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Whether wild herbivores confer biotic resistance to invasion by exotic plants remains a key question in ecology. There is evidence that wild herbivores can impede invasion by exotic plants, but it is unclear whether and how this generalises across ecosystems with varying wild herbivore diversity and functional groups of plants, particularly over long-term (decadal) time frames. Using data from three long-term (13- to 26-year) exclosure experiments in central Kenya, we tested the effects of wild herbivores on the density of exotic invasive cacti, Opuntia stricta and O. ficus-indica (collectively, Opuntia), which are among the worst invasive species globally. We also examined relationships between wild herbivore richness and elephant occurrence probability with the probability of O. stricta presence at the landscape level (6150 km2). Opuntia densities were 74% to 99% lower in almost all plots accessible to wild herbivores compared to exclosure plots. Opuntia densities also increased more rapidly across time in plots excluding wild herbivores. These effects were largely driven by megaherbivores (≥1000 kg), particularly elephants. At the landscape level, modelled Opuntia stricta occurrence probability was negatively correlated with estimated species richness of wild herbivores and elephant occurrence probability. On average, O. stricta occurrence probability fell from ~0.56 to ~0.45 as wild herbivore richness increased from 6 to 10 species and fell from ~0.57 to ~0.40 as elephant occurrence probability increased from ~0.41 to ~0.84. These multi-scale results suggest that any facilitative effects of Opuntia by wild herbivores (e.g. seed/vegetative dispersal) are overridden by suppression (e.g. consumption, uprooting, trampling). Synthesis. Our experimental and observational findings that wild herbivores confer resistance to invasion by exotic cacti add to evidence that conserving and restoring native herbivore assemblages (particularly megaherbivores) can increase community resistance to plant invasions.

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., Our study provides unique descriptive and experimental evidence linking termite mound abundance to the availability of dietary resources. Further, we showed how two ecosystem engineers in this system, large mammalian herbivores and termites, are linked in previously unexplored ways.

The high mortality and low recruitment of the myrmecophytic Acacia drepanolobium are not consistent with the demographic rates of monodominant species. The positive conspecific density dependence observed from the spatial analysis is consistent with the defensive benefits conferred by symbiotic ants to trees when they grow close to each other. Patches of savanna dominated by Acacia drepanolobium occur throughout East Africa on nutrient-rich vertisols, also known as black cotton soils. We assessed the survival and recruitment for all freestanding trees with diameter at knee height (dkh) ≥ 10 mm in one of such mono-dominated patches (47 ha) at the Mpala Research Centre, Kenya, with the aim of identifying demographic traits that might explain the dominance of this species. Over a mean 6-year interval, mortality and recruitment rates in the habitat were 4.55%/year and 1.42%/year respectively, resulting in a net loss of 17.8% of the initial individuals. Of the 30 species recorded from the first census, 11 decreased in abundance, nine increased, and the remainder 10 did not change in abundance. The monodominant A. drepanolobium had a high mortality (4.69%/year), a low recruitment (1.31%/year), and a 19% population decline. There was no evidence of conspecific negative density dependence for this species. Rather, we found a statistically significant positive correlation between the number of conspecific neighbors and individual-level probability of survival, consistent with the “shared defense” benefits that symbiotic ant colonies occupying multiple trees can confer to these latter in a small neighborhood. Thus, mortality of A. drepanolobium was higher in areas where it occurred in lower densities, which resulted in an increase in the spatial aggregation of conspecifics. Mortality increased with dkh size classes and was mostly caused by elephants and stem-boring beetles. The demographic rates during the study period in theory are inconsistent with those of monodominant species. The protection against herbivory conferred by mutualistic ants associated with this species remains the most probable explanation of its dominance in this habitat.

