such as the inner Alpine dry valleys are regarded as sensitive to climate change. In the dry Valais valley (Switzerland) the composition of the widespread, low altitude Pinus forests is shifting towards a mixed deciduous state. The sub‐boreal P. sylvestris shows high mortality rates, whereas the deciduous sub‐mediterranean Quercus pubescens is spreading. These species may act as early indicators of climate change. We evaluate this hypothesis by focusing on their differences in drought tolerance, which are hardly known, but are likely to be crucial in the current forest shift and also for future forest development. Methods: We used dendroecological methods to detect species‐specific patterns in the growth response to drought. The relationship between radial growth of 401 trees from 15 mixed stands and drought was analysed by calculating response functions using yearly tree‐ring indices and monthly drought indices. PCA was applied to the response ratios to discover spatial patterns of drought response. Results: A species‐specific response to moisture as well as a sub‐regional differentiation of the response patterns were found. While Quercus showed a response mainly to the conditions of the previous autumn and those of current spring, Pinus did not start responding before May, but showed responses throughout the whole summer. Quercus may restrict physiological activity to moist periods; growth of Pinus was much more dependent on prior growth. Conclusions: Given that the climate is changing towards (1) longer summer drought periods, (2) higher mean temperatures and (3) shifted seasonally of moisture availability, Quercus may benefit from adapting better to drier conditions. Pinus may increasingly face problems related to drought stress as it depends on summer moisture and has a smaller adaptive capacity due to its long‐lived photosynthetic tissue.     

Variations in physiological and biochemical traits of oak seedlings grown under drought and ozone stress

Despite the huge biodiversity characterizing the Mediterranean environment, environmental constraints, such as high sunlight and high temperatures alongside with dry periods, make plant survival hard. In addition, high irradiance leads to increasing ozone (O₃) concentrations in ambient air. In this era of global warming, it is necessary to understand the mechanisms that allow native species to tolerate these environmental constraints and how such mechanisms interact. Three Mediterranean oak species (Quercus ilex, Quercus pubescens and Quercus cerris) with different features (drought tolerant, evergreen or deciduous species) were selected to assess their biometrical, physiological and biochemical responses under drought and/or O₃ stress (80–100 nl l^?1 of O₃ for 5 h day^?1 for 77 consecutive days). Leaf visible injury appeared only under drought stress (alone or combined with O₃) in all three species. Drought × O₃ induced strong reductions in leaf dry weight in Q. pubescens and Q. cerris (?70 and ?75%, respectively). Alterations in physiological (i.e. decrease in maximum carboxylation rate) and biochemical parameters (i.e. increase in proline content and build‐up of malondialdehyde by‐products) occurred in all the three species, although drought represented the major determinant. Quercus ilex and Q. pubescens, which co‐occur in dry environments, were more tolerant to drought and drought × O₃. Quercus ilex was the species in which oxidative stress occurred only when drought was applied with O₃. High plasticity at a biochemical level (i.e. proline content) and evergreen habitus are likely on the basis of the higher tolerance of Q. ilex.     

The fate of recently fixed carbon after drought release: towards unravelling C storage regulation in Tilia platyphyllos and Pinus sylvestris

Carbon reserves are important for maintaining tree function during and after stress. Increasing tree mortality driven by drought globally has renewed the interest in how plants regulate allocation of recently fixed C to reserve formation. Three‐year‐old seedlings of two species (Tilia platyphyllos and Pinus sylvestris) were exposed to two intensities of experimental drought during ~10 weeks, and ¹³C pulse labelling was subsequently applied with rewetting. Tracking the ¹³C label across different organs and C compounds (soluble sugars, starch, myo‐inositol, lipids and cellulose), together with the monitoring of gas exchange and C mass balances over time, allowed for the identification of variations in C allocation priorities and tree C balances that are associated with drought effects and subsequent drought release. The results demonstrate that soluble sugars accumulated in P. sylvestris under drought conditions independently of growth trends; thus, non‐structural carbohydrates (NSC) formation cannot be simply considered a passive overflow process in this species. Once drought ceased, C allocation to storage was still prioritized at the expense of growth, which suggested the presence of ‘drought memory effects’, possibly to ensure future growth and survival. On the contrary, NSC and growth dynamics in T. platyphyllos were consistent with a passive (overflow) view of NSC formation.     

Effects of increasing root carbon investment on the mortality and resprouting of Haloxylon ammodendron seedlings under drought

• Tree mortality induced by drought is one of the most complex processes in ecology. Although two mechanisms associated with water and carbon balance are proposed to explain tree mortality, outstanding problems still exist.
• Here, in order to test how the root system benefits survival and resprouting of Haloxylon ammodendron seedlings, we examined the various water‐ and carbon‐related physiological indicators (shoot water potential, photosynthesis, dark respiration, hydraulic conductance and non‐structural carbohydrates [NSC]) of H. ammodendron seedlings, which were grown in drought and control conditions throughout a grow season in greenhouse.
• The survival time of the seedling root system (died 70 days after drought) doubled the survival time of the shoot (died at 35 days). Difference in survival time between shoot and root resulted from sustained root respiration supported by increased NSC in roots under drought. Furthermore, investment into the root contributed to resprouting following drought. Based on these results, a death criterion is proposed for this species. The time sequence of major events indicated that drought shifted carbon allocation between shoot and root and altered the flux among different sinks (growth, respiration or storage). The interaction of water and carbon processes determined death or survival of droughted H. ammodendron seedlings.
• These findings revealed that the ‘root protection’ strategy is critical in determining survival and resprouting of this species, and provided insights into the effects of carbon and water dynamics on tree mortality.

