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Coverage of living (green) vegetation influences rangeland processes and biodiversity but remains a challenge to quantify at small spatial grain. We describe a technique for rapid airborne (unmanned aerial vehicle) measurements of continuous spatial coverage of living vegetation at a resolution (spatial grain) of 8 cm ground sampling distance. We then applied this technique at the pasture (paddock) scale (tens of hectares) in two contrasting grassland ecosystems (semiarid shortgrass steppe in northeastern Colorado, United States and mesic grassland in central Texas, United States) to determine effects of locally relevant grazing treatments on spatial variability and dependence (pattern) in fractional coverage of green vegetation (fractional vegetation cover; FVC). Site-specific regression models developed using reflectance in visible and near-infrared wavebands explained 90% and 89% of variance in FVC for semiarid and mesic grassland, respectively. Mean FVC was similar among shortgrass steppe pastures differing in grazing treatment (light or heavy grazing by cattle, moderate cattle grazing with prairie dogs present). In contrast, FVC was lower with rotational compared with continuous, year-long grazing in the mesic grassland. Heavy grazing in shortgrass steppe and rotational grazing in mesic grassland increased the spatial uniformity in FVC by reducing spatial variability and increasing spatial dependence in FVC, the latter by increasing the similarity in FVC values among spatially separated patches. Unmanned aerial vehicle−enabled remote sensing provides for FVC at sufficiently small spatial grain to characterize spatial variation in FVC at the pasture scale. Results can be used to evaluate the effectiveness of management actions intended to alter spatial variation in FVC to achieve conservation or diversity objectives. Enhanced capacity to monitor FVC at small spatial grain promotes adaptive management of grasslands.

State-and-transition models (STMs) are tools used in rangeland management to describe linear and nonlinear vegetation dynamics as conceptual models. STMs can be improved by including additional ecosystem services, such as wildlife habitat, so that managers can predict how local populations might respond to state changes and to illustrate the tradeoffs in managing for different ecosystem services. Our objective was to incorporate songbird density into an STM developed for sagebrush rangelands in northwest Colorado to guide local management of sagebrush birds. The STM included two shrub-dominated community phases, a native grassland state, and a shrubland and grassland phase within an exotic-dominated state. We surveyed plots for songbirds, collected a suite of vegetation indicators at each plot, and quantified songbird habitat relationships with count-based regression models. We then used the estimated models to predict songbird density based on average vegetation conditions per state or community phase. Moderate or increasing shrub cover were important predictors for shrubland-associated species, and responses to understory components varied by species. In the STM, we predicted higher densities of shrubland-associated bird species in the shrub-dominated phases and higher densities of grassland-associated bird species in the state and phase lacking shrub cover. No single state or phase captured the highest density for all songbirds, illustrating the value of alternative states. Our results also demonstrate the utility of displaying traditional wildlife count models against the range of vegetation conditions associated with each state or phase to understand how wildlife density can vary within states and phases. Our approach can assist land managers to gauge the potential impacts of land-use decisions and natural vegetation variability on wildlife, especially for species of conservation concern.

Frequent, severe defoliation reduces grass production and can alter plant species composition in grasslands. Multipaddock rotational grazing has been proposed as a grazing strategy that may reduce the frequency and intensity of defoliation on palatable grass plants without altering stocking rates. Previous studies evaluated this hypothesis using small, homogeneous paddocks and nonadaptive rotation schedules and found small and inconsistent differences between continuous and rotational grazing systems. Using a stakeholder-driven collaborative adaptive management (CAM) framework, we conducted the first ranch-scale experimental investigation into tiller defoliation patterns in the context of adaptive multipaddock rotational grazing. We monitored tiller defoliation frequency and intensity in 10 paired 130-ha pastures assigned to either a collaborative adaptive multipaddock rotational grazing treatment (CARM, one livestock herd) or a season-long continuous grazing treatment (traditional rangeland management [TRM]; 10 separate herds) in shortgrass steppe. Consistent with previous studies, we observed that frequencies of grazing and regrazing on a palatable, cool-season grass (western wheatgrass, Pascopyrum smithii) were much more sensitive to stocking rate than grazing system. Under moderate stocking rates used in both CARM and TRM treatments, roughly two-thirds of western wheatgrass tillers remained ungrazed annually, regardless of grazing system. Thus, season-long rest is present in season-long continuous and rotational grazing systems. Frequencies of tiller regrazing were low (5−15%) and similar between CARM and TRM treatments. Although defoliation patterns were similar between treatments at the whole-ranch scale, CARM enhanced spatial and temporal heterogeneity in defoliation frequencies among individual pastures. Pastures grazed earlier in the season or for longer experienced more defoliation. Managers implementing adaptive, multipaddock rotational grazing could use this heightened and predictable variability to strategically manage impacts of grazing on western wheatgrass at the individual pasture scale. The CAM model enabled our team to identify and directly address key stakeholder hypotheses and resulted in coproduction of management-relevant research.

Fauna of North America’s Great Plains evolved strategies to contend with the region’s extreme spatiotemporal variability in weather and low annual primary productivity. The capacity for large-scale movement (migration and/or nomadism) enables many species, from bison to lark buntings, to track pulses of productivity at broad spatial scales (> 1 000 km2). Furthermore, even sedentary species often rely on metapopulation dynamics over extensive landscapes for long-term population viability. The current complex pattern of land ownership and use of Great Plains grasslands challenges native species conservation. Approaches to managing both public and private grasslands, frequently focused at the scale of individual pastures or ranches, limit opportunities to conserve landscape-scale processes such as fire, animal movement, and metapopulation dynamics. Using the US National Land Cover Database and Cropland Data Layers for 2011−2017, we analyzed land cover patterns for 12 historical grassland and savanna communities (regions) within the US Great Plains. On the basis of the results of these analyses, we highlight the critical contribution of restored grasslands to the future conservation of Great Plains biodiversity, such as those enrolled in the Conservation Reserve Program. Managing disturbance regimes at larger spatial scales will require acknowledging that, where native large herbivores are absent, domestic livestock grazing can function as a central component of Great Plains disturbance regimes if they are able move at large spatial scales and coexist with a diverse array of native flora and fauna. Opportunities to increase the scale of grassland management include 1) spatial prioritization of grassland restoration and reintroduction of grazing and fire, 2) finding creative approaches to increase the spatial scale at which fire and grazing can be applied to address watershed to landscape-scale objectives, and 3) developing partnerships among government agencies, landowners, businesses, and conservation organizations that enhance cross-jurisdiction management and address biodiversity conservation in grassland landscapes, rather than pastures.

Collaborative adaptive management (CAM) is hypothesized to benefit management of rangeland ecosystems, but the presumed benefits have seldom been quantified, and never in a multipaddock rotational grazing system. Here, we evaluated average daily weight gain (ADG) of livestock (kg steer−1 d−1) in four grazing management treatments during the summers of 2016−2018 in a semiarid shortgrass steppe. These four treatments had the same stocking rate but differed in stocking densities. The three lowest stocking densities were implemented using nonadaptive grazing management, while the highest stocking density was implemented using CAM by an 11-member Stakeholder Group. Three of the four treatments used multipaddock rotational grazing. Growing season precipitation varied from drought in 2016 to near average in 2017 and dry in 2018. During nondrought years, ADG under nonadaptive grazing declined linearly as stocking density increased from low to high. This relationship was not significant during drought (2016). CAM increased absolute livestock production by 0.13 to 0.19 kg steer−1 d−1 in nondrought years, or a 23−25% relative increase in ADG. This benefit of CAM arose from the Stakeholder Group's ability to rotate cattle in response to spatiotemporal heterogeneity across the landscape—i.e., the ability to graze the “right pastures at the right time.” Multiplying the additional grazing season livestock gains achieved through CAM by the monetary value of gains ($ kg−1) resulted in an estimated additional gross revenue return from CAM of $48.16 to $55.54 per steer annually, as compared with revenues from nonadaptive multipaddock rotational grazing under nondrought conditions. These results indicate that CAM, supported with substantial and timely monitoring data, can minimize decreases in livestock production associated with high stocking densities used in multipaddock rotation systems. However, in this experimental context, the economic benefits of increased livestock production associated with CAM were likely insufficient to offset the substantial cost of this approach.

Livestock distribution in extensive rangelands of North America can present management challenges to land managers. Understanding the role of topography on livestock distribution, within and across diverse rangeland ecosystems, could provide land managers valuable information for adaptive management of livestock to address both conservation and production goals from these ecosystems. Here, we examine the influence of topography on grazing distribution unevenness and intensity of use of beef cattle in seven rangeland ecosystems spanning arid, semiarid, and subtropical environments. We focused on grazing distribution during the late growing season (summer and autumn) periods when topographic variation in rangelands is more coupled to low and nonuniform availability of high-quality forage. Pasture size and water sources strongly influence grazing distribution across ecosystems. High unevenness of grazing occurred in pastures with extensive distances to water, low stock density, and more rugged topography. Conversely, more uniform grazing distribution occurred in smaller, well-watered pastures that support higher stock density (animals per unit area) and gentler terrain. Comparison of two topographic indices, topographic wetness index and topographic position class index, in terms of their ability to predict cattle grazing distribution, revealed that categorical topographic position classes were more effective. For most arid and semiarid rangelands, livestock grazing distributions showed affinities for lowlands and flat plains compared with open slopes and uplands. In contrast to drier rangelands, livestock grazing distributions exhibited preference for upland and sloped areas of subtropical pastures, as low-lying areas with water-inundation likely curtailed selection. Across these diverse rangeland ecosystems of North America, results provide benchmark information on livestock grazing distribution to formulate improvements in adaptive management and decision making and incorporate technological advancements in precision livestock management to integrate abiotic environmental information with spatial movements of livestock and temporal vegetation dynamics.