Analysing temporal patterns in plant communities is extremely important to quantify the extent and the consequences of ecological changes, especially considering the current biodiversity crisis. Long-term data collected through the regular sampling of permanent plots represent the most accurate resource to study ecological succession, analyse the stability of a community over time and understand the mechanisms driving vegetation change. We hereby present the LOng-Term Vegetation Sampling (LOTVS) initiative, a global collection of vegetation time-series derived from the regular monitoring of plant species in permanent plots. With 79 data sets from five continents and 7,789 vegetation time-series monitored for at least 6 years and mostly on an annual basis, LOTVS possibly represents the largest collection of temporally fine-grained vegetation time-series derived from permanent plots and made accessible to the research community. As such, it has an outstanding potential to support innovative research in the fields of vegetation science, plant ecology and temporal ecology., The authors acknowledge institutional support as follows. Nicola J. Day: Te Apārangi Royal Society of New Zealand (Rutherford Postdoctoral Fellowship). Jiří Danihelka: Czech Science Foundation (project no. 19-28491X) and Czech Academy of Sciences (project no. RVO 67985939). Francesco de Bello: Spanish Plan Nacional de I+D+i (project PGC2018-099027-B-I00). Eric Garnier: La Fage INRA experimental station. Tomáš Herben: GAČR grant 20-02901S. Anke Jentsch: German Federal Ministry of Education and Research (grant 031B0516C - SUSALPS) and Oberfrankenstiftung (grant OFS FP00237). Norbert Juergens: German Federal Ministry of Education and Research (grant 01LG1201N - SASSCAL ABC). Frédérique Louault and Katja Klumpp: AnaEE-France (ANR-11-INBS-0001). Robin J. Pakeman: Strategic Research Programme of the Scottish Government's Rural and Environment Science and Analytical Services Division. Meelis Pärtel: Estonian Research Council (PRG609) and European Regional Development Fund (Centre of Excellence EcolChange). Josep Peñuelas: Spanish Government (grant PID2019-110521GB-I00), Fundación Ramon Areces (grant ELEMENTAL-CLIMATE), Catalan Government (grant SGR 2017-1005), and European Research Council (Synergy grant ERC-SyG-2013-610028, IMBALANCE-P). Ute Schmiedel: German Federal Ministry of Education and Research (Promotion numbers 01LC0024, 01LC0024A, 01LC0624A2, 01LG1201A, 01LG1201N). Hana Skálová: GAČR grant 20-02901S. Karsten Wesche: International Institute Zittau, Technische Universität Dresden. Susan K. Wiser: New Zealand Ministry for Business, Innovation and Employment’s Strategic Science Investment Fund. Ben A. Woodcock: NERC and BBSRC (NE/N018125/1 LTS-M ASSIST - Achieving Sustainable Agricultural Systems). Enrique Valencia: Program for attracting and retaining talent of Comunidad de Madrid (no. 2017-T2/AMB-5406) and Community of Madrid and Rey Juan Carlos University (Young Researchers R&D Project. Ref. M2165 – INTRANESTI). Truman P. Young: National Science Foundation (LTREB DEB 19-31224).

Fire is a major selective force on arid grassland communities, favoring traits such as the smoke‐induced seed germination response seen in a wide variety of plant species. However, little is known about the relevance of smoke as a cue for plants beyond the seedling stage. We exposed a fire‐adapted savanna tree, Vachellia (=Acacia) drepanolobium, to smoke and compared nutrient concentrations in leaf and root tissues to unexposed controls. Experiments were performed on three age cohorts: 2‐year‐old, 9‐month‐old, and 3‐month‐old plants. For the 2‐year‐old plants exposed to smoke, carbon and nitrogen concentrations were lower in the leaves and higher in the roots than controls. Less pronounced trends were found for boron and magnesium. In contrast, smoke‐exposed 3‐month‐old plants had lower root nitrogen concentrations than controls. No significant differences were found in the 9‐month‐old plants, and no significant shifts in other nutrient concentrations were observed between plant tissues for any of the three age cohorts. Synthesis: Our findings are consistent with smoke‐induced translocation of nutrients from leaves to roots in 2‐year‐old V. drepanolobium. This could represent a novel form of fire adaptation, with variation over the course of plant development. The translocation differences between age cohorts highlight the need to investigate smoke response in older plants of other species. Accounting for this adaptation could better inform our understanding of savanna community structure and nutrient flows under fire regimes altered by anthropogenic land use and climate change.

Analysing temporal patterns in plant communities is extremely important to quantify the extent and the consequences of ecological changes, especially considering the current biodiversity crisis. Long-term data collected through the regular sampling of permanent plots represent the most accurate resource to study ecological succession, analyse the stability of a community over time and understand the mechanisms driving vegetation change. We hereby present the LOng-Term Vegetation Sampling (LOTVS) initiative, a global collection of vegetation time-series derived from the regular monitoring of vascular plants in permanent plots. With 79 datasets from five continents and 7789 vegetation time-series monitored for at least six years and mostly on an annual basis, LOTVS possibly represents the largest collection of temporally fine-grained vegetation time-series derived from permanent plots and made accessible to the research community. As such, it has an outstanding potential to support innovative research in the fields of vegetation science, plant ecology and temporal ecology.

Fire is a major selective force on arid grassland communities, favoring traits such as the smoke‐induced seed germination response seen in a wide variety of plant species. However, little is known about the relevance of smoke as a cue for plants beyond the seedling stage.We exposed a fire‐adapted savanna tree, Vachellia (=Acacia) drepanolobium, to smoke and compared nutrient concentrations in leaf and root tissues to unexposed controls. Experiments were performed on three age cohorts: 2‐year‐old, 9‐month‐old, and 3‐month‐old plants.For the 2‐year‐old plants exposed to smoke, carbon and nitrogen concentrations were lower in the leaves and higher in the roots than controls. Less pronounced trends were found for boron and magnesium.In contrast, smoke‐exposed 3‐month‐old plants had lower root nitrogen concentrations than controls. No significant differences were found in the 9‐month‐old plants, and no significant shifts in other nutrient concentrations were observed between plant tissues for any of the three age cohorts. Synthesis: Our findings are consistent with smoke‐induced translocation of nutrients from leaves to roots in 2‐year‐old V. drepanolobium. This could represent a novel form of fire adaptation, with variation over the course of plant development. The translocation differences between age cohorts highlight the need to investigate smoke response in older plants of other species. Accounting for this adaptation could better inform our understanding of savanna community structure and nutrient flows under fire regimes altered by anthropogenic land use and climate change., We exposed a fire‐adapted savanna tree, Vachellia (=Acacia) drepanolobium, to smoke and compared nutrient concentrations in leaf and root tissues to unexposed controls. Our findings suggest trees may detect smoke and respond by translocating nutrients from leaves to roots, a potential mechanism for improving survival and post‐fire resprouting.