     

The combined effects of a long‐term experimental drought and an extreme drought on the use of plant‐water sources in a Mediterranean forest

Vegetation in water‐limited ecosystems relies strongly on access to deep water reserves to withstand dry periods. Most of these ecosystems have shallow soils over deep groundwater reserves. Understanding the functioning and functional plasticity of species‐specific root systems and the patterns of or differences in the use of water sources under more frequent or intense droughts is therefore necessary to properly predict the responses of seasonally dry ecosystems to future climate. We used stable isotopes to investigate the seasonal patterns of water uptake by a sclerophyll forest on sloped terrain with shallow soils. We assessed the effect of a long‐term experimental drought (12 years) and the added impact of an extreme natural drought that produced widespread tree mortality and crown defoliation. The dominant species, Quercus ilex, Arbutus unedo and Phillyrea latifolia, all have dimorphic root systems enabling them to access different water sources in space and time. The plants extracted water mainly from the soil in the cold and wet seasons but increased their use of groundwater during the summer drought. Interestingly, the plants subjected to the long‐term experimental drought shifted water uptake toward deeper (10–35 cm) soil layers during the wet season and reduced groundwater uptake in summer, indicating plasticity in the functional distribution of fine roots that dampened the effect of our experimental drought over the long term. An extreme drought in 2011, however, further reduced the contribution of deep soil layers and groundwater to transpiration, which resulted in greater crown defoliation in the drought‐affected plants. This study suggests that extreme droughts aggravate moderate but persistent drier conditions (simulated by our manipulation) and may lead to the depletion of water from groundwater reservoirs and weathered bedrock, threatening the preservation of these Mediterranean ecosystems in their current structures and compositions.     

Co‐occurring woody species have diverse hydraulic strategies and mortality rates during an extreme drought

From 2011 to 2013, Texas experienced its worst drought in recorded history. This event provided a unique natural experiment to assess species‐specific responses to extreme drought and mortality of four co‐occurring woody species: Quercus fusiformis, Diospyros texana, Prosopis glandulosa, and Juniperus ashei. We examined hypothesized mechanisms that could promote these species' diverse mortality patterns using postdrought measurements on surviving trees coupled to retrospective process modelling. The species exhibited a wide range of gas exchange responses, hydraulic strategies, and mortality rates. Multiple proposed indices of mortality mechanisms were inconsistent with the observed mortality patterns across species, including measures of the degree of iso/anisohydry, photosynthesis, carbohydrate depletion, and hydraulic safety margins. Large losses of spring and summer whole‐tree conductance (driven by belowground losses of conductance) and shallower rooting depths were associated with species that exhibited greater mortality. Based on this retrospective analysis, we suggest that species more vulnerable to drought were more likely to have succumbed to hydraulic failure belowground.     

Impacts of long‐term precipitation manipulation on hydraulic architecture and xylem anatomy of piñon and juniper in Southwest USA

Hydraulic architecture imposes a fundamental control on water transport, underpinning plant productivity, and survival. The extent to which hydraulic architecture of mature trees acclimates to chronic drought is poorly understood, limiting accuracy in predictions of forest responses to future droughts. We measured seasonal shoot hydraulic performance for multiple years to assess xylem acclimation in mature piñon (Pinus edulis ) and juniper (Juniperus monosperma ) after 3+ years of precipitation manipulation. Our treatments consisted of water addition (+20% ambient precipitation), partial precipitation‐exclusion (?45% ambient precipitation), and exclusion‐structure control. Supplemental watering elevated leaf water potential, sapwood‐area specific hydraulic conductivity, and leaf‐area specific hydraulic conductivity relative to precipitation exclusion. Shifts in allocation of leaf area to sapwood area enhanced differences between irrigated and droughted K _L in piñon but not juniper. Piñon and juniper achieved similar K _L under ambient conditions, but juniper matched or outperformed piñon in all physiological measurements under both increased and decreased precipitation treatments. Embolism vulnerability and xylem anatomy were unaffected by treatments in either species. Absence of significant acclimation combined with inferior performance for both hydraulic transport and safety suggests piñon has greater risk of local extirpation if aridity increases as predicted in the southwestern USA.     

Germination and regeneration mechanisms in Mediterranean degenerate forests

Abstract. Germination and developmental behaviour during juvenile stages are of importance in relation to succession mechanisms acting on plant communities. This is particularly so in the Mediterranean region, where the climatic summer drought causes stress. We investigated and compared germination and development of four tree species in two parts of the French Mediterranean region where the original dominant tree species (Quercus pubescens) has been largely replaced by Quercus ilex in the southern part, and Castanea sativa in the northern part. Observed differences in germination rates and developmental behaviour, especially shootroot ratio and the role of water supply, suggest that other species may eventually replace the current dominant species.     

