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Process-based vegetation models attempt to represent the wide range of trait variation in biomes by grouping ecologically similar species into plant functional types (PFTs). This approach has been successful in representing many aspects of plant physiology and biophysics but struggles to capture biogeographic history and ecological dynamics that determine biome boundaries and plant distributions. Grass-dominated ecosystems are broadly distributed across all vegetated continents and harbour large functional diversity, yet most Land Surface Models (LSMs) summarise grasses into two generic PFTs based primarily on differences between temperate C3 grasses and (sub)tropical C4 grasses. Incorporation of species-level trait variation is an active area of research to enhance the ecological realism of PFTs, which form the basis for vegetation processes and dynamics in LSMs. Using reported measurements, we developed grass functional trait values (physiological, structural, biochemical, anatomical, phenological, and disturbance-related) of dominant lineages to improve LSM representations. Our method is fundamentally different from previous efforts, as it uses phylogenetic relatedness to create lineage-based functional types (LFTs), situated between species-level trait data and PFT-level abstractions, thus providing a realistic representation of functional diversity and opening the door to the development of new vegetation models.

Many savannas are experiencing increased cover of trees and shrubs, resulting in reduced herbaceous productivity, shifts in savanna functional structure and potential reductions in ecotourism. Clearing woody plants has been suggested as an effective management strategy to mitigate these effects and restore these systems to an open state with higher rates of grass production and herbivory. This study investigated the effectiveness of repeated shrub clearing as a tool to mitigate bush encroachment in a semi‐arid savanna in southern Africa.We present data from a 7‐year experiment in the Mthimkhulu Game Reserve bordering Kruger National Park, South Africa. Colophospermum mopane stems and resprouting shoots were basally cut 2–3 times per year (2015–2022) in three pairs of treatment and control plots of 60 × 60 m. We monitored changes in soil moisture, grass biomass and herbivore activity via dung counts. We assessed C. mopane physiological responses to repeated cutting using non‐structural carbohydrates and stable water isotopes to infer changes to energy storage and functional rooting depth, respectively.The cleared treatment had higher soil moisture and grass biomass than the control treatment. Dung counts showed impala and buffalo visited the cleared treatment more frequently than the control treatment.Repeated cutting had limited effects on C. mopane survival in the first 2–3 years after initial clearing, but 80% of individuals were dead after 7 years. Repeatedly cut C. mopane had lower belowground starch concentrations and used water from shallower soil depths than C. mopane in control plots.Synthesis and applications. Repeated cutting increased soil moisture availability and grass biomass, and attracted charismatic grazing herbivores. While more costly than once‐off clearing methods, this practice created more employment opportunities for a neighbouring rural community. Transforming portions of the ecosystem to a grass‐dominated state may increase ecotourism potential through improved game viewing in open systems. [ABSTRACT FROM AUTHOR]

Regime shifts are difficult-to-reverse transitions that occur when an ecosystem reorganizes around a new set of self-reinforcing feedbacks. Regime shifts are predicted to occur when the intensity of some exogenous driver variable, such as temperature, annual harvest rate, or nutrient addition rate, gradually approaches and crosses a threshold value, initiating a transition to an alternative state. However, many driver variables now change rapidly as presses or pulses, not gradually, requiring new conceptual frameworks for understanding and predicting regime shifts. We argue that identifying and controlling regime shifts in response to presses and pulses will require a greater focus on the duration, not just the intensity, of changes in driver variables. In ecosystems with slower dynamics, transitions to an alternative state can take years to decades and as a result, a driver press with an intensity capable of resulting in a regime shift over long time spans may fail to cause a regime shift when applied for shorter durations. We illustrate these ideas using simulations of local-scale alternative stable state models and preliminary evidence from long-term grazing and eutrophication experiments. The simulations also suggest that small changes in the duration of driver presses or pulses can determine whether an ecosystem recovers to its original state. These insights may extend to larger scales. In spatially extended simulations that included patchiness, spatial heterogeneity, and spatial connectivity, all patches recovered to their original state after shorter presses. However, once press duration exceeded a threshold, growing proportions of the landscape shifted to an alternative state as press duration increased. We observed similar patchy transitions in a catchment-scale experiment that reinstated frequent fires approximately halfway through a regime shift from grassland to shrubland, initiated by fire suppression. In both the local- and larger-scale models, the threshold duration needed to elicit regime shifts decreased as press intensity increased or when factors counteracting regime shifts weakened. These multiple lines of evidence suggest that conceptualizing regime shifts as an interactive function of the intensity and duration of driver changes will increase understanding of the varying effects of driver presses, pulses, and cycles on ecosystem dynamics.