Grassland birds have experienced some of the steepest population declines of any guild of birds in North America. The shortgrass steppe contains some of North America's most intact grasslands, which makes the region particularly important for these species. It is well known that grassland birds differentially respond to variation in vegetation structure generated by spatiotemporally varying disturbance like grazing management. However, understanding how species respond to characteristics beyond vegetation structure or grazing could better inform management for these species in the shortgrass steppe. We analyzed point count data for 5 grassland bird species breeding on the Central Plains Experimental Range in northeastern Colorado from 2013 to 2017 to examine the predictive capacity of models representing fine-scale (∼5 ha) vegetation attributes (vegetation structure and cover type) and topography, combined with interannual precipitation variability (i.e. vegetation-abiotic models). We then compared these models to models based on grazing management treatments (applied to whole pastures, ∼130 ha) and edaphic conditions (ecological sites), which represented information more generally available to rangeland managers. Precipitation, vegetation structure, and vegetation cover type influenced all species in a manner consistent with, but more nuanced than, vegetation structure alone. These models also explained more variation in abundance for species that responded to grazing management. Thus, while grazing management can be applied adaptively to improve habitat for these species, our more detailed vegetation-abiotic models identified species-specific habitat components that could be targeted for management. For example, not grazing pastures with extensive, homogenous stands of mid-height grasses (e.g., Hesperostipa comata) for an entire growing season during wet years could be one strategy to enhance Grasshopper Sparrow (Ammodramus savannarum) abundance and stockpile residual forage for future utilization by livestock. Our models provide a better understanding of and reveal nuances in the suite of environmental conditions to which grassland birds respond in shortgrass steppe rangelands. LAY SUMMARY We lack clear understanding of which environmental characteristics might support declining bird species breeding in the shortgrass steppe of the United States. These bird species often breed on lands managed for cattle grazing and are known to respond to vegetation structure generated by grazing and/or soil conditions. We sought to evaluate how precipitation and different types of vegetation cover (e.g., shrubs, shortgrasses) could explain additional variation in bird abundance in this system. We found vegetation cover, vegetation structure, and precipitation could explain more variation in abundance than cattle grazing management and/or ecological sites (i.e. unique soil types with associated plant communities) for most bird species. For bird species that did not respond to grazing management, ecological site and year effects explained more variation in abundance than vegetation characteristics and precipitation. Precipitation conditions, types of vegetation cover, and ecological sites often are not considered when developing grazing management plans to support grassland birds; incorporating these factors into management plans may help support populations of these declining species.

On the Ground • We assessed diet quality and livestock weight gains for shortgrass steppe pastures dominated by Loamy Plains or Sandy Plains ecological sites. • When growing season precipitation is “normal,” livestock gains are higher on Sandy Plains ecological sites, and diet quality is not limiting livestock production. • Conversely, when growing season precipitation declines by ≥ 20%, digestible organic matter, but not crude protein, influences livestock gains. These negative effects on livestock gains are more pronounced for the Loamy Plains ecological site. • Pastures with multiple ecological sites may provide range managers greater forage diversity for livestock and higher livestock gains during dry growing seasons.

Aboveground net primary productivity (ANPP) in grasslands is an important integrator of terrestrial ecosystem function, a key driver of global biogeochemical cycles, and a critical source of food for wild and domesticated herbivores. ANPP exhibits high spatial and temporal variability, driven by a suite of factors including precipitation amount and pattern, biotic and abiotic legacies, and topographic heterogeneity. Global climate models forecast an altered hydrological cycle due to climate change, including higher precipitation variability and more extreme events, which may further increase spatiotemporal variability in ANPP. Therefore, it is essential to understand the sensitivity of this central ecosystem function to various precipitation metrics, legacies, and topographic positions to better inform sustainable grassland management. In this study, we analyzed long‐term (36‐yr) ANPP data collected across a topographic sequence in the semiarid shortgrass steppe of North America to examine patterns and drivers of spatiotemporal variability in ANPP. We observed that (1) ANPP varied substantially by topographic position, with greater divergence during years with high production, (2) ANPP variability was higher temporally (16‐fold maximum difference across years) than spatially (4‐fold maximum difference across topographic positions), (3) warm‐season perennial grasses were the dominant plant functional type across all topographic positions and strongly influenced total ANPP dynamics, and (4) ANPP had strong sensitivities to current year precipitation amount and pattern that varied by plant functional type, as well as weaker sensitivities to precipitation and productivity legacies. Overall, the lowest topographic position had the highest sensitivity to precipitation, likely due to higher resource availability via the downhill movement of water and nutrients during years with high precipitation and large rainfall events. These results suggest that temporal and spatial ANPP variability in shortgrass steppe is primarily driven by the combined effects of precipitation amount and pattern during the current year, with the dominant warm‐season perennial grasses governing these responses.

Habitat loss and altered disturbance regimes have led to declines in many species of grassland and sagebrush birds, including the imperiled Mountain Plover (Charadrius montanus). In certain parts of their range Mountain Plovers rely almost exclusively on black-tailed prairie dog (Cynomys ludovicianus) colonies as nesting habitat. Previous studies have examined Mountain Plover nest and brood survival on prairie dog colonies, but little is known about how colony size and shape influence these vital rates or patterns of habitat selection. We examined how (1) adult habitat utilization, (2) nest-site selection, and (3) nest success responded to a suite of local- and site-level variables on large prairie dog colony complexes in northeastern Wyoming. Abundance of adult Mountain Plovers was highest on points within older, “medium”-sized (100–500 ha) colonies with high cover of annual forbs and bare ground (5.8 birds km−2), but lower on extremely large (>2,000 ha) colonies (2.1 birds km−2). Nest sites were characterized by high proportions of annual forbs and bare ground and low cactus cover and vegetation height. Nest survival was higher for older nests, and nests with lower cactus cover, and decreased with increasing temperatures. Uncertainty was high for models of daily nest survival, potentially because of 2 competing sources of nest failure: nest depredation and nest abandonment or inviability of eggs. Drivers of these 2 sources of nest failure differed, with inclement weather and higher temperatures associated with nest abandonment or egg inviability. We highlight how prairie dogs alter vegetation structure and bare ground heterogeneously across the landscape, and how this in turn influences bird abundance and nest distribution at different temporal and spatial scales. Furthermore, our work reveals how partitioning the causes of nest failure during nest survival analyses enhances understanding of survival rate covariates.

Considerable uncertainty remains regarding grazing-induced influences on soil organic carbon (SOC) sequestration in semiarid grassland ecosystems due to three important complications associated with studying such effects: (1) Ecologically meaningful shifts in SOC pools attributable to grazing are difficult to detect relative to inherently large grassland SOC pools, (2) a lack of baseline (pre-treatment) data, and (3) frequent lack of or limited replication of long-term grazing manipulations. SOC sequestration rates were determined in 74-year-old grazing exclosures and paired moderately grazed sites, established across a soil texture gradient, in the western North American shortgrass steppe in northeastern Colorado. We sampled soils (0–20 cm) from 12 exclosures and paired grazed sites to measure SOC concentration and soil radiocarbon ∆14C (‰); the latter allowed us to determine turnover of the SOC pool over a 7-decade period in the presence versus the absence of grazing. Removal of grazing for more than 7 decades substantially altered plant community composition but did not affect total soil C, SOC, soil ∆14C, SOC turnover rate, or total soil N. Grazing effect also did not interact with soil texture to influence any of those soil properties. Soil texture (silt + clay content) did influence total soil C and SOC, and total soil N, but not ∆14C or SOC turnover. Results provide evidence that long-term removal of grazing from semiarid grassland ecosystems in the western North American Great Plains does not enhance long-term SOC sequestration, despite changes in the relative dominance of C3 versus C4 grasses.

Semiarid rangelands experience substantial interannual variability in precipitation, which can determine the relative abundance of species in any given year and influence the way that fire affects plant community composition and productivity. Long-term studies are needed to examine potential interactions between fluctuating community composition and the role of fire in these ecosystems. Here, we report on an 11-year (2006 to 2016) study of annual and triennial dormant-season prescribed fires in the semiarid shortgrass steppe of Colorado, USA. Productivity of the dominant C4 shortgrasses was not reduced by dormant-season burns in any year. The C3 annual grass, six-weeks fescue (Vulpia octoflora [Walter] Rydb.) was rare during the first 7 years (2006 to 2012) but, following drought in 2012, increased dramatically in unburned plots (2013 to 2016). Both spring and autumn annual burns reduced fescue biomass during 2014 to 2016 by an average of 87%. Autumn triennial burns prior to the 2013 and 2016 growing seasons similarly reduced fescue (86%), while spring triennial burns implemented prior to the 2012 and 2015 growing seasons did not. Results indicate that burning during fescue establishment can prevent proliferation, but burning two years later when fescue had reached peak abundance was ineffective. All three burn treatments that suppressed fescue subsequently enhanced C4 grass production. We suggest that rangeland managers be aware of the potential for sixweeks fescue germination and establishment during warm, wet winters that follow drought years, and consider the use of dormant-season prescribed fire to adaptively reduce negative impacts on forage production.

