Disruption of an ant-plant mutualism shapes interactions between lions and their primary prey.

Mutualisms often define ecosystems, but they are susceptible to human activities. Combining experiments, animal tracking, and mortality investigations, we show that the invasive big-headed ant (Pheidole megacephala) makes lions (Panthera leo) less effective at killing their primary prey, plains zebra (Equus quagga). Big-headed ants disrupted the mutualism between native ants (Crematogaster spp.) and the dominant whistling-thorn tree (Vachellia drepanolobium), rendering trees vulnerable to elephant (Loxodonta africana) browsing and resulting in landscapes with higher visibility. Although zebra kills were significantly less likely to occur in higher-visibility, invaded areas, lion numbers did not decline since the onset of the invasion, likely because of prey-switching to African buffalo (Syncerus caffer). We show that by controlling biophysical structure across landscapes, a tiny invader reconfigured predator-prey dynamics among iconic species.

Assessment of multiple herbicide protection seed treatments for seed-based restoration of native perennial bunchgrasses and sagebrush across multiple sites and years.

The invasion of exotic, annual plant species is a leading contributor to ecological degradation in drylands globally, and the use of pre-emergent herbicide to control these species is common. Pre-emergent herbicides pose challenges for seed-based restoration due to toxicity to the seeds of desired species. Herbicide protection (HP) technologies pose a potential solution by using activated carbon seed treatments to protect desirable seeds from herbicide exposure. In the sagebrush steppe ecosystem of North America, we used an adaptive small plot design over three planting years to test for effects on seeding outcomes (seedling density and size) of large and small multi-seed HP pellets, several single-seed HP coatings, and carbon banding treatments at geographically dispersed sites for several perennial bunchgrasses and the keystone perennial shrub, Wyoming big sagebrush. We also compared different methods of seed delivery and litter pre-seeding management. Seeding success was low overall, especially for sagebrush, and it was clear that other, often less predictable barriers to establishment than herbicide exposure, such as inadequate spring moisture, were strong drivers of seeding outcomes. Despite this, HP treatments were associated with higher seedling density than bare seed in multiple instances, most notably for grasses. The large HP pellet occasionally outperformed the small HP pellet, and several HP coatings performed similarly to the small pellet. Surprisingly, we did not see consistent negative effects of pre-emergent herbicide on unprotected bare seed. We conclude that HP seed treatments show some promise to improve seeding success in the presence of herbicide, but that consistent success will require further improvements to HP treatments as well as integration with other innovations and approaches.

Flowering time advances since the 1970s in a sagebrush steppe community: Implications for management and restoration.

Climate change is widely known to affect plant phenology, but little is known about how these impacts manifest in the widespread sagebrush ecosystem of the Western United States, which supports a number of wildlife species of concern. Shifts in plant phenology can trigger consequences for the plants themselves as well as the communities of consumers that depend upon them. We assembled historical observations of first-flowering dates for 51 species collected in the 1970s and 1980s in a montane sagebrush community in the Greater Yellowstone Ecosystem and compared these to contemporary phenological observations targeting the same species and locations (2016-2019). We also assembled regional climate data (average spring temperature, day of spring snowmelt, and growing degree days) and tested the relationship between first-flowering time and these variables for each species. We observed the largest change in phenology in early-spring flowers, which, as a group, bloomed on average 17 days earlier, and as much as 36 days earlier, in the contemporary data set. Mid-summer flowers bloomed on average 10 days earlier, nonnative species 15 days earlier, and berry-producing shrubs 5 days earlier, while late summer flowering plants did not shift. The greatest correlates of early-spring and mid-summer flowering were average spring temperature and day of snowmelt, which was 21 days earlier, on average, in 2016-2019 relative to the 1973-1978 observations. The shifts in flowering phenology that we observed could indicate developing asynchronies or novel synchronies of these plant resources and wildlife species of conservation concern, including Greater Sage-grouse, whose nesting success is tied to availability of spring forbs; grizzly bears, which rely heavily on berries for their fall diet; and pollinators. This underscores the importance of maintaining a diverse portfolio of native plants in terms of species composition, genetics, phenological responsiveness to climatic cues, and ecological importance to key wildlife and pollinator species. Redundancy within ecological niches may also be important considering that species roles in the community may shift as climate change affects them differently. These considerations are particularly relevant to restoration and habitat-enhancement projects in sagebrush communities across western North America.

