Transplanted sagebrush "wildlings" exhibit higher survival than greenhouse-grown tubelings yet both recruit new plants.

BACKGROUND: Land uses such as crop production, livestock grazing, mining, and urban development have contributed to degradation of drylands worldwide. Loss of big sagebrush (Artemisia tridentata) on disturbed drylands across the western U.S. has prompted massive efforts to re-establish this foundational species. There has been growing interest in avoiding the severe limitations experienced by plants at the seed and seedling stages by instead establishing plants from containerized greenhouse seedlings ("tubelings"). In some settings, a potential alternative approach is to transplant larger locally-collected plants ("wildlings"). We compared the establishment of mountain big sagebrush (A. tridentata ssp. vaseyana) from tubelings vs. wildlings in southeastern Idaho. A mix of native and non-native grass and forb species was drill-seeded in a pasture previously dominated by the introduced forage grass, smooth brome (Bromus inermis). We then established 80 m x 80 m treatment plots and planted sagebrush tubelings (n = 12 plots, 1200 plants) and wildlings (n = 12 plots, 1200 plants). We also established seeded plots (n = 12) and untreated control plots (n = 6) for long-term comparison. We tracked project expenses in order to calculate costs of using tubelings vs. wildlings as modified by probability of success. RESULTS: There was high (79%) tubeling and low (10%) wildling mortality within the first year. Three years post-planting, chance of survival for wildlings was significantly higher than that of tubelings (85% and 14% respectively). Despite high up-front costs of planting wildlings, high survival rates resulted in their being < 50% of the cost of tubelings on a per-surviving plant basis. Additionally, by the third year post-planting 34% of surviving tubelings and 95% of surviving wildlings showed evidence of reproduction (presence / absence of flowering stems), and the two types of plantings recruited new plants via seed (3.7 and 2.4 plants, respectively, per surviving tubeling/wildling). CONCLUSIONS: Our results indicate that larger plants with more developed root systems (wildlings) may be a promising avenue for increasing early establishment rates of sagebrush plants in restoration settings. Our results also illustrate the potential for tubelings and wildlings to improve restoration outcomes by "nucleating" the landscape via recruitment of new plants during ideal climate conditions.

At high stocking rates, cattle do not functionally replace wild herbivores in shaping understory community composition.

Over a quarter of the world's land surface is grazed by cattle and other livestock, which are replacing wild herbivores, potentially impairing ecosystem structure, and functions. Previous research suggests that cattle at moderate stocking rates can functionally replace wild herbivores in shaping understory communities. However, it is uncertain whether this is also true under high stocking rates and the effects of wild herbivore on plant communities are moderate, enhanced, or simply additive to the effects of cattle at high stocking rates. To evaluate the influence of cattle stocking rates on the ability of cattle to functionally replace wild herbivores and test for interactive effects between cattle and wild herbivores in shaping understory vegetation, we assessed herbaceous vegetation in a long-term exclosure experiment in a semi-arid savanna in central Kenya that selectively excludes wild mesoherbivores (50-1000 kg) and megaherbivores (elephant and giraffe). We tested the effects of cattle stocking rate (zero/moderate/high) on herbaceous vegetation (diversity, composition, leafiness). We also tested how those effects depend on the presence of wild mesoherbivores and megaherbivores. We found that herbaceous community composition (primary ordination axis) was better explained by the presence/absence of herbivore types than by total herbivory, suggesting that herbivore identity is a more important determinant of community composition than total herbivory at high cattle stocking rates. The combination of wild mesoherbivores and cattle stocked at high rates led to increased bare ground and annual grass cover, reduced perennial grass cover and understory leafiness, and enhanced understory diversity. These shifts were weaker or absent when cattle were stocked at high stocking rates in the absence of wild mesoherbivores. Megaherbivores tempered the effects of cattle stocked at high rates on herbaceous community composition but amplified the effects of high cattle stocking rate on bare ground and understory diversity. Our results show that cattle at high stocking rates do not functionally replace wild herbivores in shaping savanna herbaceous communities contrary to previous findings at moderate stocking rates. In mixed-use rangelands, interactions between cattle stocking rate and wild herbivore presence can lead to non-additive vegetation responses with important implications for both wildlife conservation and livestock production.

