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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]

• 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]

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

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]

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]

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]

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]

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]

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]