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  1. Abstract

    Grassland ecosystems globally are being negatively impacted by changes in climate, disturbance regimes, nutrient flux, and consumer guilds. Changes in the trophic ecology of consumers can substantially influence local resources, contributing to shifting diversity, community turnover, and other processes of ecosystem change. Small mammals are diverse and abundant within grasslands and yet the impact of changing ecosystems on small mammals and the role of these mammals as consumers are still both under‐studied. We assessed small mammal resource use within grassland and woodland vegetation types that have resulted from landscape‐scale experimental disturbance through fire treatments within the tallgrass prairie ecoregion of the North American Great Plains. We predicted that resource use would vary significantly among grassland vs. woodland communities, in turn reducing the role of small mammals in contributing to future maintenance of native prairies. We sampled five dominant species of rodents across three years and multiple habitats. Using stable isotope analysis, we investigated isotopic niche area and overlap to infer variation in diet, both within and among species. Resource use shifted in bivariate isotopic space seasonally but not across years when combining all species and habitats. Inferred spring diet (based on fur samples) was highly diverse and overlapping. Summer isotopic values (based on liver tissue) in woody habitat treatments were narrower and overlapped less than within grassland habitats. Consumers generally shifted from C4herbivory to C3herbivory, or greater omnivory, when analyzing grassland, shrubland, and woodland habitats respectively. Within the tallgrass prairie ecosystem, small mammal populations in herbaceous‐dominated habitats use a broader variety of resources than small mammals in proximate woody‐dominated habitats. As native grasslands experience woody encroachment, small mammal assemblages experience turnover of dominant species and associated changes in diet. Ecosystem changes such as cessation of frequent fire resulting in more woody habitats may include reduced roles by native small mammals as consumers/dispersers/propagators of native grassland plants.

     
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  2. Abstract

    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.

     
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  3. Abstract

    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 onCornus drummondiiC.A. Mey, a resprouting clonal native shrub species. Browsing reducedC. drummondiicanopy 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.

     
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  4. Abstract

    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 libitumprotein 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.

     
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  5. Abstract

    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.

     
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