The extinction of 80% of megaherbivore (>1,000 kg) species towards the end of the Pleistocene altered vegetation structure, fire dynamics and nutrient cycling world-wide. Ecologists have proposed (re)introducing megaherbivores or their ecological analogues to restore lost ecosystem functions and reinforce extant but declining megaherbivore populations. However, the effects of megaherbivores on smaller herbivores are poorly understood. We used long-term exclusion experiments and multispecies hierarchical models fitted to dung counts to test (a) the effect of megaherbivores (elephant and giraffe) on the occurrence (dung presence) and use intensity (dung pile density) of mesoherbivores (2–1,000 kg), and (b) the extent to which the responses of each mesoherbivore species was predictable based on their traits (diet and shoulder height) and phylogenetic relatedness. Megaherbivores increased the predicted occurrence and use intensity of zebras but reduced the occurrence and use intensity of several other mesoherbivore species. The negative effect of megaherbivores on mesoherbivore occurrence was stronger for shorter species, regardless of diet or relatedness. Megaherbivores substantially reduced the expected total use intensity (i.e. cumulative dung density of all species) of mesoherbivores, but only minimally reduced the expected species richness (i.e. cumulative predicted occurrence probabilities of all species) of mesoherbivores (by

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.

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 remains 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 and the second a tree removal experiment– to demonstrate that patterns in termite mound abundance and cover are unexpectedly dynamic. Termite mound abundance, but not areal cover not significantly, is positively associated with experimentally controlled presence of cattle, but not wild mesoherbivores (15-1000 kg) or megaherbivores (elephants and giraffes). Herbaceous productivity and tree density, termite dietary resources that 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 is intimately tied to herbivore-driven resource availability.

Excluding large native mammals is an inverse test of rewilding. A 25-year exclosure experiment in an African savanna rangeland offers insight into the potentials and pitfalls of the rewilding endeavor as they relate to the native plant community. A broad theme that has emerged from this research is that entire plant communities, as well as individual plants, adjust to the absence of herbivores in ways that can ill-prepare them for the return of these herbivores. Three lines of evidence suggest that these “naïve” individuals, populations, and communities are likely to initially suffer from herbivore rewilding. First, plots protected from wild herbivores for the past 25 years have developed rich diversity of woody plants that are absent from unfenced plots, and presumably would disappear upon rewilding. Second, individuals of the dominant tree in this system, Acacia drepanolobium, greatly reduce their defences in the absence of browsers, and the sudden arrival of these herbivores (in this case, through a temporary fence break), resulted in far greater elephant damage than for their conspecifics in adjacent plots that had been continually exposed to herbivory. Third, the removal of herbivores favoured the most palatable grass species, and a large number of rarer species, which presumably would be at risk from herbivore re-introduction. In summary, the native communities that we observe in defaunated landscapes may be very different from their pre-defaunation states, and we are likely to see some large changes to these plant communities upon rewilding with large herbivores, including potential reductions in plant diversity. Lastly, our experimental manipulation of cattle represents an additional test of the role of livestock in rewilding. Cattle are in many ways ecologically dissimilar to wildlife (in particular their greater densities), but in other ways they may serve as ecological surrogates for wildlife, which could buffer ecosystems from some of the ecological costs of rewilding. More fundamentally, African savannah ecosystems represent a challenge to traditional Western definitions of “wilderness” as ecosystems free of human impacts. We support the suggestion that as we “rewild” our biodiversity landscapes, we redefine “wildness” in the 21st Century to be inclusive of (low impact, and sometimes traditional) human practices that are compatible with the sustainability of native (and re-introduced) biodiversity.

