Responses of terrestrial ecosystems to climate change have been explored in many regions worldwide. While continued drying and warming may alter process rates and deteriorate the state and performance of ecosystems, it could also lead to more fundamental changes in the mechanisms governing ecosystem functioning. Here we argue that climate change will induce unprecedented shifts in these mechanisms in historically wetter climatic zones, towards mechanisms currently prevalent in dry regions, which we refer to as ‘dryland mechanisms’. We discuss 12 dryland mechanisms affecting multiple processes of ecosystem functioning, including vegetation development, water flow, energy budget, carbon and nutrient cycling, plant production and organic matter decomposition. We then examine mostly rare examples of the operation of these mechanisms in non-dryland regions where they have been considered irrelevant at present. Current and future climate trends could force microclimatic conditions across thresholds and lead to the emergence of dryland mechanisms and their increasing control over ecosystem functioning in many biomes on Earth.

Water availability in vineyards varies in space and time. The spatiotemporal variability of water availability to vines is affected by terrain, meteorology, and irrigation. This study aimed to analyze the response of vegetative and reproductive attributes of vines to spatiotemporal variability in water availability. We quantified the spatial autocorrelation of terrain covariates and grapevine attributes and determined the relative influence (RI) of the covariates on these attributes. In each of four growing seasons (2017-2020), in a Vitis vinifera cv. ``Sauvignon Blanc'' vineyard, five vegetative and reproductive attributes were collected for 240 vines. Terrain covariates included elevation, slope, aspect, topographic wetness index, and categorical landforms; and meteorological covariates consisted of annual rainfall and chilling hours. A Moran's I statistic was computed for each spatial covariate and each grapevine attribute during each season to define temporal changes in spatial variability. A multivariate analysis using gradient boosted regression trees algorithm was applied to extract the RI of the covariates. The results showed high spatial autocorrelation of the terrain covariates and a strong negative shift in spatial dependency of the grapevine attributes throughout the experiment. Yield and number of clusters per vine were highly affected by seasonal precipitation (RI of 46.15% and 42.59%), while changes in inter-seasonal cluster weight and pruning weight were highly subjected to irrigation amounts (RI of 23% and 26.66%), with complementary terrain influences. The number of canes per vine was mainly affected by terrain characteristics. Long-term changes in grapevine attributes depended on meteorological shifts, while higher precipitation amounts were associated with weaker responses of vines to irrigation strategies. The spatial patterns of terrain affected water distribution in the vineyard and controlled the spatial dynamics of grapevine attributes. Knowledge regarding the space-time trends of water availability effects on grapevine attributes may assist decision making of irrigation practices in vineyards.

Lemon (Citrus limon) is a fruit tree with major agricultural importance around the Mediterranean basin and is considered to be highly drought resistant. In this study, we tested the effect of two months summer-desiccation on physiological and yield parameters of mature lemon trees growing under Mediterranean climate during three consecutive years. We also examined the efficiency of current irrigation regime, which is based on reference evapo-transpiration. We measured leaf gas exchange and water potential (Psi(l)), monitored sap flow and soil moisture and followed flowering, fruit set and fruit size. Lemon trees showed an isohydric stomatal regulation, as stomata maintained leaf water potential >-2 MPa. Summer desiccation caused a gradual decrease in diurnal tree water use, starting immediately after cessation of irrigation, with leaf gas exchange practically halted at the end of the drought period. Tree function recovered following re-irrigation, and fruit yields were not reduced, but even mildly increased during the first year. In contrast, summer desiccation during two consecutive years caused long-term effects of tree activity decrease, significantly lower yield, main branch collapse and even tree mortality. Irrigation amounts matched closely tree water-use amounts; soil moisture was maintained around 26% (v/v); and irrigation responded dynamically to meteorological changes, indicating that current irrigation regime represents highly efficient water management. The lemon desiccation protocol relied on the physiological capacity of this species to avoid short-term drought effects through stomatal closure. Still, this protocol must be managed carefully, to reduce risk to trees and save yields.

