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Questions: Grasslands provide important provisioning services worldwide and their management has consequences for these services. Management intensification is a widespread land‐use change and has accelerated across North America to meet rising demands on productivity, yet its impact on the relationship between plant diversity and productivity is still unclear. Here, we investigated the relationship between plant diversity and grassland productivity across nine ecoclimatic domains of the continental United States. We also tested the effect of management intensification on diversity and productivity in four case studies. Methods: We acquired remotely sensed gross primary productivity data (GPP, 1986–2018) and plant diversity data measured at different spatial scales (1, 10, 100, 400 m2), as well as climate variables including the Palmer drought index from two ecological networks. We used general linear mixed models to relate GPP to plant diversity across sites. For the case study analysis, we used linear mixed models to relate plant diversity to management intensity, and tested if the management intensity influenced the relationship between GPP (mean and temporal variation) and drought. Results: Across all sites, we observed positive relationships among species richness, productivity, and the temporal stability of mean annual biomass production. These relationships were not affected by the scale at which species richness was observed. In three out of the four case studies, we observed that management effects on species richness were only significant at broader scales (i.e., ≥10 m2) with no clear effect found at the commonly used 1‐m2 quadrat scale. In one case study, species‐poor, intensively managed pastures presented the highest productivity but were more sensitive to dry conditions than less intensified pastures. However, in other case studies, we did not observe significant effects of management intensity on the magnitude or stability of productivity. Conclusions: Generalization across studies may be difficult and require the development of intensification indices general enough to be applied across diverse management strategies in grazilands. Understanding how management intensification affects grassland productivity will inform the development of sustainable intensification strategies. [ABSTRACT FROM AUTHOR]

Context: Technologies are now available to continuously monitor livestock foraging behaviours, but it remains unclear whether such measurements can meaningfully inform livestock grazing management decisions. Empirical studies in extensive rangelands are needed to quantify relationships between short-term foraging behaviours (e.g. minutes to days) and longer-term measures of animal performance. Aims: The objective of this study was to examine whether four different ways of measuring daily foraging behaviour (grazing-bout duration, grazing time per day, velocity while grazing, and turn angle while grazing) were related to weight gain by free-ranging yearling steers grazing semiarid rangeland. Methods: Yearling steers were fitted with neck collars supporting a solar-powered device that measured GPS locations at 5 min intervals and used an accelerometer to predict grazing activity at 4 s intervals. These devices were used to monitor steers in four different paddocks that varied in forage biomass, and across two grazing seasons encompassing a wide range of forage conditions. Steer weight gain (kg/steer.day) was measured in each paddock during each of three ∼60 day time intervals, and daily foraging behaviours were measured during 15–21 days in each interval. Results: A model based on only two daily measurements of foraging behaviour, mean grazing bout duration (calculated at a 5 min resolution) and mean velocity while grazing explained 62% of the variation in animal weight gain. Conclusions: Daily measurements of foraging behaviour vary substantially in response to varying foraging conditions in space and time, and can effectively serve as indicators of variation in cattle weight gain. Implications: On-animal sensors that monitor foraging behaviour have the potential to transmit indicators to livestock managers in real time (e.g. daily) to help inform decisions such as when to move animals among paddocks, or when to sell or transition animals from rangeland to confined feeding operations. Empirical studies in extensive rangelands are needed to quantify relationships between short-term foraging behaviours (e.g. minutes to days) and longer-term measures of animal performance. We found that two daily measurements of foraging behaviour, mean grazing-bout duration and mean velocity while grazing (both calculated at a 5 min resolution), can explain 62% of the variation in weight gains of free-ranging yearling steers grazing semiarid rangeland. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

The conservation and management of black‐tailed prairie dogs (Cynomys ludovicianus) have been contentious issues in grasslands of central North America for much of the past century, primarily because of the perception that they compete with livestock for forage. Studies quantifying the magnitude of competition between prairie dogs and cattle are difficult to conduct because of the large spatial and long temporal scales needed to quantify how competition varies in response to interannual variation in precipitation and prairie dog abundance. We examined variation in mass gains of yearling steers in shortgrass steppe of northeastern Colorado, USA, with and without prairie dogs from 2008–2019, a period that encompassed a full cycle in prairie dog abundance from a nadir following plague‐induced population collapse, to peak abundance following population recovery, to plague‐induced population lows again. Analyses of cattle grazing distribution with global positioning system (GPS)‐collars revealed preferential grazing on colonies following a period of unusually high vegetation production, and preferential grazing off colonies following a period of rapid vegetation senescence, but these patterns were not clearly related to cattle mass gains. Across all 12 years of the study, average daily mass gain (ADG) during the growing season was 0.97 kg/steer/day in pastures where prairie dogs were controlled annually, and 0.95 kg/steer/day in pastures where they were not. Average daily mass gain was a quadradic function of precipitation and a linear function of prairie dog occupancy within a pasture, with a generalized linear mixed model predicting an 8.0% decrease in ADG as prairie dog occupancy increased from 0 to 60% of a pasture with average growing‐season precipitation. We did not detect a significant interaction between precipitation and prairie dog occupancy, but one limitation of our study is that the only drought year (2012) occurred when prairie dogs occupied low percentages (10–25%) of the study pastures. Prairie dogs had a small but detectable negative effect on cattle mass gains during the growing season in shortgrass steppe. The magnitude of this effect can be used by managers in combination with market conditions and the spatial extent of prairie dog colonies to estimate economic effects of prairie dogs on livestock operations. © 2021 The Wildlife Society. This article is a U.S. Government work and is in the public domain in the USA. : Mass gain of cattle during the growing season declined linearly by 8% as prairie dog occupancy of shortgrass pastures increased from 0 to 60%. Decisions on the need to manage prairie dog populations could be based on the magnitude of the effect documented in our study, market values of cattle, and the costs of prairie dog control, but managers should also consider the potential for drought to coincide with peak prairie dog abundance, the frequency and timing of plague epizootics in the prairie dog population, and costs of alternative forage sources during drought. [ABSTRACT FROM AUTHOR]

