Herbivory and Drought Generate Short‐Term Stochasticity and Long‐Term Stability in A Savanna Understory Community

Rainfall and herbivory are fundamental drivers of grassland plant dynamics, yet few studies have examined long‐term interactions between these factors in an experimental setting. Understanding such interactions is important, as rainfall is becoming increasingly erratic and native wild herbivores are being replaced by livestock. Livestock grazing and episodic low rainfall are thought to interact, leading to greater community change than either factor alone. We examined patterns of change and stability in herbaceous community composition through four dry periods, or droughts, over 15 years of the Kenya Long‐term Exclosure Experiment (KLEE), which consists of six different combinations of cattle, native wild herbivores (e.g., zebras, gazelles), and mega‐herbivores (giraffes, elephants). We used principal response curves to analyze the trajectory of change in each herbivore treatment relative to a common initial community and asked how droughts contributed to community change in these treatments. We examined three measures of stability (resistance, variability, and turnover) that correspond to different temporal scales and found that each had a different response to grazing. Treatments that included both cattle and wild herbivores had higher resistance (less net change over 15 years) but were more variable on shorter time scales; in contrast, the more lightly grazed treatments (no herbivores or wild herbivores only) showed lower resistance due to the accumulation of consistent, linear, short‐term change. Community change was greatest during and immediately after droughts in all herbivore treatments. But, while drought contributed to directional change in the less grazed treatments, it contributed to both higher variability and resistance in the more heavily grazed treatments. Much of the community change in lightly grazed treatments (especially after droughts) was due to substantial increases in cover of the palatable grass Brachiaria lachnantha. These results illustrate how herbivory and drought can act together to cause change in grassland communities at the moderate to low end of a grazing intensity continuum. Livestock grazing at a moderate intensity in a system with a long evolutionary history of grazing contributed to long‐term stability. This runs counter to often‐held assumptions that livestock grazing leads to directional, destabilizing shifts in grassland systems

Glade cascades: indirect legacy effects of pastoralism enhance the abundance and spatial structuring of arboreal fauna

Peer Reviewedhttp://deepblue.lib.umich.edu/bitstream/2027.42/117219/1/ecy2013944827.pd

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

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

Herbivore Effects on Productivity Vary by Guild: Cattle Increase Mean Productivity While Wildlife Reduce Variability

Wild herbivores and livestock share the majority of rangelands worldwide, yet few controlled experiments have addressed their individual, additive, and interactive impacts on ecosystem function. While ungulate herbivores generally reduce standing biomass, their effects on aboveground net primary production (ANPP) can vary by spatial and temporal context, intensity of herbivory, and herbivore identity and species richness. Some evidence indicates that moderate levels of herbivory can stimulate aboveground productivity, but few studies have explicitly tested the relationships among herbivore identity, grazing intensity, and ANPP. We used a long- term exclosure experiment to examine the effects of three groups of wild and domestic ungulate herbivores (megaherbivores, mesoherbivore wildlife, and cattle) on herbaceous productivity in an African savanna. Using both field measurements (productivity cages) and satellite imagery, we measured the effects of different herbivore guilds, separately and in different combinations, on herbaceous productivity across both space and time. Results from both productivity cage measurements and satellite normalized difference vegetation index (NDVI) demonstrated a positive relationship between mean productivity and total ungulate herbivore pressure, driven in particular by the presence of cattle. In contrast, we found that variation in herbaceous productivity across space and time was driven by the presence of wild herbivores (primarily mesoherbivore wildlife), which significantly reduced heterogeneity in ANPP and NDVI across both space and time. Our results indicate that replacing wildlife with cattle (at moderate densities) could lead to similarly productive but more heterogeneous herbaceous plant communities in rangelands

Unlikely Alliances and Their Implications for Resource Management in the American West

Collaborative, or participatory governance is an increasingly common means of addressing natural resource issues, especially in the American West where patchworks of public, private, and tribal interests characterize the region’s resources. In this context, unlikely alliances, or partnerships among diverse actors who have historically been at odds, have a growing potential to shape social and ecological outcomes, for better or worse. While these unlikely alliances have received greater attention in recent years, relatively little research has worked to synthesize the concept across diverse contexts and disciplines. Based on a review of the literature on unlikely alliances in natural resource governance, we develop a framework that synthesizes the individual motivations and contextual factors that influence their formation, as well as the social and ecological outcomes that they create. We use this framework to analyze six illustrative cases of unlikely alliances. Our analysis of these cases suggests that unlikely alliances in the American West are likely to arise in the presence of a crisis, when appropriate leadership is present, when some of the actors have interacted effectively in the past, and when actors need to pool resources. The cases also illustrate some common outcomes, including environmental improvement, transformation of social networks, policy change, and shifts in power relationships. We discuss the implications of unlikely alliances for the social-ecological future of the American West. Our paper highlights the role of unlikely alliances in shaping patterns of natural resource governance, and provides a focus for further research in this realm