State transitions are changes in ecosystem structure and self-reinforcing feedbacks that are initiated when an exogenous driver variable crosses a threshold. Reversing state transitions is difficult and costly. While some state transitions are relatively rapid, many take years to decades. Outside of theoretical models, very little is known about slower state transitions and how they unfold in time and space. We quantified changes in spatial variance as a mesic grassland ecosystem shifts to a shrub-dominated state, using long-term experiments and simulations that maintain grasslands with annual fires or initiate a state transition to shrub dominance by decreasing fire frequency. In the experiments, the susceptibility to state transitions varied substantially in space. In the less frequent fire treatment, some plots became shrub-dominated around year 20 and grass extirpations began in year 25, but a third of the plots were still grass-dominated in year 37. Variable rates of state transition resulted in increasing spatial variance of grass cover over time, whereas shrub cover variance decreased. In the annually burned treatment, grasses remained dominant and the spatial variance of grass cover declined. In a separate experiment, less frequent fires were maintained for 23 years and then switched to annual fires. The switch to annual fires occurred shortly after grass variance started to increase and a majority of these plots quickly returned to a grass dominated state. In simulations, spatial variance remained low and average grass cover was high under frequent fires. If fire frequency decreased below a threshold, the ecosystem transitioned to shrubland, with a transient increase in the spatial variance of grass cover during the transition between states. Synthesis. Spatial variability in the rate and susceptibility to state transitions is indicative of a system with a patchy spatial structure, high spatial heterogeneity and low connectivity between patches. Increases in spatial variance can serve as an indication that some patches have begun a state transition and that management interventions are needed to avoid widespread transitions. This is one of the first empirical examples where altering management after an increase in spatial variance prevented state transitions.

Water stress associated with drought-like conditions is a major factor limiting plant growth and impacts productivity of natural plant communities and agricultural crops. Molecular responses of plants to water stress have been studied most extensively in model species and crops, few of which have evolved natural drought tolerance. In the current study, we examined physiological and transcriptomic responses at multiple timepoints during increasing water stress and following initial recovery from stress in a drought-tolerant C3 species, Festuca ovina. Results demonstrated non-linear transcriptomic changes during increasing stress, but largely linear declines in physiological measurements during this same period. Transcription factors represented approximately 12.7% of all differentially expressed genes. In total, 117 F. ovina homologs of previously identified and molecularly characterized drought-responsive plant genes were identified. This information will be valuable for further investigations of the molecular mechanisms involved in drought tolerance in [C.sub.3] plants. Key words: Festuca ovina, RNA-seq, water stress, transcription factor, drought tolerant. Le stress hydrique associe a des conditions de secheresse est un facteur important limitant la croissance des plantes et affectant la productivite au sein de communautes vegetales naturelles et chez les especes cultivees. Les reponses moleculaires des plantes au stress hydrique ont ete etudiees le plus chez des especes modeles et des especes cultivees, dont certaines ont acquis une tolerance naturelle a la secheresse. Dans le present travail, les auteurs examinent les reponses physiologiques et transcriptomiques a differents stades temporels suite a l'imposition d'un stress hydrique croissant et apres retablissement post-stress chez une espece C3 tolerante a la secheresse, Festuca ovina. Les resultats ont montre une evolution non-lineaire des reponses transcriptomiques en reponse a l'accroissement du stress, alors que les declins physiologiques etaient largement lineaires au cours des memes periodes. Les facteurs de transcription constituaient environ 12,7 % des genes montrant une expression differentielle. Chez F. ovina, les auteurs ont identifie 117 homologues de genes precedemment connus au plan moleculaire pour reagir a la secheresse. Cette information sera utile pour des futurs travaux des mecanismes moleculaires impliques dans la tolerance a la secheresse chez les plantes [C.sub.3]. [Traduit par la Redaction] Mots-cles : Festuca ovina, RNA-seq, stress hydrique, facteur de transcription, tolerance a la secheresse., Introduction Drought is a major environmental stress for plants and negatively affects growth and reproduction. Drought events cause yield loss in crop systems (Brevedan and Egli 2003; van Asten et [...]