Wine quality is the final outcome of the interactions within a vineyard between meteorological conditions, terrain and soil properties, plant physiology and numerous viticultural decisions, all of which are commonly summarized as the terroir effect. Associations between wine quality and a single soil or topographic factor are usually weak, but little information is available on the effect of terrain (elevation, aspect and slope) as a compound micro-terroir factor. We used the topographic wetness index (TWI) as a steady-state hydrologic and integrative measure to delineate management zones (MZs) within a vineyard and to study the interactions between vine vigor, water status and grape and wine quality. The study was conducted in a commercial 2.5-ha Vitis vinifera `Cabernet Sauvignon' vineyard in Israel. Based on the TWI, the vineyard was divided into three MZs located along an elongate wadi that crosses the vineyard and bears water only in the rainy winter season. MZ1 was the most distant from the wadi and had low TWI values, MZ3 was closest to the wadi and had high TWI values. Remotely sensed crop water stress index (CWSI) was measured simultaneously with canopy cover (as determined by normalized difference vegetation index; NDVI) and with field measurements of midday stem water potential (psi(stem)) and leaf area index (LAI) on several days during the growing seasons of 2017 and 2018. Vines in MZ1 had narrow trunk diameter and low LAI and canopy cover on most measurement days compared to the other two MZs. MZ1 vines also exhibited the highest water stress (highest CWSI and lowest psi(stem)), lowest yield and highest wine quality. MZ3 vines showed higher LAI on most measurement days, lowest water deficit stress (psi(stem)) during phenological stage I, highest yield and lowest wine quality. Yet, in stage III, MZ3 vines exhibited a similar water deficit stress (CWSI and psi(stem)) as MZ2, suggesting that the relatively high vigor in MZ3 vines resulted in higher water deficit stress than expected towards the end of the season, possibly because of high water consumption over the course of the season. TWI and its classification into three MZs served as a reliable predictor for most of the attributes in the vineyard and for their dynamics within the season, and, thus, can be used as a key factor in delineation of MZs for irrigation. Yet, in-season remotely sensed monitoring is required to follow the vine dynamics to improve precision irrigation decisions.

Perennial plants perpetually adapt to environmental changes in complex and yet insufficiently understood manner. We aimed to separate the intra-seasonal temperature effects on structure and function from perennial and annual water stress effects. This study focused on grapevine (Vitis vinifera L. `Cabernet Sauvignon') petioles, which being a continuously produced organ, represent the current status of the plant. Field-grown mature plants subjected to multi-annual irrigation treatments (severe water stress, mild water stress and non-stressed) throughout the growing season were compared with greenhouse-grown plants under three temperature regimes (22, 28 and 34 degrees C). Physiological and functional anatomy parameters were measured. A generalized additive model (GAM) based on meteorological and lysimeter-based field data was applied to determine the relative influence of various meteorological parameters on evapotranspiration (ETc) during the growing season in the field experiment. At the beginning of the growing season, in May, petioles in the severe stress treatment showed a stress-related structure (decreased length, safer hydraulic structure and increased lignification), though having high values of stem water potential (SWP). As the season progressed and temperatures increased, all water availability treatments petioles showed similar changes, and at the end of season, in August, were structurally very similar. Those changes were independent of SWP and were comparable to high temperature-induced changes in the greenhouse. In contrast, stems hydraulic structure was strongly influenced by water availability. Regression analyses indicated a relationship between petioles xylem structure and stomatal conductance (g(s)), whereas g(s) (but not SWP) was temperature-dependent. The GAM showed that ETc was mainly dependent on temperature. Our results indicate a perennial water-stress memory response, influencing the petiole structure at the beginning of the following season. Intra-seasonally, the petiole's structure becomes independent of water status, whereas temperature drives the structural changes. Thus, ongoing climate change might disrupt plant performance by purely temperature-induced effects.