Understanding animals’ use of space can shed valuable light on multiple other aspects of behavioral ecology, including social organization, dispersal, and foraging efficiency. Home ranges, territories, core areas, and home range overlaps have been widely studied, but unless animals are directly observed or are tracked remotely on a fine temporal scale, how they actually use the space available to them and how they share (or partition) this space with a community of conspecifics over time cannot be fully understood. Using GPS technology, we tracked three adjacent groups of vervet monkeys (Chlorocebus pygerythrus) in Laikipia, Kenya, for 1 year to better understand the processes involved in territoriality and home range overlaps. Home ranges overlapped with any one neighboring group by 12.7–34.7%, but intergroup encounters only occurred in restricted areas within these zones, which defined territorial boundaries. The resources closest to the territorial boundaries were nocturnal core areas with preferred sleeping sites adjacent to short-grass areas offering fewer hiding places for ambush predators and greater visibility for predator detection. Home range overlaps were not neutral zones, the result of shifting home range boundaries over time, or based on intergroup encounters at boundaries, but resulted when groups made incursions beyond their territorial boundaries while the neighbor was far away and likely unaware of the intruders. Thus, territories can be non-exclusive but may still be perceived by the animals themselves as sole-owned, as neighbors only intrude when territory owners are absent from that area. How animals use their space is a perennial focus in behavioral ecology because space use influences nearly everything else in animals’ lives. While tracking has long been used to investigate space use, particularly for nocturnal animals, only direct observation or tracking on a fine scale can reveal the processes involved in the creation or maintenance of home ranges and territories and that cause home range overlaps to persist. We remotely tracked three adjacent groups of vervet monkeys on a fine temporal scale for 1 year. Despite behaving territorially, vervet home ranges overlapped well beyond territorial boundaries because neighbors were “absentee owners,” unable to prevent incursions when they were not nearby. Our findings suggest that exclusivity of use in the definition of territoriality may be too strict; animals may behave as if they are exclusive owners even if neighbors’ use of space does not reflect it.

Background An estimated 59,000 people die from rabies annually, with 99% of those deaths attributable to bites from domestic dogs (Canis lupus familiaris). This preventable Neglected Tropical Disease has a large impact across continental Africa, especially for rural populations living in close contact with livestock and wildlife. Mass vaccinations of domestic dogs are effective at eliminating rabies but require large amounts of resources, planning, and political will to implement. Grassroots campaigns provide an alternative method to successful implementation of rabies control but remain understudied in their effectiveness to eliminate the disease from larger regions. Methodology/Principal Findings We report on the development, implementation, and effectiveness of a grassroots mass dog rabies vaccination campaign in Kenya, the Laikipia Rabies Vaccination Campaign. During 2015–2017, a total of 13,155 domestic dogs were vaccinated against rabies in 17 communities covering approximately 1500 km2. Based on an estimated population size of 34,275 domestic dogs, percent coverages increased across years, from 2% in 2015 to 24% in 2017, with only 3 of 38 community-years of vaccination exceeding the 70% target. The average cost of vaccinating an animal was $3.44 USD with in-kind contributions and $7.44 USD without in-kind contributions. Conclusions/Significance The evolution of the Laikipia Rabies Vaccination Campaign from a localized volunteer-effort to a large-scale program attempting to eliminate rabies at the landscape scale provides a unique opportunity to examine successes, failures, and challenges facing grassroots campaigns. Success, in the form of vaccinating more dogs across the study area, was relatively straightforward to achieve. However, lack of effective post-vaccination monitoring and education programs, limited funding, and working in diverse community types appeared to hinder achievement of 70% coverage levels. These results indicate that grassroots campaigns will inevitably be faced with a philosophical question regarding the value of local impacts versus their contributions to a larger effort to eliminate rabies at the regional, country, or global scale., Author summary Given the importance of mass vaccinations of domestic dogs towards eliminating human rabies in Africa and the site-specific challenges facing such campaigns, additional studies on the development and implementation of such efforts are needed. One mechanism of mass vaccination lies in grassroots efforts that often begin at a very local scale and either develop into larger campaigns, remain local, or cease to persist past several years once interest and funding is exhausted. Here, we discuss the development of a grassroots campaign in Laikipia County, Kenya from its local inception to its development into a county-wide rabies elimination effort. Our results highlight challenges associated with achieving the targeted 70% coverage rate, including a need for consistent and systematic demographic monitoring of dog populations, limitations of the central point method, and logistical and financial challenges facing a volunteer-based effort. Serious political commitment from both the local and national governments are necessary to take the budget beyond what a crowdfunded campaign can raise, including availability and access to quality dog rabies vaccines. Without such outside support and substantial time to grow, grassroots campaigns might be better relegated to raising awareness and vaccinating dogs in small communities to protect those communities directly, without contributing to the broader ecosystem-wide transmission-stopping aim often sought by government human health and veterinary organizations.