Advances in on-animal sensor technologies to monitor location and activity have enhanced the ability to study foraging decisions of free-ranging herbivores. Sensors monitoring jaw movements that quantify ingestive behaviours, such as the RumiWatch (RW) noseband sensor system, have primarily been used in indoor animal housing systems or structurally homogeneous, small pasture (paddock) environments. Continuously monitoring these ingestive behaviours in extensive and heterogeneous rangelands has not been previously conducted. We evaluated the accuracy of the RW noseband sensor system for two grazing seasons in 130-ha pastures (paddocks) composed of native, mixed-species plant communities in a semiarid environment. The noseband sensor was used to compare ingestive behavior at different sites and seasons characterized by varying sward complexity, stocking rate, and levels of forage limitation. We evaluated the noseband sensor against direct visual observations of yearling steers grazing with two different validation studies. First, the time duration of grazing recorded by the sensor was compared to direct visual observation data (Validation Study 1). A high correlation (rs = 0.95) for hourly grazing time resulted between the RW system and visual observations. Second, we examined the ability of the RW system to measure prehension bite rates in distinct plant communities varying in height and leaf angle (Validation Study 2). The accordance between direct observation and measurement by the RW system for bite rate improved from 2019 (Concordance Correlation Coefficient (CCC) = 0.71) to 2020 (CCC = 0.80) after modifications to improve the fit of the halter supporting the noseband sensor. Correlations between the sensor and visual observations increased by ∼17 % with this modification for grazing bouts in mixed-species and midgrass-dominated swards; correlations remained ∼10 % lower in shortgrass-dominated swards. Our results show that the RW system is an effective tool for monitoring free-ranging cattle grazing activity and quantifying bite rates in a heterogeneous rangeland ecosystem. Bite rate measurements are more accurate in swards with vertically oriented stems and leaves compared to lawn-like prostrate swards. Grazing bout length and rumination chew rate may represent behavioral indicators employed in managing animal performance in semiarid rangelands. Our validation study and experimental investigation indicate that the RW noseband sensor is a useful animal-borne sensor technology for research demanding sustained ingestion measurements across mixed-species forage communities.

As grassland birds of central North America experience steep population declines with changes in land use, management of remaining tracts becomes increasingly important for population viability. The integrated use of fire and grazing may enhance vegetation heterogeneity and diversity in breeding birds, but the subsequent effects on reproduction are unknown. We examined the influence of patch-burn grazing management in shortgrass steppe in eastern Colorado on habitat use and reproductive success of 3 grassland bird species, horned lark (Eremophila alpestris), lark bunting (Calamospiza melanocorys), and McCown's longspur (Rhynchophanes mccownii), at several spatial scales during 2011 and 2012. Although no simple direct relationship to patch-burn grazing treatment existed, habitat selection depended on precipitation- and management-induced vegetation conditions and spatial scale. All species selected taller-than-expected vegetation at the nest site, whereas at the territory scale, horned larks and McCown's longspurs selected areas with low vegetation height and sparse cover of tall plants (taller than the dominant shortgrasses). Buntings nested primarily in unburned grassland under average rainfall. Larks and longspurs shifted activity from patch burns during average precipitation (2011) to unburned pastures during drought (2012). Daily survival rate (DSR) of nests varied with time in season, species, weather, and vegetation structure. Daily survival rate of McCown's longspur nests did not vary with foliar cover of relatively tall vegetation at the nest under average precipitation but declined with increasing cover during drought. At the 200-m scale, increasing cover of shortgrasses, rather than taller plant species, improved DSR of larks and longspurs. These birds experience tradeoffs in the selection of habitat at different spatial scales: tall structure at nests may reduce visual detection by predators and provide protection from sun, wind, and rain, yet taller structure surrounding territories may host nest predators. Patch-burn grazing management in combination with other strategies that retain taller-structured vegetation may help sustain a diversity of breeding habitats for shortgrass birds under varying weather conditions.Published 2017. This article is a U.S. Government work and is in the public domain in the USA.

Plant community responses to livestock grazing lack conformity across studies, even those conducted within similar ecosystems. Variability in outcomes can often be traced back to short-term or mid-term weather patterns, differences in grazing timing or intensity, or interactive effects of management and weather. Long-term experimental data are needed to determine how grazing intensity affects plant community composition in semiarid ecosystems where precipitation is low and highly variable. However, long-term grazing intensity experiments, particularly experiments with more than two grazing intensity treatment levels, are quite rare. We capitalized on one of the longest-term grazing studies, with 75 yr of sustained stocking rate treatments (none, light, moderate, and heavy), to identify long-term effects of livestock grazing on plant community composition in shortgrass steppe. Plant community composition was similar between moderately and heavily grazed pastures after 75 yr of continuous, season-long (May to October) grazing treatments, and heavy grazing did not extirpate cool-season perennial graminoids. These findings support the long-term sustainability of livestock grazing in the shortgrass steppe, which has high resistance to season-long heavy grazing. Conversely, ungrazed and lightly grazed pastures experienced relatively large shifts in plant community composition, especially in the past 25 yr. Light or no grazing was associated with increased abundance of cool-season perennial graminoids, as well as several weedy and invasive species. Moreover, across most grazing treatments, several aspects of plant community composition have been shifting directionally during the past 25 yr, which recent experiments in this grassland suggest may be a response to increasing atmospheric (CO2). The shortgrass steppe is not only tolerant of fairly high grazing intensities but also likely requires some level of grazing to resist invasion by weedy annuals and to maintain cover of blue grama, a highly drought-tolerant species.

Small-bodied terrestrial animals such as songbirds (Order Passeriformes) are especially vulnerable to hail-induced mortality; yet, hail events are challenging to predict, and they often occur in locations where populations are not being studied. Focusing on nesting grassland songbirds, we demonstrate a novel approach to estimate hail-induced mortality. We quantify the relationship between the probability of nests destroyed by hail and measured Level-III Next Generation Radar (NEXRAD) data, including atmospheric base reflectivity, maximum estimated size of hail and maximum estimated azimuthal wind shear. On 22 June 2014, a hailstorm in northern Colorado destroyed 102 out of 203 known nests within our research site. Lark bunting (Calamospiza melanocorys) nests comprised most of the sample (n = 186). Destroyed nests were more likely to be found in areas of higher storm intensity, and distributions of NEXRAD variables differed between failed and surviving nests. For 133 ground nests where nest-site vegetation was measured, we examined the ameliorative influence of woody vegetation, nest cover and vegetation density by comparing results for 13 different logistic regression models incorporating the independent and additive effects of weather and vegetation variables. The most parsimonious model used only the interactive effect of hail size and wind shear to predict the probability of nest survival, and the data provided no support for any of the models without this predictor. We conclude that vegetation structure may not mitigate mortality from severe hailstorms and that weather radar products can be used remotely to estimate potential for hail mortality of nesting grassland birds. These insights will improve the efficacy of grassland bird population models under predicted climate change scenarios.

Questions How does long-term exclusion of cattle grazing influence plant community composition in a semiarid grassland? Can spatial variation in the effects of cattle grazing exclusion be explained by variation in soil texture? Location The shortgrass steppe of northeastern Colorado, USA, located in the North American Great Plains. Method We used 16 long-term (72 yr) cattle exclosures to examine the effects of grazers on plant communities, and evaluate whether grazer effects interact with soil texture. Results Although shortgrass steppe communities are relatively unaffected by grazing in the short-term (1-2 decades), exclusion of cattle grazing for seven decades caused a compositional shift from dominance by a C4 shortgrass (Bouteloua gracilis) to co-dominance by a C3 midgrass (Pascopyrum smithii) and B. gracilis. The strength of this shift was highly variable across sites. Soil texture was correlated with the abundance of certain plant species, but did not explain variation in the magnitude of grazer effects. Grazed communities contained perennial and annual growth forms with a diversity of strategies to coexist with grazers and B. gracilis. Ungrazed communities included increased abundance of annual, ruderal forbs and three woody plant species. Grazing effects occurred against a backdrop of changing plant communities: during the past seven decades, C3 perennial graminoids and subshrubs have increased in relative abundance in both grazed and ungrazed communities. Conclusions Our long-term experiment shows that community responses to grazing in this semiarid grassland occur very slowly, but are predictable, with C4 shortgrasses eventually giving way to taller C3 grasses and ruderal forbs. Spatial variation in grazing effects across sites (and lack of a relationship with soil texture) may reflect the importance of fine-scale heterogeneity in C3 grass abundance, and the slow rate at which taller C3 grasses can coalesce into monodominant patches that outcompete C4 shortgrasses. Increased abundance of C3 species over the past seven decades, both in the presence and absence of grazing, may be related to recovery from the severe drought and dust storms of the 1930s as well as enhanced growth of C3 plants under increasing atmospheric [CO2]. This article is protected by copyright. All rights reserved.

