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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]

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]

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Understanding spatial and temporal variation in plant traits is needed to accurately predict how communities and ecosystems will respond to global change. The National Ecological Observatory Network's (NEON's) Airborne Observation Platform (AOP) provides hyperspectral images and associated data products at numerous field sites at 1 m spatial resolution, potentially allowing high‐resolution trait mapping. We tested the accuracy of readily available data products of NEON's AOP, such as Leaf Area Index (LAI), Total Biomass, Ecosystem Structure (Canopy height model [CHM]), and Canopy Nitrogen, by comparing them to spatially extensive field measurements from a mesic tallgrass prairie. Correlations with AOP data products exhibited generally weak or no relationships with corresponding field measurements. The strongest relationships were between AOP LAI and ground‐measured LAI (r = 0.32) and AOP Total Biomass and ground‐measured biomass (r = 0.23). We also examined how well the full reflectance spectra (380–2,500 nm), as opposed to derived products, could predict vegetation traits using partial least‐squares regression (PLSR) models. Among all the eight traits examined, only Nitrogen had a validation R2of more than 0.25. For all vegetation traits, validation R2 ranged from 0.08 to 0.29 and the range of the root mean square error of prediction (RMSEP) was 14–64%. Our results suggest that currently available AOP‐derived data products should not be used without extensive ground‐based validation. Relationships using the full reflectance spectra may be more promising, although careful consideration of field and AOP data mismatches in space and/or time, biases in field‐based measurements or AOP algorithms, and model uncertainty are needed. Finally, grassland sites may be especially challenging for airborne spectroscopy because of their high species diversity within a small area, mixed functional types of plant communities, and heterogeneous mosaics of disturbance and resource availability. Remote sensing observations are one of the most promising approaches to understanding ecological patterns across space and time. But the opportunity to engage a diverse community of NEON data users will depend on establishing rigorous links with in‐situ field measurements across a diversity of sites. [ABSTRACT FROM AUTHOR]

Roots are key components of terrestrial ecosystems, yet little is known about how root structure and function vary across a broad range of species, functional groups and ecological gradients in situ.We assessed how woody and grass root anatomical traits vary among soil depths and different fire frequencies to better understand the water‐use strategies exhibited by these two functional groups in tallgrass prairie experiencing woody encroachment. Specifically, we asked: (a) Do root anatomical traits differ with fire frequency or soil depth?; (b) Do relationships between anatomical traits that confer hydraulic safety versus efficiency vary by fire frequency or soil depth?; (c) Is root anatomy associated with integrative root traits (e.g. root diameter, specific root length (SRL) and root biomass)?; and (d) When scaled by root biomass, do root water‐use traits impact the capacity for water uptake?We collected grass and woody roots from 10, 30 and 50 cm deep soil in areas burned every 1, 4 and 20 years. We then measured xylem conduit diameter, conduit cell wall thickness, conduit number, conduit mechanical safety (t/b), stele area, endoderm thickness, hydraulic diameter, theoretical hydraulic conductivity and root‐system theoretical hydraulic conductance.We observed: (a) Woody roots had high hydraulic conductance in shallow soils and greater mechanical strength in deeper soils, which may provide a competitive advantage in less frequently burned, more diverse plant communities; (b) shallow grass roots had unique trait combinations at the anatomical and root‐system levels (thinner, more numerous conduits and higher root‐system hydraulic conductance compared to deeper roots) that likely allow these plants to rapidly use water but tolerate dry soils under multiple fire regimes; and (c) hydraulic safety versus efficiency trade‐offs translate between different hierarchical scales (i.e. from anatomical to integrative root traits).These results provide anatomical evidence to explain water‐use dynamics in tallgrass prairie and also provide novel insight regarding functional strategies that may facilitate the conversion from grassland to shrubland in less frequently burned tallgrass prairie. Future work should investigate these dynamics in situ, as they may explain current and future patterns of woody‐grass coexistence in tallgrass prairies. A free Plain Language Summary can be found within the Supporting Information of this article. [ABSTRACT FROM AUTHOR]

Understanding the complex and unpredictable ways ecosystems are changing and predicting the state of ecosystems and the services they will provide in the future requires coordinated, long‐term research. This paper is a product of a U.S. National Science Foundation funded Long Term Ecological Research (LTER) network synthesis effort that addressed anticipated changes in future populations and communities. Each LTER site described what their site would look like in 50 or 100 yr based on long‐term patterns and responses to global change drivers in each ecosystem. Common themes emerged and predictions were grouped into state change, connectivity, resilience, time lags, and cascading effects. Here, we report on the "state change" theme, which includes examples from the Georgia Coastal (coastal marsh), Konza Prairie (mesic grassland), Luquillo (tropical forest), Sevilleta (arid grassland), and Virginia Coastal (coastal grassland) sites. Ecological thresholds (the point at which small changes in an environmental driver can produce an abrupt and persistent state change in an ecosystem quality, property, or phenomenon) were most commonly predicted. For example, in coastal ecosystems, sea‐level rise and climate change could convert salt marsh to mangroves and coastal barrier dunes to shrub thicket. Reduced fire frequency has converted grassland to shrubland in mesic prairie, whereas overgrazing combined with drought drive shrub encroachment in arid grasslands. Lastly, tropical cloud forests are susceptible to climate‐induced changes in cloud base altitude leading to shifts in species distributions. Overall, these examples reveal that state change is a likely outcome of global environmental change across a diverse range of ecosystems and highlight the need for long‐term studies to sort out the causes and consequences of state change. The diversity of sites within the LTER network facilitates the emergence of overarching concepts about state changes as an important driver of ecosystem structure, function, services, and futures. [ABSTRACT FROM AUTHOR]