Wild herbivore populations are declining in many African savannas, which is related to replacement by livestock (mainly cattle) and the loss of megaherbivores. Although some livestock management practices may be compatible with the conservation of native savanna biodiversity, the sustainability of these integrated wild herbivore/livestock management practices is unknown. For instance, how will these herbivore mixes influence key processes for the long-term functioning of savanna ecosystems, such as soil carbon, nitrogen and phosphorus pools and cycling? The Kenya Long-term Exclosure Experiment studies the ecosystem consequences of manipulating the presence and absence of wild herbivores and cattle at moderate densities in a ‘black cotton’ savanna. Here we show that after 20 years, cattle presence decreased total soil carbon and nitrogen pools, while the presence of megaherbivores (mainly elephants) increased these pools and even reversed the negative effects of cattle. Our results suggest that a mix of cattle at moderate densities and wild herbivores can be sustainable, provided that the assemblage of wild herbivores includes the largest species. Cattle are replacing wildlife in many African savannas. This field study finds that wild megaherbivores, such as elephants, increased soil carbon and nitrogen, and hence soil fertility, normally lost when only cattle are present.

On rangelands worldwide, cattle interact with many forms of biodiversity, most obviously with vegetation 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 recently (2013) crossed these treatments with small-scale controlled burns. These replicated experimental treatments simulate different land management practices. We seek to disentangle the complex relationships between livestock and biodiversity in a biome where worldwide, uneasy coexistence is the norm. Here, we synthesize more than 20 yr of data to address three central questions about the potentially unique role of cattle in savanna ecology: 1) To what extent do cattle and wild herbivores compete with or facilitate each other? 2) Are the effects of cattle on vegetation similar to those of wildlife, or do cattle have unique effects? 3) What effects do cattle and commercial cattle management have on other savanna organisms? We found that 1) Cattle compete at least as strongly with browsers as grazers, and wildlife compete with cattle, although these negative effects are mitigated by cryptic herbivores (rodents), rainfall, fire, and elephants. 2) Cattle effects on herbaceous vegetation (composition, productivity) are similar to those of the rich mixture of ungulates they replace, differing mainly due to the greater densities of cattle. In contrast, cattle, wild mesoherbivores, and megaherbivores have strongly guild-specific effects on woody vegetation. 3) Both cattle and wild ungulates regulate cascades to other consumers, notably termites, rodents, and disease vectors (ticks and fleas) and pathogens. Overall, cattle management, at moderate stocking densities, can be compatible with the maintenance of considerable native biodiversity, although reducing livestock to these densities in African rangelands is a major challenge.

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.

Summary Prescribed burning is used in tropical savannas to improve habitat conditions for domestic and wild herbivores, but its effects on the ecological interactions between these herbivore guilds have never been assessed experimentally. Understanding such effects will contribute towards more informed management of both guilds in landscapes where they share habitats. We investigated the effects of burning on the nutritional outcomes for cattle sharing habitat with wildlife in a Kenyan savanna ecosystem. We compared forage availability and cattle forage and nutrient intake rates across burned and unburned areas cattle accessed exclusively, and those they shared with medium-sized wild ungulates, both with and without megaherbivores (elephants and giraffes). We performed these measurements in May 2013 (wet period, 2 months post-burning) and February 2014 (dry period, 11 months post-burning). Additionally, we monitored wildlife use of these areas. Prescribed burning enhanced cattle nutrition, but only in areas cattle did not share with wildlife. Shared foraging with wildlife reduced cattle forage and nutrient intake rates by 37–97% in burned areas (burns), but not in unburned areas; these reductions corresponded with reduced herbage availability in the shared burns. In May (the wet period), cattle met their nutrient intake requirements in burns, regardless of whether they were sharing these areas with wildlife. However, in February (the dry period), nutrient requirements were unmet or tended to be unmet in burns shared with wildlife; requirements were met or significantly exceeded in the unshared burns. Experimental exclusion of megaherbivores did not moderate these effects, suggesting that they were primarily caused by medium-sized wild ungulates which were highly attracted to burns. Synthesis and applications. Prescribed burning produces negative nutritional outcomes for cattle when sharing habitat with wild ungulates. Because these effects could negatively influence livestock–wildlife coexistence, burning should be applied prudently in such human-occupied savanna landscapes. Specifically, because unburned areas serve as refuge foraging areas during the dry season, interspersing burns with unburned areas could minimize fire-driven negative interactions between cattle and wild ungulates. Conversely, burning could be used to draw wildlife away from valuable cattle foraging areas, such as those near available water.