Questions: What are the rate, reversibility, and degree of symmetry in plant species compositional change in response to the addition and removal of cattle grazing in the shortgrass steppe? Specifically, how does the imposition and removal of grazing affect the abundance of perennial C4 shortgrasses and C3 midgrasses that are of primary importance for livestock production in the region? Location: Shortgrass steppe, northeastern Colorado, USA, in the North American Great Plains. Methods: We evaluate rates and magnitude of basal cover change in newly ungrazed and newly grazed sites (established in 1991), relative to change in long‐term (grazed and ungrazed) comparison treatments (established in 1939) over 25 years. We also compare shifts in species basal cover in newly implemented treatments relative to baseline community composition measured at the start of the study. Results: Unlike the limited change observed in long‐term treatments between 1939 and 1991, we documented more rapid, reversible and largely symmetric effects of both the imposition and removal of grazing between 1992 and 2017. This was primarily due to differences in the rate of increase in basal cover of C3 midgrasses, litter, and bare ground. However, the rate and direction of change differed when assessed relative to continuously evaluated and (early‐treatment) baseline cover data. Conclusions: Studies of plant community change in response to large‐herbivore grazing under varying ecological conditions and management regimes are needed to effectively guide the development and revision of state‐and‐transition models (STMs) for grassland managers, and to better detect and plan for dynamic ecological conditions. Effective adaptive management and STM development under a changing climate will recognize that effects of grazing and removal of grazing on shortgrass steppe can be reversible in a relatively symmetrical pattern, occurring within 6–16 years. [ABSTRACT FROM AUTHOR]

North American sagebrush (Artemisia spp.)‐obligate birds are experiencing steep population declines due in part to increased disturbance, mainly human‐caused, across their range. At the eastern edge of the sagebrush steppe, this issue may potentially be exacerbated because of natural disturbance by black‐tailed prairie dogs (Cynomys ludovicianus). Our goal was to compare local and landscape models of habitat use by greater sage‐grouse (Centrocercus urophasianus), Brewer's sparrow (Spizella breweri), and sage thrasher (Oreoscoptes montanus) with models including effects of natural (i.e., prairie dog) and anthropogenic disturbance. We used a combination of field data collection, and state and national datasets for the Thunder Basin National Grassland, eastern Wyoming, USA, to understand the factors that influence lek attendance by sage‐grouse and habitat use by 2 passerines in this system. For all 3 species, models including big sagebrush (Artemisia tridentata) cover at local and landscape scales were the most competitive among univariate models, supporting the paradigm that sagebrush is key for these species. Models including anthropogenic disturbance (well density, road density) explained more variation than models of prairie dog disturbance alone for 2 of the 3 species, but long‐term disturbance by prairie dogs did reduce abundance of Brewer's sparrows. Although long‐term prairie dog disturbance has the potential to reduce sagebrush cover for sagebrush‐obligate birds, such events are likely rare because outbreaks of plague (Yersina pestis) and lethal control on borders with private land reduce prairie dog disturbance. Conversely, anthropogenic disturbance is slated to increase in this system, suggesting potentially accelerated declines for sagebrush birds into the future. © 2020 The Wildlife Society. Imperiled sagebrush songbirds and greater sage‐grouse are less likely to occupy disturbed habitat in northeastern Wyoming. Although these species are sensitive to both natural (prairie dog) and anthropogenic disturbance, the latter appears to play a larger role in habitat use in this region. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Learning is recognized as central to collaborative adaptive management (CAM), yet few longitudinal studies examine how learning occurs in CAM or apply the science of learning to interpret this process. We present an analysis of decision-making processes within the collaborative adaptive rangeland management (CARM) experiment, in which 11 stakeholders use a structured CAM process to make decisions about livestock grazing and vegetation management for beef, vegetation, and wildlife objectives. We analyzed four years of meeting transcripts, stakeholder communications, and biophysical monitoring data to ask what facilitated and challenged stakeholder decision making, how challenges affected stakeholder learning, and whether CARM met theorized criteria for effective CAM. Despite thorough monitoring and natural resource agency commitment to implementing collaborative decisions, CARM participants encountered multiple decision-making challenges born of ecological and social complexity. CARM was effective in achieving several of its management objectives, including reduced ecological uncertainty, knowledge coproduction, and multiple-loop social learning. CARM revealed limitations of the idealized CAM cycle and challenged conceptions of adaptive management that separate reduction of scientific uncertainty from participatory and management dimensions. We present a revised, empirically grounded CAM framework that depicts CAM as a spiral rather than a circle, where feedback loops between monitoring data and management decisions are never fully closed. Instead, complexities including time-lags, trade-offs, path-dependency, and tensions among stakeholders' differing types of knowledge and social worlds both constrain decision making and foster learning by creating disorienting dilemmas that challenge participants' pre-existing mental models and relationships. Based on these findings, we share recommendations for accelerating learning in CAM processes. [ABSTRACT FROM AUTHOR]

Abstract: 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. [ABSTRACT FROM AUTHOR]

ABSTRACT: 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. [ABSTRACT FROM AUTHOR]