In anticipation of a drier climate and to project future changes in forest dynamics, it is imperative to understand species-specific differences in drought resistance. The objectives of this study were to form a comprehensive understanding of the drought resistance strategies adopted by Eastern Mediterranean woodland species, and to elaborate specific ecophysiological traits that can explain the observed variation in survival among these species. We examined leaf water potential (Psi), gas exchange and stem hydraulics during 2-3 years in mature individuals of the key woody species Phillyrea latifolia L., Pistacia lentiscus L. and Quercus calliprinos Webb that co-exist in a dry woodland experiencing similar to 6 rainless summer months. As compared with the other two similarly functioning species, Phillyrea displayed considerably lower Psi (minimum Psi of -8.7 MPa in Phillyrea vs -4.2 MPa in Pistacia and Quercus), lower Psi at stomatal closure and lower leaf turgor loss point (Psi(TLP)), but reduced hydraulic vulnerability and wider safety margins. Notably, Phillyrea allowed Psi to drop below Psi(TLP) under severe drought, whereas the other two species maintained positive turgor. These results indicate that Phillyrea adopted a more anisohydric drought resistance strategy, while Pistacia and Quercus exhibited a more isohydric strategy and probably relied on deeper water reserves. Unlike the two relatively isohydric species, Phillyrea reached complete stomatal closure at the end of the dry summer. Despite assessing a large number of physiological traits, none of them could be directly related to tree mortality. Higher mortality was observed for Quercus than for the other two species, which may result from higher water consumption due to its 2.5-10 times larger crown volume. The observed patterns suggest that similar levels of drought resistance in terms of survival can be achieved via different drought resistance strategies. Conversely, similar resistance strategies in terms of isohydricity can lead to different levels of vulnerability to extreme drought.

Despite the important role of tropical forest ecosystems in the uptake and storage of atmospheric carbon dioxide (CO2), the carbon (C) dynamics of tropical tree species remains poorly understood, especially regarding belowground roots. This study assessed the allocation of newly assimilated C in the fast-growing pioneer tropical tree species Ceiba pentandra (L.), with a special focus on different root categories. During a 5-day pulse-labelling experiment, 9-month-old (~3.5-m-tall) saplings were labelled with 13CO2 in a large-scale aeroponic facility, which allowed tracing the label in bulk biomass and in non-structural carbohydrates (sugars and starch) as well as respiratory CO2 from the canopy to the root system, including both woody and non-woody roots. A combined logistic and exponential model was used to evaluate 13C mean transfer time and mean residence time (MRT) to the root systems. We found 13C in the root phloem as early as 2 h after the labelling, indicating a mean C transfer velocity of 2.4 ± 0.1 m h−1. Five days after pulse labelling, 27% of the tracers taken up by the trees were found in the leaves and 13% were recovered in the woody tissue of the trunk, 6% in the bark and 2% in the root systems, while 52% were lost, most likely by respiration and exudation. Larger amounts of 13C were found in root sugars than in starch, the former also demonstrating shorter MRT than starch. Of all investigated root categories, non-woody white roots (NRW) showed the largest 13C enrichment and peaked in the deepest NRW (2–3.5 m) as early as 24 ± 2 h after labelling. In contrast to coarse woody brown roots, the sink strength of NRW increased with root depth. The findings of this study improve the understanding of C allocation in young tropical trees and provide unique insights into the changing contributions of woody and non-woody roots to C sink strengths with depth.

Abstract Drought-related tree mortality is increasing globally, but the sequence of events leading to it remains poorly understood. To identify this sequence, we used a 2016 tree mortality event in a semi-arid pine forest where dendrometry and sap flow measurements were carried out in 31 trees, of which seven died. A comparative analysis revealed three stages leading to mortality. First, a decrease in tree diameter in all dying trees, but not in the surviving trees, 8?months ?prior to the visual signs of mortality? (PVSM; e.g., near complete canopy browning). Second, a decay to near zero in the diurnal stem swelling/shrinkage dynamics, reflecting the loss of stem radial water flow in the dying trees, 6?months PVSM. Third, cessation of stem sap flow 3?months PVSM. Eventual mortality could therefore be detected long before visual signs were observed, and the three stages identified here demonstrated the differential effects of drought on stem growth, water storage capacity and soil water uptake. The results indicated that breakdown of stem radial water flow and phloem function is a critical element in defining the ?point of no return? in the sequence of events leading to mortality of mature trees.