Made available in DSpace on 2018-11-26T17:44:52Z (GMT). No. of bitstreams: 0 Previous issue date: 2018-01-26 Robert Bosch Foundation Animal movement is fundamental for ecosystem functioning and species survival, yet the effects of the anthropogenic footprint on animal movements have not been estimated across species. Using a unique GPS-tracking database of 803 individuals across 57 species, we found that movements of mammals in areas with a comparatively high human footprint were on average one-half to one-third the extent of their movements in areas with a low human footprint. We attribute this reduction to behavioral changes of individual animals and to the exclusion of species with long-range movements from areas with higher human impact. Global loss of vagility alters a key ecological trait of animals that affects not only population persistence but also ecosystem processes such as predator-prey interactions, nutrient cycling, and disease transmission. 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Fitotecn, P-7002554 Evora, Portugal Univ Evora, ICAAM, Inst Mediterranean Agr & Environm Sci, Evora, Portugal Univ Montpellier, Univ Paul Valery Montpellier, CNRS,EPHE, Ctr Ecol Fonctionnelle & Evolut,UMR 5175, F-34293 Montpellier 5, France Phillip Isl Nat Parks, Cowes, Vic, Australia Monash Univ, Sch Biol Sci, Melbourne, Vic, Australia Ohio State Univ, Dept Civil Environm & Geodet Engn, Columbus, OH 43210 USA Fiji Natl Univ, Dept Biol, POB 5529, Natabua, Lautoka, Fiji Univ Massachusetts, US Geol Survey, Massachusetts Cooperat Fish & Wildlife Res Unit, Amherst, MA 01003 USA Penn State Univ, US Geol Survey, Penn Cooperat Fish & Wildlife Res Unit, University Pk, PA 16802 USA Univ Oxford, Dept Zool, Oxford OX1 3PS, England Giraffe Conservat Fdn, POB 86099, Eros, Namibia German Primate Ctr, Behav Ecol & Sociobiol Unit, D-37077 Gottingen, Germany Tech Univ Munich, Dept Ecol & Ecosyst Management, Restorat Ecol, D-85354 Freising Weihenstephan, Germany Cary Inst Ecosyst Studies, Millbrook, NY 12545 USA Univ British Columbia, Irving K Barber Sch Arts & Sci, Unit 2, Biol, Okanagan Campus, Kelowna, BC V1V 1V7, Canada Univ Zurich, Dept Evolutionary Biol & Environm Studies, CH-8057 Zurich, Switzerland Univ Wyoming, Dept Zool & Physiol, Laramie, WY 82071 USA Univ Washington, Sch Environm & Forest Sci, Seattle, WA 98195 USA Univ Montana, Coll Forestry & Conservat, Dept Ecosyst & Conservat Sci, Wildlife Biol Program, Missoula, MT 59812 USA Bavarian Forest Natl Pk, Dept Conservat & Res, D-94481 Grafenau, Germany Albert Ludwigs Univ Freiburg, Wildlife Ecol & Management, D-79106 Freiburg, Germany Univ Toulouse, INRA, CEFS, Castanet Tolosan, France Univ Calif Davis, Anim Behav Grad Grp, Davis, CA 95616 USA Bionet Natr Onderzoek, NL-6171 EL Stein, Netherlands Univ Vet Med Vienna, Res Inst Wildlife Ecol, A-1160 Vienna, Austria Univ Coll Cork, Sch Biol Earth & Environm Sci, Cork, Ireland Univ Wyoming, US Geol Survey, Dept Zool & Physiol, Wyoming Cooperat Fish & Wildlife Res Unit, Laramie, WY USA North Carolina Museum Nat Sci, Raleigh, NC 27601 USA North Carolina State Univ, Dept Forestry & Environm Resources, Raleigh, NC 27695 USA Karatina Univ, Dept Nat Resource Management, POB 195710101, Karatina, Kenya Univ Lethbridge, Dept Psychol, Lethbridge, AB T1K 3M4, Canada Columbia Univ, Lamont Doherty Earth Observ, Palisades, NY 10964 USA Univ Valencia, Terr Vertebrates Grp, Cavanilles Inst Biodivers & Evolutionary Biol, E-46980 Valencia, Spain SUNY Stony Brook, Dept Anthropol, Stony Brook, NY 11794 USA TSG, SSC, IUCN, Rua Licuala 622,Damha 1,Campo Grande, BR-79046150 Mato Grosso Sul, Brazil IPE Inst Pesquisas Ecol Inst Ecol Res, Caixa Postal 47, BR-12960000 Sao Paulo, Brazil Univ Alicante, Dept Ciencias Ambient & Recursos Natur, Vertebrates Zool Res Grp, Alicante, Spain Embrapa Pantanal, BR-79320900 Corumba, MS, Brazil Chico Mendes Inst Conservat Biodivers, Natl Res Ctr Carnivores Conservat, BR-12952011 Atiba, SP, Brazil Univ Glasgow, Inst Biodivers Anim Hlth & Comparat Med, Glasgow, Lanark, Scotland NYU, Dept Biol, Ctr Genom & Syst Biol, New York, NY 10003 USA Univ Calif Davis, Dept Microbiol & Mol Genet, Davis, CA 95616 USA Univ Oslo, Dept Biosci, Ctr Ecol & Evolutionary Synth, NO-0316 Oslo, Norway Hebrew Univ Jerusalem, Alexander Silberman Inst Life Sci, Dept Ecol Evolut & Behav, Movement Ecol Lab, IL-91904 Jerusalem, Israel Norwegian Inst Nat Res, NO-0349 Oslo, Norway Norwegian Univ Sci & Technol NTNU, Dept Math Sci & Ctr Biodivers Dynam, N-7491 Trondheim, Norway Univ Fed Mato Grosso do Sul, Dept Ecol, BR-79070900 Campo Grande, MS, Brazil Field Museum Nat Hist, Integrat Res Ctr, Chicago, IL 60605 USA Consorzio Parco Nazl Stelvio, Bormio, Sondrio, Italy Univ Grenoble Alpes, Irstea, UR LESSEM, BP 76, F-38402 St Martin Dheres, France Univ Bayreuth, BayCEER, D-95447 Bayreuth, Germany Natl Park Schwarzwald, D-77889 Seebach, Germany Univ New South Wales, Evolut & Ecol Res Ctr, Sydney, NSW 2052, Australia Univ New South Wales, Sch Biol Earth & Environm Sci, 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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.

