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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]

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]

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]

Woody encroachment has impacted grassland ecohydrology worldwide, prompting management strategies aimed at woody vegetation removal to prevent or mitigate loss of water yield. We measured stream discharge following sustained cutting of riparian trees (2010–2020) in a native tallgrass prairie (northeastern Kansas, USA). Discharge has declined at this site since the 1980's despite a concurrent increase in precipitation. This decline has been previously attributed to increased transpiration of stream water by riparian vegetation. We used water stable isotopes (δ18O and δ2H) to determine whether riparian grasses, shrubs, and trees primarily used stream/groundwater or soil water. Additionally, we quantified the increase in riparian and non-riparian woody cover (1978–2020) and combined it with sap-flux data to estimate changes in transpirative water loss. Sustained cutting of riparian trees did not result in increased discharge. Rather than stream/groundwater, the largest proportion of water used by riparian trees (Quercus spp.) was deep soil water. Cornus drummondii (clonal woody shrub) used a higher proportion of stream water and had greater overall variability in water-use. Riparian shrub cover increased about 57% from 1978 to 2020. Over the same time period, shrub cover increased about 20% in areas outside the riparian zone, resulting in an estimated 25% increase in daily transpirative water loss. Although stream water use was less than 50% for all riparian zone species, the total increase in shrub cover on this watershed, coupled with higher transpiration rates of shrubs, suggests that these woody species—within and outside the riparian zone—are key contributors to observed declines in stream flow in this system. [ABSTRACT FROM AUTHOR]

Grassland ecosystems are historically shaped by climate, fire, and grazing which are essential ecological drivers. These grassland drivers influence morphology and productivity of grasses via physiological processes, resulting in unique water and carbon-use strategies among species and populations. Leaf-level physiological responses in plants are constrained by the underlying anatomy, previously shown to reflect patterns of carbon assimilation and water-use in leaf tissues. However, the magnitude to which anatomy and physiology are impacted by grassland drivers remains unstudied. To address this knowledge gap, we sampled from three locations along a latitudinal gradient in the mesic grassland region of the central Great Plains, USA during the 2018 (drier) and 2019 (wetter) growing seasons. We measured annual biomass and forage quality at the plot level, while collecting physiological and anatomical traits at the leaf-level in cattle grazed and ungrazed locations at each site. Effects of ambient drought conditions superseded local grazing treatments and reduced carbon assimilation and total productivity in A. gerardii. Leaf-level anatomical traits, particularly those associated with water-use, varied within and across locations and between years. Specifically, xylem area increased when water was more available (2019), while xylem resistance to cavitation was observed to increase in the drier growing season (2018). Our results highlight the importance of multi-year studies in natural systems and how trait plasticity can serve as vital tool and offer insight to understanding future grassland responses from climate change as climate played a stronger role than grazing in shaping leaf physiology and anatomy. [ABSTRACT FROM AUTHOR]

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]

• The accuracy of indirect LAI measurements was evaluated in dense shrub canopies. • Indirect methods were compared against direct LAI measurements. • Ceptometer was accurate in non-browsed canopies but overestimated browsed canopies. • The NEON airborne observatory platform spectral LAI underestimated LAI of canopies. • A handheld 3D scanner had high precision but underestimated large leaf area values. Leaf area index (LAI) is a key variable describing ecosystem structure and influencing the exchange of carbon, water, and energy. LAI is often evaluated with indirect methods. However, the accuracy of indirect measurements can vary with canopy structure and is not always generalizable across ecosystems. Previous research has characterized the accuracy of indirect methods for woody plants in forest ecosystems, but it is not well established for woody plants in open ecosystems—despite having large differences in canopy structure. In this study, we compared direct LAI measurements in clonal grassland shrub canopies to three indirect methods: a ceptometer, a handheld 3D scanner (processed using EXScanPro-3.6 software), and NEON's LAI product obtained from airborne hyperspectral imaging (derived from SAVI). To our knowledge, this is the first study to assess the accuracy of leaf area data in woody plants using a handheld 3D scanner and one of few studies assessing the accuracy of the National Ecological Observation Network's (NEON) Airborne Observation Platform—our source of airborne hyperspectral imaging. Data were collected in tallgrass prairie undergoing woody encroachment and three treatments: no herbivore disturbance, bison present, and simulated browsing. We found that direct LAI measurements of control and grazed C. drummondii canopies averaged ∼8.0. The ceptometer accurately estimated LAI in non-browsed canopies but overestimated LAI of browsed canopies by 38 %. One-sided leaf area of ramets measured with a handheld 3D scanner was strongly related to direct measurements (r2=0.86), but underestimated leaf area at greater values. LAI estimated from airborne spectral data underestimated LAI by 55 %. We conclude that a ceptometer was adequate for measuring LAI in dense shrub canopies when browsing was not present, the handheld 3D scanner was adequate for measuring leaf area of individual ramets, and the airborne spectral data was not suitable for estimating LAI of dense, grassland shrub canopies. [ABSTRACT FROM AUTHOR]