Sustainable management of grazinglands depends upon understanding how management practices influence livestock movements in space and time. We conducted a ranch-scale (2,600-ha) social-ecological experiment to examine how foraging behavior of cattle differs between a single large herd rotated adaptively among paddocks (collaborative, adaptive rangeland management; CARM) versus continuous, season-long grazing of paddocks by small non-rotational herds (traditional rangeland management; TRM). We analyzed how differences in cattle movement patterns may be linked to reductions in cattle growth rates and diet quality in the CARM treatment, relative to TRM. Cattle in the CARM treatment exhibited more linear grazing pathways, moved at lower velocity while grazing, and exhibited longer grazing bouts early in the growing season compared to TRM cattle. Grazing time within any given 10 × 10 m area was distributed more unevenly across TRM vs. CARM paddocks in years with average or above-average precipitation, but not in dry years. In all years, areas of high and low grazing intensity were more spatially clustered in TRM than CARM paddocks. Movement patterns of cattle managed using adaptive, multi-paddock rotations at high stock density (CARM) are consistent with less selective foraging. Such cattle form a "grazing front" that moves across the paddock and distributes grazing pressure in a more spatially homogeneous fashion. In years with substantial forage production, TRM cattle spent more time than CARM cattle in preferred areas of the paddock and foraged in more circular patches. In dry years, however, both treatments resulted in similarly even grazing distribution, likely due to limited intra-paddock variation in forage quality and quantity. At the ranch scale, these different intra-paddock movement patterns led to reductions in animal growth rates and no overall effect of grazing management on forage production. • Rotational herds altered grazing patterns relative to season-long grazing herds. • Rotational herds grazed more linear pathways, at increased velocity and length. • Rotational herds grazed across pastures more evenly in wet years but not dry years. • Foraging indicated less diet selectivity, consistent with reduced protein intake. • Rotational management reduced animal growth, without boosting forage production. [ABSTRACT FROM AUTHOR]

Questions How does long-term exclusion of cattle grazing influence plant community composition in a semi-arid grassland? Can spatial variation in the effects of cattle grazing exclusion be explained by variation in soil texture? Location Shortgrass steppe, northeast Colorado, USA, 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 (one to two 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 co-exist 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 sub-shrubs have increased in relative abundance in both grazed and ungrazed communities. Conclusions Our long-term experiment shows that community responses to grazing in this semi-arid 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 mono-dominant 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]. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

African savannas are complex socio-ecological systems with diverse wild and domestic herbivore assemblages, which adapt spatially to intra- and interannual variation in forage quantity and quality, predation and disease risks., As African savannas become increasingly fragmented by growing human populations and their associated ecological impacts, adaptive foraging options for wild and domestic herbivore populations are correspondingly limited, resulting in declining wildlife populations and impoverished pastoral societies. In addition, competition for grazing by expanding domestic herbivore populations threatens the viability of wild herbivore populations occupying similar grazing niches., Conservation initiatives are further impacted by conflicts between wildlife and local communities of people who often receive little benefit from adjacent protected areas, creating conflict between the livelihood-orientated goals of communities and the conservation-oriented goals of the international community and those with vested interests in wildlife. Conservation strategies facilitating the alignment of these opposing goals of communities and conservationists are needed., Synthesis and applications. Key to understanding facilitative and competitive interactions between wild and domestic herbivores are the concepts of niche differentiation and functional resource heterogeneity. Uncontrolled incursions of burgeoning domestic herbivore populations into protected areas ( PAs) threaten the conservation of wild herbivore biodiversity. However, domestic herbivores can be managed to minimize competition with wild herbivores and to enhance habitat by maximizing grassland structural heterogeneity (greater adaptive foraging options), creation of nutrient hotspots in the landscape and facilitation of high-quality grazing. Ecosystem service benefits to communities through controlled access to grazing resources in PAs, associated with appropriate disease management, can provide a conservation payment to promote communities' support of conservation of key wildlife migratory ranges and corridors outside PAs. [ABSTRACT FROM AUTHOR]

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 C3grassesversuswarm-season C4grasses, and with prior research suggesting that early season production by C3grasses is more responsive to a given amount of precipitation than mid-summer growth of C4shortgrasses. 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. [ABSTRACT FROM PUBLISHER]

ABSTRACT 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 (<4 cm), sparse vegetation on recent burns. Although cattle selectively grazed recent burns, this did not alter vegetation structure in unburned portions of patch-burned pastures relative to controls. Of the 6 grassland bird species we examined, mountain plovers ( Charadrius montanus) occurred exclusively on recent burns, whereas grasshopper sparrows ( Ammodramus savannarum) occurred exclusively in grassland not burned for ≥3 years. Two species (lark bunting [ Calamospiza melanocorys] and western meadowlark [ Sturnella neglecta]) were 2-3 times less abundant on recent burns compared to controls, whereas densities of horned larks ( Eremophila alpestris) and McCown's longspur ( Rhynchophanes mccownii) were unaffected by burning. Lark bunting, western meadowlark, and grasshopper sparrow densities varied substantially among years. In the years when they were abundant, all 3 species increased in density across the time-since-burning gradient. Consistent with this pattern, patch-burn grazing management reduced the abundance of all 3 species at the whole-pasture scale. We found no evidence that unburned patches within the patch-burned pastures differed from unburned pastures in terms of the abundance of any bird species. Patch-burn grazing management was an effective strategy to create breeding habitat for mountain plovers. However, our findings suggest that in the shortgrass steppe, additional strategies that generate taller, more dense vegetation than occurs under moderate cattle grazing need to be considered in combination with patch-burn grazing management to sustain breeding habitat for the full suite of native grassland birds. Published 2015. This article is a U.S. Government work and is in the public domain in the USA. [ABSTRACT FROM AUTHOR]

Abstract: 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. [Copyright &y& Elsevier]

ABSTRACT 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 (5.9 birds/km2; 95% CI = 4.7-7.4) and sites burned during the preceding dormant season (6.7 birds/km2; 95% CI = 4.6-9.6), 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 of disturbances (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. [ABSTRACT FROM AUTHOR]

