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Soil carbon flux rates are a crucial metric of carbon cycling that contribute to calculating an ecosystem's carbon budget, and thus whether it is a source or sink of atmospheric carbon dioxide. However, soil carbon flux datasets are frequently low‐resolution across either space or time, limiting our abilities to identify small‐scale ecological contexts that influence soil carbon dynamics. Existing datasets are distributed unevenly, with some soil carbon‐rich regions (like tropical grasslands) significantly understudied. We developed an autonomous, inexpensive, do‐it‐yourself (DIY) soil carbon flux chamber (a “fluxbot”) and data processing software. We deployed a distributed array of 12 fluxbots in a long‐term experiment in a central Kenyan savanna where it has been logistically impossible to collect high‐resolution soil carbon flux data. With this array we collected over 10,000 individual flux estimates over almost two months, spanning the end of a dry season and the start of a wet season. With our successful deployment in situ, we demonstrate the potential for low‐cost, autonomous, DIY sensors in improving resolution of soil carbon flux datasets (particularly in under‐studied or logistically challenging systems). If implemented widely, such an improvement in data collection capacities could improve our understanding of ecological and climatic drivers of soil carbon flux dynamics on the local to global scale.

 

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 strictaandO. 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 ofO. strictapresence at the landscape level (6150 km²).

Opuntiadensities were 74% to 99% lower in almost all plots accessible to wild herbivores compared to exclosure plots.Opuntiadensities 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, modelledOpuntia strictaoccurrence probability was negatively correlated with estimated species richness of wild herbivores and elephant occurrence probability. On average,O. strictaoccurrence 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 ofOpuntiaby 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.

 

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 onAcacia drepanolobiumsapling 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.

 

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 inOdontotermestermite 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.

 

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.

 

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 <1 species).

Simulated extirpation of megaherbivores altered use intensity by mesoherbivores, which should be considered during (re)introductions of megaherbivores or their ecological proxies. Species' traits (in this case shoulder height) may be more reliable predictors of mesoherbivores' responses to megaherbivores than phylogenetic relatedness, and may be useful for predicting responses of data‐limited species.

 

Cattle and other livestock graze more than a quarter of the world's terrestrial area and are widely regarded to be drivers of global biodiversity declines. Studies often compare the effects of livestock presence/absence but, to our knowledge, no studies have tested for interactive effects between large wild herbivores and livestock at varying stocking rates on small‐bodied wild vertebrates.

We investigated the effects of cattle stocking rates (none/moderate/high) on the diversity of wildlife 0.05–1,000 kg using camera traps at a long‐term exclosure experiment within a semi‐arid savanna ecosystem in central Kenya. In addition, by selectively excluding wild ‘mesoherbivores’ (50–1,000 kg) and ‘megaherbivores’ (>1,000 kg; elephant and giraffe), we tested whether the presence of these two wild herbivore guilds (collectively, ‘larger wild herbivores’) mediates the effect of cattle stocking rate on habitat use and diversity of ‘smaller wildlife’ (mammals ranging between 10 and 70 cm shoulder height and birds).

Our results show that cattle enhance alpha diversity of smaller wildlife (with or without larger wild herbivore presence) and of all wildlife 0.05–1,000 kg (with or without megaherbivore presence), by altering vegetation structure. However, for smaller wildlife, this effect is less pronounced in the presence of larger wild herbivores, which also shorten grass. In the absence of cattle, mesoherbivore‐accessible sites showed higher alpha diversity of smaller wildlife than sites excluding mesoherbivores.

Smaller wildlife habitat use was increased by high cattle stocking rates and wild mesoherbivores more in the presence of the other.

Synthesis and applications. Our findings imply that grazing, whether by livestock or wildlife, can enhance local savanna wildlife diversity. The biodiversity benefits of localised increases in herbivory are likely to be due to shortened grass and associated visibility improvements (for predator avoidance/foraging). This suggests that land managers can increase local biodiversity by shortening grass, with wild or domestic herbivores (or both), at least in patches within a taller grass matrix.