Rangelands are temporally and spatially complex socioecological systems on which the predominant land use is livestock production. In North America, rangelands also contain approximately 80% of remaining habitat for grassland birds, a guild of species that has experienced precipitous declines since the 1970s. While livestock grazing management may benefit certain grassland bird species by generating the vegetation structure and density they prefer, these outcomes are poorly understood for avian species breeding in the shortgrass steppe. We evaluated how two grazing management systems, continuous, season-long grazing and adaptive, rest-rotational grazing, affected grassland bird abundance from 2013 to 2017 in Colorado's shortgrass steppe. We examined grazing impacts in conjunction with ecological sites, which constitute unique soil and plant communities. When grazing management was evaluated in conjunction with spatial variation in ecological sites, we found three of our five focal bird species responded to grazing management. McCown's Longspur abundance decreased in pastures rested from grazing the previous year. The effect of grazing on Horned Lark and Grasshopper Sparrow depended on ecological site: Horned Lark density was highest in pastures that were intensively grazed and Grasshopper Sparrow density was highest in pastures that were rested the previous year in the least productive ecological site. In addition, densities of all species varied across ecological sites. Our results suggest consideration of soil and vegetation characteristics can inform how adaptive management is applied on a landscape to benefit the full suite of breeding grassland birds, including species that have seemingly contrasting habitat needs. For example, a manager could target adaptive drought mitigation practices, such as resting pastures for 1 yr to generate grassbanks, in less productive soils to benefit grassland birds that prefer taller/denser vegetation structure, or could apply intensive, short-duration grazing on less productive soils to benefit species preferring shorter/sparser vegetation. A single year of intensive, short-duration grazing (i.e., one component of our rotational treatment) across the landscape, however, might not create sufficient habitat for species that prefer short/sparse vegetation in our system (e.g., McCown's Longspur). Ultimately, our study indicates how cattle production on rangelands can congruently support grassland bird populations in the shortgrass steppe.

Rangeland ecosystems worldwide are characterized by a high degree of uncertainty in precipitation, both within and across years. Such uncertainty creates challenges for livestock managers seeking to match herbivore numbers with forage availability to prevent vegetation degradation and optimize livestock production. Here, we assess variation in annual large herbivore production (LHP, kg/ha) across multiple herbivore densities over a 78-yr period (1940-2018) in a semiarid rangeland ecosystem (shortgrass steppe of eastern Colorado, USA) that has experienced several phase changes in global-level sea surface temperature (SST) anomalies, as measured by the Pacific Decadal Oscillation (PDO) and the El Nino-Southern Oscillation (ENSO). We examined the influence of prevailing PDO phase, magnitude of late winter (February-April) ENSO, prior growing-season precipitation (prior April to prior September) and precipitation during the six months (prior October to current April) preceding the growing season on LHP. All of these are known prior to the start of the growing season in the shortgrass steppe and could potentially be used by livestock managers to adjust herbivore densities. Annual LHP was greater during warm PDO irrespective of herbivore density, while variance in LHP increased by 69% (moderate density) and 91% (high density) under cold-phase compared to warm-phase PDO. No differences in LHP attributed to PDO phase were observed with low herbivore density. ENSO effects on LHP, specifically La Nina, were more pronounced during cold-phase PDO years. High herbivore density increased LHP at a greater rate than at moderate and low densities with increasing fall and winter precipitation. Differential gain, a weighted measure of LHP under higher relative to lower herbivore densities, was sensitive to prevailing PDO phase, ENSO magnitude, and precipitation amounts from the prior growing season and current fall-winter season. Temporal hierarchical approaches using PDO, ENSO, and local-scale precipitation can enhance decision-making for flexible herbivore densities. Herbivore densities could be increased above recommended levels with lowered risk of negative returns for managers during warm-phase PDO to result in greater LHP and less variability. Conversely, during cold-phase PDO, managers should be cognizant of the additional influences of ENSO and prior fall-winter precipitation, which can help predict when to reduce herbivore densities and minimize risk of forage shortages.

Context Understanding how grazing management decisions influence the productivity and composition of rangeland plant communities is essential for the development of effective strategies to sustainably produce multiple ecosystem goods and services. Informed with experimental measurements, simulation models can advance our understanding and stewardship of rangeland ecosystems. Objective Our main objective was to evaluate the APEX (Agricultural Policy/Environmental eXtender) plant growth modules and grazing animal selectivity in simulating forage production using experimental data collected from both traditional season-long grazing and adaptive rotational grazing management on western rangelands. Specifically, we evaluated APEX's capability to simulate forage productivity and its response to soil types and climate conditions under grazing management options. Methods Capitalizing on a comparative field study with 20 large pastures (> 123 ha each), APEX modifications were evaluated by comparing simulated forage production with experimental data. The field study evaluated traditional grazing (season long grazing on a single pasture) and an alternative grazing system that utilized collaborative adaptive rangeland management with stakeholders engaged in decision making (such as when and where to rotate a single herd). APEX was modified to include rotational grazing based on a user-defined sequence and automatic rotational grazing based on user-defined forage grazing limits and minimum/maximum grazing durations. Results and conclusions The APEX model was able to simulate the relative differences in forage production between grazing treatments, across years, and among soil types; however, APEX underestimated forage production in 2015 and 2017 due to overestimating drought stress for the warm season perennial grass functional group. Simulation of grazing management scenarios showed that the collaborative adaptive management decision criteria resulted in grazing durations that produced more forage than consistent 7- or 14-day rotation intervals. Significance These modifications were needed to capture the complexity of semiarid environments and thus enhance APEX to better assess grazing management decisions on forage production in regions such as the Western US Great Plains.

Knowledge of climatic and management influences on large herbivore production (LHP, kg ha−1) is needed for low productivity, semiarid grasslands to address potential consequences of both increasing climate variability and the need to increase animal protein for human consumption. Here, we evaluate the influence of climatic variability and herbivore density on LHP in semiarid grassland using a unique long-term (80 years: 1939–2018) grazing study with three grazing intensities based on forage utilization (light, moderate and heavy). Seasonal variation in precipitation, but not temperature, was the primary influence on LHP. Winter (October-March) and spring (April-June), but not summer (July-September), precipitation during the current year positively influenced LHP across the 3 grazing intensities, whereas prior growing season (prior April-September) precipitation was consistently a negative influence. Although spring precipitation was the most influential seasonal weather variable for LHP, the effect of winter precipitation closely followed under all three grazing intensities, suggesting that non-growing season precipitation is essential for soil water storage to initiate production of sufficient high-quality forage in the subsequent grazing season, resulting in a positive feedback on LHP. A key finding from our analysis was that the effect of summer precipitation is smaller than the combined effects of winter and spring precipitation. As such, much of the variation in LHP can be predicted by seasonal weather parameters that are known early in the growing season. The magnitude of seasonal precipitation effects on LHP was greatest for heavy grazing; consequently LHP with heavy grazing is more reliant on primary production produced in the current year to increase LHP as forage quantity is more limiting than forage quality. Moreover, stability of LHP across years (range: 7.5 to 34.6 kg ha−1) was less with heavy grazing, which results in “boom-bust” economics that threaten sustainability of operations. Management adaptations to mitigate climatic variability, therefore, will be most necessary and advantageous when land managers employ heavy grazing intensities. Despite the substantial interannual variability in precipitation that characterizes semiarid grasslands, our results show that proactive flexibility by land managers in adjusting grazing management decisions to seasonal precipitation amounts forecasted for the winter and spring seasons would reduce enterprise risk and improve confidence in decision-making, profitability, production efficiency and environmental sustainability from semiarid grasslands.

Predicting impacts on phenology of the magnitude and seasonal timing of rainfall pulses in water-limited grassland ecosystems concerns ecologists, climate scientists, hydrologists, and a variety of stakeholders. This report describes a simple, effective procedure to emulate the seasonal response of grassland biomass, represented by the satellite-based normalized difference vegetation index NDVI, to daily rainfall. The application is a straightforward adaptation of a staged linear reservoir that simulates the pulse-like entry of rainwater into the soil and its redistribution as soil moisture, the uptake of water by plant roots, short-term biomass development, followed by the subsequent transpiration of water through foliage. The algorithm precludes the need for detailed, site specific information on soil moisture dynamics, plant species, and the local hydroclimate, while providing a direct link between discrete rainfall events and consequential biomass responses throughout the growing season. We applied the algorithm using rainfall data from the Central Plains Experimental Range to predict vegetation growth dynamics in the semi-arid shortgrass steppe of North America. The mean annual rainfall is 342 mm, which is strongly bifurcated into a dominantly ‘wet’ season, where during the three wettest months May, June and July the mean monthly rainfall is approximately 55 mm month−1; and a ‘dry’ season, where during the three driest months December, January and February, the mean monthly rainfall is approximately 7 mm month−1. NDVI data from the Moderate Resolution Imaging Spectroradiometer MODIS MOD13Q1 16 day, 250 m × 250 m product were used as a proxy for grassland phenology for the period-of-record 2000–2013. Allowing for temporal changes in basic parameters of the response function over the growing season, the predicted response of the model tracks the observed NDVI metric with correlation coefficients exceeding 0.92. A two-stage series reservoir is preferred, whereby the characteristic time for transfer of a rainfall event to the peak response of NDVI decreases from 24 days early growing season to 12 days late growing season, while the efficiency of a given volume of rainfall to produce a correspondingly similar amount of aboveground biomass decreases by a factor of 40% from April to October. Behaviours of the characteristic time of greenup and loss of rainfall efficiency with progression of the growing season are consistent with physiological traits of cool-season C3 grasses versus warm-season C4 grasses, and with prior research suggesting that early season production by C3 grasses is more responsive to a given amount of precipitation than mid-summer growth of C4 shortgrasses. Our model explains >90% of seasonal biomass dynamics. We ascribe a systematic underprediction of observed early season greenup following drought years to a lagged or ‘legacy’ effect, as soil inorganic nitrogen, accumulated during drought, becomes available for future plant uptake.