Combining active restoration and targeted grazing to establish native plants and reduce fuel loads in invaded ecosystems

Many drylands have been converted from perennial-dominated ecosystems to invaded, annual-dominated, fire-prone systems. Innovative approaches are needed to disrupt fire-invasion feedbacks. Targeted grazing can reduce invasive plant abundance and associated flammable fuels, and fuelbreaks can limit fire spread. Restored strips of native plants (native greenstrips) can function as fuelbreaks while also providing forage and habitat benefits. However, methods for establishing native greenstrips in invaded drylands are poorly developed. Moreover, if fuels reduction and greenstrip establishment are to proceed simultaneously, it is critical to understand how targeted grazing interacts with plant establishment. We determined how targeted grazing treatments interacted with seed rate, spatial planting arrangement (mixtures vs. monoculture strips), seed coating technology, and species identity (five native grasses) to affect standing biomass and seeded plant density in experimental greenstrips. We monitored for two growing seasons to document effects during the seedling establishment phase. Across planting treatments, ungrazed paddocks had the highest second-year seeded plant densities and the highest standing biomass. Paddocks grazed in fall of the second growing season had fewer seedlings than paddocks grazed in spring, five months later. High seed rates minimized negative effects of grazing on plant establishment. Among seeded species, Elymus trachycaulus and Poa secunda had the highest second-year densities, but achieved this via different pathways. Elymus trachycaulus produced the most first-year seedlings, but declined in response to grazing, whereas P. secunda had moderate first-year establishment but high survival across grazing treatments. We identified clear tradeoffs between reducing fuel loads and establishing native plants in invaded sagebrush steppesimilar tradeoffs may exist in other invaded drylands. In our system, tradeoffs were minimized by boosting seed rates, using grazing-tolerant species, and delaying grazing. In invaded ecosystems, combining targeted grazing with high-input restoration may create opportunities to limit wildfire risk while also shifting vegetation toward more desirable species

Elephants Mitigate the Effects of Cattle on Wildlife and Other Ecosystem Traits: Experimental Evidence

On rangelands worldwide, cattle interact with many ecosystem components, most obviously with soils, plants, and other large herbivores. Since 1995, we have been manipulating the presence of cattle, mesoherbivores, and megaherbivores (elephants and giraffes) in a series of eighteen 4-ha (10-acre) plots at the Kenya Long-term Exclosure Experiment. We have demonstrated a wide array of cattle effects on this savanna rangeland, including their reduction of grass cover, wildlife use, and soil nitrogen and phosphorus pools, but their increase of primary productivity and termite abundance. Strikingly, we demonstrate that the presence of mega-herbivores (elephants, mainly) reduces the sizes of these cattle. We provide further experimental evidence that this may be because the elephants are reducing the most desirable (N-rich) forage, causing cattle to slow their extraction of (low-N) grasses, while simultaneously reducing tree cover

Using APAR to Predict Aboveground Plant Productivity in Semi-Arid Rangelands: Spatial and Temporal Relationships Differ

