Secondary production of the central rangeland region of the United States.

Rangelands are the dominant land use across a broad swath of central North America where they span a wide gradient, from <350 to >900 mm, in mean annual precipitation. Substantial efforts have examined temporal and spatial variation in aboveground net primary production (ANPP) to precipitation (PPT) across this gradient. In contrast, net secondary productivity (NSP, e.g., primary consumer production) has not been evaluated analogously. However, livestock production, which is a form of NSP or primary consumer production supported by primary production, is the dominant non-cultivated land use and an integral economic driver in these regions. Here, we used long-term (mean length = 19 years) ANPP and NSP data from six research sites across the Central Great Plains with a history of a conservative stocking to determine resource (i.e., PPT)-productivity relationships, NSP sensitivities to dry-year precipitation, and regional trophic efficiencies (e.g., NSP:ANPP ratio). PPT-ANPP relationships were linear for both temporal (site-based) and spatial (among site) gradients. The spatial PPT-NSP model revealed that PPT mediated a saturating relationship for NSP as sites became more mesic, a finding that contrasts with many plant-based PPT-ANPP relationships. A saturating response to high growing-season precipitation suggests biogeochemical rather than vegetation growth constraints may govern NSP (i.e., large herbivore production). Differential sensitivity in NSP to dry years demonstrated that the primary consumer production response heightened as sites became more xeric. Although sensitivity generally decreased with increasing precipitation as predicted from known PPT-ANPP relationships, evidence suggests that the dominant species' identity and traits influenced secondary production efficiency. Non-native northern mixed-grass prairie was outperformed by native Central Great Plains rangeland in sensitivity to dry years and efficiency in converting ANPP to NSP. A more comprehensive understanding of the mechanisms leading to differences in producer and consumer responses will require multisite experiments to assess biotic and abiotic determinants of multi-trophic level efficiency and sensitivity.

A big data-model integration approach for predicting epizootics and population recovery in a keystone species.

Infectious diseases pose a significant threat to global health and biodiversity. Yet, predicting the spatiotemporal dynamics of wildlife epizootics remains challenging. Disease outbreaks result from complex nonlinear interactions among a large collection of variables that rarely adhere to the assumptions of parametric regression modeling. We adopted a nonparametric machine learning approach to model wildlife epizootics and population recovery, using the disease system of colonial black-tailed prairie dogs (BTPD, Cynomys ludovicianus) and sylvatic plague as an example. We synthesized colony data between 2001 and 2020 from eight USDA Forest Service National Grasslands across the range of BTPDs in central North America. We then modeled extinctions due to plague and colony recovery of BTPDs in relation to complex interactions among climate, topoedaphic variables, colony characteristics, and disease history. Extinctions due to plague occurred more frequently when BTPD colonies were spatially clustered, in closer proximity to colonies decimated by plague during the previous year, following cooler than average temperatures the previous summer, and when wetter winter/springs were preceded by drier summers/falls. Rigorous cross-validations and spatial predictions indicated that our final models predicted plague outbreaks and colony recovery in BTPD with high accuracy (e.g., AUC generally >0.80). Thus, these spatially explicit models can reliably predict the spatial and temporal dynamics of wildlife epizootics and subsequent population recovery in a highly complex host-pathogen system. Our models can be used to support strategic management planning (e.g., plague mitigation) to optimize benefits of this keystone species to associated wildlife communities and ecosystem functioning. This optimization can reduce conflicts among different landowners and resource managers, as well as economic losses to the ranching industry. More broadly, our big data-model integration approach provides a general framework for spatially explicit forecasting of disease-induced population fluctuations for use in natural resource management decision-making.

Disease and weather induce rapid shifts in a rangeland ecosystem mediated by a keystone species (Cynomys ludovicianus).