The expansion of woody species into grasslands has altered community structure and ecosystem function of grasslands worldwide. In tallgrass prairie of the Central Great Plains, USA, decreased fire frequency and intensity have increased the cover and abundance of woody species. In particular, clonal shrub cover has increased at accelerated rates due to vegetative reproduction and resprouting after disturbance. We measured the intra-clonal stem demography and relative growth rates (estimated change in woody biomass) of the shrub Cornus drummondii in response to fire frequency (4 vs 20 year burn intervals) and simulated browsing during the 2018 and 2019 growing seasons at Konza Prairie Biological Station (Manhattan, Kansas). Overall, infrequent fire (4 year burn interval) increased intra-clonal stem relative growth rates and shrub relative growth rates. Intra-clonal stem relative growth rates were reduced in unbrowsed clones in 2018 due to drought and simulated browsing reduced intra-clonal stem relative growth rates in 2019. Additionally, simulated browsing nearly eliminated flower production within clones but did not affect intra-clonal stem mortality or recruitment within a growing season. Fire in conjunction with simulated browsing reduced estimated relative growth rates for entire shrub clones. Browsed shrubs that experienced prescribed fire in 2017 had reduced intra-clonal stem densities compared to unbrowsed shrubs and stem densities of browsed shrubs did not recover in 2018 or 2019. These results illustrate that infrequent fire alone promotes the expansion of clonal shrubs in tallgrass prairie and multiple interacting disturbances (e.g., fire and browsing) are required to control the spread of clonal shrubs into grasslands. [ABSTRACT FROM AUTHOR]

Background and Aims Andropogon gerardii is a highly productive C4 grass species with a large geographic range throughout the North American Great Plains, a biome characterized by a variable temperate climate. Plant traits are often invoked to explain growth rates and competitive abilities within broad climate gradients. For example, plant competition models typically predict that species with large geographic ranges benefit from variation in traits underlying high growth potential. Here, we examined the relationship between climate variability and leaf-level traits in A. gerardii , emphasizing how leaf-level microanatomical traits serve as a mechanism that may underlie variation in commonly measured traits, such as specific leaf area (SLA). Methods Andropogon gerardii leaves were collected in August 2017 from Cedar Creek Ecosystem Science Reserve (MN), Konza Prairie Biological Station (KS), Platte River Prairie (NE) and Rocky Mountain Research Station (SD). Leaves from ten individuals from each site were trimmed, stained and prepared for fluorescent confocal microscopy to analyse internal leaf anatomy. Leaf microanatomical data were compared with historical and growing season climate data extracted from PRISM spatial climate models. Key Results Microanatomical traits displayed large variation within and across sites. According to AICc (Akaike's information criterion adjusted for small sample sizes) selection scores, the interaction of mean precipitation and temperature for the 2017 growing season was the best predictor of variability for the anatomical and morphological traits measured here. Mesophyll area and bundle sheath thickness were directly correlated with mean temperature (annual and growing season). Tissues related to water-use strategies, such as bulliform cell and xylem area, were significantly correlated with one another. Conclusions The results indicate that (1) microanatomical trait variation exists within this broadly distributed grass species, (2) microanatomical trait variability appears likely to impact leaf-level carbon and water use strategies, and (3) microanatomical trait values vary across climate gradients, and may underlie variation in traits measured at larger ecological scales. [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]

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]

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 Climate variability is a key driver of physiological responses in common grass species in grasslands of North America. Differences in microanatomical traits among coexisting species may influence physiological responses to climate variability over large geographic scales. The goal of this research was to determine leaf-level physiological and microanatomical trait variability among four dominant C 4 grass species across a natural precipitation gradient. Physiological traits were observed to vary significantly across the gradient with greater variability than microanatomical traits. Microanatomical traits were shown to predict physiological responses in A. gerardii and P. virgatum , but the nature of the relationships varied between species. These results illustrate that microanatomical and physiological traits vary across a precipitation gradient, there are clear linkages between microanatomy and physiology in grass species, and this evidence underscores the need for further investigation using phylogenetically diverse assemblages. Highlights • Physiology and microanatomy in C 4 grasses varies across precipitation gradients. • Within species, trait variability is low within individual locations. • Microanatomical traits did not vary significantly over the growing season. • Grass physiology can be predicted by species – specific microanatomical traits. [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]