Euclea divinorum, a fast establishing, unpalatable, and fire resistant bush is considered an invasive species in some parts of its range. In Ol Pejeta Conservancy (OPC), Kenya, E. divinorum bushes cover ˜27% of the total area (˜9470 ha.) and has been expanding in coverage and encroaching into A. drepanolobium woodlands, a key woody habitat for the endangered black rhino. Between 2006 and 2010, we assessed the spatial distribution, annual rates of spread and recruitment of E. divinorum in OPC. We used data from satellite imagery and belt transects laid at the transition between E. divinorum habitat and other habitats. Density of E. divinorum seedlings increased by 27% annually over five years, with more seedlings establishing in grassland habitat (56.6%) than in A. drepanolobium woodland (43.4%). Within the infection frontier, the number of seedlings was high at the ecotone and reduced predictably with an increase in distance into the infection zone. Increase in rainfall facilitated recruitment and survival of E. divinorum seedlings and also reduced damage on trees and seedlings by mega-herbivores, especially elephants. This study confirms the encroachment of E. divinorum bushland into other habitats. This has been accelerated by burning and damage to A. drepanolobium habitat thus opening it up to encroachment by E. divinorum. Resume Euclea divinorum est un buisson qui s'installe rapidement, d'un gout desagreable et resistant au feu. Il est considere comme une espece invasive dans certaines parties de son aire de repartition. Dans l'Ol Pejeta Conservancy (OPC), au Kenya, les buissons d'E. divinorum couvrent quelque 27% de la superficie totale (env. 9470 ha) et ils s'etendent encore et envahissent les forets a A. drepanolobium, un habitat forestier crucial pour le rhinoceros noir qui est une espece en danger. Entre 2006 et 2010, nous avons evalue la distribution spatiale, le taux de dispersion annuel et le recrutement d'E. divinorum dans l'OPC. Nous avons utilise des donnees de l'imagerie satellitaire et des belt-transects traces sur la ligne de transition entre l'habitat d'E. divinorum et les autres habitats. La densite des jeunes plants d'E. divinorum a augmente de 27% par an durant ces cinq ans, et il y avait plus de jeunes plants qui s'etablissaient dans un habitat de prairie (56.6%) que dans la foret a A. drepanolobium (43.4%). A la frontiere des envahissements, le nombre de jeunes plants est eleve dans l'ecotone et se reduit comme attendu avec l'augmentation de la distance par rapport a la zone d'infection. Une augmentation des chutes de pluie facilitait le recrutement et la survie des jeunes plants d'E. divinorum et reduisait aussi les degâts causes aux arbres et aux jeunes plants par les mega-herbivores, specialement par les elephants. Cette etude confirme l'envahissement des buissons d'E. divinorum dans d'autres habitats. Ce phenomene a ete accelere par les feux et par les degâts causes a l'habitat a A. drepanolobium qui s'est ainsi ouvert a l'envahissement d'E. divinorum.

Data collected on a free ranging group of Tana River mangabeys (Cercocebus galeritus) indicates that this endangered primate species, which has previously been regarded as a seed predator, plays an important role in seed dispersal and do contribute to the regeneration of a highly fragmented gallery forest. We observed fruit handling behavior and the post-dispersal fate of seeds ingested by the mangabeys. The two main fruit handling behaviors observed, fruit swallowing and processing fruits in cheek pouches, positively contributed to seed dispersal. Seed predation was not common during this study, apart from a few particular plant species such as Acacia robusta and Alangium salviifolium with non-fleshy fruits. We found a correlation between seed predation and fruit availability, respectively seasonal differential specific dispersal efficiency. The role of the mangabeys in dispersing seeds and facilitating forest regeneration is enhanced by their movement across forest patches through non-forested matrix, which contributes to the deposition of seeds and regeneration in these habitat gaps.