Habitat loss and changing climate have direct impacts on native species but can also interact with disease pathogens to influence wildlife communities. In the North American Great Plains, black-tailed prairie dogs (Cynomys ludovicianus) are a keystone species that create important grassland habitat for numerous species and serve as prey for predators, but lethal control driven by agricultural conflict has severely reduced their abundance. Novel disease dynamics caused by epizootic plague (Yersinia pestis) within prairie dog colonies have further reduced prairie dog abundances, in turn destabilizing associated wildlife communities. We capitalized on a natural experiment, collecting data on prairie dog distributions, vegetation structure, avian abundance, and mesocarnivore and ungulate occupancy before (2015-2017) and after (2018-2019) a plague event in northeastern Wyoming, USA. Plague decimated black-tailed prairie dog populations in what was then the largest extant colony complex, reducing colony cover in the focal area from more than 10,000 ha to less than 50 ha. We documented dramatic declines in mesocarnivore occupancy and raptor abundance post-plague, with probability of occupancy or abundance approaching zero in species that rely on prairie dogs for a high proportion of their diet (e.g., ferruginous hawk [Buteo regalis], American badger [Taxidea taxus], and swift fox [Vulpes velox]). Following the plague outbreak, abnormally high precipitation in 2018 hastened vegetation recovery from prairie dog disturbance on colonies in which constant herbivory had formerly maintained shortgrass structure necessary for certain colony-associates. As a result, we observed large shifts in avian communities on former prairie dog colonies, including near-disappearance of mountain plovers (Charadrius montanus) and increases in mid-grass associated songbirds (e.g., lark bunting [Calamospiza melanocorys]). Our research highlights how precipitation can interact with disease-induced loss of a keystone species to induce drastic and rapid shifts in wildlife communities. Although grassland taxa have co-evolved with high spatiotemporal variation, fragmentation of the remaining North American rangelands paired with higher-than-historical variability in climate and disease dynamics are likely to destabilize these systems in the future.

Predicting spatial-temporal patterns of diet quality and large herbivore performance using satellite time series.

Adaptive management of large herbivores requires an understanding of how spatial-temporal fluctuations in forage biomass and quality influence animal performance. Advances in remote sensing have yielded information about the spatial-temporal dynamics of forage biomass, which in turn have informed rangeland management decisions such as stocking rate and paddock selection for free-ranging cattle. However, less is known about the spatial-temporal patterns of diet quality and their influence on large herbivore performance. This is due to infrequent concurrent ground observations of forage conditions with performance (e.g., mass gain), and previously limited satellite data at fine spatial and temporal scales. We combined multi-temporal field observations of diet quality (weekly) and mass gain (monthly) with satellite-derived phenological metrics (pseudo-daily, using data fusion and interpolation) to model daily mass gains of free-ranging yearling cattle in shortgrass steppe. We used this model to predict grazing season (mid-May to October) mass gains, a key management indicator, across 40 different paddocks grazed over a 10-year period (n = 138). We found strong relationships between diet quality and the satellite-derived phenological metrics, especially metrics related to the timing and rate of green-up and senescence. Satellite-derived diet quality estimates were strong predictors of monthly mass gains (R2  = 0.68) across a wide range of aboveground net herbaceous production. Season-long predictions of average daily gain and cattle off-mass had mean absolute errors of 8.9% and 2.9%, respectively. The model performed better temporally (across repeated observations in the same paddock) than spatially (across all paddocks within a given year), highlighting the need for accurate vegetation maps and robust field data collection across both space and time. This study demonstrates that free-ranging cattle performance in rangelands is strongly affected by diet quality, which is related to the timing of vegetation green-up and senescence. Senescing vegetation suppressed mass gains, even if adequate forage was available. The satellite-based pseudo-daily approach presented here offers new opportunities for adaptive management of large herbivores, such as identifying within-season triggers to move livestock among paddocks, predicting wildlife herd health, or timing the grazing season to better match earlier spring green-up caused by climate change and plant species invasion.

Large herbivores suppress liana infestation in an African savanna.