A significant fraction of the terrestrial biosphere comprises biomes containing tree-grass mixtures. Forecasting vegetation dynamics in these environments requires a thorough understanding of how trees and grasses use and compete for key belowground resources. There is disagreement about the extent to which tree-grass vertical root separation occurs in these ecosystems, how this overlap varies across large-scale environmental gradients, and what these rooting differences imply for water resource availability and tree-grass competition and coexistence. To assess the extent of tree-grass rooting overlap and how tree and grass rooting patterns vary across resource gradients, we examined landscape-level patterns of tree and grass functional rooting depth along a mean annual precipitation (MAP) gradient extending from ~ 450 to ~ 750 mm year in Kruger National Park, South Africa. We used stable isotopes from soil and stem water to make inferences about relative differences in rooting depth between these two functional groups. We found clear differences in rooting depth between grasses and trees across the MAP gradient, with grasses generally exhibiting shallower rooting profiles than trees. We also found that trees tended to become more shallow-rooted as a function of MAP, to the point that trees and grasses largely overlapped in terms of rooting depth at the wettest sites. Our results reconcile previously conflicting evidence for rooting overlap in this system, and have important implications for understanding tree-grass dynamics under altered precipitation scenarios. [ABSTRACT FROM AUTHOR]

Hydraulic lift, the passive movement of water through plant roots from wet to dry soil, is an important ecohydrological process in a wide range of water-limited ecosystems. This phenomenon may also alter plant functioning, growth, and survival in mesic grasslands, where soil moisture is spatially and temporally variable. Here, we monitored diurnal changes in the isotopic signature of soil and plant xylem water to assess (1) whether hydraulic lift occurs in woody and herbaceous tallgrass prairie species ( Rhus glabra, Amorpha canescens, Vernonia baldwinii, and Andropogon gerardii), (2) if nocturnal transpiration or grazing by large ungulates limits hydraulic lift, and (3) if a dominant grass, A. gerardii, utilizes water lifted by other tallgrass prairie species. Broadly, the results shown here suggest that hydraulic lift does not appear to be widespread or common in this system, but isolated instances suggest that this process does occur within tallgrass prairie. The isolated instance of hydraulic lift did not vary by grazing treatment, nor did they result in facilitation for neighboring grasses. We suggest that the topographic complexity of this tallgrass prairie and the high rates of nocturnal transpiration observed in this study likely limit the frequency and occurrence of hydraulic lift. These results suggest that hydraulic lift can be a patchy process, particularly in heterogeneous landscapes. [ABSTRACT FROM AUTHOR]

Background and Aims: Fire and grazing are important disturbances in grasslands, yet we know little about how they impact a variety of plant physiological processes such as plant ecohydrology. Here, we assessed the impact of fire history and grazing by Bison bison on the source of water uptake and niche overlap in common grassland species at the Konza Prairie Biological Station, a temperate mesic grassland located in northeastern Kansas, USA. Methods: We used the stable isotopic signature of soil and xylem water to evaluate water uptake in Andropogon gerardii, Vernonia baldwinii, Amorpha canescens, and Rhus glabra within varying grazing (grazed, ungrazed), fire (0,1,2 or 3 years since last burn), topography (upland, lowland), and month (July, August) contrasts over 3 years (2013-2015). Results: The presence of grazers, not fire history, altered water uptake patterns in these common grassland species. Particularly, grazing increased the proportion of shallow water utilized by A. gerardii and R. glabra, reducing niche overlap with other co-occurring species. However, these responses varied intra-annually and were often modulated by topography. Conclusions: These results suggest that grazing can alter aspects of grassland ecohydrology at small scales, which may extend to impact community and ecosystem processes at larger spatial scales. [ABSTRACT FROM AUTHOR]

Global change drivers are altering climatic and edaphic conditions of ecosystems across the globe, and we expect novel plant communities to become more common as a result. In the Colorado Front Range, compositional changes have occurred in the mixed-grass prairie plant community in conjunction with shifts in winter precipitation and atmospheric nitrogen (N) deposition. To test whether these environmental changes have been responsible for the observed plant community change, we conducted an in situ manipulative experiment in a mixed-grass meadow near Boulder, CO. We simulated historical conditions by reducing N availability (+500 g C m year) and winter precipitation (with rainout shelters) for 2 years (2013-2014) and compared vegetation response to these treatments with that of ambient conditions. The site experienced an extreme precipitation event in autumn 2013 which allowed comparison of an exceptionally wet year with an average year. We measured pre-treatment species composition in 2012, and treatment responses in the spring and summer of 2013 and 2014. As predicted, simulating historical low N-winter dry conditions resulted in a plant community dominated by historically abundant species. Cool-season introduced species were significantly reduced in low N-winter dry plots, particularly the annual plants Bromus tectorum and Alyssum parviflorum. These same species responded strongly to the extreme precipitation event with large increases, while native grasses and forbs showed little change in productivity or composition under varying climatic or edaphic conditions. This work provides clear evidence linking on-going global change drivers to altered plant community composition in an otherwise relatively undisturbed grassland ecosystem. [ABSTRACT FROM AUTHOR]