Hysteresis is a fundamental characteristic of alternative stable state theory, yet evidence of hysteresis is rare. In mesic grasslands, fire frequency regulates transition from grass‐ to shrub‐dominated system states. It is uncertain, however, if increasing fire frequency can reverse shrub expansion, or if grass‐shrub dynamics exhibit hysteresis. We implemented annual burning in two infrequently burned grasslands and ceased burning in two grasslands burned annually. With annual fires, grassland composition converged on that of long‐term annually burned vegetation due to rapid recovery of grass cover, although shrubs persisted. When annual burning ceased, shrub cover increased, but community composition did not converge with a long‐term infrequently burned reference site because of stochastic and lagged dispersal by shrubs, reflecting hysteresis. Our results demonstrated that annual burning can slow, but not reverse, shrub encroachment. In addition, reversing fire frequencies resulted in hysteresis because vegetation trajectories from grassland to shrubland differed from those of shrubland to grassland. [ABSTRACT FROM AUTHOR]

Grassland ecosystems globally are being negatively impacted by changes in climate, disturbance regimes, nutrient flux, and consumer guilds. Changes in the trophic ecology of consumers can substantially influence local resources, contributing to shifting diversity, community turnover, and other processes of ecosystem change. Small mammals are diverse and abundant within grasslands and yet the impact of changing ecosystems on small mammals and the role of these mammals as consumers are still both under‐studied. We assessed small mammal resource use within grassland and woodland vegetation types that have resulted from landscape‐scale experimental disturbance through fire treatments within the tallgrass prairie ecoregion of the North American Great Plains. We predicted that resource use would vary significantly among grassland vs. woodland communities, in turn reducing the role of small mammals in contributing to future maintenance of native prairies. We sampled five dominant species of rodents across three years and multiple habitats. Using stable isotope analysis, we investigated isotopic niche area and overlap to infer variation in diet, both within and among species. Resource use shifted in bivariate isotopic space seasonally but not across years when combining all species and habitats. Inferred spring diet (based on fur samples) was highly diverse and overlapping. Summer isotopic values (based on liver tissue) in woody habitat treatments were narrower and overlapped less than within grassland habitats. Consumers generally shifted from C4 herbivory to C3 herbivory, or greater omnivory, when analyzing grassland, shrubland, and woodland habitats respectively. Within the tallgrass prairie ecosystem, small mammal populations in herbaceous‐dominated habitats use a broader variety of resources than small mammals in proximate woody‐dominated habitats. As native grasslands experience woody encroachment, small mammal assemblages experience turnover of dominant species and associated changes in diet. Ecosystem changes such as cessation of frequent fire resulting in more woody habitats may include reduced roles by native small mammals as consumers/dispersers/propagators of native grassland plants. [ABSTRACT FROM AUTHOR]

In savannas, partitioning of below‐ground resources by depth could facilitate tree–grass coexistence and shape vegetation responses to changing rainfall patterns. However, most studies assessing tree versus grass root‐niche partitioning have focused on one or two sites, limiting generalization about how rainfall and soil conditions influence the degree of rooting overlap across environmental gradients.We used two complementary stable isotope techniques to quantify variation (a) in water uptake depths and (b) in fine‐root biomass distributions among dominant trees and grasses at eight semi‐arid savanna sites in Kruger National Park, South Africa. Sites were located on contrasting soil textures (clayey basaltic soils vs. sandy granitic soils) and paired along a gradient of mean annual rainfall.Soil texture predicted variation in mean water uptake depths and fine‐root allocation. While grasses maintained roots close to the surface and consistently used shallow water, trees on sandy soils distributed roots more evenly across soil depths and used deeper soil water, resulting in greater divergence between tree and grass rooting on sandy soils. Mean annual rainfall predicted some variation among sites in tree water uptake depth, but had a weaker influence on fine‐root allocation.Synthesis. Savanna trees overlapped more with shallow‐rooted grasses on clayey soils and were more distinct in their use of deeper soil layers on sandy soils, consistent with expected differences in infiltration and percolation. These differences, which could allow trees to escape grass competition more effectively on sandy soils, may explain observed differences in tree densities and rates of woody encroachment with soil texture. Differences in the degree of root‐niche separation could also drive heterogeneous responses of savanna vegetation to predicted shifts in the frequency and intensity of rainfall. [ABSTRACT FROM AUTHOR]

Summary: 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. [ABSTRACT FROM AUTHOR]