An experiment in variable-rate drip irrigation (VRDI) was aimed at reducing variance of midday stem water potential (SWP) and in yield parameters by applying VRDI in a highly variable vineyard. During 2018, irrigation was separated into VRDI and uniform irrigation (UI) blocks. Each block was delineated to 10 management cells, and results of 2018 season were compared to 2017 season under UI. Standard error (SE) of SWP in the last 3 measurements before harvest decreased in the VRDI block in 2018 compared to 2017 by 14-44%. In contrast, in the UI block, SE in 2018 was higher by 11-42% compared to 2017. SE of fruit yield showed a similar trend. Applying principles of precision irrigation may lead to a more homogeneous vineyard in the parameters described above and to improved wine quality. © Wageningen Academic Publishers 2019

The rate of change in atmospheric CO2 is significantly affected by the terrestrial carbon sink, but the size and spatial distribution of this sink, and the extent to which it can be enhanced to mitigate climate change are highly uncertain. We combined carbon stock (CS) and eddy covariance (EC) flux measurements that were collected over a period of 15 years (2001–2016) in a 55 year old 30 km2 pine forest growing at the semiarid timberline (with no irrigating or fertilization). The objective was to constrain estimates of the carbon (C) storage potential in forest plantations in such semiarid lands, which cover  18% of the global land area. The forest accumulated 145–160 g C m−2 year−1 over the study period based on the EC and CS approaches, with a mean value of 152.5 ± 30.1 g C m−2 year−1 indicating 20% uncertainty in carbon uptake estimates. Current total stocks are estimated at 7,943 ± 323 g C/m2 and 372 g N/m2. Carbon accumulated mostly in the soil ( 71% and 29% for soil and standing biomass carbon, respectively) with long soil carbon turnover time (59 years). Regardless of unexpected disturbances beyond those already observed at the study site, the results support a considerable carbon sink potential in semiarid soils and forest plantations, and imply that afforestation of even 10% of semiarid land area under conditions similar to that of the study site, could sequester  0.4 Pg C/year over several decades. © 2019 John Wiley & Sons Ltd

Climate change is a world-wide threat to biodiversity and ecosystem structure, functioning and services. To understand the underlying drivers and mechanisms, and to predict the consequences for nature and people, we urgently need better understanding of the direction and magnitude of climate change impacts across the soil–plant–atmosphere continuum. An increasing number of climate change studies are creating new opportunities for meaningful and high-quality generalizations and improved process understanding. However, significant challenges exist related to data availability and/or compatibility across studies, compromising opportunities for data re-use, synthesis and upscaling. Many of these challenges relate to a lack of an established ‘best practice’ for measuring key impacts and responses. This restrains our current understanding of complex processes and mechanisms in terrestrial ecosystems related to climate change. To overcome these challenges, we collected best-practice methods emerging from major ecological research networks and experiments, as synthesized by 115 experts from across a wide range of scientific disciplines. Our handbook contains guidance on the selection of response variables for different purposes, protocols for standardized measurements of 66 such response variables and advice on data management. Specifically, we recommend a minimum subset of variables that should be collected in all climate change studies to allow data re-use and synthesis, and give guidance on additional variables critical for different types of synthesis and upscaling. The goal of this community effort is to facilitate awareness of the importance and broader application of standardized methods to promote data re-use, availability, compatibility and transparency. We envision improved research practices that will increase returns on investments in individual research projects, facilitate second-order research outputs and create opportunities for collaboration across scientific communities. Ultimately, this should significantly improve the quality and impact of the science, which is required to fulfil society's needs in a changing world. © 2019 The Authors. Methods in Ecology and Evolution published by John Wiley & Sons Ltd on behalf of British Ecological Society.

Nutrient cycling in most terrestrial ecosystems is controlled by moisture-dependent decomposer activity. In arid ecosystems, plant litter cycling exceeds rates predicted based on precipitation amounts, suggesting that additional factors are involved. Attempts to reveal these factors have focused on abiotic degradation, soil–litter mixing and alternative moisture sources. Our aim was to explore an additional hypothesis that macro-detritivores control litter cycling in deserts. We quantified the role different organisms play in clearing plant detritus from the desert surface, using litter baskets with different mesh sizes that allow selective entry of micro-, meso- or macrofauna. We also measured soil nutrient concentrations in increasing distances from the burrows of a highly abundant macro-detritivore, the desert isopod Hemilepistus reaumuri. Macro-detritivores controlled the clearing of plant litter in our field site. The highest rates of litter removal were measured during the hot and dry summer when isopod activity peaks and microbial activity is minimal. We also found substantial enrichment of inorganic nitrogen and phosphorous near isopod burrows. We conclude that burrowing macro-detritivores are important regulators of litter cycling in this arid ecosystem, providing a plausible general mechanism that explains the unexpectedly high rates of plant litter cycling in deserts.