Nitrogen (N) and phosphorus (P) are limiting nutrients for many plant communities worldwide. Foliar N and P along with leaf area are among the most important controls on photosynthesis and hence productivity. However, foliar N and P are typically assessed as species level traits, whereas productivity is often measured at the community scale. Here, we compared the community-level traits of leaf area index (LAI) to total foliar nitrogen (TFN) and total foliar phosphorus (TFP) across nearly three orders of magnitude LAI in grazed and ungrazed tallgrass prairie in north-eastern Kansas, USA. LAI was strongly correlated with both TFN and TFP across communities, and also within plant functional types (grass, forb, woody, and sedge) and grazing treatments (bison or cattle, and ungrazed). Across almost the entire range of LAI values and contrasting communities, TFN:TFP ratios indicated co-limitation by N and P in almost all communities; this may further indicate a community scale trend of an optimal N and P allocation per unit leaf area for growth. Previously, results from the arctic showed similar tight relationships between LAI:TFN, suggesting N is supplied to canopies to maximize photosynthesis per unit leaf area. This tight coupling between LAI, N, and P in tallgrass prairie suggests a process of optimal allocation of N and P, wherein LAI remains similarly constrained by N and P despite differences in species composition, grazing, and canopy density. [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]

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]

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]

Soil moisture is a critical variable in grassland function, yet how fire regimes influence ecohydrology is poorly understood. By altering productivity, species composition, and litter accumulation, fire can indirectly increase or decrease soil water depletion on a range of time scales and depths in the soil profile. To better understand how fire influences soil moisture in grasslands, we analyzed 28 years of soil moisture data from two watersheds in a central North American grassland which differ in their long-term fire frequency. Across 28 years, cessation of prescribed burning initially led to wetter soils, likely as litter accumulated and both transpiration and evaporation were suppressed. Long-term, cessation of burning led to soils drying more, especially at depths greater than 75 cm. The long-term drying of deep soils is consistent with the increase in woody species in the infrequently burned grassland as woody species likely have a greater reliance on soil water from deeper soil layers compared to co-occurring herbaceous species. Despite the ecohydrological changes associated with the cessation of prescribed burning, watersheds with different burn regimes responded similarly to short-term variation in climate variation. In both watersheds, low precipitation and high temperatures led to drier soils with greater responses in soil moisture to climate variation later in the season than earlier. There is no current evidence that the cessation of burning in this ecosystem will qualitatively alter how evapotranspiration responds to climate variation, but the use of deeper soil water by woody plants has the potential for greater transpiration during dry times. In all, modeling the depth-specific responses of soil moisture and associated ecosystem processes to changes in burn regimes will likely require including responses of plant community composition over short and long time scales. [ABSTRACT FROM AUTHOR]

Shrub encroachment of grasslands is a transformative ecological process by which native woody species increase in cover and frequency and replace the herbaceous community. Mechanisms of encroachment are typically assessed using temporal data or experimental manipulations, with few large spatial assessments of shrub physiology. In a mesic grassland in North America, we measured inter- and intra-annual variability in leaf δ13C in Cornus drummondii across a grassland landscape with varying fire frequency, presence of large grazers and topographic variability. This assessment of changes in individual shrub physiology is the largest spatial and temporal assessment recorded to date. Despite a doubling of annual rainfall (in 2008 versus 2011), leaf δ13C was statistically similar among and within years from 2008-11 (range of −28 to −27‰). A topography*grazing interaction was present, with higher leaf δ13C in locations that typically have more bare soil and higher sensible heat in the growing season (upland topographic positions and grazed grasslands). Leaf δ13C from slopes varied among grazing contrasts, with upland and slope leaf δ13C more similar in ungrazed locations, while slopes and lowlands were more similar in grazed locations. In 2011, canopy greenness (normalized difference vegetation index – NDVI) was assessed at the centroid of individual shrubs using high-resolution hyperspectral imagery. Canopy greenness was highest mid-summer, likely reflecting temporal periods when C assimilation rates were highest. Similar to patterns seen in leaf δ13C, NDVI was highest in locations that typically experience lowest sensible heat (lowlands and ungrazed). The ability of Cornus drummondii to decouple leaf physiological responses from climate variability and fire frequency is a likely contributor to the increase in cover and frequency of this shrub species in mesic grassland and may be generalizable to other grasslands undergoing woody encroachment. [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]