Frequent fire and grazing by megafauna are important determinants of tallgrass prairie plant community structure. However, fire suppression and removal of native grazers have altered these natural disturbance regimes and changed grassland plant communities with potential long‐term consequences for soil carbon (C) and nitrogen (N) storage. We investigated multidecade changes in soil C and N pools in response to contrasting long‐term burning and grazing treatments. Fire suppression with or without grazers and exclusion of grazers in annually burned prairie increased soil C content and shifted the δ13C signature of soil C over time, concomitant with changes in plant community composition. Soil δ13C values indicated that increased soil C content was associated with an increased contribution from plants using a C3 photosynthetic pathway (i.e., woody shrubs) under fire suppression. Soil N content also increased when fire was suppressed, relative to frequently burned grassland, but the rate of increase was slower when grazers were present. Additionally, changes in δ15N values suggested that grazing increased the openness of the N cycle, presumably due to greater N losses. By coupling long‐term fire and grazing treatments with plant community data and soil samples archived over three decades, we demonstrate that human‐caused changes to natural disturbance regimes in a tallgrass prairie significantly alter soil C and N cycles through belowground changes associated with shifts in the plant community. Since natural disturbance regimes have been altered in grasslands across the world, our results are relevant for understanding the long‐term biogeochemical consequences of these ongoing land use changes. Plain Language Summary: The tallgrass prairie of the central United States was historically maintained by frequent fire and bison grazing. However, human‐caused fire suppression and bison removal has altered the plant community composition with cascading effects on elemental cycling. By analyzing soil collected over a 30‐year period, we investigated the long‐term effects of contrasting fire and grazing regimes on soil carbon and nitrogen. Soil carbon content increased over time if fire was suppressed and/or bison were absent. Soil nitrogen content increased only if fire was suppressed. Plant community data and soil isotopic evidence indicated that soil carbon was highest when woody plants were contributing more carbon to the soil in the fire suppression treatments. On the other hand, soil carbon was highest in the annually burned and grazed treatment when warm‐season grasses were contributing more carbon to the soil. Since many grasslands around the world are subject to similar land use changes, our results are important for predicting the long‐term effects of those changes on the ecosystem. Additionally, while our results indicate that the encroachment of woody plants into grasslands may increase soil carbon, land managers should consider the negative trade‐offs of woody encroachment on grassland diversity and ecosystem services. Key Points: We identified the longitudinal effects of 30 years of contrasting burning and grazing regimes on soil C and N in tallgrass prairieIncreased soil C concentrations were associated with increased woody plants under fire suppression or C4 grasses under annual burningSoil N concentration increased over time when fire was suppressed; N was recycled less tightly if bison were present [ABSTRACT FROM AUTHOR]

RNA sequencing (RNA‐seq) is a widely used approach to investigate gene expression and increasingly is used in time‐course studies to characterize transcriptomic changes over time. Two primary options are available as controls in time‐course experiments: samples collected at the first sampling time are used as controls (temporal control, TC) and samples collected in parallel at each individual sampling time are used as controls (biological control, BC). While both approaches are used in experimental studies, we know of no analyses performed to date that directly compare effects of control type choices on identifying differentially expressed genes (DEGs) and subsequent functional analysis. In the current study, we compare experimental results using these different control types for time‐course RNA‐seq drought stress experiments in two wild grass species in the genus Paspalum. Our results showed BC assemblies gave a higher number of loci in both species. The number of DEGs increased with increasing stress and then decreased dramatically at the recovery time point using both control types. Expression levels of the same DEGs were highly correlated between control types in both species, ranging from r =.653 to r =.852. We also observed similar rank orders of shared enriched Gene Ontology term lists using the two different control types. Collectively, our findings suggest similar results in differential gene expression and functional annotation between control types. The ultimate choice of control type will rely on the experimental length and organism type, with labour time and sequencing costs as additional factors to be considered. [ABSTRACT FROM AUTHOR]

Predicting the hydrological consequences following changes in grassland vegetation type (i.e., woody encroachment) requires an understanding of water flux dynamics at high spatiotemporal resolution for predominant species within grassland communities. However, grassland fluxes are typically measured at the leaf or landscape scale, which inhibits our ability to predict how individual species contribute to changing ecosystem fluxes. We used external heat balance sap flow sensors and a hierarchical Bayesian state‐space modeling approach to bridge this "flux gap" and estimate continuous species‐level water flux in common tallgrass prairie species. Specifically, we asked the following: (1) How do diurnal and nocturnal water fluxes differ among woody and herbaceous plants? (2) How sensitive are woody and herbaceous species to environmental drivers of diurnal and nocturnal water flux? We highlight three results: (1) Cornus drummondii, the primary woody encroacher in this grassland, exhibited the greatest canopy‐level water loss; (2) nocturnal transpiration was a large component of the water lost in this ecosystem and was driven primarily by C4 grasses and C. drummondii; and (3) the sensitivity of canopy transpiration to environmental drivers varies among plant functional types and throughout a 24‐hr period. Our data reveal important insights regarding the water use strategies of woody versus herbaceous species in tallgrass prairies and about the potential hydrological consequences of ongoing woody encroachment. We suggest that the high, static flux rates observed in woody species will likely deplete deep water stores over time, potentially creating hydrological deficits in grasslands experiencing woody encroachment and concomitantly increasing the vulnerability of these ecosystems to drought. Plain Language Summary: Woody encroachment, the process through which woody plants (trees and shrubs) replace grasses and wildflowers in grassland ecosystems, is occurring globally. Because woody plants use water differently (e.g., rates, soil depths of uptake, and time periods of activity) than grasses, woody encroachment could potentially change the water cycle in grasslands. In order to understand how grassland water use might change with ongoing woody encroachment, we measured continuous rates of water loss in common plant species in a North American grassland. We found that the main encroaching shrub in this grassland used more water over time than grasses or wildflowers. Additionally, this shrub and grasses used a large amount of water at night compared to wildflowers. Finally, we found that different weather conditions had different impacts on how water was used by woody plants, grasses, and wildflowers. We suggest that woody encroachment might reduce the amount of water stored in grasslands, which could alter the composition of grassland plants in the future. Key Points: Cornus drummondii, the primary woody encroacher in this grassland, exhibited the greatest canopy‐level water lossNocturnal transpiration was a large component of the water lost in this ecosystem and was driven primarily by C4 grasses and C. drummondiiThe sensitivity of canopy transpiration to environmental drivers varies among plant functional types and throughout a 24‐hr period [ABSTRACT FROM AUTHOR]