Browsing wildlife and heavy grazing indirectly facilitate sapling recruitment in an East African savanna.

Management of tree cover, either to curb bush encroachment or to mitigate losses of woody cover to over-browsing, is a major concern in savanna ecosystems. Once established, trees are often "trapped" as saplings, since interactions among disturbance, plant competition, and precipitation delay sapling recruitment into adult size classes. Saplings can be directly suppressed by wildlife browsing and competition from adjacent plants, and indirectly facilitated by grazers, such as cattle, which feed on neighboring grasses. Yet few experimental studies have simultaneously quantified the effects of cattle and wildlife on sapling growth, particularly over long time scales. We used a series of replicated 4-ha herbivore-manipulation plots to investigate the net effects of wildlife and moderate cattle grazing on Acacia drepanolobium sapling growth over 10 years that encompassed extended wet and dry periods. We also simulated more intense cattle grazing using grass removal treatments (0.5-m radius around saplings), and we quantified the role of intraspecific tree competition using neighborhood tree surveys (trees within a 3-m radius). Wildlife, which included elephants, had a positive effect on sapling growth. Wildlife also reduced neighbor tree density during the 10-yr study, which likely caused the positive effect of wildlife on saplings. Although moderate cattle grazing did not affect sapling growth, grass removal treatments simulating heavy grazing increased sapling growth. Both grass removal and neighbor tree effects on saplings were strongest during above-average rainfall years following drought. This highlights that livestock-driven reductions in grass cover and catastrophic wildlife damage to trees during droughts present a need, or an opportunity, for targeted management of sapling growth and woody plant cover during ensuing wet periods.

Termite mound cover and abundance respond to herbivore-mediated biotic changes in a Kenyan savanna.

Both termites and large mammalian herbivores (LMH) are savanna ecosystem engineers that have profound impacts on ecosystem structure and function. Both of these savanna engineers modulate many common and shared dietary resources such as woody and herbaceous plant biomass, yet few studies have addressed how they impact one another. In particular, it is unclear how herbivores may influence the abundance of long-lived termite mounds via changes in termite dietary resources such as woody and herbaceous biomass. While it has long been assumed that abundance and areal cover of termite mounds in the landscape remain relatively stable, most data are observational, and few experiments have tested how termite mound patterns may respond to biotic factors such as changes in large herbivore communities. Here, we use a broad tree density gradient and two landscape-scale experimental manipulations-the first a multi-guild large herbivore exclosure experiment (20 years after establishment) and the second a tree removal experiment (8 years after establishment)-to demonstrate that patterns in Odontotermes termite mound abundance and cover are unexpectedly dynamic. Termite mound abundance, but areal cover not significantly, is positively associated with experimentally controlled presence of cattle, but not wild mesoherbivores (15-1,000 kg) or megaherbivores (elephants and giraffes). Herbaceous productivity and tree density, termite dietary resources that are significantly affected by different LMH treatments, are both positive predictors of termite mound abundance. Experimental reductions of tree densities are associated with lower abundances of termite mounds. These results reveal a richly interacting web of relationships among multiple savanna ecosystem engineers and suggest that termite mound abundance and areal cover are intimately tied to herbivore-driven resource availability.

Synergistic effects of long-term herbivory and previous fire on fine-scale heterogeneity of prescribed grassland burns.

Grassland and savanna ecosystems, important for both livelihoods and biodiversity conservation, are strongly affected by ecosystem drivers such as herbivory, fire, and drought. Interactions among fire, herbivores and vegetation produce complex feedbacks in these ecosystems, but these have rarely been studied in the context of fuel continuity and resultant fire heterogeneity. We carried out 36 controlled burns within replicated experimental plots that had allowed differential access by wild and domestic large herbivores since 1995 in a savanna ecosystem in Kenya. Half of these were reburns of plots burned 5 yr previously. We show here that the fine-scale spatial heterogeneity of fire was greater in plots (1) previously burned, (2) accessible to large herbivores, and especially (3) these two in combination. An additional embedded experiment demonstrated that even small experimental burn-free patches can have strong positive effects on tree saplings, which experienced less damage during controlled burns and quicker postfire recovery. This work highlights the importance of simultaneously examining the interactions between fire and herbivory on fuel heterogeneity, which can have important impacts on the growth of woody saplings in savanna grasslands.