Whole-plant hydraulics provide important information about responses to water limitation and can be used to understand how plant communities may change in a drier climate when measured on multiple species. Here, we measured above- and belowground hydraulic traits in Cornus drummondii, an encroaching shrub within North American tallgrass prairies, and Andropogon gerardii, a dominant C4 grass, to assess the potential hydraulic responses to future drought as this region undergoes woody expansion. Shelters that reduced precipitation by 50% and 0% were built over shrubs and grasses growing in sites that are burned at 1-year and 4-year frequencies. We then measured aboveground (Kshoot), belowground (Kroot), and whole-plant maximum hydraulic conductance (Kplant) in C. drummondii and Kroot in A. gerardii. We also measured vulnerability to embolism (P50) in C. drummondii stems. Overall, we show that: (1) A. gerardii had substantially greater Kroot than C. drummondii; (2) belowground hydraulic functioning was linked with aboveground processes; (3) above- and belowground C. drummondii hydraulics were not negatively impacted by the rainfall reductions imposed here. These results suggest that a multi-year drought will not ameliorate rates of woody expansion and highlight key differences in aboveground and belowground hydraulics for dominant species within the same ecosystem. [ABSTRACT FROM AUTHOR]

Hyperspectral remote sensing has the potential to map numerous attributes of the Earth's surface, including spatial patterns of biological diversity. Grasslands are one of the largest biomes on Earth. Accurate mapping of grassland biodiversity relies on spectral discrimination of endmembers of species or plant functional types. We focused on spectral separation of grass lineages that dominate global grassy biomes: Andropogoneae (C4), Chloridoideae (C4), and Pooideae (C3). We examined leaf reflectance spectra (350–2,500 nm) from 43 grass species representing these grass lineages from four representative grassland sites in the Great Plains region of North America. We assessed the utility of leaf reflectance data for classification of grass species into three major lineages and by collection site. Classifications had very high accuracy (94%) that were robust to site differences in species and environment. We also show an information loss using multispectral sensors, that is, classification accuracy of grass lineages using spectral bands provided by current multispectral satellites is much lower (accuracy of 85.2% and 61.3% using Sentinel 2 and Landsat 8 bands, respectively). Our results suggest that hyperspectral data have an exciting potential for mapping grass functional types as informed by phylogeny. Leaf‐level hyperspectral separability of grass lineages is consistent with the potential increase in biodiversity and functional information content from the next generation of satellite‐based spectrometers. Plain Language Summary: Understanding and identifying changes in plant diversity along broad environmental gradients requires scalable and reliable data. Spectroscopy has been shown to provide data across scales with the ability to measure plant reflectance at various extents (e.g., leaf, plot, and landscapes) with high spectral resolution and broad coverage of the electromagnetic spectrum (350–2,500 nm). In grasses, evolutionary lineage captures major axes in plant biodiversity and functional variation. We show that identifying grass evolutionary lineages from spectroscopy is possible based on common characteristics in their leaf‐level spectra. We classified 43 grass species from four sites in North America into their respective evolutionary lineages with very high accuracy (>90%) based on similarities in their leaf spectra. Classifying grass species into lineages using coarser spectral resolution data, similar to existing multispectral satellites, Sentinel 2 and Landsat 8, resulted in lower accuracy due to a loss of information from decreasing the spectral resolution. Grass lineages likely have similar spectra because of common leaf traits that evolved under similar ecological contexts. The importance of these distinctions found in the spectral reflectance of dominant grass lineages, should help our efforts in mapping and understanding grassland ecosystem function and patterns of biodiversity. Key Points: Globally dominant grass lineages are identifiable using hyperspectral leaf signaturesKey wavelengths for separating grass lineages were visible, red‐edge and shortwave infrared regions, rarely measured by multispectral sensorsHyperspectral sensors have the potential to improve remote sensing identification of grass functional types over multispectral sensors [ABSTRACT FROM AUTHOR]