Biodiversity enhances a variety of ecosystem processes, and yet the underlying mechanisms through which these relationships occur remain a critical knowledge gap. Here, we used the natural abundance of stable isotopes to measure depth of water uptake in five common grassland species (Asclepias tuberosa, Lespedeza capitata, Liatris aspera, Schizachyrium scoparium and Sorghastrum nutans) growing across an experimental grassland diversity gradient. Using this approach, we addressed the following questions: 1) does the depth‐specific provenance of water uptake differ among species and/or do interspecific differences in water source manifest with increasing community diversity? 2) Does the isotopic niche space occupied by plants change with increasing diversity? 3) Is plasticity in water uptake depth across a diversity gradient associated with functional plant responses? We found that the depth of soil water used by plants was inherently different among species when grown in monocultures. All species used less shallow soil water and more intermediate‐depth soil water in mixed assemblages than in monocultures, resulting in similar interspecific differences in water source across the diversity gradient. However, plasticity in the locations of water used were positively associated with increases in plant growth in higher diversity treatments. These results indicate that plasticity in water‐use may contribute to positive biodiversity–productivity relationships commonly observed in temperate grasslands. [ABSTRACT FROM AUTHOR]

Abstract: Nocturnal transpiration can impact water balance from the local community to earth‐atmosphere fluxes. However, the dynamics and drivers of nocturnal transpiration among coexisting plant functional groups in herbaceous ecosystems are unknown. Here, we addressed the following questions: (1) How do nocturnal (Enight) and diurnal (Eday) transpiration vary among coexisting grasses, forbs, and shrubs in a tallgrass prairie? (2) What environmental variables drive Enight and do these differ from the drivers of Eday? (3) Is Enight associated with daytime physiological processes? We measured diurnal and nocturnal leaf gas exchange on perennial grass, forb and woody species in a North American tallgrass prairie. Measurements were made periodically across two growing seasons (May–August 2014–2015) on three C4 grasses (Andropogon gerardii, Sorghastrum nutans and Panicum virgatum), two C3 forbs (Vernonia baldwinii and Solidago canadensis), one C3 sub‐shrub (Amorpha canescens) and two C3 shrubs (Cornus drummondii and Rhus glabra). By extending our study to multiple functional groups, we were able to make several key observations: (1) Enight was variable among co‐occurring plant functional groups, with the highest rates occurring in C4 grasses, (2) Enight and Eday exhibited different responses to vapour pressure deficit and other environmental drivers, and (3) rates of Enight were strongly related to predawn leaf water potential for grasses and woody species, and were likely modulated by small‐scale changes in soil moisture availability. Our results provide novel insight into an often‐overlooked portion of ecosystem water balance. Considering the high rates of Enight observed in C4 grasses, as well as the widespread global occurrence of C4 grasses, nocturnal water loss might constitute a greater proportion of global evapotranspiration than previously estimated. Additionally, future predictions of nocturnal water loss may be complicated by stomatal behaviour that differs between the day and at night. Finally, these data suggest a water‐use strategy by C4 grasses wherein the high rates of Enight occurring during wet periods may confer a competitive advantage to maximize resource consumption during periods of greater availability. A plain language summary is available for this article. [ABSTRACT FROM AUTHOR]

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. [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. [ABSTRACT FROM AUTHOR]

Recent models suggest that herbivores optimize nutrient intake by selecting patches of low to intermediate vegetation biomass. We assessed the application of this hypothesis to plains bison ( Bison bison) in an experimental grassland managed with fire by estimating daily rates of nutrient intake in relation to grass biomass and by measuring patch selection in experimental watersheds in which grass biomass was manipulated by prescribed burning. Digestible crude protein content of grass declined linearly with increasing biomass, and the mean digestible protein content relative to grass biomass was greater in burned watersheds than watersheds not burned that spring (intercept; F1,251 = 50.57, P < 0.0001). Linking these values to published functional response parameters, ad libitum protein intake, and protein expenditure parameters, Fryxell's ( Am. Nat., 1991, 138, 478) model predicted that the daily rate of protein intake should be highest when bison feed in grasslands with 400-600 kg/ha. In burned grassland sites, where bison spend most of their time, availability of grass biomass ranged between 40 and 3650 kg/ha, bison selected foraging areas of roughly 690 kg/ha, close to the value for protein intake maximization predicted by the model. The seasonal net protein intake predicted for large grazers in this study suggest feeding in burned grassland can be more beneficial for nutrient uptake relative to unburned grassland as long as grass regrowth is possible. Foraging site selection for grass patches of low to intermediate biomass help explain patterns of uniform space use reported previously for large grazers in fire-prone systems. [ABSTRACT FROM AUTHOR]