The degree to which large herbivores select and forage within recently burned areas is a key driver of vegetation heterogeneity in rangeland ecosystems. However, few studies have quantified the strength and timing of herbivore selection for burned areas or examined how selection strength varies among ecosystems differing in precipitation and primary productivity., We conducted a 4-year patch-burning experiment in semi-arid rangeland of Colorado, USA, where 25% of the area available to cattle was burned each year and burned patches were shifted annually. We used GPS collars with activity sensors to quantify the distribution of free-ranging cattle at a high temporal resolution (5-min intervals) during the growing season each year. We used a classification tree model to discriminate between cattle grazing vs. non-grazing locations, which significantly increased precision in quantifying burn selection strength. We fit generalized linear models predicting the frequency of cattle use of a given location within each study area and month, enabling comparisons between the relative influence of burns and topography on grazing distribution., Across multiple growing seasons, cattle selectively spent 31% of grazing time on recently burnt areas, which comprised 25% of the landscape; this selection strength was half as strong as that documented in mesic rangeland., At a monthly temporal scale, strong cattle selection for burned areas occurred during periods of rapid vegetation growth regardless of when during the growing season this greening occurred. Outside these intervals, burn selection strength was inconsistent and cattle grazing distribution was primarily influenced by topography. Thus, the relative importance of fire and topography in controlling grazer distribution was temporally contingent upon the timing and size of precipitation pulses., Synthesis and applications. Spatiotemporal interactions between fire and herbivores are a consistent feature of both semi-arid and mesic rangelands, with interaction strength varying across gradients of precipitation and primary productivity. Management of semi-arid ecosystems to sustain ecological processes should include strategies that allow ungulate herbivores to shift their grazing distribution seasonally in response to fire, topoedaphic variation and precipitation patterns. Combined management of fire and grazing for conservation objectives can be consistent with, and even complementary to, livestock production goals. [ABSTRACT FROM AUTHOR]

Large mammalian herbivores are well recognized to play important roles in regulating woody cover and biomass in savannas, but the extent to which browsing ungulates are capable of regulating woody populations in the absence of other disturbances such as fire is unclear. Moreover, the degree to which browser effects on savannas operate through effects on woody plant recruitment vs. mortality has rarely been examined., We conducted a 10-year, replicated herbivore exclusion experiment in a semi-arid savanna in East Africa (mean annual rainfall = 514 mm), where fires have been actively suppressed for decades. Browsers dramatically influenced recruitment, growth and mortality of all size classes of woody vegetation. A decade of herbivore exclusion resulted in a sevenfold increase in recruitment, a 2.5-fold decrease in mortality and a threefold increase in woody biomass inside exclosures, while biomass outside exclosures remained relatively unchanged., At the plant community level, extensive browsing of seedlings by small-bodied ungulates suppressed woody recruitment in this semi-arid system, generating a 'browsing trap' comparable to the 'fire trap' reported for mesic systems. Browsing by large- and medium-bodied ungulates reduced both growth and survival of individuals in larger size classes., At the plant species level, browser impacts were variable. Although browsers negatively influenced recruitment of all species, they had little to no impact on the mortality of some dominant species, resulting in a long-term, browser-driven shift in woody species composition that was largely mediated via their differential effects on plant mortality rates rather than recruitment., Synthesis. Our results demonstrate unequivocally that, even in the absence of fire, native browsing ungulates can exert dramatic 'top-down' controls in semi-arid savannas, influencing all aspects of woody plant demography. Besides suppressing woody plant recruitment, browsers can also have substantial cumulative long-term impacts on growth and mortality rates of woody plants, including adults, which can differ between species in ways that fundamentally alter the structure and function of woody vegetation. In semi-arid rangelands, intact communities of native browsing ungulates thus provide a critical ecosystem service by regulating woody cover, and their removal (or extinction) from these systems can lead to rapid woody encroachment. [ABSTRACT FROM AUTHOR]

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 América [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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 effects 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. The 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 effects 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 effects on grass cover, which may reflect the dominance of grazing-tolerant shortgrasses at both complexes. Variation in modification of vegetation structure may be understood in terms of the responses of different dominant perennial grasses to intense defoliation and differences 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. Thus, ecosystem engineering can have a spatial component not evident from single-place studies. [ABSTRACT FROM AUTHOR]

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 (<4 cm tall) vegetated patches interspersed with >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 management to provide multiple ecosystem goods and services. © 2012 The Wildlife Society. [ABSTRACT FROM AUTHOR]

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 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. Both species were grown at ambient (400 μL L) and elevated (780 μL L) atmospheric [CO] 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 both reduced carbon inputs to the rhizosphere of Bouteloua gracilis (C) and Pascopyrum smithii (C). 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. [ABSTRACT FROM AUTHOR]

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 prairie dog colonies affected by epizootic plague in the past 1-2 years, and 4) rangeland with no recent history of fire or prairie dogs. Mountain plover densities were similar on active black-tailed prairie dog colonies (X¯ = 6.8 birds/km², 95% CI = 4.3-10.6) and prescribed burns (X&#x00AF = 5.6 birds/km², 95% CI = 3.5-9.1). In contrast, no plovers were detected at randomly selected rangeland sites grazed by cattle but lacking recent disturbance by prairie dogs or fire, even though survey effort was highest for this rangeland habitat. Mountain plover densities were intermediate (2.0 birds/km², 95% CI = 0.8-5.0) on sites where black-tailed prairie dogs had recently been extirpated by plague. These findings suggest that prescribed burns and active black-tailed prairie dog colonies may enhance breeding habitat for mountain plovers in shortgrass steppe and illustrate the potential for suppressed or altered disturbance processes to influence habitat availability for declining wildlife species. [ABSTRACT FROM AUTHOR]

The distribution and abundance of native ungulates were measured on commercially managed, semi-arid rangeland in central Kenya over a 3-year period that encompassed severe drought and above-average rainfall. Native ungulate biomass density averaged 5282 kg km over the study and was dominated by elephant ( Loxodonta africana), impala ( Aepyceros melampus) and dik-dik ( Madoqua kirkii). Biomass density of domestic cattle ( Bos taurus) averaged 2280 kg km during the study. Responses of native ungulates to severe drought were variable. Impala densities were similar to or greater than densities for similar habitat in protected areas, and varied from 12 to 16 km during and following the drought to 24-29 km following above-average rainfall. Dik-dik densities were also greater than densities reported for protected areas and were surprisingly stable throughout the study despite the wide annual fluctuations in rainfall. Elephant migrated out of the region during drought but were present at high densities (2.9-5.2 km) during wet seasons, consistent with telemetry studies emphasizing the importance of Acacia bushland habitat on commercial rangelands for the migratory portion of the Laikipia-Samburu elephant population. Results show that substantial densities of native browsing and mixed-feeding ungulates can occur on rangeland managed for commercial beef production and suggest that the capacity for ungulates to move over large spatial scales (>100 km) and to shift distributions in response to locally variable thunderstorms may be important for sustaining these populations. [ABSTRACT FROM AUTHOR]