Anthropogenic changes to disturbance regimes in grasslands, and associated homogenization of vegetation structure, have been implicated as factors contributing to declines in populations of grassland birds in North America. We examined the influence of patch-burn grazing management, which employs spatiotemporal interactions between fire and livestock grazing guided by historical disturbance patterns, on vegetation structure and bird abundance in shortgrass steppe in northeastern Colorado, USA. All study pastures were grazed by cattle at moderate stocking rates from May to October each year. In the patch-burn treatment, we burned 25% of each pasture in autumn (Oct or Nov) each year during 2007–2010; control pastures were not burned. Patch-burn grazing management increased vegetation heterogeneity by generating short (

Management strategies that allow for spatiotemporal interactions between fire and herbivores can potentially achieve multiple management goals related to livestock production and wildlife conservation, but little is known about such interactions in semiarid grasslands where fire has traditionally been viewed as having few management applications. We studied patch burn grazing management in the shortgrass steppe of northeastern Colorado, comparing unburned pastures to pastures where 25% of the area was burned in October or November each year over 4 years. Our objective was to examine the interactive effects of patch burns and the subsequent response by pronghorn (Antilocapra americana) on plains pricklypear (Opuntia polyacantha) and wind erosion rates. We monitored abundance of plains pricklypear and wind erosion rates throughout the experiment and quantified seasonal pronghorn densities and postburn damage to plains pricklypear cladodes during the latter 2 years of the study. Pronghorn density was 26 times greater in winter and 7 times greater in spring on patch burns compared with unburned pastures. By late winter, densities of bitten or uprooted plains pricklypear cladodes were five times greater on patch burns compared with unburned pastures. Patch burns, as well as the subsequent response of pronghorn, reduced plains pricklypear density by 54–71% during the first year after the burns, and density remained suppressed for up to 6 years after burns. Wind erosion rates on patch burns were greater compared with unburned pastures but were two orders of magnitude lower than rates measured on fallow croplands in the region. Autumn patch burns can be a valuable means to suppress plains pricklypear and thereby increase grass available for livestock consumption in the shortgrass steppe. These outcomes can be achieved without increasing wind erosion in a manner that threatens long-term soil sustainability and without negative consequences for livestock weight gains.

State-and-transition models for semiarid grasslands in the North American Great Plains suggest that the presence of herbivorous black-tailed prairie dogs (Cynomys ludovicianus) on a site 1) creates a vegetation state characterized by increased dominance of annual forbs and unpalatable bunchgrasses and increased bare soil exposure and 2) requires long-term (> 40 yr) prairie dog removal to transition back to a vegetation state dominated by palatable perennial grasses. Here, we examine 1) how the recent history of prairie dog occupancy on a site (1–10 yr) influences the magnitude of prairie dog effects on vegetation composition and 2) how occupancy history affects vegetation dynamics following extirpation of prairie dogs. We used a natural experiment in the shortgrass steppe of northeastern Colorado, USA, where prairie dogs were extirpated from multiple sites during an outbreak of epizootic plague. On sites occupied by prairie dogs for 1–4 yr prior to extirpation, plant cover and composition recovered to conditions similar to unoccupied sites within a single growing season. Larger reductions in perennial C4 grasses occurred on sites occupied for the prior 7–10 yr compared to sites with shorter occupancy histories (

We studied land-cover associations at nest sites and reproductive success of two Buteo species of conservation concern on the southern Great Plains, USA. The study area was in Cimarron County, Oklahoma, where land use is dominated by row-crop agriculture, livestock grazing, and grasslands enrolled in the Conservation Reserve Program (CRP). Ferruginous hawks (B. regalis) were uncommon and nested primarily in and around the Rita Blanca National Grassland. Swainson's hawks (B. swainsoni) were common and nested throughout the study area. Territories of ferruginous hawks contained more sandsage (Artemisia filifolia) habitat and less cropland and CRP lands than random sites, whereas territories of Swainson's hawks mirrored proportions of available landcover. Our results suggest that proportion of nearby sandsage habitat is an important factor in determining Swainson's hawk reproductive success. In addition, ferruginous hawk nest sites were located in areas that contained significantly more sandsage habitat than randomly selected sites in the study area. Nest-site availability may have constrained the distribution of buteos in our study area and is probably a major factor limiting nesting density of ferruginous hawks. Ferruginous hawks typically nest on man-made platforms (e.g., nest platforms and windmills) that in the study area were most common in and around the Rita Blanca National Grassland. Our results suggest that conversion of native grasslands to cropland may have negative consequences for ferruginous and Swainson's hawks. This relationship has been previously demonstrated in several studies of ferruginous hawks, but not for Swainson's hawks. In particular, loss of sandsage habitat on the southern Great Plains may have contributed to range declines in ferruginous hawks and decreased breeding success for Swainson's hawks.

In semi-arid grasslands of the North American Great Plains, fire has traditionally been viewed as having few management applications, and quantitative measurements of fire behavior in the low fuel loads characteristic of this region are lacking. More recently, land managers have recognized potential applications of prescribed fire to control undesirable plant species and to manage habitat for wildlife in this region. Working in the shortgrass steppe of northeastern Colorado over a 7-year period, we quantified peak temperatures, heating duration, and heat dosage produced near ground level during prescribed burns conducted under a wide range of fuel loads and weather conditions. We use an information theoretic approach to develop models that predict peak temperature and heat dosage as a function of weather parameters and fuel loads. Under the weather conditions that we examined, successful burns (>80 % of target area burnt) occurred with fuel loads varying from 350 kg ha−1 to 1175 kg ha−1, while burns with fuel loads

In arid and semiarid ecosystems, fire can potentially affect ecosystem dynamics through changes in soil moisture, temperature, and nitrogen cycling, as well as through direct effects on plant meristem mortality. We examined effects of annual and triennial prescribed fires conducted in early spring on soil moisture, temperature, and N, plant growth, and plant N content in semiarid shortgrass steppe. Annual burning increased soil inorganic N availability throughout the growing season, which was associated with increased soil temperature and a reduction in aboveground N in C3 plants. Furthermore, the increase in soil inorganic N pools with annual burning was modest and did not facilitate success of ruderal species. Negative fire effects on C3 plant production could be due to increased soil temperature, reduced soil moisture, or direct negative effects on C3 plant meristems, although fuel loads and fire temperatures were low relative to other grasslands. Triennial burning had intermediate effects on N availability and C3 plant production compared to annual burning and unburned controls. Results show that prescribed burns can be used in the management of this semiarid grassland without facilitating annual plant invasion, but excessively frequent burning can reduce production of C3 plants.

Disturbed xeric grasslands with short, sparse vegetation provide breeding habitat for mountain plovers (Charadrius montanus) across the western Great Plains. Maintaining local disturbance regimes through prairie dog conservation and prescribed fire may contribute to the sustainability of recently declining mountain plover populations, but these management approaches can be controversial. We estimated habitat- specific mountain plover densities and nest survival rates on black-tailed prairie dog (Cynomys ludovicianus) colonies and burns in the shortgrass steppe of northeastern Colorado. Mountain plover densities were similar on prairie dog colonies (5.9birds/km 2 ; 95% CI ¼4.7-7.4) and sites burned during the preceding dormant season (6.7birds/km 2 ; 95% CI ¼4.6-9.6), whereas the 29-day nest survival rate was greater on prairie dog colonies (0.81 in 2011 and 0.39 in 2012) compared to the burned sites (0.64 in 2011 and 0.17 in 2012). Reduced nest survival in 2012 compared to 2011 was associated with higher maximum daily temperatures in 2012, consistent with a previous weather-based model of mountain plover nest survival in the southern Great Plains. Measurements of mountain plover density relative to time since disturbance showed that removal of prairie dog disturbance by sylvatic plague reduced mountain plover density by 70% relative to active prairie dog colonies after 1 year. Plover densities declined at a similar rate (by 78%) at burned sites between the first and second post-burn growing season. Results indicate that black-tailed prairie dog colonies are a particularly important nesting habitat for mountain plovers in the southern Great Plains. In addition, findings suggest that prescribed burning can be a valuable means to create nesting habitat in landscapes where other types ofdisturbances (such as prairie dog colonies) are limited in distribution and size. Published 2014. This article is a U.S. Government work and is in the public domain in the USA.

Th e response of semiarid grasslands to small, non-colonial herbivores has received little attention, focusing primarily on the eff ects of granivore assemblages on annual plant communities. We studied the long-term eff ects of both small and large herbivores on vegetation structure and species diversity of shortgrass steppe, a perennial semiarid grassland considered marginal habitat for small mammalian herbivores. We hypothesized that 1) large generalist herbivores would aff ect more abundant species and proportions of litter-bare ground-vegetation cover through non-selective herbivory, 2) small herbivores would aff ect less common species through selective but limited consumption, and 3) herbivore eff ects on plant richness would increase with increasing aboveground net primary production (ANPP). Plant community composition was assessed over a 14-year period in pastures grazed at moderate intensities by cattle and in exclosures for large (cattle) and large-plus-small herbivores (additional exclusion of rabbits and rodents). Exclusion of large herbivores aff ected litter and bare ground and basal cover of abundant, common and uncommon species. Additional exclusion of small herbivores did not aff ect uncommon components of the plant community, but had indirect eff ects on abundant species, decreased the cover of the dominant grass Bouteloua gracilis and total vegetation, and increased litter and species diversity. Th ere was no relationship between ANPP and the intensity of eff ects of either herbivore body size on richness. Exclusion of herbivores of both body sizes had complementary and additive eff ects which promoted changes in vegetation composition and physiognomy that were linked to increased abundance of tall and decreased abundance of short species. Our fi ndings show that small mammalian herbivores had disproportionately large eff ects on plant communities relative to their small consumption of biomass. Even in smallseeded perennial grasslands with a long history of intensive grazing by large herbivores, non-colonial small mammalian herbivores should be recognized as an important driver of grassland structure and diversity.