Nitrogen (N) metabolism is a major research target for increasing productivity in crop plants. In maize (Zea mays L.), yield gain over the last few decades has been associated with increased N absorption and utilization efficiency (i.e. grain biomass per unit of N absorbed). However, a dynamical framework is still needed to unravel the role of internal processes such as uptake, allocation, and translocation of N in these adaptations. This study aimed to 1) characterize how genetic enhancement in N efficiency conceals changes in allocation and translocation of N, and 2) quantify internal fluxes behind grain N sources in two historical genotypes under high and low N supply. The genotypes 3394 and P1197, landmark hybrids representing key eras of genetic improvement (1990s and 2010s), were grown under high and low N supply in a two-year field study. Using stable isotope 15N labelling, post-silking nitrogen fluxes were modeled through Bayesian estimation by considering the external N (exogenous-N) and the pre-existing N (endogenous-N) supply across plant organs. Regardless of N availability, P1197 exhibited greater exogenous-N accumulated in leaves and cob-husks. This response was translated to a larger amount of N mobilized to grains (as endogenous-N) during grain-filling in this genotype. Furthermore, the enhanced N supply to leaves in P1197 was associated with increased post-silking carbon accumulation. The overall findings suggest that increased N utilization efficiency over time in maize genotypes was associated with an increased allocation of N to leaves and subsequent translocation to the grains. • Bayesian modelling and 15N analysis were combined to estimate N allocation in maize. • Modern maize plants evidenced an improved post-silking N partitioning to leaves. • Greater N allocation to leaves resulted in an increased post-silking carbon fixation. • Internal translocation to the grains was improved due to larger supply from leaves. [ABSTRACT FROM AUTHOR]

Background: C grass species in the mesic tallgrass prairie of central North America can exhibit both high root production and deep rooting in the soil profile (>2 m). Differences in root growth and the types of roots produced vary according to local environmental gradients and management practices. The production of deep roots in tallgrass prairie has been historically presumed as a mechanism for water uptake when surface soils are dry. Methods: We examined changes in root biomass, total root length, root width, and theoretical hydraulic conductivity using roots collected from deep soil cores in upland and lowland topographic positions in grazed and ungrazed watersheds of the Konza Prairie Biological Station in north-eastern Kansas, USA. Results: Root biomass, total root length, and theoretical hydraulic conductivity were highest in roots found in the top 20 cm of the soil profile, and then declined exponentially with increasing soil depth. Compared to grazed areas, ungrazed locations had more root biomass and total root length of roots in the most superficial soil layers. No differences in rooting profiles were present among topographic contrasts. Theoretical hydraulic conductivity of axial root xylem did not vary by topographic position or grazing contrasts, and declines in conductivity by depth were driven by changes in the number of vessels per stele, rather than changes in vessel size. Conclusions: Irrespective of differences by grazing treatment or topographic position, significant reductions in root biomass, total root length, and theoretical hydraulic conductivity of grass roots at soil depths greater than 1 m suggest deep roots in this grassland have limited functional significance for water uptake. [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]

Future climates are forecast to include greater precipitation variability and more frequent heat waves, but the degree to which the timing of climate variability impacts ecosystems is uncertain. In a temperate, humid grassland, we examined the seasonal impacts of climate variability on 27 y of grass productivity. Drought and high-intensity precipitation reduced grass productivity only during a 110-d period, whereas high temperatures reduced productivity only during 25 d in July. The effects of drought and heat waves declined over the season and had no detectable impact on grass productivity in August. If these patterns are general across ecosystems, predictions of ecosystem response to climate change will have to account not only for the magnitude of climate variability but also for its timing. [ABSTRACT FROM AUTHOR]