Thresholds are in the eye of the beholder: plants and wildlife respond differently to short-term cattle corrals.

Rangelands are governed by threshold dynamics, and factors such as drought, wildfire, and herbivory can drive change across thresholds and between ecological states. Most work on this topic has focused on shifts in a single response variable, vegetation, and little research has considered how to reconcile responses of more than one variable to determine whether a system has undergone a genuine state change. In sub-Saharan Africa, mobile overnight livestock corrals (bomas) can be used by managers to precipitate ecological transitions from areas dominated by bare ground to productive ecosystem hotspots (glades) that are attractive to wild herbivores. We asked how long bomas must be occupied by cattle before undergoing a state change, considering both plant and animal response variables, to glade ecosystem hotspots. We tested five durations of boma occupation: 0, 4, 7, 14, and 28 days. Each treatment was replicated five times, and we assessed vegetation as well as herbivore dung (as a proxy of use) at multiple time points over 3 yr following boma abandonment. Vegetation in 7-, 14-, and 28-d boma duration treatments appeared to undergo a complete transition to glade-like plant communities, whereas the shortest 4-d treatment had not converted to a glade plant community by year 3. Wildlife responses appeared to lag behind vegetation responses, with transitions to glade-like herbivore use occurring only in the longest duration (14- and 28-d) treatments. Our results show that different response variables, when considered individually, may provide incomplete or misleading information about state changes. Although shorter-occupied bomas might be effective for reducing bare ground, they may not attract enough wild herbivores to constitute crossing into an alternative state. Understanding threshold dynamics associated not only with vegetation responses but with a broader suite of response variables is challenging, but will provide a more complete representation of ecosystem function and greater opportunity for more successful ecosystem management.

Competition and coexistence in plant communities: intraspecific competition is stronger than interspecific competition.

Theory predicts that intraspecific competition should be stronger than interspecific competition for any pair of stably coexisting species, yet previous literature reviews found little support for this pattern. We screened over 5400 publications and identified 39 studies that quantified phenomenological intraspecific and interspecific interactions in terrestrial plant communities. Of the 67% of species pairs in which both intra- and interspecific effects were negative (competitive), intraspecific competition was, on average, four to five-fold stronger than interspecific competition. Of the remaining pairs, 93% featured intraspecific competition and interspecific facilitation, a situation that stabilises coexistence. The difference between intra- and interspecific effects tended to be larger in observational than experimental data sets, in field than greenhouse studies, and in studies that quantified population growth over the full life cycle rather than single fitness components. Our results imply that processes promoting stable coexistence at local scales are common and consequential across terrestrial plant communities.

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

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

Herbivore removal reduces influence of arbuscular mycorrhizal fungi on plant growth and tolerance in an East African savanna.

The functional relationship between arbuscular mycorrhizal fungi (AMF) and their hosts is variable on small spatial scales. Here, we hypothesized that herbivore exclusion changes the AMF community and alters the ability of AMF to enhance plant tolerance to grazing. We grew the perennial bunchgrass, Themeda triandra Forssk in inoculum from soils collected in the Kenya Long-term Exclosure Experiment where treatments representing different levels of herbivory have been in place since 1995. We assessed AMF diversity in the field, using terminal restriction fragment length polymorphism and compared fungal diversity among treatments. We conducted clipping experiments in the greenhouse and field and assessed regrowth. Plants inoculated with AMF from areas accessed by wild herbivores and cattle had greater biomass than non-inoculated controls, while plants inoculated with AMF from where large herbivores were excluded did not benefit from AMF in terms of biomass production. However, only the inoculation with AMF from areas with wild herbivores and no cattle had a positive effect on regrowth, relative to clipped plants grown without AMF. Similarly, in the field, regrowth of plants after clipping in areas with only native herbivores was higher than other treatments. Functional differences in AMF were evident despite little difference in AMF species richness or community composition. Our findings suggest that differences in large herbivore communities over nearly two decades has resulted in localized, functional changes in AMF communities. Our results add to the accumulating evidence that mycorrhizae are locally adapted and that functional differences can evolve within small geographical areas.