Demographic consequences of mutualism disruption: Browsing and big-headed ant invasion drive acacia population declines.

Across the globe, biological invasions have disrupted mutualisms, producing reverberating consequences for ecosystems. Although invasive species frequently trigger mutualism disruptions, few studies have quantified the demographic mechanisms by which mutualism breakdown may generate population effects. In a Kenyan savanna, the invasive big-headed ant (Pheidole megacephala) has disrupted a foundational mutualism between the monodominant whistling-thorn tree (Acacia drepanolobium) and native ants (Crematogaster spp.) that deter browsing by large mammalian herbivores. We conducted experiments to quantify the demographic consequences of this mutualism disruption in the presence and absence of large mammalian herbivores. Invasion by P. megacephala exacerbated population declines of A. drepanolobium, primarily through decreased survival and reproduction of adult trees. However, these fitness reductions were small compared to those resulting from the presence of large mammalian herbivores, which negatively impacted growth and survival. Contrary to expectation, the expulsion of metabolically costly Crematogaster mutualists by P. megacephala did not result in higher population growth rates for trees protected from large mammalian herbivores. Our results suggest that invasive P. megacephala may impose a direct metabolic cost to trees exceeding that of native mutualists while providing no protection from browsing by large mammalian herbivores. Across landscapes, we expect that invasion by P. megacephala will reduce A. drepanolobium populations, but that the magnitude and demographic pathways of this effect will hinge on the presence and abundance of browsers.

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.

Mutualism disruption by an invasive ant reduces carbon fixation for a foundational East African ant-plant.

Invasive ants shape assemblages and interactions of native species, but their effect on fundamental ecological processes is poorly understood. In East Africa, Pheidole megacephala ants have invaded monodominant stands of the ant-tree Acacia drepanolobium, extirpating native ant defenders and rendering trees vulnerable to canopy damage by vertebrate herbivores. We used experiments and observations to quantify direct and interactive effects of invasive ants and large herbivores on A. drepanolobium photosynthesis over a 2-year period. Trees that had been invaded for ≥ 5 years exhibited 69% lower whole-tree photosynthesis during key growing seasons, resulting from interaction between invasive ants and vertebrate herbivores that caused leaf- and canopy-level photosynthesis declines. We also surveyed trees shortly before and after invasion, finding that recent invasion induced only minor changes in leaf physiology. Our results from individual trees likely scale up, highlighting the potential of invasive species to alter ecosystem-level carbon fixation and other biogeochemical cycles.

Density dependence and the spread of invasive big-headed ants (Pheidole megacephala) in an East African savanna.

Supercolonial ants are among the largest cooperative units in nature, attaining extremely high densities. How these densities feed back into their population growth rates and how abundance and extrinsic factors interact to affect their population dynamics remain open questions. We studied how local worker abundance and extrinsic factors (rain, tree density) affect population growth rate and spread in the invasive big-headed ant, which is disrupting a keystone mutualism between acacia trees and native ants in parts of East Africa. We measured temporal changes in big-headed ant (BHA) abundance and rates of spread over 20 months along eight transects, extending from areas behind the front with high BHA abundances to areas at the invasion front with low BHA abundances. We used models that account for negative density dependence and incorporated extrinsic factors to determine what variables best explain variation in local population growth rates. Population growth rates declined with abundance, however, the strength of density dependence decreased with abundance. We suggest that weaker density dependence at higher ant abundances may be due to the beneficial effect of cooperative behavior that partially counteracts resource limitation. Rainfall and tree density had minor effects on ant population dynamics. BHA spread near 50 m/year, more than previous studies reported and comparable to rates of spread of other supercolonial ants. Although we did not detect declines in abundance in areas invaded a long time ago (> 10 years), continued monitoring of abundance at invaded sites may help to better understand the widespread collapse of many invasive ants.