Summary: Evolutionary history plays a key role driving patterns of trait variation across plant species. For scaling and modeling purposes, grass species are typically organized into C3 vs C4 plant functional types (PFTs). Plant functional type groupings may obscure important functional differences among species. Rather, grouping grasses by evolutionary lineage may better represent grass functional diversity.We measured 11 structural and physiological traits in situ from 75 grass species within the North American tallgrass prairie. We tested whether traits differed significantly among photosynthetic pathways or lineages (tribe) in annual and perennial grass species.Critically, we found evidence that grass traits varied among lineages, including independent origins of C4 photosynthesis. Using a rigorous model selection approach, tribe was included in the top models for five of nine traits for perennial species. Tribes were separable in a multivariate and phylogenetically controlled analysis of traits, owing to coordination of important structural and ecophysiological characteristics.Our findings suggest grouping grass species by photosynthetic pathway overlooks variation in several functional traits, particularly for C4 species. These results indicate that further assessment of lineage‐based differences at other sites and across other grass species distributions may improve representation of C4 species in trait comparison analyses and modeling investigations. [ABSTRACT FROM AUTHOR]

Summary: Models of tree–grass coexistence in savannas make different assumptions about the relative performance of trees and grasses under wet vs dry conditions. We quantified transpiration and drought tolerance traits in 26 tree and 19 grass species from the African savanna biome across a gradient of soil water potentials to test for a trade‐off between water use under wet conditions and drought tolerance.We measured whole‐plant hourly transpiration in a growth chamber and quantified drought tolerance using leaf osmotic potential (Ψosm). We also quantified whole‐plant water‐use efficiency (WUE) and relative growth rate (RGR) under well‐watered conditions.Grasses transpired twice as much as trees on a leaf‐mass basis across all soil water potentials. Grasses also had a lower Ψosm than trees, indicating higher drought tolerance in the former. Higher grass transpiration and WUE combined to largely explain the threefold RGR advantage in grasses.Our results suggest that grasses outperform trees under a wide range of conditions, and that there is no evidence for a trade‐off in water‐use patterns in wet vs dry soils. This work will help inform mechanistic models of water use in savanna ecosystems, providing much‐needed whole‐plant parameter estimates for African species. [ABSTRACT FROM AUTHOR]

Riparian zones and the streams they border provide vital habitat for organisms, water quality protection, and other important ecosystem services. These areas are under pressure from local (land use/land cover change) to global (climate change) processes. Woody vegetation is expanding in grassland riparian zones worldwide. Here we report on a decade‐long watershed‐scale mechanical removal of woody riparian vegetation along 4.5 km of stream channel in a before–after control impact experiment. Prior to this removal, woody plants had expanded into grassy riparian areas, associated with a decline in streamflow, loss of grassy plant species, and other ecosystem‐scale impacts. We confirmed some expected responses, including rapid increases in stream nutrients and sediments, disappearance of stream mosses, and decreased organic inputs to streams via riparian leaves. We were surprised that nutrient and sediment increases were transient for 3 years, that there was no recovery of stream discharge, and that areas with woody removal did not shift back to a grassland state, even when reseeded with grassland species. Rapid expansion of shrubs (Cornus drummondii, Prunus americana) in the areas where trees were removed allowed woody vegetation to remain dominant despite repeating the cutting every 2 years. Our results suggest woody expansion can fundamentally alter terrestrial and aquatic habitat connections in grasslands, resulting in inexorable movement toward a new ecosystem state. Human pressures, such as climate change, atmospheric CO2 increases, and elevated atmospheric nitrogen deposition, could continue to push the ecosystem along a trajectory that is difficult to change. Our results suggest that predicting relationships between riparian zones and the streams they border could be difficult in the face of global change in all biomes, even in well‐studied sites. [ABSTRACT FROM AUTHOR]