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.

Interacting effects of land use and climate on rodent-borne pathogens in central Kenya.

Understanding the effects of anthropogenic disturbance on zoonotic disease risk is both a critical conservation objective and a public health priority. Here, we evaluate the effects of multiple forms of anthropogenic disturbance across a precipitation gradient on the abundance of pathogen-infected small mammal hosts in a multi-host, multi-pathogen system in central Kenya. Our results suggest that conversion to cropland and wildlife loss alone drive systematic increases in rodent-borne pathogen prevalence, but that pastoral conversion has no such systematic effects. The effects are most likely explained both by changes in total small mammal abundance, and by changes in relative abundance of a few high-competence species, although changes in vector assemblages may also be involved. Several pathogens responded to interactions between disturbance type and climatic conditions, suggesting the potential for synergistic effects of anthropogenic disturbance and climate change on the distribution of disease risk. Overall, these results indicate that conservation can be an effective tool for reducing abundance of rodent-borne pathogens in some contexts (e.g. wildlife loss alone); however, given the strong variation in effects across disturbance types, pathogen taxa and environmental conditions, the use of conservation as public health interventions will need to be carefully tailored to specific pathogens and human contexts.This article is part of the themed issue 'Conservation, biodiversity and infectious disease: scientific evidence and policy implications'.

Fire disturbance disrupts an acacia ant-plant mutualism in favor of a subordinate ant species.

Although disturbance theory has been recognized as a useful framework in examining the stability of ant-plant mutualisms, very few studies have examined the effects of fire disturbance on these mutualisms. In myrmecophyte-dominated savannas, fire and herbivory are key drivers that could influence ant-plant mutualisms by causing complete colony mortality and/or decreasing colony size, which potentially could alter dominance hierarchies if subordinate species are more fire resilient. We used a large-scale, replicated fire experiment to examine long-term effects of fire on acacia-ant community composition. To determine if fire shifted ant occupancy from a competitive dominant to a subordinate ant species, we surveyed the acacia-ant community in 6-7 yr old burn sites and examined how the spatial scale of these burns influenced ant community responses. We then used two short-term fire experiments to explore possible mechanisms for the shifts in community patterns observed. Because survival of ant colonies is largely dependent on their ability to detect and escape an approaching fire, we first tested the evacuation response of all four ant species when exposed to smoke (fire signal). Then to better understand how fire and its interaction with large mammal herbivory affect the density of ants per tree, we quantified ant worker density in small prescribed burns within herbivore exclusion plots. We found clear evidence suggesting that fire disturbance favored the subordinate ant Crematogaster nigriceps more than the dominant and strong mutualist ant C. mimosae, whereby C. nigriceps (1) was the only species to occupy a greater proportion of trees in 6-7 yr old burn sites compared to unburned sites, (2) had higher burn/unburn tree ratios with increasing burn size, and (3) evacuated significantly faster than C. mimosae in the presence of smoke. Fire and herbivory had opposite effects on ant density per meter of branch for both C. nigriceps and C. mimosae, with fire decreasing ant densities per meter of branch and the presence of large herbivores increasing ant density. Taken together, these experiments suggest that major ecosystem disturbances like fire can disrupt mutualistic associations and maintain diversity in partner quality and identity despite a clear dominance hierarchy.

Patterns in Greater Sage-grouse population dynamics correspond with public grazing records at broad scales.