A central goal of ecosystem ecology is to understand how the cycling of nutrients and the growth of organisms are linked. Ecologists have repeatedly observed that nutrient mineralization and plant production are closely coupled in time in many terrestrial ecosystems. Typically, mineralization rates of limiting nutrients, particularly of nitrogen, during the growing season determine nutrient availability while pools of mineral nutrients remain low and relatively constant. Although several previous reports suggest nitrogen mineralization has the potential lo vary seasonally and out of phase with plant production, such a phenomenon has been poorly documented. Here we report results from a semiarid savanna ecosystem characterized by distinct temporal asynchrony in rates of soil nitrogen cycling and plant production. Periods of positive plant growth following the onset of rains coincide with periods of low N turnover rates, whereas higher rates occur late in the wet season following plant senescence and throughout dry seasons. Plant uptake from the substantial mineral N pool present early in the growing season is sufficient to explain most of the N allocation to aboveground plant biomass during the growing season, even in the absence of any wet-season mineralization. The mineral N pool is subsequently recharged by late wet- and dry-season mineralization, plus urine inputs at sites with high levels of ungulate activity. These findings suggest fundamental changes in the quality of substrates available to decomposers over a seasonal cycle, with significant implications for the partitioning of limiting nutrients by plant species, the seasonal pattern of nutrient limitations of aboveground production, and the effective use of N fertilizers in semiarid ecosystems. [ABSTRACT FROM AUTHOR]

In sub-Saharan Africa, the widespread practice of corralling livestock overnight in thorn-scrub "bomas" creates nutrient-enriched patches within rangelands that can subsequently support unique plant communities for decades to centuries after boma abandonment. These nutrient-rich patches (glades) may be preferentially used by native ungulates that coexist with livestock. To evaluate the potential link between cattle management via bomas and habitat for impala (Aepyceros melampus), I examined seasonal patterns of habitat selection by impala and landscape variation in grass nutrient content on a commercial cattle ranch in central Laikipia, Kenya. Studies using automated, infrared camera monitors showed that impala selected nutrient-rich glades 2.6 times more frequently than surrounding Acacia bushland habitat during dry seasons, and 9.6 times more frequently during wet seasons. Significantly greater impala presence in glade versus bushland habitat during dry seasons suggests that impala presence may be related to reduced predation risk in shrub-free glades. The large, significant increase in impala presence in glades from dry to wet seasons suggests that impala distribution also is linked to the availability of nutrient-rich forage. In particular, grass nutrient analyses showed that wet-season phosphorus (P) concentrations in grasses throughout the bushland landscape (x = 2,125 mg P/kg dry matter, varying from 1,789 to 2,922 mg P/kg across topographic positions and from 1,508 to 3,215 mg P/kg among grass species) were below recommended levels for pregnant and lactating ruminants, while mean P concentrations in glade grasses (x ± SE = 5,346 ± 2.92 mg P/kg) exceeded recommended levels. Results suggest that management to increase the relocation rate and distribution of current cattle bomas can have a positive, long-term effect on the local distribution and abundance of impala. [ABSTRACT FROM AUTHOR]

Ecosystem-level studies of producer-decomposer interactions have focused primarily on plant production and soil texture as regulators of decomposer abundance but have rarely considered the role of grazers in mediating such interactions. Here, we conducted replicated exclosure experiments at both high and low levels of soil fertility to investigate the effects of large, mammalian grazers on decomposer biomass and activity patterns in a semiarid grazing ecosystem in Kenya. Within only two years of grazer exclusion, microbial biomass was greater in soil of fenced grassland across all levels of soil fertility. This consistent negative effect of grazers on microbial biomass occurred despite the fact that grazers stimulated aboveground plant production in nutrient-rich sites and depressed it in nutrient-poor sites. A consideration of all the potential pathways by which grazers influence decomposer populations suggests that observed grazing-induced reductions in microbial biomass were predominantly associated with a depression in the amount of plant carbon inputs to soils. Finally, across all study sites, microbial biomass was highly correlated with soil carbon content, suggesting that landscape-scale constraints on soil organic matter content and plant production overarch grazer effects on microbial abundance. Our results support previous ecosystem-level studies showing that microbial biomass and growth are constrained by plant production and soil C availability. In addition, our findings demonstrate that decomposer abundance can be influenced by an ecosystem's trophic structure, with significant reductions in microbial biomass occurring as a result of herbivores diverting plant carbon away from soils. [ABSTRACT FROM AUTHOR]

Journal of Applied Ecology (2004) 41, 45–58 [ABSTRACT FROM AUTHOR]

Summary 1. The effect of livestock on African rangelands has been a major focus of recent research, but little attention has been paid to the way livestock affects the distribution and availability of soil nutrients. In East African savannas, overnight containment of livestock in thorn-scrub corrals or ‘bomas’ concentrates large quantities of nutrients into small areas, potentially altering the landscape distribution of nitrogen (N) and phosphorus (P) in soils and plants. 2. This study was designed to (i) measure the density, turnover rates and soil nutrient concentrations of abandoned cattle bomas on nutrient-poor rangeland in central Kenya; (ii) determine whether long-term glades dominated by Cynodon plectostachyus are derived from abandoned bomas; and (iii) evaluate the effect of cattle bomas on the landscape-level distribution of N and P. 3. In the study area, glades (> 39 years old) averaged 0·71 ha in size and occurred at a density of 0·71 km-2 . Abandoned bomas (1–39 years since abandonment) averaged 0·39 ha and occurred at a density of 1·21 km-2 . During 1961–2000, no glades reverted to bushland vegetation, while 53 bomas were abandoned. 4. All characteristics of soils measured across a boma–glade chronosequence indicated glades were indeed derived from abandoned bomas. Soil N, P and organic matter quality in the surface (0–15 cm) layer were similar for glades and 30–39-year-old bomas, but were significantly enriched relative to surrounding bushland. In contrast, at 40–65 cm depth beneath bomas, glades and bushland, soil N was similar. The texture of surface soils from bomas, glades and bushland was similar, indicating glades were not derived from a unique parent material. 5. Leaves of C. plectostachyus from 12–24-year bomas and long-term glades were enriched in P, calcium (Ca) and N relative to leaves of Cynodon dactylon from nearby bushland sites.... [ABSTRACT FROM AUTHOR]

Examines the effects of migratory herbivores on spatial heterogeneity of soil nitrogen properties in a grassland ecosystem at the Yellow Stone National Park in the United States. Spatchiness of ungrazed grasslands; Importance of the recycling of nitrogen; Diversification of plant species effects on soils.