Carbon dynamics in canopy and roots influence whole tree carbon fluxes, but little is known about canopy regulation of tree-root activity. Here, we assess the patterns and dynamics of canopy-root carbon coupling in tropical trees. We used large aeroponics to directly study the root systems of Ceiba pentandra and Khaya anthotheca saplings at different light intensities. In Ceiba, root respiration (R ) co-varied with photosynthesis (A ) in large saplings (3-to-7-m canopy-root axis) at high-light, but showed no consistent pattern at low-light. At medium-light and in small saplings (c. 1-m axis), R tended to decrease transiently towards midday. Proximal roots had higher R and non-structural carbohydrate concentrations than distal roots, but canopy-root coupling was unaffected by root location. In medium-sized Khaya, no R pattern was observed, and in both species, R was unrelated to temperature. The early-afternoon increase in R suggests canopy-root coupling is based on mass flow of newly fixed carbon in the phloem, while the early-morning rise in R with A indicates an additional coupling signal that travels faster than the phloem sap. In large saplings and potentially also in higher trees, light and possibly additional environmental factors control the diurnal patterns of canopy-root coupling, irrespective of root location. This article is protected by copyright. All rights reserved.

Abstract Drought-related tree mortality had become a widespread phenomenon in forests around the globe. This process leading to these events and its complexity is not fully understood. Trees in the dry timberline are exposed to ongoing drought, and the available water for transpiration in the soil can determine their survival chances. Recent drought years led to 5%–10% mortality in the semi-arid pine forest of Yatir (Israel). The distribution of dead trees was, however, highly heterogeneous with parts of the forest showing >80% dead trees (D plots) and others with mostly live trees (L plots). At the tree level, visible stress was associated with low pre-dawn leaf water potential at the dry season (−2.8 MPa vs. −2.3 MPa in non-stressed trees), shorter needles (5.5 vs. 7.7 mm) and lower chlorophyll content (0.6 vs. 1 mg/g dw). Trends in tree-ring widths reflected differences in stress intensity (30% narrower rings in stressed compared with unstressed trees), which could be identified 15–20 years prior to mortality. At the plot scale, no differences in topography, soil type, tree age or stand density could explain the mortality difference between the D and L plots. It could only be explained by the higher surface rock cover and in stoniness across the soil profile in the L plots. Simple bucket model simulations using the site’s long-term hydrological data supported the idea that these differences could result in higher soil water concentration (m3/m3) in the L plots and extend the time above wilting point by several months across the long dry season. Accounting for subsurface heterogeneity may therefore critical to assessing stand-level response to drought and projecting tree survival, and can be used in management strategies in regions undergoing drying climate trends. A plain language summary is available for this article.

This study investigated the impact of canopy cover and seasonality on litter decay in Mediterranean pine forests to enhance climate predictions.

Abstract Decomposition of organic matter in semi-arid ecosystems is a key component of the terrestrial carbon (C) cycle. The well-known inaccuracies in predicting litter decay in water-limited regions were lessened by considering solar radiation as an abiotic decay driver of photodegradation. Moreover, exposure to high solar irradiance in dry periods often led to massive facilitation of litter decay in subsequent wet periods (“photoacceleration”), though in many studies this effect was absent. Recently, water vapour and dew were identified as modulators enabling substantial microbial degradation during rainless periods. Here, we investigated, (1) whether the activity of micro-organisms modifies litter traits, such as litter quality and microbial community in dry periods, consequently altering the loss of litter mass and nitrogen (N) in wet periods, and (2) whether it can co-occur with photoacceleration. By successively introducing litter to the field at the beginning and the end of the dry season, we found that microbial activity during the dry season affected litter mass and N loss during the wet season. Low microbial activity in the dry season led to inhibition of mass loss in the wet season, while high microbial activity led to facilitation of mass loss. Microbial activity during the dry season also caused strong inhibition of N loss from litter during the wet season, likely by enhancing the dry-season N loss. A microclimate manipulation experiment using radiation filters showed that microbial activity and exposure to solar radiation jointly modified the litter during the dry season and affected subsequent decay in the wet season. Knowledge of biotic and abiotic modifications of litter during dry periods and their implication for wet periods enhances our understanding of litter decay in semi-arid regions. Furthermore, it can improve biogeochemical model predictions of C and N cycling in drylands and in the many regions that are projected to experience a drier climate during the coming decades. A plain language summary is available for this article.