A common theme in plant physiological research is the trade-off between stress tolerance and growth; an example of this trade-off at the tissue level is the safety vs efficiency hypothesis, which suggests that plants with the greatest resistance to hydraulic failure should have low maximum hydraulic conductance., Here, we quantified the leaf-level drought tolerance of nine C4 grasses as the leaf water potential at which plants lost 50% ( P50 × RR) of maximum leaf hydraulic conductance ( Ksat), and compared this trait with other leaf-level and whole-plant functions., We found a clear trade-off between Ksat and P50 × RR when Ksat was normalized by leaf area and mass ( P = 0.05 and 0.01, respectively). However, no trade-off existed between P50 × RR and gas-exchange rates; rather, there was a positive relationship between P50 × RR and photosynthesis ( P = 0.08). P50 × RR was not correlated with species distributions based on precipitation ( P = 0.70), but was correlated with temperature during the wettest quarter of the year ( P < 0.01)., These results suggest a trade-off between safety and efficiency in the hydraulic system of grass leaves, which can be decoupled from other leaf-level functions. The unique physiology of C4 plants and adaptations to pulse-driven systems may provide mechanisms that could decouple hydraulic conductance from other plant functions. [ABSTRACT FROM AUTHOR]

Leguminous cover crops are considered part of sustainable agricultural systems. With the development of no-till cropping systems, cover crops have been recognized for their ability to provide N for succeeding crops. The objectives of this study were: (i) to determine the N contribution of summer cover crops and double-cropped grain crops following winter wheat (Triticum aestivum L.) and N rates to subsequent maize (Zea mays L.) crops' physiological traits and yield, (ii) to calculate the fertilizer N replacement value, and (iii) to perform economic analyses of the cropping systems. Field experiments were conducted during the 2012-2013 and 2013-2014 growing seasons in Kansas. The grain yield of maize in all the cover crop systems and the double- cropped soybean [Glycine max (L.) Merr.] system was similar to maize in the fallow system with 45 kg N ha-1. The mean increase in grain yield as a result of including cowpea [Vigna unguiculata (L.) Walp.], pigeonpea [Cajanus cajan (L.) Millsp.], sunn hemp (Crotalaria juncea L.), double-cropped soybean, and double-cropped grain sorghum [Sorghum bicolor (L.) Moench] in the rotation over the fallow system with 0 kg N ha-1 was 78, 91, 66, 72, and 12%, respectively. Fertilizer N replacement values for cowpea, pigeonpea, sunn hemp, double-cropped soybean, and double-cropped grain sorghum were 53, 64, 43, 47, and -5 kg N ha-1, respectively. We conclude that the inclusion of summer leguminous cover crops in a cropping system has the potential to reduce or supplement N requirements and increase the grain yield of subsequent maize crops. [ABSTRACT FROM AUTHOR]

Leguminous cover crop systems have been envisaged as a critical component of sustainable agriculture because of their potential to increase soil productivity by cycling C and N in agricultural systems. Our objectives were to: (i) determine the effects of including summer cover crops and N rates in the cropping system on the growth and yield of the succeeding grain sorghum [Sorghum bicolor (L.) Moench] crop, (ii) calculate the N fertilizer replacement value (NFRV) and evaluate economic returns, and (iii) determine the most cost-effective cropping system. Field experiments were conducted for two seasons in Kansas. Leguminous summer cover crops and double-cropped soybean [Glycine max (L.) Merr.] residues improved soil N availability; N was subsequently used by the succeeding crop. Across years and cropping systems, the mean increases in grain yield as a result of including cowpea [Vigna unguiculata (L.) Walp.], pigeonpea [Cajanus cajan (L.) Millsp.], sunn hemp (Crotalaria juncea L.), double-cropped soybean, and double-cropped grain sorghum in the rotation compared with a fallow system with 0 kg N ha-1 were 56, 62, 43, 32, and 3%, respectively, and NFRVs across the years were 53, 64, 36, 27, and -3 kg N ha-1, respectively. Across years, grain sorghum in a double-cropped soybean system and a fallow system with 90 kg N ha-1 gave profitable economic net returns. We conclude that including leguminous cover crops in a cropping system has the potential to reduce N requirements and improve the N availability and grain yield of the succeeding grain sorghum crop. [ABSTRACT FROM AUTHOR]

For many grassland and savanna ecosystems, water limitation is a key regulator of individual plant, community and ecosystem processes. Maximum rooting depth is commonly used to characterize the susceptibility of plant species to drought. This rests on the assumption that deep-rooted plant species would have a greater total volume of soil water to exploit and should be less susceptible to episodic changes in water availability., Independent of maximum rooting depth, rooting strategies based on differences in biomass allocation with depth, uptake plasticity in relation to water availability and variation in water transport capability may all influence growth responses and susceptibility to drought. Many examples from grasslands and savannas reflect these rooting strategies among coexisting grass, forb and woody species., Here, we use a dynamic model of plant water uptake and growth to show how changes in root distribution, functional plasticity and root hydraulic conductivity have the potential to influence aboveground biomass and competitive outcomes, even when maximum rooting depth remains constant. We also show theoretically that shifts in root distribution to surface soils without changes in maximum depth can potentially outweigh the benefits of increased maximum rooting depth., Combining our current reliance on biogeographic descriptions of maximum rooting depth with insights about other, more subtle aspects of root structure and function are likely to improve our understanding of ecosystem responses to dynamic water limitation. [ABSTRACT FROM AUTHOR]