Human land use, such as livestock grazing, can have profound yet varied effects on wildlife interacting within common ecosystems, yet our understanding of land-use effects is often generalized from short-term, local studies that may not correspond with trends at broader scales. Here we used public land records to characterize livestock grazing across Wyoming, USA, and we used Greater Sage-grouse (Centrocercus urophasianus) as a model organism to evaluate responses to livestock management. With annual counts of male Sage-grouse from 743 leks (breeding display sites) during 2004-2014, we modeled population trends in response to grazing level (represented by a relative grazing index) and timing across a gradient in vegetation productivity as measured by the Normalized Vegetation Difference Index (NDVI). We found grazing can have both positive and negative effects on Sage-grouse populations depending on the timing and level of grazing. Sage-grouse populations responded positively to higher grazing levels after peak vegetation productivity, but populations declined when similar grazing levels occurred earlier, likely reflecting the sensitivity of cool-season grasses to grazing during peak growth periods. We also found support for the hypothesis that effects of grazing management vary with local vegetation productivity. These results illustrate the importance of broad-scale analyses by revealing patterns in Sage-grouse population trends that may not be inferred from studies at finer scales, and could inform sustainable grazing management in these ecosystems.

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.

Influence of cattle on browsing and grazing wildlife varies with rainfall and presence of megaherbivores.

In many savanna ecosystems worldwide, livestock share the landscape and its resources with wildlife. The nature of interactions between livestock and wildlife is a subject of considerable interest and speculation, yet little controlled experimental research has been carried out. Since 1995, we have been manipulating the presence and absence of cattle and large mammalian herbivore wildlife in a Kenyan savanna in order to better understand how different herbivore guilds influence habitat use by specific wildlife species. Using dung counts as a relative assay of herbivore use of the different experimental plots, we found that cattle had a range of effects, mostly negative, on common mesoherbivore species, including both grazers and mixed feeders, but did not have significant effects on megaherbivores. The effect of cattle on most of the mesoherbivore species was contingent on both the presence of megaherbivores and rainfall. In the absence of megaherbivores, wild mesoherbivore dung density was 36% lower in plots that they shared with cattle than in plots they used exclusively, whereas in the presence of megaherbivores, wild mesoherbivore dung density was only 9% lower in plots shared with cattle than plots used exclusively. Cattle appeared to have a positive effect on habitat use by zebra (a grazer) and steinbuck (a browser) during wetter periods of the year but a negative effect during drier periods. Plots to which cattle had access had lower grass and forb cover than plots from which they were excluded, while plots to which megaherbivores had access had more grass cover but less forb cover. Grass cover was positively correlated with zebra and oryx dung density while forb cover was positively correlated with eland dung density. Overall these results suggest that interactions between livestock and wildlife are contingent on rainfall and herbivore assemblage and represent a more richly nuanced set of interactions than the longstanding assertion that cattle simply compete with (grazing) wildlife. Specifically, rainfall and megaherbivores seemed to moderate the negative effects of cattle on some mesoherbivore species. Even if cattle tend to reduce wildlife use of the landscape, managing simultaneously for livestock production (at moderate levels) and biodiversity conservation is possible.

Are cattle surrogate wildlife? Savanna plant community composition explained by total herbivory more than herbivore type.

The widespread replacement of wild ungulate herbivores by domestic livestock in African savannas is composed of two interrelated phenomena: (1) loss or reduction in numbers of individual wildlife species or guilds and (2) addition of livestock to the system. Each can have important implications for plant community dynamics. Yet very few studies have experimentally addressed the individual, combined, and potentially interactive effects of wild vs. domestic herbivore species on herbaceous plant communities within a single system. Additionally, there is little information about whether, and in which contexts, livestock might functionally replace native herbivore wildlife or, alternatively, have fundamentally different effects on plant species composition. The Kenya Long-term Exclosure Experiment, which has been running since 1995, is composed of six treatment combinations of mega-herbivores, meso-herbivore ungulate wildlife, and cattle. We sampled herbaceous vegetation 25 times between 1999 and 2013. We used partial redundancy analysis and linear mixed models to assess effects of herbivore treatments on overall plant community composition and key plant species. Plant communities in the six different herbivore treatments shifted directionally over time and diverged from each other substantially by 2013. Plant community composition was strongly related (R2  = 0.92) to residual plant biomass, a measure of herbivore utilization. Addition of any single herbivore type (cattle, wildlife, or mega-herbivores) caused a shift in plant community composition that was proportional to its removal of plant biomass. These results suggest that overall herbivory pressure, rather than herbivore type or complex interactions among different herbivore types, was the main driver of changes in plant community composition. Individual plant species, however, did respond most strongly to either wild ungulates or cattle. Although these results suggest considerable functional similarity between a suite of native wild herbivores (which included grazers, browsers, and mixed feeders) and cattle (mostly grazers) with respect to understory plant community composition, responses of individual plant species demonstrate that at the plant-population-level impacts of a single livestock species are not functionally identical to those of a diverse group of native herbivores.