Anthropogenic climate change is projected to alter precipitation patterns, resulting in novel environments for plants. The responses of dominant plant species (e.g. Panicum virgatum L. (switchgrass)) to climate changes can drive broader ecosystem processes such as primary productivity. Using a rainfall mesocosm facility, three ecotypes of P. virgatum (collected from Kansas, Oklahoma and Texas, USA) were subjected to three precipitation regimes (average, -25%, +25%) to determine the physiological and growth responses to altered precipitation in a common garden setting. Results showed mean maximum photosynthetic rates, stomatal conductance, transpiration, midday water potential and dark-adapted chlorophyll fluorescence were lowest in the Kansas ecotypes. Increased precipitation treatments raised the mean midday water potentials and lowered water-use efficiency. Aboveground biomass responded positively to changes in precipitation, but flowering initiation was later and rates were lower for Texas ecotypes. In general, ecotype origin was a better predictor of differences in physiological responses and flowering, whereas the precipitation treatments had greater control over biomass production. Depending on the growth variable measured, these results show responses for P. virgatum are under varying ecotypic or environmental control with few interactions, suggesting that future predictions to climate change need not inherently consider localised adaptations in this economically important and widely distributed species. [ABSTRACT FROM AUTHOR]

oveground biomass in grasslands varies according to landscape gradients in resource availability and seasonal patterns of growth. Using a transect spanning a topographic gradient in annually burned ungrazed tallgrass prairie, we measured changes in the height of four abundant C grass species, LAI, biomass, and cumulative carbon flux using two closely located eddy flux towers. We hypothesized that seasonal patterns of plant growth would be similar across the gradient, but the magnitude of growth and biomass accumulation would vary by topographic position, reflecting spatial differences in microclimate, slope, elevation, and soil depth. Thus, identifying and measuring local growth responses according to topographic variability should significantly improve landscape predictions of aboveground biomass. For most of the growth variables measured, classifying topography into four positions best captured the inherent spatial variability. Biomass produced, seasonal LAI and species height increased from the upland and break positions to the slope and lowland. Similarly, cumulative carbon flux in 2008 was greater in lowland versus upland tower locations (difference of 64 g m by DOY 272). Differences in growth by topographic position reflected increased production of flowering culms by Andropogon gerardii and Sorghastrum nutans in lowland. Varying growth responses by these species may be a significant driver of biomass and carbon flux differences by topographic position, at least for wet years. Using a digital elevation model to classify the watershed into topographic positions, we performed a geographically weighted regression to predict landscape biomass. The minimum and maximum predictions of aboveground biomass for this watershed had a large range (86-393 t per 40.4 ha), illustrating the drastic spatial variability in growth within this annually-burned grassland. [ABSTRACT FROM AUTHOR]

Abstract: Switchgrass (Panicum virgatum L.) is a broadly adapted warm-season grass species native to most of the central and eastern United States. Switchgrass has been identified as a potential biofuel species because it is a native species that requires minimal management, and has a large potential to sequester carbon underground. Since the 1990’s, switchgrass has been bred to produce cultivars with increased biomass and feedstock quality. This review addresses potential ecological consequences of widespread switchgrass cultivation for biofuel production in the central United States. Specifically, this review address the ecological implications of changing use of marginal and CRP land, impacts on wildlife, potentials for disease and invasions, and changes in soil quality through reductions in erosion, decomposition rates, and carbon sequestrations. A central theme of the review is the utility of maintaining landscape heterogeneity during switchgrass biofuel production. This includes implementing harvest rotations, no till farming, and mixed species composition. If negative ecological consequences of switchgrass cultivation are minimized, biofuel production using this species has economical and environmental benefits. [Copyright &y& Elsevier]

Future climate change is likely to reduce the floristic diversity of grasslands. Yet the potential consequences of climate-induced plant species losses for the functioning of these ecosystems are poorly understood. We investigated how climate change might alter the functional composition of grasslands for Konza Prairie, a diverse tallgrass prairie in central North America. With species-specific climate envelopes, we show that a reduction in mean annual precipitation would preferentially remove species that are more abundant in the more productive lowland positions at Konza. As such, decreases in precipitation could reduce productivity not only by reducing water availability but by also removing species that inhabit the most productive areas and respond the most to climate variability. In support of this prediction, data on species abundance at Konza over 16 years show that species that are more abundant in lowlands than uplands are preferentially reduced in years with low precipitation. Climate change is likely to also preferentially remove species from particular functional groups and clades. For example, warming is forecast to preferentially remove perennials over annuals as well as Cyperaceae species. Despite these predictions, climate change is unlikely to unilaterally alter the functional composition of the tallgrass prairie flora, as many functional traits such as physiological drought tolerance and maximum photosynthetic rates showed little relationship with climate envelope parameters. In all, although climatic drying would indirectly alter grassland productivity through species loss patterns, the insurance afforded by biodiversity to ecosystem function is likely to be sustained in the face of climate change. [ABSTRACT FROM AUTHOR]