Advances in global positioning system (GPS) technology have dramatically enhanced the ability to track and study distributions of free-ranging livestock. Understanding factors controlling the distribution of free-ranging livestock requires the ability to assess when and where they are foraging. For four years (2008–2011), we periodically collected GPS and activity sensor data together with direct observations of collared cattle grazing semiarid rangeland in eastern Colorado. From these data, we developed classification tree models that allowed us to discriminate between grazing and non-grazing activities. We evaluated: (1) which activity sensor measurements from the GPS collars were most valuable in predicting cattle foraging behavior, (2) the accuracy of binary (grazing, non-grazing) activity models vs. models with multiple activity categories (grazing, resting, traveling, mixed), and (3) the accuracy of models that are robust across years vs. models specific to a given year. A binary classification tree correctly removed 86.5% of the non-grazing locations, while correctly retaining 87.8% of the locations where the animal was grazing, for an overall misclassification rate of 12.9%. A classification tree that separated activity into four different categories yielded a greater misclassification rate of 16.0%. Distance travelled in a 5 minute interval and the proportion of the interval with the sensor indicating a head down position were the two most important variables predicting grazing activity. Fitting annual models of cattle foraging activity did not improve model accuracy compared to a single model based on all four years combined. This suggests that increased sample size was more valuable than accounting for interannual variation in foraging behavior associated with variation in forage production. Our models differ from previous assessments in semiarid rangeland of Israel and mesic pastures in the United States in terms of the value of different activity sensor measurements for identifying grazing activity, suggesting that the use of GPS collars to classify cattle grazing behavior will require calibrations specific to the environment and vegetation being studied.

Colonial burrowing herbivores can modify vegetation structure, create belowground refugia, and generate landscape heterogeneity, thereby affecting the distribution and abundance of associated species. Black-tailed prairie dogs (Cynomys ludovicianus) are such a species, and they may strongly affect the abundance and composition of grassland bird communities. We examined how prairie dog colonies in the North American Great Plains affect bird species and community composition. Areas occupied by prairie dogs, characterized by low percent cover of grass, high percent cover of bare soil, and low vegetation height and density, supported a breeding bird community that differed substantially from surrounding areas that lacked prairie dogs. Bird communities on colony sites had significantly greater densities of large-bodied carnivores (Burrowing Owls [Athene cunicularia], Mountain Plovers, [Charadrius montanus], and Killdeer [Charadrius vociferus]) and omnivores consisting of Horned Larks (Eremophila alpestris) and McCown's Longspurs (Rhynchophanes mccownii) than bird communities off colony sites. Bird communities off colony sites were dominated by small-bodied insectivorous sparrows (Ammodramus spp.) and omnivorous Lark Buntings (Calamospiza melanocorys), Vesper Sparrows (Pooecetes gramineus), and Lark Sparrows (Chondestes grammacus). Densities of 3 species of conservation concern and 1 game species were significantly higher on colony sites than off colony sites, and the strength of prairie dog effects was consistent across the northern Great Plains. Vegetation modification by prairie dogs sustains a diverse suite of bird species in these grasslands. Collectively, our findings and those from previous studies show that areas in the North American Great Plains with prairie dog colonies support higher densities of at least 9 vertebrate species than sites without colonies. Prairie dogs affect habitat for these species through multiple pathways, including creation of belowground refugia, supply of prey for specialized predators, modification of vegetation structure within colonies, and increased landscape heterogeneity. Asociaciones de Comunidades de Aves de Pastizales con Perros de la Pradera en la Gran Llanura de Norte America Resumen Los herbivoros excavadores coloniales pueden modificar la estructura de la vegetacion, crear refugios subterraneos y generar heterogeneidad en el paisaje, por lo tanto afectan la distribucion y abundancia de especies asociadas. Los perros de la pradera (Cynomys ludovicianus) son una de esas especies, y pueden afectar significativamente la abundancia y composicion de comunidades de aves de pastizales. Examinamos como afectan las colonias de perros de la pradera a las especies de aves y la composicion de la comunidad en la Gran Llanura de Norte America. Las areas ocupadas por perros de la pradera, caracterizadas por un bajo porcentaje de cobertura de pasto, alto porcentaje de cobertura de suelo desnudo y baja altura y densidad de la vegetacion, soportaban una comunidad de aves que difirio sustancialmente en areas que no tenian perros de la pradera. Las comunidades de aves en sitios con colonias tuvieron significativamente mayores densidades de carnivoros de talla grande (Athene cunicularia,Charadrius montanusyCharadrius vociferus) y omnivoros (Eremophila alpestrisyRhynchophanes mccownii) que las comunidades de aves en sitios sin colonias. Las comunidades de aves en sitios sin colonias fueron dominadas por gorriones insectivoros de talla pequena (Ammodramus spp.) y omnivoros (Calamospiza melanocorys,Pooecetes gramineusyChondestes grammacus). Las densidades de 3 especies de preocupacion para la conservacion y una especie cinegetica fueron significativamente mayores en los sitios con colonias, y la fuerza de los efectos de los perros de la pradera fue consistente en el norte de Gran Llanura. La modificacion de la vegetacion por los perros de la pradera sostiene a un conjunto diverso de especies de aves estos pastizales. Colectivamente, nuestros resultados y los de estudios previos muestran que las areas con colonias de perros de la pradera en la Gran Llanura de Norte America soportan densidades mas altas de por lo menos 9 especies de vertebrados que en los sitios sin colonias. Los perros de la pradera afectan el habitat de esas especies por diferentes vias, incluyendo la creacion de refugios subterraneos, proveyendo presas para depredadores especializados, modificando la estructura de vegetacion en las colonias e incrementando la heterogeneidad del paisaje.

Nitrogen (N) availability can strongly influence forage quality and the capacity for semiarid rangelands to respond to increasing atmospheric CO2. Although many pathways of nitrogen input and loss from rangelands have been carefully quantified, cattlemediated N losses are often poorly understood. We used measurements of cattle N consumption rate, weight gains, and spatial distribution in shortgrass rangeland of northeastern Colorado to evaluate the influence of cattle on rangeland N balance. Specifically, we estimated annual rates of N loss via cattle weight gains and spatial redistribution of N into pasture corners and areas near water tanks, and used previous studies to calculate ammonia volatilization from urine patches. Using measurements of plant biomass and N content inside and outside grazing cages over 13 yr, we estimate that cattle stocked at 0.65 animal unit

Aerial surveys using line-intercept methods are one approach to estimate the extent of prairie dog colonies in a large geographic area. Although black-tailed prairie dogs (Cynomys ludovicianus) construct conspicuous mounds at burrow openings, aerial observers have difficulty discriminating between areas with burrows occupied by prairie dogs (colonies) versus areas of uninhabited burrows (uninhabited colony sites). Consequently, aerial line-intercept surveys may overestimate prairie dog colony extent unless adjusted by an on-the-ground inspection of a sample of intercepts. We compared aerial line-intercept surveys conducted over 2 National Grasslands in Colorado, USA, with independent ground-mapping of known black-tailed prairie dog colonies. Aerial line-intercepts adjusted by ground surveys using a single activity category adjustment overestimated colonies by ≥94% on the Comanche National Grassland and ≥58% on the Pawnee National Grassland. We present a ground-survey technique that involves 1) visiting on the ground a subset of aerial intercepts classified as occupied colonies plus a subset of intercepts classified as uninhabited colony sites, and 2) based on these ground observations, recording the proportion of each aerial intercept that intersects a colony and the proportion that intersects an uninhabited colony site. Where line-intercept techniques are applied to aerial surveys or remotely sensed imagery, this method can provide more accurate estimates of black-tailed prairie dog abundance and trends. Published 2012. This article is a U.S. Government work and is in the public domain in the USA.

Precipitation pulses in arid ecosystems can lead to temporal asynchrony in microbial and plant processing of nitrogen (N) during drying/wetting cycles causing increased N loss. In contrast, more consistent availability of soil moisture in mesic ecosystems can synchronize microbial and plant processes during the growing season, thus minimizing N loss. We tested whether microbial N cycling is asynchronous with plant N uptake in a semiarid grassland. Using (15)N tracers, we compared rates of N cycling by microbes and N uptake by plants after water pulses of 1 and 2 cm to rates in control plots without a water pulse. Microbial N immobilization, gross N mineralization, and nitrification dramatically increased 1-3 days after the water pulses, with greatest responses after the 2-cm pulse. In contrast, plant N uptake increased more after the 1-cm than after the 2-cm pulse. Both microbial and plant responses reverted to control levels within 10 days, indicating that both microbial and plant responses were short lived. Thus, microbial and plant processes were temporally synchronous following a water pulse in this semiarid grassland, but the magnitude of the pulse substantially influenced whether plants or microbes were more effective in acquiring N. Furthermore, N loss increased after both small and large water pulses (as shown by a decrease in total (15)N recovery), indicating that changes in precipitation event sizes with future climate change could exacerbate N losses from semiarid ecosystems.