Monitoring of aboveground net primary production (ANPP) is critical for effective management of rangeland ecosystems but is problematic due to the vast extent of rangelands globally, and the high costs of ground-based measurements. Remote sensing of absorbed photosynthetically active radiation (APAR) can be used to predict ANPP, potentially offering an alternative means of quantifying ANPP at both high temporal and spatial resolution across broad spatial extents. The relationship between ANPP and APAR has often been quantified based on either spatial variation across a broad region or temporal variation at a location over time, but rarely both. Here we assess: (i) if the relationship between ANPP and APAR is consistent when evaluated across time and space; (ii) potential factors driving differences between temporal versus spatial models, and (iii) the magnitude of potential errors relating to space for time transformations in quantifying productivity. Using two complimentary ANPP datasets and remotely sensed data derived from MODIS and a Landsat/MODIS fusion data product, we find that slopes of spatial models are generally greater than slopes of temporal models. The abundance of plant species with different structural attributes, specifically the abundance of C4 shortgrasses with prostrate canopies versus taller, more productive C3 species with more vertically complex canopies, tended to vary more dramatically in space than over time. This difference in spatial versus temporal variation in these key plant functional groups appears to be the primary driver of differences in slopes among regression models. While the individual models revealed strong relationships between ANPP to APAR, the use of temporal models to predict variation in space (or vice versa) can increase error in remotely sensed predictions of ANPPEEA Concepción del UruguayFil: Gaffney, Rowan. United States Department of Agriculture–Agricultural Research Service. Rangeland Resources and Systems Research Unit; Estados UnidosFil: Porensky, Lauren M. United States Department of Agriculture–Agricultural Research Service. Rangeland Resources and Systems Research Unit; Estados UnidosFil: Feng, Gao. United States Department of Agriculture–Agricultural Research Service. Hydrology and Remote Sensing Laboratory; Estados UnidosFil: Irisarri, Jorge Gonzalo Nicolás. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Instituto de Investigaciones Fisiológicas y Ecológicas Vinculadas a la Agricultura. Universidad de Buenos Aires. Facultad de Agronomía. Instituto de Investigaciones Fisiológicas y Ecológicas Vinculadas a la Agricultura; ArgentinaFil: Durante, Martin. Instituto Nacional de Tecnología Agropecuaria (INTA). Estación Experimental Agropecuaria Concepción del Uruguay; ArgentinaFil: Derner, Justin D. United States Department of Agriculture-Agricultural Research Service. Rangeland Resources Research Unit; Estados UnidosFil: Augustine, David J.. United States Department of Agriculture–Agricultural Research Service. Rangeland Resources and Systems Research Unit; Estados Unido

Apparent Competition, Lion Predation, and Managed Livestock Grazing: Can Conservation Value Be Enhanced?

Predator restorations often result in apparent competition, where co-occurring prey populations experience asymmetric predation pressure driven by predator preferences. In many rangeland ecosystems, livestock share the landscape with wildlife, including ungulates and the large carnivores that consume them. We examined whether apparent competition reorganized prey communities following restoration of lions (Panthera leo) to a savanna ecosystem, and whether and how livestock management could alter this indirect interaction between lions and their prey. Three lines of evidence supported the hypothesis that Jackson's hartebeest (Alcelaphus bucelaphus lelwel; an ungulate of conservation concern) are suppressed via lion-mediated apparent competition. First, hartebeest exhibited an Allee effect where they were exposed to lions, but displayed negative density-dependent population growth where they were protected from lions. Second, spatial overlap between plains zebra (Equus burchelli; the primary prey of lions) and hartebeest further exacerbated lion predation on hartebeest. Finally, hartebeest were killed selectively by lions, whereas zebra were killed by lions in proportion to their abundance. We then tested whether glades [nutrient-rich hotspots created by abandoned cattle (Bos indicus) corrals] could be used to manipulate top-down control of hartebeest via their influence on the spatial distribution of zebra. Zebra aggregated at glades, and survival of hartebeest increased with increasing distance from glades, suggesting that corrals may be placed on the landscape away from hartebeest to create spatial refuges from lions. Our findings demonstrate how informed placement of livestock corrals can be used to manipulate the spatial distribution of primary prey (zebra), thereby reducing apparent competition suffered by hartebeest. Our work further provides an example of how integrating apparent competition theory with proactive livestock management can improve conservation efforts in multiple-use landscapes

Complex Consequences of Herbivory and Interplant Cues in Three Annual Plants

Information exchange (or signaling) between plants following herbivore damage has recently been shown to affect plant responses to herbivory in relatively simple natural systems. In a large, manipulative field study using three annual plant species (Achyrachaena mollis, Lupinus nanus, and Sinapis arvensis), we tested whether experimental damage to a neighboring conspecific affected a plant's lifetime fitness and interactions with herbivores. By manipulating relatedness between plants, we assessed whether genetic relatedness of neighboring individuals influenced the outcome of having a damaged neighbor. Additionally, in laboratory feeding assays, we assessed whether damage to a neighboring plant specifically affected palatability to a generalist herbivore and, for S. arvensis, a specialist herbivore. Our study suggested a high level of contingency in the outcomes of plant signaling. For example, in the field, damaging a neighbor resulted in greater herbivory to A. mollis, but only when the damaged neighbor was a close relative. Similarly, in laboratory trials, the palatability of S. arvensis to a generalist herbivore increased after the plant was exposed to a damaged neighbor, while palatability to a specialist herbivore decreased. Across all species, damage to a neighbor resulted in decreased lifetime fitness, but only if neighbors were closely related. These results suggest that the outcomes of plant signaling within multi-species neighborhoods may be far more context-specific than has been previously shown. In particular, our study shows that herbivore interactions and signaling between plants are contingent on the genetic relationship between neighboring plants. Many factors affect the outcomes of plant signaling, and studies that clarify these factors will be necessary in order to assess the role of plant information exchange about herbivory in natural systems