It is still far from clear whether and to what extent trees and grasses partition soil moisture in tropical savannas. A major reason for this is that we don't know how snapshot data on rooting differences translate into ecologically relevant patterns of water use at seasonal scales. We used stable isotopes in soil and stem water to quantify functional rooting profiles in grasses and two tree species in a South African savanna. Concurrently, we measured tree sap-flow velocity, grass canopy temperature (a transpiration correlate), and soil moisture content at multiple depths over the course of a growing season. We used lasso regression to identify the dominant soil moisture layers driving daily variation in tree and grass water-use metrics while controlling for weather variables. We found clear rooting depth differences between grasses (shallow) and trees (deep) from the isotopic data, and these patterns were strongly supported by the water-use data, which showed that grasses and trees predominantly responded to soil moisture availability at 5 and 40 cm depth, respectively. Our results provide a rare example of mechanistic support for the resource partitioning hypothesis in savannas, with important implications for our understanding of tree-grass dynamics under altered precipitation regimes. [ABSTRACT FROM AUTHOR]

This review synthesizes evidence that altered fire frequency drives discontinuous ecosystem transitions from mesic grasslands to shrublands or woodlands in the Central Great Plains, USA., Long-term fire manipulations reveal that grassland to shrubland transitions are triggered when fire-free intervals increase from 1-3 years to ≥ 3-8 years, and longer fire returns (˜10 years or more) result in transitions to woodlands. Grazing and soil properties alter these fire thresholds., Grassland to shrubland transitions are abrupt and exhibit nonlinear relationships between driver and state variables. Transitions to shrublands and woodlands exhibit hysteresis, where reintroducing frequent fires does not reverse transitions in management-relevant time-scales (decades)., Nonlinear transitions and hysteresis emerge because grasses generate positive feedbacks with fire that create strong demographic barriers for shrub and tree establishment. Fire-free intervals allow shrubs and trees to reach a size sufficient to survive fire, reproduce and disrupt the fire feedback loop through competition., Synthesis. Mesic grasslands, shrublands and woodlands constitute self-reinforcing states (alternative attractors) separated by critical fire frequency thresholds. Even without major shifts in climate, altered fire frequency can produce dramatic state changes, highlighting the importance of fire for predicting future ecosystem states. Local management should focus on prevention of unwanted transitions rather than post hoc restoration. [ABSTRACT FROM AUTHOR]

Ecosystems with alternative attractors are susceptible to abrupt regime shifts that are often difficult to predict and reverse. In this study, we quantify multiple system dynamics to determine whether the transition of mesic grassland to shrubland, a widespread phenomenon, represents a linear reversible process, a nonlinear but reversible threshold process, or a transition between alternative attractors that is nonlinear and prone to hysteresis. Using a 28-yr data set with annual resolution and extensive spatial replication, we found that shrub cover is correlated with distinct thresholds of fire and C4 grass cover, resulting in temporal bimodality of shrub cover and abrupt shifts of shrub cover despite gradual changes in grass cover. These abrupt increases in shrub cover are the most rapid ever reported in grasslands, and illustrate internal thresholds that separate grasslands and shrublands. Nonlinear transitions from low to high shrub cover were also closely associated with positive feedback mechanisms that alter fire and competition (r² = 0.65), suggesting that grasslands and shrublands could show hysteresis, and by definition exist as alternative attractors. Thus, the response of this ecosystem to anthropogenic activity should tend to be rapid, nonlinear, and perhaps difficult to reverse. Regime shifts in this mesic grassland were predictable: we found that grassland and shrubland attractors were differentiated by critical thresholds of ~50-70% grass cover, 5-10% shrub cover, and a fire return interval of ~3 yr. These thresholds may provide adaptive potential for managing nonlinear behavior in socio-ecological systems in a changing environment. [ABSTRACT FROM AUTHOR]

Many North American grasslands are receiving atmospheric nitrogen (N) deposition at rates above what are considered critical eutrophication thresholds. Yet, potential changes in grassland function due to anthropogenic N deposition are poorly resolved, especially considering that other dynamic factors such as land use and precipitation can also affect N availability. To better understand whether elevated N deposition has altered ecosystem structure or function in North American grasslands, we analyzed a 27-year record of ecophysiological, community, and ecosystem metrics for an annually burned Kansas tallgrass prairie. Over this time, despite increasing rates of N deposition that are within the range of critical loads for grasslands, there was no evidence of eutrophication. Plant N concentrations did not increase, soil moisture did not decline, forb diversity did not decHne, and the relative abundance of dominant grasses did not shift toward more eutrophic species. Neither aboveground primary productivity nor N availability to plants increased. The fates of deposited N in grasslands are still uncertain, and could include management losses through burning and grazing. However, evidence from this grassland indicates that eutrophication of North American grassland ecosystems is not an inevitable consequence of current levels of N deposition. [ABSTRACT FROM AUTHOR]

In the central Great Plains of North America, climate change predictions include increases in mean annual temperature of 1.5-5.5 °C by 2100. Ecosystem responses to increased temperatures are likely to be regulated by dominant plant species, such as the potential biofuel species Panicum virgatum (switchgrass) in the tallgrass prairie. To describe the potential physiological and whole-plant responses of this species to future changes in air temperatures, we used louvered open-sided chambers (louvered OSC; 1 × 1 m, adjustable height) to passively alter canopy temperature in native stands of P. virgatum growing in tallgrass prairie at varying topographic positions (upland/lowland). The altered temperature treatment decreased daily mean temperatures by 1 °C and maximum temperatures by 4 °C in May and June, lowered daytime stomatal conductance and transpiration, decreased tiller density, increased specific leaf area, and delayed flowering. Among topographic contrasts, aboveground biomass, flowering tiller density, and tiller weight were greater in lowland sites compared to upland sites, with no temperature treatment interactions. Differences in biomass production responded more to topography than the altered temperature treatment, as soil water status varied considerably between topographic positions. These results indicate that while water availability as a function of topography was a strong driver of plant biomass, many leaf-level physiological processes were responsive to the small decreases in daily mean and maximum temperature, irrespective of landscape position. The varying responses of leaf-level gas exchange and whole-plant growth of P. virgatum in native stands to altered air temperature or topographic position illustrate that accurately forecasting yields for P. virgatum in mixed communities will require greater integration of physiological responses to simulated climate change (increased temperature) and resource availability over natural environmental gradients (soil moisture). [ABSTRACT FROM AUTHOR]