Rangelands are composed of patchy, highly dynamic herbaceous plant communities that are difficult to quantify across broad spatial extents at resolutions relevant to their characteristic spatial scales. Furthermore, differentiation of these plant communities using remotely sensed observations is complicated by their similar spectral absorption profiles. To better quantify the impacts of land management and weather variability on rangeland vegetation change, we analyzed high resolution hyperspectral data produced by the National Ecological Observatory Network (NEON) at a 6500-ha experimental station (Central Plains Experimental Range) to map vegetation composition and change over a 5-year timescale. The spatial resolution (1 m) of the data was able to resolve the plant community type at a suitable scale and the information-rich spectral resolution (426 bands) was able to differentiate closely related plant community classes. The resulting plant community class map showed strong accuracy results from both formal quantitative measurements (F1 75% and Kappa 0.83) and informal qualitative assessments. Over a 5-year period, we found that plant community composition was impacted more strongly by weather than by the rangeland management regime. Our work displays the potential to map plant community classes across extensive areas of herbaceous vegetation and use resultant maps to inform rangeland ecology and management. Critical to the success of the research was the development of computational methods that allowed us to implement efficient and flexible analyses on the large and complex data. [ABSTRACT FROM AUTHOR]

Aims: Livestock production is the most widespread land use globally and occurs across a diverse set of ecosystems. Variability in long‐term livestock grazing impacts across ecosystems is poorly characterized, particularly at larger spatial scales, despite strong relationships with various ecosystem services related to soil fertility and stabilization and vegetation productivity. Here we examine the effects of grazing on vegetation and the implications for resistance and resilience to global change. Methods: We use six long‐term research stations in the western United States, spanning two ecoregions, multiple ecosystems, and 311 total site‐years of research. Across these sites we evaluate convergence and divergence of vegetation response to grazing vs grazing removal, focusing on interactions with drivers of global change. Results: We found that at long time scales (multiple decades), grazing has numerous convergent and divergent effects across ecoregions and ecosystems. Similarity among precipitation patterns and plant traits linked to grazing and production timing were key elements explaining convergence or divergence in long‐term patterns of livestock grazing response. Ecosystem differences across western US rangelands are also associated with variable effects of grazing on resistance and resilience to invasive species and climate change. Conclusions: These results suggest that unique ecosystem or ecoregion responses to future global change may result from complex interactions between grazing and environmental factors, such as precipitation timing and plant traits. Adapting livestock and grazing management to specific ecosystem vegetation and climate variability is needed to manage for the myriad global changes affecting rangeland production and diversity.Livestock grazing has numerous long‐term convergent and divergent effects across western US rangeland ecoregions and ecosystems, due to complex interactions with environmental factors, such as precipitation and plant traits. Variability in grazing effects on resistance and resilience suggests that ecosystem‐specific adaptation strategies are needed to sustain rangeland production and diversity in response to global change drivers. [ABSTRACT FROM AUTHOR]

Examines studies from a wide range of ecosystems concerning the effects of ungulate herbivores on plant community composition. Effects of herbivore foraging selectivity; Effects of plant tolerance; Underlying influences on foraging selectivity and plant tolerance; Conclusions.

Studies how white-tailed deer can cause a change in the structure and composition of the forest in southcentral Minnesota. Methodology; Understory forbs; Statistical analysis; Discussion on the results of the study.

The effects of grazing by white-tailed deer (Odocoileus virginianus) on populations of Trillium spp. were examined in remnant, old-growth patches of the highly fragmented Big Woods forest ecosystem in southeastern Minnesota. We conducted three separate studies involving an exclosure experiment, transplant experiments, and comparisons of Trillium populations among study sites. The highest grazing intensity was observed where deer occurred at high overwinter concentrations (~25–35/km2); significantly lower grazing intensities occurred at low overwinter density (~5–10/km2). Deer focused their grazing on large, reproductive plants; at sites with high deer density, Trillium population structure was skewed toward small plants, and deer consistently caused over 50% reduction in reproduction during the growing season. Protection of individual plants from deer for two growing seasons resulted in dramatically increased flowering rates and significantly greater leaf area compared to control plants. No significant impact of current-year herbivory on reproduction in the following year was detected. Nevertheless, flowering rates at one site with high overwinter deer densities for at least the past 5 years suggest that the cumulative effects of grazing over several years can reduce reproduction in subsequent years. Transplant experiments with Trillium grandiflorum also showed that deer had significant effects on growth and reproduction where deer occur at high density. Our results suggest that changes in landscape structure and local deer abundance have altered plant-deer relationships such that grazing can lead to the local extirpation of sensitive forbs such as Trillium spp. As a result, active, long-term management of deer at low densities appears necessary for the conservation and restoration of fragmented forest communities in eastern North America. Efectos del Venado Cola Blanca en Poblaciones de Trillium spp. en Bosques Decidous... [ABSTRACT FROM AUTHOR]

Examines the interactions between the Chlamydia psittaci infection and koalas. Sensitivity of koala population persistence to reduced annual survivorship of adults; Result of habitat fragmentation; Coexistence of host and parasite in transmission parameter values of koala.