African savannas are the last stronghold of diverse large-mammal communities, and a major focus of savanna ecology is to understand how these animals affect the relative abundance of trees and grasses. However, savannas support diverse plant life-forms, and human-induced changes in large-herbivore assemblages-declining wildlife populations and their displacement by livestock-may cause unexpected shifts in plant community composition. We investigated how herbivory affects the prevalence of lianas (woody vines) and their impact on trees in an East African savanna. Although scarce (<2% of tree canopy area) and defended by toxic latex, the dominant liana, Cynanchum viminale (Apocynaceae), was eaten by 15 wild large-herbivore species and was consumed in bulk by native browsers during experimental cafeteria trials. In contrast, domesticated ungulates rarely ate lianas. When we experimentally excluded all large herbivores for periods of 8 to 17 y (simulating extirpation), liana abundance increased dramatically, with up to 75% of trees infested. Piecewise exclusion of different-sized herbivores revealed functional complementarity among size classes in suppressing lianas. Liana infestation reduced tree growth and reproduction, but herbivores quickly cleared lianas from trees after the removal of 18-y-old exclosure fences (simulating rewilding). A simple model of liana contagion showed that, without herbivores, the long-term equilibrium could be either endemic (liana-tree coexistence) or an all-liana alternative stable state. We conclude that ongoing declines of wild large-herbivore populations will disrupt the structure and functioning of many African savannas in ways that have received little attention and that may not be mitigated by replacing wildlife with livestock.

Grasses continue to trump trees at soil carbon sequestration following herbivore exclusion in a semiarid African savanna.

Although studies have shown that mammalian herbivores often limit aboveground carbon storage in savannas, their effects on belowground soil carbon storage remain unclear. Using three sets of long-term, large herbivore exclosures with paired controls, we asked how almost two decades of herbivore removal from a semiarid savanna in Laikipia, Kenya affected aboveground (woody and grass) and belowground soil carbon sequestration, and determined the major source (C3 vs. C4 ) of belowground carbon sequestered in soils with and without herbivores present. Large herbivore exclusion, which included a diverse community of grazers, browsers, and mixed-feeding ungulates, resulted in significant increases in grass cover (~22%), woody basal area (~8 m2 /ha), and woody canopy cover (31%), translating to a ~8.5 t/ha increase in aboveground carbon over two decades. Herbivore exclusion also led to a 54% increase (20.5 t/ha) in total soil carbon to 30-cm depth, with ~71% of this derived from C4 grasses (vs. ~76% with herbivores present) despite substantial increases in woody cover. We attribute this continued high contribution of C4 grasses to soil C sequestration to the reduced offtake of grass biomass with herbivore exclusion together with the facilitative influence of open sparse woody canopies (e.g., Acacia spp.) on grass cover and productivity in this semiarid system.

Large-scale and local climatic controls on large herbivore productivity: implications for adaptive rangeland management.

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 Niño-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 Niña, 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.

Adaptive rangeland management benefits grassland birds utilizing opposing vegetation structure in the shortgrass steppe.

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.

Large herbivores maintain a two-phase herbaceous vegetation mosaic in a semi-arid savanna.

Many arid and semi-arid rangelands exhibit distinct spatial patterning of vegetated and bare soil-dominated patches. The latter potentially represent a grazing-induced, degraded ecosystem state, but could also arise via mechanisms related to feedbacks between vegetation cover and soil moisture availability that are unrelated to grazing. The degree to which grazing contributes to the formation or maintenance of degraded patches has been widely discussed and modeled, but empirical studies of the role of grazing in their formation, persistence, and reversibility are limited.We report on a long-term (17 years) grazing removal experiment in a semi-arid savanna where vegetated patches composed of perennial grasses were interspersed within large (>10 m2) patches of bare soil.Short-term (3 years) grazing removal did not allow bare patches to become revegetated, whereas following long-term (17 years) grazing removal, bare soil patches were revegetated by a combination of stoloniferous grasses and tufted bunchgrasses. In the presence of grazers, stoloniferous grasses partially recolonized bare patches, but this did not lead to full recovery or to the establishment of tufted bunchgrasses.These results show that grazers alter both the balance between bare and vegetated patches, as well as the types of grasses dominating both patch types in this semiarid savanna.Synthesis: Large herbivores fundamentally shaped the composition and spatial pattern of the herbaceous layer by maintaining a two-phase herbaceous mosaic. However, bare patches within this mosaic can recover given herbivore removal over sufficiently long time scales, and hence do not represent a permanently degraded ecosystem state.