Changes in precipitation amount and variability have the potential to alter the structure and function of grasslands, but we know little about how changes in the timing of precipitation might affect grasslands. Here, we analyze long-term records from a tallgrass prairie to show that shifts in the timing of precipitation during the growing season have little effect on primary productivity or grass reproduction, but can greatly affect grazer performance. While greater late-season precipitation increases the weight gain of adult and young bison, greater mid-season precipitation decreases their weight gain. In addition, calving rates are lower after years with greater mid-season precipitation and higher after years with greater late-season precipitation. As well-timed drought can actually increase grazer weight gain and reproduction, it will be necessary to generate predictions of within-season distribution of precipitation to successfully forecast future grazer performance. [ABSTRACT FROM AUTHOR]

Woody encroachment is a widespread and acute phenomenon affecting grasslands and savannas worldwide. We performed a meta-analysis of 29 studies from 13 different grassland/savanna communities in North America to determine the consequences of woody encroachment on plant species richness. In all 13 communities, species richness declined with woody plant encroachment (average decline = 45%). Species richness declined more in communities with higher precipitation (r2 = 0.81) and where encroachment was associated with a greater change in annual net primary productivity (ANPP; r2 = 0.69). Based on the strong positive correlation between precipitation and ANPP following encroachment (r2 = 0.87), we hypothesize that these relationships occur because water-limited woody plants experience a greater physiological and demographic release as precipitation increases. The observed relationship between species richness and ANPP provides support for the theoretical expectation that a trade-off occurs between richness and productivity in herbaceous communities. We conclude that woody plant encroachment leads to significant declines in species richness in North American grassland/savanna communities. [ABSTRACT FROM AUTHOR]

Summary [ABSTRACT FROM AUTHOR]

In plants, ecologically important life history traits often display clinal patterns of population divergence. Such patterns can provide strong evidence for spatially varying selection across environmental gradients but also may result from nonselective processes, such as genetic drift, population bottlenecks and spatially restricted gene flow. Comparison of population differentiation in quantitative traits (measured as Q) with neutral molecular markers (measured as F) provides a useful tool for understanding the relative importance of adaptive and nonadaptive processes in the formation and maintenance of clinal variation. Here, we demonstrate the existence of geographic variation in key life history traits in the diploid perennial sunflower species Helianthus maximiliani across a broad latitudinal transect in North America. Strong population differentiation was found for days to flowering, growth rate and multiple size-related traits. Differentiation in these traits greatly exceeds neutral predictions, as determined both by partial Mantel tests and by comparisons of global Q values with theoretical F distributions. These findings indicate that clinal variation in these life history traits likely results from local adaptation driven by spatially heterogeneous environments. [ABSTRACT FROM AUTHOR]

The flowering of grasses is a process critical to plant population dynamics and genetics, herbivore performance, and human health. To better understand the climate factors governing grass flowering, we analyzed the patterns of culm production over 25 years for three perennial tallgrass prairie species at Konza Prairie in Kansas, USA. The three species (Andropogon gerardii, Sorghastrum nutans, and Schizachyrium scoparium) all utilize the C4 photosynthetic pathway and were measured annually at the same locations for the past 25 years in an annually burned watershed. Culm production of all three species increased with higher growing-season soil moisture and precipitation but differed in their responses to water availability at different times during the growing season. Relative to Andropogon, Sorghastrum responded more to precipitation early in the growing season, and Schizachyrium responded more to precipitation late in the growing season. Flowering by each species also revealed a threshold relationship with late-season soil moisture at ∼1 m depth, which likely is a proxy for season-long water balance. Although flowering can be influenced by conditions antecedent to the current growing season, neither soil moisture nor precipitation during the previous year influenced flowering over the 25-year period. Flowering culm production averaged 9% and 7% of total graminoid aboveground net primary production (ANPP) in the uplands and lowlands, respectively. Interannual variation in ANPP correlated only with Sorghastrum flowering, suggesting a predominant role of the species in ANPP responses to climate. [ABSTRACT FROM AUTHOR]

1. If we are to understand the mechanisms underlying species responses to climate change in natural systems, studies are needed that focus on responses of non-model species under field conditions. We measured transcriptional profiles of individuals of Andropogon gerardii, a C4 grass native to North American grasslands, in a field experiment in which both temperature and precipitation were manipulated to simulate key aspects of forecasted climate change. 2. By using microarrays developed for a closely related model species, Zea mays, we were able to compare the relative influence of warming versus altered soil moisture availability on expression levels of over 7000 genes, identify responsive functional groups of genes and correlate changes in gene transcription with physiological responses. 3. We observed more statistically significant shifts in transcription levels of genes in response to thermal stress than in response to water stress. We also identified candidate genes that demonstrated transcription levels closely associated with physiological variables, in particular chlorophyll fluorescence. 4. Synthesis. These results suggest that an ecologically important species responds differently to different environmental aspects of forecast climate change. These translational changes have the potential to influence phenotypic characters and ultimately adaptive responses. [ABSTRACT FROM AUTHOR]