The accurate estimation of aboveground net herbaceous production (ANHP) is crucial in rangeland management and monitoring. Remote and rural rangelands typically lack direct observation infrastructure, making satellite-derived methods essential. When ground data are available, a simple and effective way to estimate ANHP from satellites is to derive the empirical relationship between ANHP and plant-absorbed photosynthetically active radiation (APAR), which can be estimated from the normalized difference vegetation index (NDVI). While there is some evidence that this relationship will differ across rangeland vegetation types, it is unclear whether this relationship will change across grazing management regimes. This study aimed to assess the impact of grazing management on the relationship between ground-observed ANHP and satellite-derived APAR, considering variations in plant communities across ecological sites in the shortgrass steppe of northeastern Colorado. Additionally, we compared satellite-predicted biomass production from the process-based Rangeland Analysis Platform (RAP) model to our empirical APAR-based model. We found that APAR could be used to predict ANHP in the shortgrass steppe, with the relationship being influenced by ecosite characteristics rather than grazing management practices. For each unit of added APAR (MJ m−2 day−1), ANHP increased by 9.39 kg ha−1, and ecosites with taller structured herbaceous vegetation produced, on average, 3.92–5.71 kg ha−1 more ANHP per unit APAR than an ecosite dominated by shorter vegetation. This was likely due to the increased allocation of plant resources aboveground for C3 mid-grasses in taller structured ecosites compared to the C4 short-grasses that dominate the shorter structured ecosites. Moreover, we found that our locally calibrated empirical model generally performed better than the continentally calibrated process-based RAP model, though RAP performed reasonably well for the dominant ecosite. For our empirical models, R2 values varied by ecosite ranging from 0.49 to 0.67, while RAP R2 values ranged from 0.07 to 0.4. Managers in the shortgrass steppe can use satellites to estimate herbaceous production even without detailed information on short-term grazing management practices. The results from our study underscore the importance of understanding plant community composition for enhancing the accuracy of remotely sensed predictions of ANHP. [ABSTRACT FROM AUTHOR]

Recent work suggests that savanna woody plant species utilise two different strategies based on their defences against herbivory; a low nutrient/high chemical defence strategy and a nutrition paired with mostly architectural defences strategy. The concept that chemical and structural defences can augment each other and do not necessarily trade‐off has emanated from this work. In this study, we examine woody plant defence strategies, how these respond to herbivore removal and how they affect plant growth in an East African savanna.At three paired long‐term exclosure sites with high browser and mixed‐feeder densities at Mpala Ranch, Kenya, we investigated: (a) whether defences employed by the dominant fine‐ and broad‐leaved woody savanna species form defence strategies and if these align with previously proposed strategies, (b) how nine key plant defence traits respond to herbivore removal and (c) how effective the different defence strategies are at protecting against intense herbivory (by measuring plant growth with and without herbivores present).We identified three defence strategies. We found a group (a) with high N, short spines and high N‐free secondary metabolites, a group (b) with high N, long spines and low N‐free secondary metabolites and a group (c) with moderate N, no spines and low N‐free secondary metabolites (most likely defended by unmeasured chemical defences). Structural defences (spine length, branching) were generally found to be induced by herbivory, leaf available N increased or did not respond, and N‐free secondary metabolites decreased or did not respond to herbivory. Species with long spines combined with increased "caginess" (dense canopy architecture arising from complex arrangement of numerous woody and spiny axis categories) of branches, maintained the highest growth under intense browsing, compared to species with short spines and high N‐free secondary metabolites and species with no spines and low N‐free secondary metabolites.Synthesis. At our study site, structural traits (i.e. spines, increased caginess) were the most inducible and effective defences against intense mammalian herbivory. We propose that high levels of variability in the way that nutrient and defence traits combine may contribute to the coexistence of closely related species comprising savanna woody communities. [ABSTRACT FROM AUTHOR]

• Deep learning (CNN) can accurately detect prairie dog burrows in imagery. • CNN-predicted burrow density was very strongly correlated with validation data. • Active prairie dog colony areas can be estimated from burrow density maps. • Recently inactive colony area was over-estimated by the CNN method. • Satellite time series data were useful for identifying inactive colonies. Monitoring wildlife is fundamental to managing the health of rangelands but challenging due to the extensive and dynamic nature of these ecosystems. The black-tailed prairie dog (Cynomys ludovicianus) is considered both a keystone species of conservation concern and an agricultural pest. This animal is an example of a wildlife species for which detailed monitoring is both high priority and difficult to accomplish cost-effectively using ground-based methods. In this study, we conducted a robust evaluation of the potential to use deep learning to detect prairie dog burrows from remotely sensed imagery acquired from unoccupied aerial systems (UAS). We processed UAS imagery to create RGB, topographic position index (TPI) and normalized difference vegetation index (NDVI) products at varying spatial resolutions (2–30 cm). We then evaluated the minimum set of inputs and image resolution required to train a deep convolutional neural network (CNN) for burrow detection and scale this up to identify entire colonies. We validated results at the scale of individual burrows, sub-colony burrow density and range-wide colony area using ground and digitized observations. We found the 2 cm imagery proved computationally impractical for scaling, but performance did not decline between 2 and 5 cm imagery, and models performed well up to 10–15 cm. The top models always included TPI and the combination of RGB + TPI tended to perform best across spatial resolutions. Adding NDVI generally did not improve model performance. At 5 cm resolution, the top models achieved high precision and recall for detecting individual burrows (F-score 0.84–0.87) and burrow density was strongly correlated with validation data (r = 0.94–0.95). In pastures with active colonies, overlap between predicted and ground delineated colonies was high (60–94%). The CNN-based approach could not distinguish between currently active colonies and a colony that had recently become inactive due to a sylvatic plague (Yersinia pestis) epizootic. However, further analysis showed that CNN-derived burrow density was related to colony age and satellite-derived vegetation conditions in active colonies, and that the plague-affected colony deviated from expected vegetation trends. We conclude that a deep learning algorithm can accurately detect prairie dog burrows from UAS imagery acquired at 5–10 cm resolution, and that scaling from individual burrows to entire colonies is achievable but warrants further research. Combining CNN-derived burrow density maps with satellite-derived vegetation conditions may help identify recent colony abandonment, despite ongoing presence of burrows. [ABSTRACT FROM AUTHOR]