Grass-Shrub Associations over a Precipitation Gradient and Their Implications for Restoration in the Great Basin, USA.

As environmental stress increases positive (facilitative) plant interactions often predominate. Plant-plant associations (or lack thereof) can indicate whether certain plant species favor particular types of microsites (e.g., shrub canopies or plant-free interspaces) and can provide valuable insights into whether "nurse plants" will contribute to seeding or planting success during ecological restoration. It can be difficult, however, to anticipate how relationships between nurse plants and plants used for restoration may change over large-ranging, regional stress gradients. We investigated associations between the shrub, Wyoming big sagebrush (Artemisia tridentata ssp. wyomingensis), and three common native grasses (Poa secunda, Elymus elymoides, and Pseudoroegneria spicata), representing short-, medium-, and deep-rooted growth forms, respectively, across an annual rainfall gradient (220-350 mm) in the Great Basin, USA. We hypothesized that positive shrub-grass relationships would become more frequent at lower rainfall levels, as indicated by greater cover of grasses in shrub canopies than vegetation-free interspaces. We sampled aerial cover, density, height, basal width, grazing status, and reproductive status of perennial grasses in canopies and interspaces of 25-33 sagebrush individuals at 32 sites along a rainfall gradient. We found that aerial cover of the shallow rooted grass, P. secunda, was higher in sagebrush canopy than interspace microsites at lower levels of rainfall. Cover and density of the medium-rooted grass, E. elymoides were higher in sagebrush canopies than interspaces at all but the highest rainfall levels. Neither annual rainfall nor sagebrush canopy microsite significantly affected P. spicata cover. E. elymoides and P. spicata plants were taller, narrower, and less likely to be grazed in shrub canopy microsites than interspaces. Our results suggest that exploring sagebrush canopy microsites for restoration of native perennial grasses might improve plant establishment, growth, or survival (or some combination thereof), particularly in drier areas. We suggest that land managers consider the nurse plant approach as a way to increase perennial grass abundance in the Great Basin. Controlled experimentation will provide further insights into the life stage-specific effectiveness and practicality of a nurse plant approach for ecological restoration in this region.

Synergistic effects of fire and elephants on arboreal animals in an African savanna.

Disturbance is a crucial determinant of animal abundance, distribution and community structure in many ecosystems, but the ways in which multiple disturbance types interact remain poorly understood. The effects of multiple-disturbance interactions can be additive, subadditive or super-additive (synergistic). Synergistic effects in particular can accelerate ecological change; thus, characterizing such synergies, the conditions under which they arise, and how long they persist has been identified as a major goal of ecology. We factorially manipulated two principal sources of disturbance in African savannas, fire and elephants, and measured their independent and interactive effects on the numerically dominant vertebrate (the arboreal gekkonid lizard Lygodactylus keniensis) and invertebrate (a guild of symbiotic Acacia ants) animal species in a semi-arid Kenyan savanna. Elephant exclusion alone (minus fire) had negligible effects on gecko density. Fire alone (minus elephants) had negligible effects on gecko density after 4 months, but increased gecko density twofold after 16 months, likely because the decay of fire-damaged woody biomass created refuges and nest sites for geckos. In the presence of elephants, fire increased gecko density nearly threefold within 4 months of the experimental burn; this occurred because fire increased the incidence of elephant damage to trees, which in turn improved microhabitat quality for geckos. However, this synergistic positive effect of fire and elephants attenuated over the ensuing year, such that only the main effect of fire was evident after 16 months. Fire also altered the structure of symbiotic plant-ant assemblages occupying the dominant tree species (Acacia drepanolobium); this influenced gecko habitat selection but did not explain the synergistic effect of fire and elephants. However, fire-driven shifts in plant-ant occupancy may have indirectly mediated this effect by increasing trees' susceptibility to elephant damage. Our findings confirm the importance of fire × elephant interactions in structuring arboreal wildlife populations. Where habitat modification by megaherbivores facilitates co-occurring species, fire may amplify these effects in the short term by increasing the frequency or intensity of herbivory, leading to synergy. In the longer term, tree mortality due to both top kill by fire and toppling by large herbivores may reduce overall microhabitat availability, eliminating the synergy.