Embolism spread in the primary xylem of Polystichum munitum: implications for water transport during seasonal drought

Xylem network structure and function have been characterized for many woody plants, but less is known about fern xylem, particularly in species endemic to climates where water is a limiting resource for months at a time. We characterized seasonal variability in soil moisture and frond water status in a common perennial fern in the redwood understory of a costal California, and then investigated the consequences of drought‐induced embolism on vascular function. Seasonal variability in air temperature and soil water content was minimal, and frond water potential declined slowly over the observational period. Our data show that Polystichum munitum was protected from significant drought‐induced hydraulic dysfunction during this growing season because of a combination of cavitation resistant conduits (Air‐seeding threshold (ASP) = ?1.53 MPa; xylem pressure inducing 50% loss of hydraulic conductivity (P₅₀) = ?3.02 MPa) and a soil with low moisture variability. High resolution micro‐computed tomography (MicroCT) imaging revealed patterns of embolism formation in vivo for the first time in ferns providing insight into the functional status of the xylem network under drought conditions. Together with stomatal conductance measurements, these data suggest that P. munitum is adapted to tolerate drier conditions than what was observed during the growing season.     

Dynamics ofOstrya carpinifolia woods in the Southern Alps (N-Italy)

Forest regeneration in the vegetation complex of the submontane belt in the Southern Alps involves the active participation ofOstrya carpinifolia woods and their wood edge communities. The corresponding syndynamical processes are described by employing phytosociological, phytogeographical and ecological methods. It is concluded that:

1. The communities of the grassland-wood transition are of major importance in the successional developments in this man-made vegetation complex.
2. Many species occurring in the region of the deciduous forests of Eurasia find their refuge in such transitional communities and are supposed to play an important part in the succession.
3. Ostrya carpinifolia is considered as an early successional tree species.

     

Species‐specific stomatal response of trees to drought – a link to vegetation dynamics?

Question: Is stomatal regulation specific for climate and tree species, and does it reveal species‐specific responses to drought? Is there a link to vegetation dynamics? Location: Dry inner alpine valley, Switzerland Methods: Stomatal aperture (θ_E) of Pinus sylvestris, Quercus pubescens, Juniperus communis and Picea abies were continuously estimated by the ratio of measured branch sap flow rates to potential transpiration rates (adapted Penman‐Monteith single leaf approach) at 10‐min intervals over four seasons. Results: θ_E proved to be specific for climate and species and revealed distinctly different drought responses: Pinus stomata close disproportionately more than neighbouring species under dry conditions, but has a higher θ_E than the other species when weather was relatively wet and cool. Quercus keeps stomata more open under drought stress but has a lower θ_E under humid conditions. Juniperus was most drought‐tolerant, whereas Picea stomata close almost completely during summer. Conclusions: The distinct microclimatic preferences of the four tree species in terms of θ_E strongly suggest that climate (change) is altering tree physiological performances and thus species‐specific competitiveness. Picea and Pinus currently live at the physiological limit of their ability to withstand increasing temperature and drought intensities at the sites investigated, whereas Quercus and Juniperus perform distinctly better. This corresponds, at least partially, with regional vegetation dynamics: Pinus has strongly declined, whereas Quercus has significantly increased in abundance in the past 30 years. We conclude that θ_E provides an indication of a species' ability to cope with current and predicted climate.     

Growth and physiological responses to successional water deficit and recovery in four warm‐temperate woody species

Plant responses to drought and their subsequent rehydration can provide evidence for forest dynamics within the context of climate change. In this study, the seedlings of two native species (Vitex negundo var. heterophylla, Quercus acutissima) and two exotic species (Robinia pseudoacacia, Amorpha fruticosa) to China were selected in a greenhouse experiment. The gas exchange, stem hydraulic parameters, plant osmoprotectant contents and antioxidant activities of the seedlings that were subjected to sustained drought and rehydration (test group) as well as those of well‐irrigated seedlings (control group) were measured. The two native species exhibited a greater degree of isohydry with drought because they limited the stomatal opening timely from the onset of the drought. However, the two exotic species showed a more ‘water spender’‐like strategy with R. pseudoacacia showing anisohydric responses and A. fruticosa showing isohydrodynamic responses to drought. Severe drought significantly decreased the leaf gas exchange rates and hydraulic properties, whereas the instantaneous water use efficiency and osmoprotectant contents increased markedly. Most of the physiological parameters recovered rapidly after mild drought rehydration, but the water potential and/or supply of nonstructural carbohydrates did not recover after severe drought rehydration. The results demonstrate that the xylem hydraulic conductivity and shoot water potential jointly play a crucial role in the drought recovery of woody plants. In brief, the native species may play a dominant role in the future in warm‐temperate forests because they employ a better balance between carbon gain and water loss than the alien species under extreme drought conditions.