Colonial, burrowing herbivores can be engineers of grassland and shrubland ecosystems worldwide. Spatial variation in landscapes suggests caution when extrapolating single-place studies of single species, but lack of data and the need to generalize often leads to ‘ model system ’ thinking and application of results beyond appropriate statistical inference. Generalizations about the engineering eff ects of prairie dogs ( Cynomys sp.) developed largely from intensive study at a single complex of black-tailed prairie dogs C. ludovicianus in northern mixed prairie, but have been extrapolated to other ecoregions and prairie dog species in North America, and other colonial, burrowing herbivores. We tested the paradigm that prairie dogs decrease vegetation volume and the cover of grasses and tall shrubs, and increase bare ground and forb cover. We sampled vegetation on and off 279 colonies at 13 complexes of 3 prairie dog species widely distributed across 5 ecoregions in North America. Th e paradigm was generally supported at 7 black-tailed prairie dog complexes in northern mixed prairie, where vegetation volume, grass cover, and tall shrub cover were lower, and bare ground and forb cover were higher, on colonies than at paired off -colony sites. Outside the northern mixed prairie, all 3 prairie dog species consistently reduced vegetation volume, but their eff ects on cover of plant functional groups varied with prairie dog species and the grazing tolerance of dominant perennial grasses. White-tailed prairie dogs C. leucurus in sagebrush steppe did not reduce shrub cover, whereas black-tailed prairie dogs suppressed shrub cover at all complexes with tall shrubs in the surrounding habitat matrix. Black-tailed prairie dogs in shortgrass steppe and Gunnison ’ s prairie dogs C. gunnisoni in Colorado Plateau grassland both had relatively minor eff ects on grass cover, which may refl ect the dominance of grazing-tolerant shortgrasses at both complexes. Variation in modifi cation of vegetation structure may be understood in terms of the responses of diff erent dominant perennial grasses to intense defoliation and diff erences in foraging behavior among prairie dog species. Spatial variation in the engineering role of prairie dogs suggests spatial variation in their keystone role, and spatial variation in the roles of other ecosystem engineers. Th us, ecosystem engineering can have a spatial component not evident from single-place studies.

Restoring historical disturbance regimes to enhance habitat for grassland birds can conflict with livestock production goals and has been controversial because of uncertainty in the frequency and pattern of different disturbances prior to European settlement. We studied nesting habitat for the mountain plover (Charadrius montanus) in relation to prescribed fire, grazing by large herbivores (cattle), and grazing by black- tailed prairie dogs (Cynomys ludovicianus) in the shortgrass steppe of northeastern Colorado. Breeding mountain plovers primarily occurred on black-tailed prairie dog colonies or areas burned during the previous dormant season. Vegetation surrounding mountain plover nests and foraging locations was characterized by a fine-scale mosaic of prostrate ( 35% bare soil in a given square meter, with this fine-scale pattern distributed over a broad (>100-m radius) area. Mountain plovers rarely occupied grassland lacking prairie dogs or recent fire, but those that did selected sites with similar vegetation height and bare soil exposure as sites on burns and prairie dog colonies. Vegetation structure at mountain plover-occupied sites was also similar to random sites on burns and prairie dog colonies, but differed substantially from sites managed only with cattle. Intensive cattle grazing at twice the recommended stocking rate during spring (Mar-May) or summer (May-Oct) for 6 years produced significantly less bare soil than burns and prairie dog colonies, particularly following years with average or above-average precipitation. Thus, intensive cattle grazing did not substitute for prairie dog grazing or fire in terms of effects on vegetation structure and mountain plover habitat. Both prescribed burning and increased size and distribution of black- tailed prairie dog colonies appear to be effective and complementary means to manage for mountain plover breeding habitat in shortgrass steppe. Provision of mountain plover habitat has tradeoffs with traditional management for livestock production. Thus, managers need to clearly define desired outcomes for manage- ment to provide multiple ecosystem goods and services. 2012 The Wildlife Society.

We used very large scale aerial (VLSA) photography to quantify spatial patterns in bare soil in the northeastern Colorado shortgrass steppe. Using three pairs of pastures stocked at moderate (0.6 animal unit months lAUM] · ha -1 ) versus very heavy (1.2 AUM · ha -1 ) rates, we detected greater bare soil under very heavy (mean = 22.5%) versus moderate stocking (mean = 13.5%; P = 0.053) and a lower coefficient of variation across pastures under very heavy (0.48) versus moderate stocking (0.75; P = 0.032). Bare soil exhibited significant positive spatial autocorrelation across distances of 60–120 m under moderate stocking (Moran's I = 0.14), while patchiness at this scale was eliminated under very heavy grazing ( I = -0.05). Across distances of 120–480 m, we observed no spatial autocorrelation with either stocking rate. Spatial autocorrelation was greatest at a separation distance of 2 m ( I = 0.48–0.58) but was unaffected by stocking rate at this scale. Thus, very heavy grazing did not increase spatial autocorrelation in bare soil across scales of 2–480 m. Means and variability in the distribution of bare soil were not influenced by ecological site. Bare soil increased primarily at the scale of individual plant clusters through both increases in the density of small (2–20 cm) bare patch intercepts and increases in the frequency of bare patch intercepts of 20–60 cm (rather than s 20 cm). Our approach demonstrates the utility of VLSA for analyzing interactions between grazing and other landscape features and highlights the importance of spatially explicit sampling across broad scales (pastures) while testing for potential shifts in patchiness of bare soil at the scale of plant interspaces.

Much of the breeding range for the mountain plover (Charadrius montanus) occurs in shortgrass steppe and mixed-grass prairie in the western Great Plains of North America. Studies of mountain plovers in shortgrass steppe during the 1970s and 1990s were focused in Weld County, Colorado, which was considered a key breeding area for the species. These studies, however, did not include habitats influenced by black-tailed prairie dogs (Cynomys ludovicianus) or prescribed fire. The role of these 2 rangeland disturbance processes has increased substantially over the past 15 years. During 2008–2009, I used radial distance point count surveys to estimate mountain plover densities early in the nesting season in 4 habitats on public lands in Weld County, Colorado. All 4 habitats were grazed by cattle during the growing season at moderate stocking rates but had different additional disturbances consisting of 1) dormant-season prescribed burns, 2) active black-tailed prairie dog colonies, 3) black-tailed prair...

We examined the independent and combined effects of prescribed fire and livestock grazing on herbaceous plant production in shortgrass steppe of northeastern Colorado in the North American Great Plains. Burning was implemented in March, before the onset of the growing season. During the first postburn growing season, burning had no influence on soil moisture, nor did it affect soil nitrogen (N) availability in spring (April–May), but it significantly enhanced soil N availability in summer (June–July). Burning had no influence on herbaceous plant production in the first postburn growing season but enhanced in vitro dry matter digestibility of blue grama (Bouteloua gracilis [Willd. ex Kunth] Lag. ex Griffiths) forage sampled in late May. For the second postburn growing season, we found no difference in herbaceous plant production between sites that were burned and grazed in the previous year versus sites that were burned and protected from grazing in the previous year. Our results provide further evidence that prescribed burns conducted in late winter in dormant vegetation can have neutral or positive consequences for livestock production because of a neutral effect on forage quantity and a short-term enhancement of forage quality. In addition, our results indicate that with conservative stocking rates, deferment of grazing during the first postburn growing season may not be necessary to sustain plant productivity.

Spatial and temporal variations in precip- itation are central features of semiarid ecosystems, influencing patterns of plant productivity and the distribution of native fauna. Although temporal variation in precipitation has been studied exten- sively, far less is known about the spatial scale and pattern of precipitation variability in semiarid regions. I used long-term precipitation records to examine spatial variation across the 63 km 2 Central Plains Experimental Range in northeastern Colorado, and across the 117,000 km 2 region of shortgrass steppe in eastern Colorado. Relative to temporal variation, spatial variation was low at scales\10 km, increased linearly across scales of 40-120 km, and was nearly equal in magnitude to temporal variation across distances of 120-160 km. Although I hypoth- esized that most spatial variation would be generated by early-summer convective thunderstorms in June, I found that the magnitude and spatial pattern of variation was similar for precipitation received in June compared to cumulative precipitation received during the full growing season. The degree of spatial autocorrelation in precipitation across all distances that I evaluated was similar for drought, dry, above- average and wet years. Across distances of 10- 120 km, spatial variation within a single growing season was approximately two times greater than spatial variation in long-term mean growing-season precipitation, indicating spatial shifting in the loca- tions of patches of high and low precipitation over multiple years. Overall, these findings suggest spatial variation at scales of 10-160 km may have been an important factor influencing vegetation patterns and migratory fauna of the shortgrass steppe, and have implications for livestock producers and future assessments of climate change.

Over the past century, fire has been widely suppressed in the western Great Plains, in part because of the potential negative effects on forage production for livestock. More recently, interest in the use of prescribed fire in shortgrass steppe has increased because of the potential applications for wildlife management, control of unpalatable plant species, and restoration of historic disturbance regimes. We studied the effects of prescribed burns conducted during late winter on herbaceous production, forage nitrogen content, and plant species composition of shortgrass steppe on the Pawnee National Grassland in northeastern Colorado. Late-winter burns conducted in moderately grazed sites under a wide range of precipitation conditions during 1997–2001 did not negatively affect herbaceous production in either the first or the second postburn growing season. Burning followed by a severe drought in 2002 reduced production by 19% in the second postburn growing season of 2003. Burns temporarily suppressed the abundance of broom snakeweed (Gutierrezia sarothrae) and prickly pear cactus (Opuntia polyacantha) and enhanced forage nitrogen content during May and June of the first postburn growing season. These findings suggest that, except following severe drought, prescribed burns conducted during late winter in grazed shortgrass steppe for objectives unrelated to livestock production can also have neutral or positive consequences for livestock.