 

Ecological theory posits that temporal stability patterns in plant populations are associated with differences in species' ecological strategies. However, empirical evidence is lacking about which traits, or trade-offs, underlie species stability, especially across different biomes. We compiled a worldwide collection of long-term permanent vegetation records (greater than 7000 plots from 78 datasets) from a large range of habitats which we combined with existing trait databases. We tested whether the observed inter-annual variability in species abundance (coefficient of variation) was related to multiple individual traits. We found that populations with greater leaf dry matter content and seed mass were more stable over time. Despite the variability explained by these traits being low, their effect was consistent across different datasets. Other traits played a significant, albeit weaker, role in species stability, and the inclusion of multi-variate axes or phylogeny did not substantially modify nor improve predictions. These results provide empirical evidence and highlight the relevance of specific ecological trade-offs, i.e. in different resource-use and dispersal strategies, for plant populations stability across multiple biomes. Further research is, however, necessary to integrate and evaluate the role of other specific traits, often not available in databases, and intraspecific trait variability in modulating species stability. 

Small-mammal faunas of tropical savannas consist of endemic assemblages of murid rodents, small marsupials, and insectivores on four continents. Small mammals in tropical savannas are understudied compared to other tropical habitats and other taxonomic groups (e.g., Afrotropical megafauna or Neotropical rainforest mammals). Their importance as prey, ecosystem engineers, disease reservoirs, and declining members of endemic biodiversity in tropical savannas compels us to understand the factors that regulate their abundance and diversity. We reviewed field studies published in the last 35 years that examined, mostly experimentally, the effects of varying three primary endogenous disturbances in tropical savanna ecosystems—fire, large mammalian herbivory (LMH), and drought—on abundance and diversity of non-volant small mammals. These disturbances are most likely to affect habitat structure (cover or concealment), food availability, or both, for ground-dwelling small mammalian herbivores, omnivores, and insectivores. Of 63 studies (included in 55 published papers) meeting these criteria from the Afrotropics, Neotropics, and northern Australia (none was found from southern Asia), 29 studies concluded that small mammals responded (mostly negatively) to a loss of cover (mostly from LMH and fire); four found evidence of increased predation on small mammals in lower-cover treatments (e.g., grazed or burned). Eighteen studies concluded a combination of food- and cover-limitation explained small-mammal responses to endogenous disturbances. Only two studies concluded small-mammal declines in response to habitat-altering disturbance were caused by food limitation and not related to cover reduction. Evidence to date indicates that abundance and richness of small savanna mammals, in general (with important exceptions), is enhanced by vegetative cover (especially tall grass, but sometimes shrub cover) as refugia for these prey species amid a “landscape of fear,” particularly for diurnal, non-cursorial, and non-fossorial species. These species have been called “decreasers” in response to cover reduction, whereas a minority of small-mammal species have been shown to be “increasers” or disturbance-tolerant. Complex relationships between endogenous disturbances and small-mammal food resources are important secondary factors, but only six studies manipulated or measured food resources simultaneous to habitat manipulations. While more such studies are needed, designing effective ones for cryptic consumer communities of omnivorous dietary opportunists is a significant challenge. 

There has been a long-standing interest in understanding how interactions between fire and herbivory influence woody vegetation dynamics in savanna ecosystems. However, controlled, replicated experiments examining how different fire regimes interact with different herbivore groups are rare. We tested the effects of single and repeated burns, crossed with six replicated herbivore treatments, on the mortality and growth of woody vegetation in the Kenya Long-term Exclosure Experiment plots located in a semi-arid savanna system in central Kenya. Burned plots experienced higher tree mortality overall, but differences between burns and non-burns were only significant in plots excluding all wild herbivores and in plots accessible to megaherbivores. Cattle ameliorated the negative effects of repeat burns on tree mortality, perhaps by suppressing fuel load accumulation. Across all herbivore treatments, trees experienced a significant reduction in height within the first two years after fire (top-kill), which was followed by a gradual recovery. Saplings and coppices subjected to repeated burns regrew faster than those that were burned once, except in the presence of megaherbivores. This study highlights strong context-dependent interactions between fire and different herbivore groups, and extends previous approaches to understanding fire–herbivory interactions, which have tended to lump the effects of different herbivore groups, or study them separately.