Frenemy at the gate: Invasion by Pheidole megacephala facilitates a competitively subordinate plant ant in Kenya.

Biological invasions can lead to the reassembly of communities and understanding and predicting the impacts of exotic species on community structure and functioning are a key challenge in ecology. We investigated the impact of a predatory species of invasive ant, Pheidole megacephala, on the structure and function of a foundational mutualism between Acacia drepanolobium and its associated acacia-ant community in an East African savanna. Invasion by P. megacephala was associated with the extirpation of three extrafloral nectar-dependent Crematogaster acacia ant species and strong increases in the abundance of a competitively subordinate and locally rare acacia ant species, Tetraponera penzigi, which does not depend on host plant nectar. Using a combination of long-term monitoring of invasion dynamics, observations and experiments, we demonstrate that P. megacephala directly and indirectly facilitates T. penzigi by reducing the abundance of T. penzigi's competitors (Crematogaster spp.), imposing recruitment limitation on these competitors, and generating a landscape of low-reward host plants that favor colonization and establishment by the strongly dispersing T. penzigi. Seasonal variation in use of host plants by P. megacephala may further increase the persistence of T. penzigi colonies in invaded habitat. The persistence of the T. penzigi-A. drepanolobium symbiosis in invaded areas afforded host plants some protection against herbivory by elephants (Loxodonta africana), a key browser that reduces tree cover. However, elephant damage on T. penzigi-occupied trees was higher in invaded than in uninvaded areas, likely owing to reduced T. penzigi colony size in invaded habitats. Our results reveal the mechanisms underlying the disruption of this mutualism and suggest that P. megacephala invasion may drive long-term declines in tree cover, despite the partial persistence of the ant-acacia symbiosis in invaded areas.

Large herbivore loss has complex effects on mosquito ecology and vector-borne disease risk.

Loss of biodiversity can affect transmission of infectious diseases in at least two ways: by altering host and vector abundance or by influencing host and vector behaviour. We used a large herbivore exclusion experiment to investigate the effects of wildlife loss on the abundance and feeding behaviour of mosquito vectors and to explore consequences for vector-borne disease transmission. Large herbivore loss affected both mosquito abundance and blood-feeding behaviour. For Aedes mcintoshi, the dominant mosquito species in our study and a primary vector of Rift Valley fever virus (RVFV), abundance decreased with large herbivore loss, while blood feeding on humans increased. Despite an elevated human biting rate in the absence of large herbivores, we estimated that the potential for RVFV transmission to humans doubles in the presence of large herbivores. These results demonstrate that multiple effects of biodiversity loss on vectors can lead to counterintuitive outcomes for human disease risk.

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.

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.

Influence of neighboring plants on the dynamics of an ant-acacia protection mutualism.