Swift foxes (Vulpes velox) are shortgrass specialists and as a result are heavily dependent upon grassland disturbance regimes to maintain high-quality habitat. To better understand this relationship, we monitored the move- ment and spatial ecology of resident swift foxes before and after a 2005 prescribed burn in southeastern Colorado. We hypothesized that foxes would shift home ranges into and increase foraging activity within the prescribed burn area. Foxes did appear to alter their space-use patterns in response to the burn, although the response was constrained by ter- ritoriality. Foxes whose core-use areas overlapped the burn increased their use of the burn area for foraging and den- ning, but we did not observe shifts of individual home ranges to encompass more of the burned area. Foxes whose core- use areas did not overlap the burn did not alter their space use or change home range boundaries in response to the burn. Because we observed only positive or neutral responses to prescribed burning, we recommend this tool as an appropriate method to maintain high-quality swift fox habitat.

Black-tailed prairie dogs (Cynomys ludovicianus) are a key component of the disturbance regime in semi-arid grasslands of central North America. Many studies have compared community and ecosystem characteristics on prairie dog colonies to grasslands without prairie dogs, but little is known about landscape-scale patterns of disturbance that prairie dog colony complexes may impose on grasslands over long time periods. We examined spatiotemporal dynamics in two prairie dog colony complexes in southeastern Colorado (Comanche) and northcentral Montana (Phillips County) that have been strongly influenced by plague, and compared them to a complex unaffected by plague in northwestern Nebraska (Oglala). Both plague-affected complexes exhibited substantial spatiotemporal variability in the area occupied during a decade, in contrast to the stability of colonies in the Oglala complex. However, the plague-affected complexes differed in spatial patterns of colony movement. Colonies in the Comanche complex in shortgrass steppe shifted locations over a decade. Only 10% of the area occupied in 1995 was still occupied by prairie dogs in 2006. In 2005 and 2006 respectively, 74 and 83% of the total area of the Comanche complex occurred in locations that were not occupied in 1995, and only 1% of the complex was occupied continuously over a decade. In contrast, prairie dogs in the Phillips County complex in mixed-grass prairie and sagebrush steppe primarily recolonized previously occupied areas after plague-induced colony declines. In Phillips County, 62% of the area occupied in 1993 was also occupied by prairie dogs in 2004, and 12% of the complex was occupied continuously over a decade. Our results indicate that plague accelerates spatiotemporal movement of prairie dog colonies, and have significant implications for landscape-scale effects of prairie dog disturbance on grassland composition and productivity. These findings highlight the need to combine landscape-scale measures of habitat suitability with long-term measures of colony locations to understand the role of plague-affected prairie dogs as a grassland disturbance process.

Factors influencing the distribution and abundance of black-tailed prairie dog (Cynomys ludovicianus) colonies are of interest to rangeland managers because of the significant influence prairie dogs can exert on both livestock and biodiversity. We examined the influence of 4 prescribed burns and one wildfire on the rate and direction of prairie dog colony expansion in shortgrass steppe of southeastern Colorado. Our study was conducted during 2 years with below-average precipitation, when prairie dog colonies were expanding throughout the study area. Under these dry conditions, the rate of black-tailed prairie dog colony expansion into burned grassland (X ¯ 5 2.6 ha ? 100-m perimeter 21 ? y 21 ; range 5 0.8–5.9 ha ? 100-m perimeter 21 ? y 21 ; N 5 5 colonies) was marginally greater than the expansion rate into unburned grassland (X ¯ 5 1.3 ha ? 100-m perimeter 21 ? y 21 ; range 5 0.2–4.9 ha ? 100-m perimeter 21 ? y 21 ; N 5 23 colonies; P 5 0.066). For 3 colonies that were burned on only a portion of their perimeter, we documented consistently high rates of expansion into the adjacent burned grassland (38%–42% of available burned habitat colonized) but variable expansion rates into the adjacent unburned grassland (2%–39% of available unburned habitat colonized). While our results provide evidence that burning can increase colony expansion rate even under conditions of low vegetative structure, this effect was minor at the scale of the overall colony complex because some unburned colonies were also able to expand at high rates. This result highlights the need to evaluate effects of fire on colony expansion during aboveaverage rainfall years, when expansion into unburned grassland may be considerably lower.

Grazing intensity elicits changes in the composition of plant functional groups in both shortgrass steppe (SGS) and northern mixed-grass prairie (NMP) in North America. How these grazing intensity-induced changes control aboveground net primary production (ANPP) responses to precipitation remains a central open question, especially in light of predicted climate changes. Here, we evaluated effects of four levels (none, light, moderate, and heavy) of long-term (>30 yr) grazing intensity in SGS and NMP on: (1) ANPP; (2) precipitation-use efficiency (PUE, ANPP : precipitation); and (3) precipitation marginal response (PMR; slope of a linear regression model between ANPP and precipitation). We advance prior work by examining: (1) the consequences of a range of grazing intensities (more grazed vs. ungrazed); and (2) how grazing-induced changes in ANPP and PUE are related both to shifts in functional group composition and physiological responses within each functional group. Spring (April-June) precipitation, the primary determinant of ANPP, was only 12% higher in NMP than in SGS, yet ANPP and PUE were 25% higher. Doubling grazing intensity in SGS and nearly doubling it in NMP reduced ANPP and PUE by only 24% and 33%, respectively. Increased grazing intensity reduced C3 graminoid biomass and increased C4 grass biomass in both grasslands. Functional group shifts affected PUE through biomass reductions, as PUE was positively associated with the relative abundance of C3 species and negatively with C4 species across both grasslands. At the community level, PMR was similar between grasslands and unaffected by grazing intensity. However, PMR of C3 graminoids in SGS was eightfold higher in the ungrazed treatment than under any grazed level. In NMP, PMR of C3 graminoids was only reduced under heavy grazing intensity. Knowing the ecological consequences of grazing intensity provides valuable information for mitigation and adaptation strategies in response to predicted climate change. For example, moderate grazing (the recommended rate) in SGS would sequester the same amount of aboveground carbon as light grazing because ANPP was nearly the same. In contrast, reductions in grazing intensity in NMP from moderate to light intensity would increase the amount of aboveground carbon sequestrated by 25% because of increased ANPP.

Large herbivores can both positively and negatively affect primary productivity and rates of nutrient cycling in different ecosystems. Positive effects of grazers in grasslands have been attributed to migratory behavior of the dominant ungulate species and soil fertility. We studied the effects of grazers on aboveground net primary productivity (ANPP) and N cycling on central Kenyan rangeland characterized by intense, chronic grazing by a mixed community of cattle and resident native ungulates. Exclosure studies conducted at high and low levels of soil fertility showed that both soil fertility and annual rainfall patterns mediate the effects of grazers on ANPP and N cycling. In a low-rainfall year with short (1 month) growing seasons, grazers reduced aboveground productivity regardless of soil nutrient availability. However, in a high-rainfall year with a 5-month growing season, grazers increased ANPP on nutrient-rich glades and suppressed ANPP on nutrient-poor bushland sites. Concomitant studies of grazer effects on N cycling revealed complex interactions with the seasonal pattern of N-mineralization and inorganic N availability. Grazers increased the size of the inorganic N pool available to plants at the onset of the growing season, particularly in nutrient-rich glades. However, grazers also decreased N mineralization rates at all sites early in the growing season. Measures of N availability via ion-exchange resin bags suggested that the combined effects of grazers on inorganic N pool fluctuations and N-mineralization rates resulted in a net increase in N availability at glade sites and a net decrease in N availability at bushland sites. The net effect of grazers on soil N availability mirrored grazer effects on ANPP in the high-rainfall year. Overall, our results suggest that grazer effects on N dynamics are closely linked to effects on productivity and resilience to drought. Furthermore, even under optimal conditions of high soil fertility and above-average rainfall, grazer promotion of ANPP in this chronically grazed system dominated by resident ungulates was small compared to systems dominated by migratory ungulates.

Herbivores, edaphic features and fire are primary factors regulating the balance between woody and herbaceous vegetation in savannas. Many observational studies have evaluated the potential effects of browsing herbivores on woody plant dynamics in African savannas, but few experimental studies have compared the dynamics of African savannas with and without browsers. 2. A replicated herbivore exclusion experiment was used to assess the role that native ungulates play in regulating woody plant dynamics on commercial rangeland in central Kenya, where the indigenous fauna have been allowed to coexist with cattle. 3. Exclusion of native browsing ungulates for just 3 years showed that they have dramatic effects at every scale from individual twig growth rates to overall rates of woody biomass accumulation in the ecosystem. 4. At the scale of individual Acacia twigs, browsers significantly reduced leaf density, leaf biomass and growth rates of twigs 2·5 m tall, such that Acacias in intermediate height classes (0·5-2·5 m) experienced minimal browser impacts. Elephants influenced shrubland dynamics by altering shrub height-class dis- tributions, shifting species composition from broad-leaved Grewia tenax to fine-leaved Acacia species, and suppressing woody biomass accumulation; but elephants had little influence on changes in shrub density. 7. Synthesis and applications . Our results suggest that a community of native browsers that includes both small, selective species (e.g. dik-diks) and large, bulk-feeding species (elephants) can provide an important ecosystem service by suppressing shrub encroach- ment on commercial rangeland.