Conservation lessons from large-mammal manipulations in East African savannas: the KLEE, UHURU, and GLADE experiments.

African savannas support an iconic fauna, but they are undergoing large-scale population declines and extinctions of large (>5 kg) mammals. Long-term, controlled, replicated experiments that explore the consequences of this defaunation (and its replacement with livestock) are rare. The Mpala Research Centre in Laikipia County, Kenya, hosts three such experiments, spanning two adjacent ecosystems and environmental gradients within them: the Kenya Long-Term Exclosure Experiment (KLEE; since 1995), the Glade Legacies and Defaunation Experiment (GLADE; since 1999), and the Ungulate Herbivory Under Rainfall Uncertainty experiment (UHURU; since 2008). Common themes unifying these experiments are (1) evidence of profound effects of large mammalian herbivores on herbaceous and woody plant communities; (2) competition and compensation across herbivore guilds, including rodents; and (3) trophic cascades and other indirect effects. We synthesize findings from the past two decades to highlight generalities and idiosyncrasies among these experiments, and highlight six lessons that we believe are pertinent for conservation. The removal of large mammalian herbivores has dramatic effects on the ecology of these ecosystems; their ability to rebound from these changes (after possible refaunation) remains unexplored.

Elevated CO2 induces substantial and persistent declines in forage quality irrespective of warming in mixedgrass prairie.

Increasing atmospheric [CO2 ] and temperature are expected to affect the productivity, species composition, biogeochemistry, and therefore the quantity and quality of forage available to herbivores in rangeland ecosystems. Both elevated CO2 (eCO2 ) and warming affect plant tissue chemistry through multiple direct and indirect pathways, such that the cumulative outcomes of these effects are difficult to predict. Here, we report on a 7-yr study examining effects of CO2 enrichment (to 600 ppm) and infrared warming (+1.5°C day/3°C night) under realistic field conditions on forage quality and quantity in a semiarid, mixedgrass prairie. For the three dominant forage grasses, warming effects on in vitro dry matter digestibility (IVDMD) and tissue [N] were detected only in certain years, varied from negative to positive, and were relatively minor. In contrast, eCO2 substantially reduced IVDMD (two most abundant grasses) and [N] (all three dominant grass species) in most years, except the two wettest years. Furthermore, eCO2 reduced IVDMD and [N] independent of warming effects. Reduced IVDMD with eCO2 was related both to reduced [N] and increased acid detergent fiber (ADF) content of grass tissues. For the six most abundant forage species (representing 96% of total forage production), combined warming and eCO2 increased forage production by 38% and reduced forage [N] by 13% relative to ambient climate. Although the absolute magnitude of the decline in IVDMD and [N] due to combined warming and eCO2 may seem small (e.g., from 63.3 to 61.1% IVDMD and 1.25 to 1.04% [N] for Pascopyrum smithii), such shifts could have substantial consequences for the rate at which ruminants gain weight during the primary growing season in the largest remaining rangeland ecosystem in North America. With forage production increases, declining forage quality could potentially be mitigated by adaptively increasing stocking rates, and through management such as prescribed burning, fertilization at low rates, and legume interseeding to enhance forage quality.

Spatial vegetation patterns and neighborhood competition among woody plants in an East African savanna.