Ant-plant protection symbioses, in which plants provide food and/or shelter for ants in exchange for protection from herbivory, are model systems for understanding the ecology of mutualism. While interactions between ants, host plants, and herbivores have been intensively studied, we know little about how plant-plant interactions influence the dynamics of these mutualisms, despite strong evidence that plants compete for resources, that hosting ants can be costly, and that host-plant provisioning to ants can therefore be constrained by resource availability. We used field experiments in a semiarid Kenyan savanna to examine interactions between the ant-plant Acacia drepanolobium, neighboring grasses, and two species of symbiotic acacia-ants with divergent behaviors: Crematogaster mimosae, an aggressive symbiont that imposes high costs to host trees via consumption of extrafloral nectar, and Tetraponera penzigi, a less-protective symbiont that imposes lower costs because it does not consume nectar. We hypothesized that by competing with acacias for resources, neighboring grasses (1) reduce hosts' ability to support costly C. mimosae, while having little or no effect on the ability of hosts to support low-cost T. penzigi, and (2) reduce sapling growth rates irrespective of ant occupant. We factorially manipulated the presence/absence of grasses and the identity of ant occupants on saplings and evaluated effects on colony survivorship and sapling growth rates over 40 weeks. Contrary to prediction, the high-cost/high-reward nectar-dependent mutualist C. mimosae had higher colony-survival rates on saplings with grass neighbors present. Grasses appear to have indirectly facilitated the survival of C. mimosae by reducing water stress on host plants; soils under saplings shaded by grasses had higher moisture content, and these saplings produced more active nectaries than grass-removal saplings. Consistent with prediction, survival of low-cost/low-reward T. penzigi did not differ significantly between grass-removal treatments. Saplings occupied by low-cost/low-reward T. penzigi grew 100% more on average than saplings occupied by high-cost/high-reward C. mimosae, demonstrating that mutualist-partner identity strongly and differentially influences demographic rates of young plants. In contrast, contrary to prediction, grass neighbors had no significant net impact on sapling growth rates. Our results suggest that neighboring plants can exert strong and counterintuitive effects on ant-plant protection symbioses, highlighting the need to integrate plant-plant interactions into our understanding of these mutualisms.

Parasite responses to large mammal loss in an African savanna.

Biodiversity loss can alter disease transmission; however, the magnitude and direction of these effects vary widely across ecosystems, scales, and pathogens. Here we experimentally examine the effects of one of the most globally pervasive patterns of biodiversity decline, the selective loss of large wildlife, on infection probability, intensity and population size of a group of common rodent-borne parasites - macroparasitic helminths. Consistent with previous work on vector-borne pathogens, we found that large wildlife removal causes strong and systematic increases of rodent-borne parasites, largely due to increases in rodent density, as rodents are released from competition with larger herbivores. Although we predicted that increased host density would also increase per capita infection among all directly transmitted parasites, this additional amplification occurred for only two of three examined parasites. Furthermore, the actual effects of large mammal loss on per capita infection were mediated by the complex suite of abiotic and biotic factors that regulate parasite transmission. Thus, while these results strongly suggest that large wildlife loss will cause systematic increases in rodent parasite populations, they also underscore the difficulty of making more specific predictions for a given parasite based on simple attributes such as transmission mode or life history strategy. Instead, detailed information on the ecology of each parasite species would be necessary to make more accurate predictions of how biodiversity loss will affect infection.

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.

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.

Disruption of a protective ant-plant mutualism by an invasive ant increases elephant damage to savanna trees.

Invasive species can indirectly affect ecosystem processes via the disruption of mutualisms. The mutualism between the whistling thorn acacia (Acacia drepanolobium) and four species of symbiotic ants is an ecologically important one; ants strongly defend trees against elephants, which can otherwise have dramatic impacts on tree cover. In Laikipia, Kenya, the invasive big-headed ant (Pheidole megacephala) has established itself at numerous locations within the last 10-15 years. In invaded areas on five properties, we found that three species of symbiotic Crematogaster ants were virtually extirpated, whereas Tetraponera penzigi co-occurred with P. megacephala. T. penzigi appears to persist because of its nonaggressive behavior; in a whole-tree translocation experiment, Crematogaster defended host trees against P. megacephala, but were extirpated from trees within hours. In contrast, T. penzigi retreated into domatia and withstood invading ants for >30 days. In the field, the loss of defensive Crematogaster ants in invaded areas led to a five- to sevenfold increase in the number of trees catastrophically damaged by elephants compared to uninvaded areas. In savannas, tree cover drives many ecosystem processes and provides essential forage for many large mammal species; thus, the invasion of big-headed ants may strongly alter the dynamics and diversity of East Africa's whistling thorn savannas by disrupting this system's keystone acaciaant mutualism.