Survival and recruitment of the whistling thorn (Acacia drepanolobium Sjostedt) seedlings was studied in Olpejeta Conservancy, Kenya, between 1999 and 2009. Seedlings were monitored for damage by drought and herbivores as well as survival and recruitment into adult trees. We determined variability in seedling damage, survival and recruitment before and after two management interventions; the elephant translocation in 2001 and the expansion of the reserve in 2007. Seedling survival over 8 years was low, with mortalities being initially high between 2000 and 2001. Reduction in elephant numbers was followed by a gradual reduction in seedling mortality. We recorded a 47% survival over 8 years; 63% of the mortality having been caused by browsers, drought or other factors. Only 8 (0.75%) of the seedlings had grown into trees. Seedlings experienced greater damage at higher densities and exhibited a storage effect underneath grass cover. Survival and recruitment of A. drepanolobium seedlings is very low and of concern. The expansion of conservation area and the opening of wildlife migration corridors have, however, resulted in a steady recovery for seedlings. We recommend enclosing sections of the conservancy to exclude grazers and browsers such as elephants and to allow seedling survival and recruitment. Resume La survie et le recrutement des jeunes plants d’Acacia drepanolobium Sjostedt ont eteetudies au Ol Pejeta Conservancy, au Kenya, entre 1999 et 2009. Les jeunes plants furent suivis tant au point de vue des dommages causes par la secheresse et par les herbivores qu’a celui de la survie et du recrutement jusqu’au stade adulte. Nous avons determine la variabilite des dommages causes aux plants, de leur survie et de leur recrutement avant et apres deux interventions de gestion: la translocation d’elephants en 2001 et l’extension de la reserve en 2007. La survie des jeunes plants fut faible sur ces huit annees, la mortaliteetant particulierement forte entre 2000 et 2001. La reduction du nombre d’elephants fut suivie d’une reduction progressive de la mortalite des plants. Nous avons enregistre un taux de survie de 47% sur huit ans, 63% de la mortalite ayant ete causee par les herbivores, la secheresse ou d’autres facteurs. Seuls huit (0.75%) jeunes plants sont devenus des arbres. Les jeunes plants subissaient plus de degâts lorsque leur densiteetait plus forte et presentaient un effet de stockage sous la couverture herbacee. La survie et le recrutement des jeunes plants d’Acacia drepanolobium sont tres faibles, voire inquietants. L’extension de la superficie de la Conservancy et l’ouverture de corridors de migration pour la faune sauvage ont cependant abouti a une restauration prononcee des jeunes plants. Nous recommandons de cloturer certaines sections de la reserve pour en exclure les herbivores, comme les elephants, afin de permettre la survie et le recrutement des jeunes plants.

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.

Both wild and prescribed fire in savanna ecosystems influence habitat use by herbivores by creating or maintaining spatial and temporal heterogeneity in forage quality and vegetation cover. Yet little is known about how spatial scales influence long-term persistence of fire effects. We examined changes over a 6-year period in herbivore preference for experimentally burned patches that varied in spatial extent and grain. Avoidance for the burns by elephants and preference for the burns by impala and Grant’s gazelle decreased significantly. For the rest of the species (zebra, eland, oryx, hartebeest, warthog and hare), there were no significant changes in preference for the burns. Changes in preference for the burned areas depended on the spatial extent and grain of the burn, with intermediate-size (9-ha) burns and large (8-ha) patchy burns being more preferred 6–7 years after fire. Grain, but not the spatial extent of the burned area, influenced changes in grass height. Fire resulted in a delayed reduced tree density irrespective of the spatial scale of the burn. Results of this study indicate that, depending on the scale of fire prescription, the impacts of fire on herbivores may last longer than previous studies suggest.

Although livestock and wildlife share most of their ranges worldwide, little controlled experimental research has been done on their interactions. Since 1995 we have been manipulating the presence of cattle and large wild ungulates in a Kenyan savanna rangeland in order to better understand the nature of competition and coexistence between these two guilds of herbivores and how they affect biodiversity. In a replicated experiment in which different combinations of cattle and wild herbivores are allowed access to large-scale plots, we have been monitoring the impacts of these herbivores on vegetation, on the wild herbivores, and cattle themselves, and on a variety of other taxa. We have also been conducting experimental research to examine other ways in which livestock management in eastern Africa might affect biodiversity. These include studies on the impacts of fire, livestock corrals, and changes in tree density. This research has revealed the following patterns. (1) Cattle suppress many species of wild herbivores, presumably through competition for their shared resources. The nature of this competition, however, is contingent on rainfall and the presence of other herbivores. (2) Wild herbivores both compete with and facilitate cattle, depending on rainfall. (3) The pastoral practice of housing livestock nightly in protective corral enclosures (“bomas”) over time produces long-lived nutrient hotspots preferred by both livestock and wild herbivores. (4) Fire, frequently used by pastoralists in the past, is valuable for improving grass quality, with benefits for many species of wild herbivores. (5) Pastoral practices that reduce woody cover, including burning and boma construction, create local habitat patches that are preferred by wild herbivores, apparently for their greater anti-predator visibility. (6) Despite competition between livestock and wild herbivores, coexistence between these two guilds can be managed, and there are several positive (facilitative) pathways between livestock husbandry and wild herbivores and other biodiversity.