The majority of research on savanna vegetation dynamics has focused on the coexistence of woody and herbaceous vegetation. Interactions among woody plants in savannas are relatively poorly understood. We present data from a 10-yr longitudinal study of spatially explicit growth patterns of woody vegetation in an East African savanna following exclusion of large herbivores and in the absence of fire. We examined plant spatial patterns and quantified the degree of competition among woody individuals. Woody plants in this semiarid savanna exhibit strongly clumped spatial distributions at scales of 1-5 m. However, analysis of woody plant growth rates relative to their conspecific and heterospecific neighbors revealed evidence for strong competitive interactions at neighborhood scales of up to 5 m for most woody plant species. Thus, woody plants were aggregated in clumps despite significantly decreased growth rates in close proximity to neighbors, indicating that the spatial distribution of woody plants in this region depends on dispersal and establishment processes rather than on competitive, density-dependent mortality. However, our documentation of suppressive effects of woody plants on neighbors also suggests a potentially important role for tree-tree competition in controlling vegetation structure and indicates that the balanced-competition hypothesis may contribute to well-known patterns in maximum tree cover across rainfall gradients in Africa.

Grazing intensity differentially regulates ANPP response to precipitation in North American semiarid grasslands.

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.

Recovery of African wild dogs suppresses prey but does not trigger a trophic cascade.

Increasingly, the restoration of large carnivores is proposed as a means through which to restore community structure and ecosystem function via trophic cascades. After a decades-long absence, African wild dogs (Lycaon pictus) recolonized the Laikipia Plateau in central Kenya, which we hypothesized would trigger a trophic cascade via suppression of their primary prey (dik-dik, Madoqua guentheri) and the subsequent relaxation of browsing pressure on trees. We tested the trophic-cascade hypothesis using (1) a 14-year time series of wild dog abundance; (2) surveys of dik-dik population densities conducted before and after wild dog recovery; and (3) two separate, replicated, herbivore-exclusion experiments initiated before and after wild dog recovery. The dik-dik population declined by 33% following wild dog recovery, which is best explained by wild dog predation. Dik-dik browsing suppressed tree abundance, but the strength of suppression did not differ between before and after wild dog recovery. Despite strong, top-down limitation between adjacent trophic levels (carnivore-herbivore and herbivore-plant), a trophic cascade did not occur, possibly because of a time lag in indirect effects, variation in rainfall, and foraging by herbivores other than dik-dik. Our ability to reject the trophic-cascade hypothesis required two important approaches: (1) temporally replicated herbivore exclusions, separately established before and after wild dog recovery; and (2) evaluating multiple drivers of variation in the abundance of dik-dik and trees. While the restoration of large carnivores is often a conservation priority, our results suggest that indirect effects are mediated by ecological context, and that trophic cascades are not a foregone conclusion of such recoveries.

Functional response of U.S. grasslands to the early 21st-century drought.

Grasslands across the United States play a key role in regional livelihood and national food security. Yet, it is still unclear how this important resource will respond to the prolonged warm droughts and more intense rainfall events predicted with climate change. The early 21st-century drought in the southwestern United States resulted in hydroclimatic conditions that are similar to those expected with future climate change. We investigated the impact of the early 21st-century drought on aboveground net primary production (ANPP) of six desert and plains grasslands dominated by C4 (warm season) grasses in terms of significant deviations between observed and expected ANPP. In desert grasslands, drought-induced grass mortality led to shifts in the functional response to annual total precipitation (P(T)), and in some cases, new species assemblages occurred that included invasive species. In contrast, the ANPP in plains grasslands exhibited a strong linear function of the current-year P(T) and the previous-year ANPP, despite prolonged warm drought. We used these results to disentangle the impacts of interannual total precipitation, intra-annual precipitation patterns, and grassland abundance on ANPP, and thus generalize the functional response of C4 grasslands to predicted climate change. This will allow managers to plan for predictable shifts in resources associated with climate change related to fire risk, loss of forage, and ecosystem services.

Competition and facilitation between a native and a domestic herbivore: trade-offs between forage quantity and quality.