1. Temporal changes in semi-arid ecosystems can include transitions between alternative stable states, involving thresholds and multiple domains of attraction, but can also include relatively continuous, symmetric and reversible shifts within a single stable state. Conceptual state-and-transition models (STMs) describe both types of ecosystem dynamics by including state transitions (plant community changes difficult-to-reverse without substantial input or effort) and phase shifts (easily reversible community changes) as consequences of management practices and environmental variability. Grazing management is purported to be the primary driver of state transitions in current STMs for North American grasslands, but there is limited empirical evidence from these grasslands showing that grazing can cause difficult-toreverse transitions between alternate stable states. 2. In a northern mixed-grass prairie in Wyoming, USA, we examined plant community responses to (i) long-term (33-year) grazing intensity treatments (none, light, moderate and heavy stocking rates) and (ii) 8 years of light or no grazing in pastures that were grazed heavily for the previous 25 years. 3. Long-term grazing treatments were associated with distinct, but not stable, plant communities. From year 22 to 33, heavier stocking rates decreased cover of dominant C3 grasses and increased cover of the dominant C4 grass Bouteloua gracilis. 4. Reversing stocking rates from heavy to light or no grazing resulted in reversal of changes induced by prior heavy stocking for dominant C3 grasses, but not for B. gracilis. For both groups, rates of change following grazing treatment reversals were consistent with rates of change during the initial years of the experiment (1982-1990). 5. Synthesis and applications. In a semi-arid rangeland with a long evolutionary history of grazing, different long-term grazing intensity treatments caused slow, continuous and directional changes with important management implications, but did not appear to induce alternative stable states. For this and similar ecosystems, quantifying the time-scales and compositional gradients associated with key phase shifts may be more important than identifying thresholds between alternative stable states. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Adaptive management requires rangeland managers to respond to changing forage conditions (e.g., standing herbaceous biomass) within the grazing season at the scale of individual pastures. While within-season biomass can be measured or estimated in the field, it is often impractical to make field measurements in extensive rangeland systems with adequate frequency and spatial representation for responsive decision-making by rangeland managers. We sought to develop a single model to accurately predict daily herbaceous biomass across seasonally and annually varying conditions from the Harmonized Landsat-Sentinel satellite time series. We also sought to assess how information about plant community composition derived from a high-spatial resolution map would improve model performance. We used herbaceous biomass data from 1764 ground observations collected over 8 years in North American shortgrass steppe for training in a cross-validated model selection approach to evaluate (1) predictive performance of candidate models both spatially and temporally, (2) relative variable importance of individual spectral bands, vegetation indices, and recently developed broadband spectral angle indices, and (3) the degree to which including plant community composition improved model performance. All 11 candidate models identified in the model selection procedure contained a band angle index and an individual spectral band, and 6 contained one of each input feature type, demonstrating the benefit of combining spectral features for predicting herbaceous biomass across varying conditions. The spatial and temporal cross-validation and selection procedures produced the same top model with similar performance (mean absolute error = 151–182 kg ha−1; R2 = 0.75–0.79), suggesting that a single model performs well over space and time. Including plant community composition in the model reduced mean absolute error by 11–13%. Bootstrapping revealed that –six to seven years of training data were required to achieve consistent model performance across years with varying environmental conditions (e.g., wet, average, dry). The top model could accurately detect (70–87% accuracy) the week that biomass dropped below management-related thresholds as low as 450 kg ha−1 in an independent dataset (n = 950) with modest commission error (10–26%). We discuss how maps showing the probability that herbaceous biomass is below a given threshold can support adaptive management in extensive semiarid rangelands across differing scenarios of risk perception and avoidance. In addition to producing maps to support precision rangeland management strategies, this study demonstrates the importance of combining complementary vegetation indices and acquiring long-term training datasets to achieve reliable predictions of herbaceous standing biomass in highly variable systems. • Herbaceous standing biomass in rangelands can be estimated from Landsat-Sentinel. • 6–7 years of training data was needed to predict across variable conditions. • Broadband spectral angle indices were important for models across space and time. • Plant community data from high-resolution maps improved model performance. • Final models detected the week biomass dropped below management-related thresholds. [ABSTRACT FROM AUTHOR]

1. Savanna ecosystems – defined by the coexistence of trees and grasses – cover more than one-fifth the world’s land surface and harbour most of the world’s rangelands, livestock and large mammal diversity. Savanna trees can have a variety of effects on grasses, with consequences for the wild and domestic herbivores that depend on them. 2. Studies of these effects have focused on two different spatial scales. At the scale of individual trees, many studies have shown net positive effects of trees on sub-canopy grass nutrient concentrations and biomass. At the landscape scale, other studies have shown negative effects of high tree densities on grass productivity. These disparate results have led to different conclusions about the effects of trees on forage quality and ungulate nutrition in savannas. 3. We integrate these approaches by examining the effects of trees on grasses at both spatial scales and across a range of landscape-scale tree densities. We quantified grass biomass, species composition and nutrient concentrations in these different contexts in an Acacia drepanolobium savanna in Laikipia, Kenya. 4. Individual trees had positive effects on grass biomass, most likely because trees enrich soil nitrogen. Grass leaf phosphorus in sub-canopy areas, however, was depressed. The effects of individual trees could explain the effects of increasing landscape-scale tree cover for the biomass of only two of the four dominant grass species. 5. The negative effects of trees on grass and soil phosphorus, combined with depressed grass productivity in areas of high tree cover, suggest that ungulate nutrition may be compromised in areas with many trees. 6. Synthesis. We conclude that few, isolated trees may have positive local effects on savanna grasses and forage, but in areas of high tree density the negative landscape-scale effects of trees are likely to outweigh these positive effects. In savannas and other patchy landscapes, attempts to predict the consequences of changes in patch abundances for ecosystem services (e.g. rangeland productivity and carbon sequestration) will depend on our understanding of the extent to which local, patch-scale dynamics do or do not predict landscape-scale dynamics. [ABSTRACT FROM AUTHOR]