Summary: Savannas cover a significant fraction of the Earth's land surface. In these ecosystems, C3 trees and C4 grasses coexist persistently, but the mechanisms explaining coexistence remain subject to debate. Different quantitative models have been proposed to explain coexistence, but these models make widely contrasting assumptions about which mechanisms are responsible for savanna persistence. Here, we show that no single existing model fully captures all key elements required to explain tree–grass coexistence across savanna rainfall gradients, but many models make important contributions. We show that recent empirical work allows us to combine many existing elements with new ideas to arrive at a synthesis that combines elements of two dominant frameworks: Walter's two‐layer model and demographic bottlenecks. We propose that functional rooting separation is necessary for coexistence and is the crux of the coexistence problem. It is both well‐supported empirically and necessary for tree persistence, given the comprehensive grass superiority for soil moisture acquisition. We argue that eventual tree dominance through shading is precluded by ecohydrological constraints in dry savannas and by fire and herbivores in wet savannas. Strong asymmetric grass–tree competition for soil moisture limits tree growth, exposing trees to persistent demographic bottlenecks. [ABSTRACT FROM AUTHOR]

A frequently expressed viewpoint across the Earth science community is that global soil moisture estimates from satellite L‐band (1.4 GHz) measurements represent moisture only in a shallow surface layer (0–5 cm) and consequently are of limited value for studying global terrestrial ecosystems because plants use water from deeper rootzones. Using this argumentation, many observation‐based land surface studies avoid satellite‐observed soil moisture. Here, based on peer‐reviewed literature across several fields, we argue that such a viewpoint is overly limiting for two reasons. First, microwave soil emission depth considerations and statistical considerations of vertically correlated soil moisture information together indicate that L‐band measurements carry information about soil moisture extending below the commonly referenced 5 cm in many conditions. However, spatial variations of effective depths of representation remain uncertain. Second, in reviewing isotopic tracer field studies of plant water uptake, we find a prevalence of vegetation that primarily draws moisture from these upper soil layers. This is especially true for grasslands and croplands covering more than a third of global vegetated surfaces. Even some deeper‐rooted species (i.e., shrubs and trees) preferentially or seasonally draw water from the upper soil layers. Therefore, L‐band satellite soil moisture estimates are more relevant to global vegetation water uptake than commonly appreciated (i.e., relevant beyond only shallow soil processes like soil evaporation). Our commentary encourages the application of satellite soil moisture across a broader range of terrestrial hydrosphere and biosphere studies while urging more rigorous estimates of its effective depth of representation. Key Points: L‐band satellite soil moisture effective depth of representation is often deeper than the commonly referenced 5 cm limitIsotopic tracer studies reveal common preferential plant water use of moisture in these upper soil layers either primarily or seasonallyThe optimal soil moisture product changes in time and space, with satellite soil moisture often being optimal for many vegetated surfaces [ABSTRACT FROM AUTHOR]

We use the Multiple Element Limitation (MEL) model to examine responses of 12 ecosystems to elevated carbon dioxide (CO2), warming, and 20% decreases or increases in precipitation. Ecosystems respond synergistically to elevated CO2, warming, and decreased precipitation combined because higher water‐use efficiency with elevated CO2 and higher fertility with warming compensate for responses to drought. Response to elevated CO2, warming, and increased precipitation combined is additive. We analyze changes in ecosystem carbon (C) based on four nitrogen (N) and four phosphorus (P) attribution factors: (1) changes in total ecosystem N and P, (2) changes in N and P distribution between vegetation and soil, (3) changes in vegetation C:N and C:P ratios, and (4) changes in soil C:N and C:P ratios. In the combined CO2 and climate change simulations, all ecosystems gain C. The contributions of these four attribution factors to changes in ecosystem C storage varies among ecosystems because of differences in the initial distributions of N and P between vegetation and soil and the openness of the ecosystem N and P cycles. The net transfer of N and P from soil to vegetation dominates the C response of forests. For tundra and grasslands, the C gain is also associated with increased soil C:N and C:P. In ecosystems with symbiotic N fixation, C gains resulted from N accumulation. Because of differences in N versus P cycle openness and the distribution of organic matter between vegetation and soil, changes in the N and P attribution factors do not always parallel one another. Differences among ecosystems in C‐nutrient interactions and the amount of woody biomass interact to shape ecosystem C sequestration under simulated global change. We suggest that future studies quantify the openness of the N and P cycles and changes in the distribution of C, N, and P among ecosystem components, which currently limit understanding of nutrient effects on C sequestration and responses to elevated CO2 and climate change. [ABSTRACT FROM AUTHOR]