Potential competition between native and domestic herbivores is a major consideration influencing the management and conservation of native herbivores in rangeland ecosystems. In grasslands of the North American Great Plains, black-tailed prairie dogs (Cynomys ludovicianus) are widely viewed as competitors with cattle but are also important for biodiversity conservation due to their role in creating habitat for other native species. We examined spatiotemporal variation in prairie dog effects on growing-season forage quality and quantity using measurements from three colony complexes in Colorado and South Dakota and from a previous study of a fourth complex in Montana. At two complexes experiencing below-average precipitation, forage availability both on and off colonies was so low (12-54 g/m2) that daily forage intake rates of cattle were likely constrained by instantaneous intake rates and daily foraging time. Under these dry conditions, prairie dogs (1) substantially reduced forage availability, thus further limiting cattle daily intake rates, and (2) had either no or a small positive effect on forage digestibility. Under such conditions, prairie dogs are likely to compete with cattle in direct proportion to their abundance. For two complexes experiencing above-average precipitation, forage quantity on and off colonies (77-208 g/m2) was sufficient for daily forage intake of cattle to be limited by digestion rather than instantaneous forage intake. At one complex where prairie dogs enhanced forage digestibility and [N] while having no effect on forage quantity, prairie dogs are predicted to facilitate cattle mass gains regardless of prairie dog abundance. At the second complex where prairie dogs enhanced digestibility and [N] but reduced forage quantity, effects on cattle can vary from competition to facilitation depending on prairie dog abundance. Our findings show that the high spatiotemporal variation in vegetation dynamics characteristic of semiarid grasslands is paralleled by variability in the magnitude of competition between native and domestic grazers. Competitive interactions evident during dry periods may be partially or wholly offset by facilitation during periods when forage digestibility is enhanced and forage quantity does not limit the daily intake rate of cattle.

Assessing herbivore foraging behavior with GPS collars in a semiarid grassland.

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.

Associations of grassland bird communities with black-tailed prairie dogs in the North American Great Plains.

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.

Nitrogen cycling and water pulses in semiarid grasslands: are microbial and plant processes temporally asynchronous?

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.

Rhizosphere interactions, carbon allocation, and nitrogen acquisition of two perennial North American grasses in response to defoliation and elevated atmospheric CO2.

Carbon allocation and N acquisition by plants following defoliation may be linked through plant-microbe interactions in the rhizosphere. Plant C allocation patterns and rhizosphere interactions can also be affected by rising atmospheric CO(2) concentrations, which in turn could influence plant and microbial responses to defoliation. We studied two widespread perennial grasses native to rangelands of western North America to test whether (1) defoliation-induced enhancement of rhizodeposition would stimulate rhizosphere N availability and plant N uptake, and (2) defoliation-induced enhancement of rhizodeposition, and associated effects on soil N availability, would increase under elevated CO(2). Both species were grown at ambient (400 µL L(-1)) and elevated (780 µL L(-1)) atmospheric [CO(2)] under water-limiting conditions. Plant, soil and microbial responses were measured 1 and 8 days after a defoliation treatment. Contrary to our hypotheses, we found that defoliation and elevated CO(2) both reduced carbon inputs to the rhizosphere of Bouteloua gracilis (C(4)) and Pascopyrum smithii (C(3)). However, both species also increased N allocation to shoots of defoliated versus non-defoliated plants 8 days after treatment. This response was greatest for P. smithii, and was associated with negative defoliation effects on root biomass and N content and reduced allocation of post-defoliation assimilate to roots. In contrast, B. gracilis increased allocation of post-defoliation assimilate to roots, and did not exhibit defoliation-induced reductions in root biomass or N content. Our findings highlight key differences between these species in how post-defoliation C allocation to roots versus shoots is linked to shoot N yield, but indicate that defoliation-induced enhancement of shoot N concentration and N yield is not mediated by increased C allocation to the rhizosphere.