During a record drought (2006) in southwest Kansas, USA, we assessed groundwater dynamics in a shallow, unconfined aquifer, along with plant water sources and physiological responses of the invasive riparian shrub Tamarix ramosissima. In early May, diel water table fluctuations indicated evapotranspirative consumption of groundwater by vegetation. During the summer drought, the water table elevation dropped past the lowest position previously recorded. Concurrent with this drop, water table fluctuations abruptly diminished at all wells at which they had previously been observed despite increasing evapotranspirative demand. Following reductions in groundwater fluctuations, volumetric water content declined corresponding to the well-specific depths of the capillary fringe in early May, suggesting a switch from primary dependence on groundwater to vadose-zone water. In at least one well, the fluctuations appear to re-intensify in August, suggesting increased groundwater uptake by Tamarix or other non-senesced species from a deeper water table later in the growing season. Our data suggest that Tamarix can rapidly shift water sources in response to declines in the water table. The use of multiple water sources by Tamarix minimized leaf-level water stress during drought periods. This study illustrates the importance of the previous hydrologic conditions experienced by site vegetation for controlling root establishment at depth and demonstrates the utility of data from high-frequency hydrologic monitoring in the interpretation of plant water sources using isotopic methods. [ABSTRACT FROM AUTHOR]

Abstract: Ecosystem responses to climate change will largely be driven by responses of the dominant species. However, if co-dominant species have traits that lead them to differential responses, then predicting how ecosystem structure and function will be altered is more challenging. We assessed differences in response to climate change factors for the two dominant C4 grass species in tallgrass prairie, Andropogon gerardii and Sorghastrum nutans, by measuring changes in a suite of plant ecophysiological traits in response to experimentally elevated air temperatures and increased precipitation variability over two growing seasons. Maximum photosynthetic rates, stomatal conductance, water-use efficiency, chlorophyll fluorescence, and leaf water potential varied with leaf and canopy temperature as well as with volumetric soil water content (0–15cm). Both species had similar responses to imposed changes in temperature and water availability, but when differences occurred, responses by A. gerardii were more closely linked with changes in air temperature whereas S. nutans was more sensitive to changes in water availability. Moreover, S. nutans was more responsive overall than A. gerardii to climate alterations. These results indicate both grass species are responsive to forecast changes in temperature and precipitation, but their differential sensitivity to temperature and water availability suggest that future population and community structure may vary based on the magnitude and scope of an altered climate. [Copyright &y& Elsevier]

Water availability strongly governs grassland primary productivity, yet this resource varies dramatically in time (seasonally) and space (with soil depth and topography). It has long been assumed that co-occurring species differ in their partitioning of water use by depth, but direct evidence is lacking. We report data from two growing seasons (2004–2005) in which we measured the isotopic signature of plant xylem water from seven species (including C3 forbs and shrubs and C4 grasses) growing along a topographic gradient at the Konza Prairie Biological Station. Plant xylem stable oxygen isotope ratio (δ18O) values were compared to soil water δ18O profiles, recent rainfall events, and groundwater. Species varied in both their temporal patterns of water use and their responses to seasonal droughts in both years. During wet periods, species differences in water use were minimal, with common dependency on recent rainfall events stored in the upper soil layers. However, during dry periods, most C3 species used proportionally more water from deeper portions of the soil profile relative to the C4 grasses. Plants in uplands used more shallow soil water compared to those in lowlands, with the greatest differences across the topographic gradient occurring during dry periods. While the documented vertical root distribution varies by species and growth form in this grassland, each of the species we measured appeared to compete for the same surface layer soil moisture when water was not limiting. Thus, our results suggest that variation in precipitation history and landscape positions are greater determinants of water-use patterns than would be expected based on absolute rooting depth. [ABSTRACT FROM AUTHOR]

Abstract: The North American tallgrass prairie is composed of a diverse mix of C3 and C4 plant species that are subject to multiple resource limitations. C4 grasses dominate this ecosystem, purportedly due to greater photosynthetic capacity and resource-use efficiency associated with C4 photosynthesis. We tested the hypothesis that intrinsic physiological differences between C3 and C4 species are consistent with C4 grass dominance by comparing leaf gas exchange and chlorophyll fluorescence variables for seven C4 and C3 herbaceous species (legumes and non-legumes) in two different settings: experimental mesocosms and natural grassland sites. In the mesocosms, C4 grasses had higher photosynthetic rates, water potentials and water-use efficiency than the C3 species. These differences were absent in the field, where photosynthetic rates declined similarly among non-leguminous species. Thus, intrinsic photosynthetic advantages for C4 species measured in resource-rich mesocosms could not explain the dominance of C4 species in the field. Instead, C4 dominance in this ecosystem may depend more on the ability of the grasses to grow rapidly when resources are plentiful and to tolerate multiple limitations when resources are scarce. [Copyright &y& Elsevier]

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]

