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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]

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]

In highly disturbed environments, clonality facilitates plant survival via resprouting after disturbance, resource sharing among interconnected stems and vegetative reproduction. These traits likely contribute to the encroachment of deep-rooted clonal shrubs in tallgrass prairie. Clonal shrubs have access to deep soil water and are typically thought of as relatively insensitive to environmental variability. However, how leaf physiological traits differ among stems within individual clonal shrubs (hereafter 'intra-clonal') in response to extreme environmental variation (i.e. drought or fire) is unclear. Accounting for intra-clonal differences among stems in response to disturbance is needed to more accurately parameterize models that predict the effects of shrub encroachment on ecosystem processes. We assessed intra-clonal leaf-level physiology of the most dominant encroaching shrub in Kansas tallgrass prairie, Cornus drummondii , in response to precipitation and fire. We compared leaf gas exchange rates from the periphery to centre within shrub clones during a wet (2015) and extremely dry (2018) year. We also compared leaf physiology between recently burned shrubs (resprouts) with unburned shrubs in 2018. Resprouts had higher gas exchange rates and leaf nitrogen content than unburned shrubs, suggesting increased rates of carbon gain can contribute to recovery after fire. In areas recently burned, resprouts had higher gas exchange rates in the centre of the shrub than the periphery. In unburned areas, leaf physiology remained constant across the growing season within clonal shrubs (2015 and 2018). Results suggest single measurements within a shrub are likely sufficient to parameterize models to understand the effects of shrub encroachment on ecosystem carbon and water cycles, but model parameterization may require additional complexity in the context of fire. [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]

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]

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]

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]

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]

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]

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]