Contrasting effects of different mammalian herbivores on sagebrush plant communities.

Herbivory by both grazing and browsing ungulates shapes the structure and functioning of terrestrial ecosystems worldwide, and both types of herbivory have been implicated in major ecosystem state changes. Despite the ecological consequences of differences in diets and feeding habits among herbivores, studies that experimentally distinguish effects of grazing from spatially co-occurring, but temporally segregated browsing are extremely rare. Here we use a set of long-term exclosures in northern Utah, USA, to determine how domestic grazers vs. wild ungulate herbivores (including browsers and mixed feeders) affect sagebrush-dominated plant communities that historically covered ~62 million ha in North America. We sampled plant community properties and found that after 22 years grazing and browsing elicited perceptible changes in overall plant community composition and distinct responses by individual plant species. In the woody layer of the plant community, release from winter and spring wild ungulate herbivory increased densities of larger Wyoming big sagebrush (Artemisia tridentata, ssp. wyomingensis) at the expense of small sagebrush, while disturbance associated with either cattle or wild ungulate activity alone was sufficient to increase bare ground and reduce cover of biological soil crusts. The perennial bunchgrass, bottlebrush squirretail (Elymus elymoides), responded positively to release from summer cattle grazing, and in turn appeared to competitively suppress another more grazing tolerant perennial grass, Sandberg's blue grass (Poa secunda). Grazing by domestic cattle also was associated with increased non-native species biomass. Together, these results illustrate that ungulate herbivory has not caused sagebrush plant communities to undergo dramatic state shifts; however clear, herbivore-driven shifts are evident. In a dry, perennial-dominated system where plant community changes can occur very slowly, our results provide insights into potential long-term trajectories of these plant communities under different large herbivore regimes. Our results can be used to guide long-term management strategies for sagebrush systems and improve habitat for endemic wildlife species such as sage-grouse (Centrocercus spp.).

Native and domestic browsers and grazers reduce fuels, fire temperatures, and acacia ant mortality in an African savanna.

Despite the importance of fire and herbivory in structuring savanna systems, few replicated experiments have examined the interactive effects of herbivory and fire on plant dynamics. In addition, the effects of fire on associated ant-tree mutualisms have been largely unexplored. We carried out small controlled burns in each of 18 herbivore treatment plots of the Kenya Long-term Exclosure Experiment (KLEE), where experimentally excluding elephants has resulted in 42% greater tree densities. The KLEE design includes six different herbivore treatments that allowed us to examine how different combinations of megaherbivore wildlife, mesoherbivore wildlife, and cattle affect fire temperatures and subsequent loss of ant symbionts from Acacia trees. Before burning, we quantified herbaceous fuel loads and plant community composition. We tagged all trees, measured their height and basal diameter, and identified the resident ant species on each. We recorded weather conditions during the burns and used ceramic tiles painted with fire-sensitive paints to estimate fire temperatures at different heights and in different microsites (under vs. between trees). Across all treatments, fire temperatures were highest at 0-50 cm off the ground and hotter in the grass under trees than in the grassy areas between trees. Plots with more trees burned hotter than plots with fewer trees, perhaps because of greater fine woody debris. Plots grazed by wildlife and by cattle prior to burning had lower herbaceous fuel loads and experienced lower burn temperatures than ungrazed plots. Many trees lost their ant colonies during the burns. Ant survivorship differed by ant species and at the plot level was positively associated with previous herbivory (and lower fire temperatures). Across all treatments, ant colonies on taller trees were more likely to survive, but even some of the tallest trees lost their ant colonies. Our study marks a significant step in understanding the mechanisms that underlie the interactions between fire and herbivory in savanna ecosystems.