Plant litter decomposition in drylands is not well understood, and even less is known about decay of the abundant standing dead residues. Here, we followed decomposition of standing and surface litter, and assessed the underlying drivers and mechanisms.

Abstract The arid and semi-arid drylands of the world are increasingly recognized for their role in the terrestrial net carbon dioxide (CO2) uptake, which depends largely on plant litter decomposition and the subsequent release of CO2 back to the atmosphere. Observed decomposition rates in drylands are higher than predictions by biogeochemical models, which are traditionally based on microbial (biotic) degradation enabled by precipitation as the main mechanism of litter decomposition. Consequently, recent research in drylands has focused on abiotic mechanisms, mainly photochemical and thermal degradation, but they only partly explain litter decomposition under dry conditions, suggesting the operation of an additional mechanism. Here we show that in the absence of precipitation, absorption of dew and water vapor by litter in the field enables microbial degradation at night. By experimentally manipulating solar irradiance and nighttime air humidity, we estimated that most of the litter CO2 efflux and decay occurring in the dry season was due to nighttime microbial degradation, with considerable additional contributions from photochemical and thermal degradation during the daytime. In a complementary study, at three sites across the Mediterranean Basin, litter CO2 efflux was largely explained by litter moisture driving microbial degradation and ultraviolet radiation driving photodegradation. We further observed mutual enhancement of microbial activity and photodegradation at a daily scale. Identifying the interplay of decay mechanisms enhances our understanding of carbon turnover in drylands, which should improve the predictions of the long-term trend of global carbon sequestration.

Agricultural land use imposes a major disturbance on ecosystems worldwide, thus greatly modifying the taxonomic and functional composition of plant communities. However, mechanisms of community assembly, as assessed by plant functional traits, are not well known for dryland ecosystems under agricultural disturbance. Here we investigated trait responses to disturbance intensity and availability of resources to identify the main drivers of changes in composition of semiarid communities under diverging land use intensities. The eastern Mediterranean study region is characterized by an extended rainless season and by very diverse, mostly annual communities. At 24 truly replicated sites, we recorded the frequency of 241 species and the functional traits of the 53 most common species, together with soil resources and disturbance intensity across a land use gradient ranging from ungrazed shrubland to intensively managed cropland (six land use types). Multivariate RLQ analysis (linking functional traits, sites and environmental factors in a three-way ordination) and fourth corner analysis (revealing significant relations between traits and environmental factors) were used in a complementary way to get insights into trait-environment relations. Results revealed that traits related to plant size (reflecting light absorption and competitive ability) increased with resource availability, such as soil phosphorus and water holding capacity. Leaf economic traits, such as specific leaf area (SLA), leaf nitrogen content (LNC), and leaf dry matter content showed low variation across the disturbance gradient and were not related to environmental variables. In these herbaceous annual communities where plants grow and persist for just 3–5 months, SLA and LNC were unrelated, which together with relatively high SLA values might point to strategies of drought escape and grazing avoidance. Seed mass was high both at higher and lower resource availability, whereas seed number increased with the degree of disturbance. The strong response of size and reproduction traits, and the missing response of leaf economic traits reveal light interception and resource competition rather than resource acquisition and litter decomposition as drivers of plant community composition. Deviations from trait relationships observed in commonly studied temperate ecosystems confirm that climatic conditions play a fundamental role by filtering species with particular life forms and ecological strategies.