Aims: As woody plants encroach into grassland ecosystems, we expect altered plant-soil interactions to change the microbial processes that affect soil carbon storage and nutrient cycling. Specifically, this research aimed to address how (1) soil chemistry, (2) microbial nutrient demand, and (3) the rate and source of potential soil C mineralization vary spatially under clonal woody shrubs of varying size within a mesic grassland. Methods: We collected soil samples from the center, the midpoint between the center and edge, the edge, and the shrub-grass ecotone of multiple Cornus drummondii shrubs across a shrub-size gradient in infrequently burned tallgrass prairie. Results: Total soil carbon and nitrogen increased with shrub size at every sampling location but the edge. Microbial demand for nitrogen also increased as shrubs increased in size. Across all shrub sizes and sampling locations, potential soil carbon mineralization rates were higher when microbes broke down proportionally more shrub-derived (C3) organic matter than grass-derived (C4) organic matter. Conclusions: Our results suggest that the spatio-temporal context of woody encroachment is critical for understanding its impact on belowground microbial processes. In this ecosystem, a longer period of occupancy by woody plants increases potentially mineralizable soil carbon. [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]

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]

Aims: Global change agents are creating novel climatic and edaphic conditions that may favor introduced species. We attempted to identify mechanisms and impacts of Bromus tectorum invasion in the Colorado Front Range mixed-grass prairie under changing conditions. Methods: We conducted an in-situ experiment with three removal treatments (removal of B. tectorum, removal of Pascopyrum smithii, and no removal) and two nitrogen treatments (ambient and low N), and measured plant community response. We used isotopic analysis of δO and δD in plants and soils to identify seasonal source water of B. tectorum and P. smithii. Results: We found that dominance of B. tectorum was greatest under high resource conditions (ambient N, wet winter) now common due to increased N deposition and climate change. However, its removal had little impact on native plant abundance or composition. Isotopic results show B. tectorum and P. smithii partitioning water use between shallow versus deeper soil layers during the dry summer season. Conclusions: Our results suggest that changing environmental conditions favor the introduced grass Bromus tectorum over native species, but probably not due to altered competitive relationships. Instead, B. tectorum appears to be opportunistically responding to expansion of a phenological niche and increased nutrient availability. [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]

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]

Abstracts of comments on the article "Global Resilience of Tropical Forest and Savanna to Critical Transitions," by Egbert H. Van Nes, Milena Holmgren, Marina Hirota, and Marten Scheffer that appeared in the October 14, 2011 issue of "Science" are presented.

Over the past century of maize (Zea mays L.) breeding, grain yield progress has been the result of improvements in several other intrinsic physiological and morphological traits. In this study, we describe (i) the contribution of kernel weight (KW) to yield genetic gain across multiple agronomic settings and breeding programs, and (ii) the physiological bases for improvements in KW for US hybrids. A global-scale literature review concludes that rates of KW improvement in US hybrids were similar to those of other commercial breeding programs but extended over a longer period of time. There is room for a continued increase of kernel size in maize for most of the genetic materials analysed, but the trade-off between kernel number and KW poses a challenge for future yield progress. Through phenotypic characterization of Pioneer Hi-Bred ERA hybrids in the USA, we determine that improvements in KW have been predominantly related to an extended kernel-filling duration. Likewise, crop improvement has conferred on modern hybrids greater KW plasticity, expressed as a better ability to respond to changes in assimilate availability. Our analysis of past trends and current state of development helps to identify candidate targets for future improvements in maize. [ABSTRACT FROM AUTHOR]

Carbon (C) allocation strategy plays a critical role in plant adaptability, which are also important to assess the productivity stability under environmental change. Based on optimal partitioning theory, we asked two questions: (1) How is plant C allocation within tissues affected by nutrient enrichment (N addition)? And (2) does long‐term N addition alter how plants allocate C under drought?To address these questions, we conducted a greenhouse experiment using the widespread perennial C3 grass, Leymus chinensis, under four treatments: 'Watered', 'Dry', 'Watered + N' and 'Dry + N'. 13CO2 pulse labelling was used to trace C transport through the plant–soil system.We found that drought and N addition resulted in additive effects on C allocation. Greater above‐ground biomass under N addition resulted in higher C loss via above‐ground plant respiration, even under drought, which plays a more important role in the adjustment of root:shoot ratio than does the trade‐off between above‐ and below‐ground organs.Compared to the concept of active phenotype adjustment for maximized growth rate in traditional optimal partitioning theory, our results imply that pre‐drought allometry, which changes under long‐term resource addition, also determines how plants respond to drought and their adaptability to changing environmental conditions. Read the free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]