Climate change is causing measurable changes in rainfall patterns, and will likely cause increases in extreme rainfall events, with uncertain implications for key processes in ecosystem function and carbon cycling. We examined how variation in rainfall total quantity ( Q), the interval between rainfall events ( I), and individual event size ( SE) affected soil water content (SWC) and three aspects of ecosystem function: leaf photosynthetic carbon gain ( ), aboveground net primary productivity (ANPP), and soil respiration ( ). We utilized rainout shelter-covered mesocosms (2.6 m3) containing assemblages of tallgrass prairie grasses and forbs. These were hand watered with 16 I× Q treatment combinations, using event sizes from 4 to 53 mm. Increasing Q by 250% (400–1000 mm yr−1) increased mean soil moisture and all three processes as expected, but only by 20–55% ( P≤0.004), suggesting diminishing returns in ecosystem function as Q increased. Increasing I (from 3 to 15 days between rainfall inputs) caused both positive ( ) and negative ( ) changes in ecosystem processes (20–70%, P≤0.01), within and across levels of Q, indicating that I strongly influenced the effects of Q, and shifted the system towards increased net carbon uptake. Variation in SE at shorter I produced greater response in soil moisture and ecosystem processes than did variation in SE at longer I, suggesting greater stability in ecosystem function at longer I and a priming effect at shorter I. Significant differences in ANPP and between treatments differing in I and Q but sharing the same SE showed that the prevailing pattern of rainfall influenced the responses to a given event size. Grassland ecosystem responses to extreme rainfall patterns expected with climate change are, therefore, likely to be variable, depending on how I, Q, and SE combine, but will likely result in changes in ecosystem carbon cycling. [ABSTRACT FROM AUTHOR]

The majority of tallgrass prairie root biomass is located in the upper soil layers (0–25 cm), but species differences exist in reliance on soil water at varying depths. These differences have led to the hypothesis that resource partitioning belowground facilitates species co-existence in this mesic grassland. To determine if plant water relations can be linked to soil water partitioning as a potential mechanism allowing C3 species to persist among the more dominant C4 grasses, we measured differences in the source of water-use using the isotopic signature of xylem water, volumetric soil water content at 4 depths, and leaf water potentials. Data were collected for seven species representing C4 grasses, C3 forbs and C3 shrubs over three growing seasons at the Konza Prairie (Kansas, USA) to encompass a range of natural climatic conditions. C4 grasses relied on shallow soil water (5 cm) across the growing season and had midday leaf water potentials that were highly correlated with shallow soil water regardless of soil water availability at other portions of the soil profile (20, 40 and 90 cm). In contrast, C3 species only used shallow soil water when plentiful at this depth; these species increased their dependence on soil water from greater depths as the upper soil layers dried. Structural equation models describing plant water relations were very similar for the three C4 species, whereas a unique set of models and drivers were identified for each of the C3 species. These results support soil water partitioning as a mechanism for species coexistence, as C4 species in this grassland have relatively consistent dependence on water in shallow soil layers, whereas C3 species show niche differentiation in water use strategies to avoid competition with C4 grasses for water in shallow soil layers when this resource is limiting and leaf water stress is high. [ABSTRACT FROM AUTHOR]

North American grasslands have experienced increased relative abundance of shrubs and trees over the last 150 yr. Alterations in herbivore composition, abundance, and grazing pressure along with changes in fire frequency are drivers that can regulate the transition from grassland to shrubland or woodland (a process known as woody encroachment). Historically, North American grasslands had a suite of large herbivores that grazed and/or browsed (i.e., bison, elk, pronghorn, deer), as well as frequent and intense fires. In the tallgrass prairie, many large native ungulates were extirpated by the 1860s, corresponding with increased homesteading (which led to decreased fire frequencies and intensities). Changes in the frequency and intensity of these two drivers (browsing and fire) have coincided with woody encroachment in tallgrass prairie. Within tallgrass prairie, woody encroachment can be categorized in to two groups: non‐resprouting species that can be killed with fire and resprouting species that cannot be killed with fire. Resprouting species require additional active management strategies to decrease abundance and eventually be removed from the ecosystem. In this study, we investigated plant cover, ramet density, and physiological effects of continuous simulated browsing and prescribed fire on Cornus drummondii C.A. Mey, a resprouting clonal native shrub species. Browsing reduced C. drummondii canopy cover and increased grass cover. We also observed decreased ramet density, which allowed for more infilling of grasses. Photosynthetic rates between browsed and unbrowsed control shrubs did not increase in 2015 or 2016. In 2017, photosynthetic rates for browsed shrubs were higher in the unburned site than the unbrowsed control shrubs at the end of the growing season. Additionally, after the prescribed fire, browsed shrubs had ~90% decreased cover, ~50% reduced ramet density, and grass cover increased by ~80%. In the roots of browsed shrubs after the prescribed fire, nonstructural carbohydrates (NSC) experienced a twofold reduction in glucose and a threefold reduction in both sucrose and starch. The combined effects of browsing and fire show strong potential as a successful management tool to decrease the abundance of clonal‐resprouting woody plants in mesic grasslands and illustrate the potential significance of browsers as a key driver in this ecosystem. [ABSTRACT FROM AUTHOR]

Summary [ABSTRACT FROM AUTHOR]