Interplay of competition and facilitation in grazing succession by migrant Serengeti herbivores.

Competition, facilitation, and predation offer alternative explanations for successional patterns of migratory herbivores. However, these interactions are difficult to measure, leaving uncertainty about the mechanisms underlying body-size-dependent grazing-and even whether succession occurs at all. We used data from an 8-year camera-trap survey, GPS-collared herbivores, and fecal DNA metabarcoding to analyze the timing, arrival order, and interactions among migratory grazers in Serengeti National Park. Temporal grazing succession is characterized by a "push-pull" dynamic: Competitive grazing nudges zebra ahead of co-migrating wildebeest, whereas grass consumption by these large-bodied migrants attracts trailing, small-bodied gazelle that benefit from facilitation. "Natural experiments" involving intense wildfires and rainfall respectively disrupted and strengthened these effects. Our results highlight a balance between facilitative and competitive forces in co-regulating large-scale ungulate migrations.

Effects of migratory animals on resident parasite dynamics.

Migratory animals can bring parasites into resident animal (i.e., non-migratory) home ranges (transport effects) and exert trophic effects that either promote or reduce parasite exposure to resident hosts. Here, we examine the importance of these transport and trophic effects and their interactions for resident parasite dynamics. We propose that migrant transport and trophic effects are impacted by the number of migratory animals entering a resident's home range (migration intensity), the amount of time that migrants spend within a resident's home range (migration duration), and the timing of migrant-resident interactions. We then incorporate migration intensity, duration, and timing into a framework for exploring the net impact of migrant trophic and transport effects on resident animal parasite prevalence.

The generality of cryptic dietary niche differences in diverse large-herbivore assemblages.

Ecological niche differences are necessary for stable species coexistence but are often difficult to discern. Models of dietary niche differentiation in large mammalian herbivores invoke the quality, quantity, and spatiotemporal distribution of plant tissues and growth forms but are agnostic toward food plant species identity. Empirical support for these models is variable, suggesting that additional mechanisms of resource partitioning may be important in sustaining large-herbivore diversity in African savannas. We used DNA metabarcoding to conduct a taxonomically explicit analysis of large-herbivore diets across southeastern Africa, analyzing ∼4,000 fecal samples of 30 species from 10 sites in seven countries over 6 y. We detected 893 food plant taxa from 124 families, but just two families-grasses and legumes-accounted for the majority of herbivore diets. Nonetheless, herbivore species almost invariably partitioned food plant taxa; diet composition differed significantly in 97% of pairwise comparisons between sympatric species, and dissimilarity was pronounced even between the strictest grazers (grass eaters), strictest browsers (nongrass eaters), and closest relatives at each site. Niche differentiation was weakest in an ecosystem recovering from catastrophic defaunation, indicating that food plant partitioning is driven by species interactions, and was stronger at low rainfall, as expected if interspecific competition is a predominant driver. Diets differed more between browsers than grazers, which predictably shaped community organization: Grazer-dominated trophic networks had higher nestedness and lower modularity. That dietary differentiation is structured along taxonomic lines complements prior work on how herbivores partition plant parts and patches and suggests that common mechanisms govern herbivore coexistence and community assembly in savannas.

Fire, grazers, and browsers interact with grass competition to determine tree establishment in an African savanna.

In savanna ecosystems, fire and herbivory alter the competitive relationship between trees and grasses. Mechanistically, grazing herbivores favor trees by removing grass, which reduces tree-grass competition and limits fire. Conversely, browsing herbivores consume trees and limit their recovery from fire. Herbivore feeding decisions are in turn shaped by risk-resource trade-offs that potentially determine the spatial patterns of herbivory. Identifying the dominant mechanistic pathways by which fire and herbivores control tree cover remains challenging, but is essential for understanding savanna dynamics. We used an experiment in the Serengeti ecosystem and a simple simulation driven by experimental results to address two main aims: (1) determine the importance of direct and indirect effects of grass, fire, and herbivory on seedling establishment and (2) establish whether predators determine the spatial pattern of successful seedling establishment via effects on mesoherbivore distribution. We transplanted tree seedlings into plots with a factorial combination of grass and herbivores (present/absent) across a lion-kill-risk gradient in the Serengeti, burning half of the plots near the end of the experiment. Ungrazed grass limited tree seedling survival directly via competition, indirectly via fire, and by slowing seedling growth, which drove higher seedling mortality during fires. These effects restricted seedling establishment to below 18% and, in conjunction with browsing, resulted in seedling establishment dropping below 5%. In the absence of browsing and fire, grazing drove a 7.5-fold increase in seedling establishment. Lion predation risk had no observable impact on herbivore effects on seedling establishment. The severe negative effects of grass on seedling mortality suggests that regional patterns of tree cover and fire may overestimate the role of fire in limiting tree cover, with regular fires representing a proxy for the competitive effects of grass.

Anthropogenic modifications to fire regimes in the wider Serengeti-Mara ecosystem.

Fire is a key driver in savannah systems and widely used as a land management tool. Intensifying human land uses are leading to rapid changes in the fire regimes, with consequences for ecosystem functioning and composition. We undertake a novel analysis describing spatial patterns in the fire regime of the Serengeti-Mara ecosystem, document multidecadal temporal changes and investigate the factors underlying these patterns. We used MODIS active fire and burned area products from 2001 to 2014 to identify individual fires; summarizing four characteristics for each detected fire: size, ignition date, time since last fire and radiative power. Using satellite imagery, we estimated the rate of change in the density of livestock bomas as a proxy for livestock density. We used these metrics to model drivers of variation in the four fire characteristics, as well as total number of fires and total area burned. Fires in the Serengeti-Mara show high spatial variability-with number of fires and ignition date mirroring mean annual precipitation. The short-term effect of rainfall decreases fire size and intensity but cumulative rainfall over several years leads to increased standing grass biomass and fuel loads, and, therefore, in larger and hotter fires. Our study reveals dramatic changes over time, with a reduction in total number of fires and total area burned, to the point where some areas now experience virtually no fire. We suggest that increasing livestock numbers are driving this decline, presumably by inhibiting fire spread. These temporal patterns are part of a global decline in total area burned, especially in savannahs, and we caution that ecosystem functioning may have been compromised. Land managers and policy formulators need to factor in rapid fire regime modifications to achieve management objectives and maintain the ecological function of savannah ecosystems.

Alternate Grassy Ecosystem States Are Determined by Palatability-Flammability Trade-Offs.

Fire and mammalian grazers both consume grasses, and feedbacks between grass species, their functional traits, and consumers have profound effects on grassy ecosystem structure worldwide, such that savanna and grassland states determined by fire or grazing can be considered alternate states. These parallel savanna-forest alternate states, which likewise have myriad cascading ecosystem impacts.

Pyrodiversity interacts with rainfall to increase bird and mammal richness in African savannas.

Fire is a fundamental process in savannas and is widely used for management. Pyrodiversity, variation in local fire characteristics, has been proposed as a driver of biodiversity although empirical evidence is equivocal. Using a new measure of pyrodiversity (Hempson et al.), we undertook the first continent-wide assessment of how pyrodiversity affects biodiversity in protected areas across African savannas. The influence of pyrodiversity on bird and mammal species richness varied with rainfall: strongest support for a positive effect occurred in wet savannas (> 650 mm/year), where species richness increased by 27% for mammals and 40% for birds in the most pyrodiverse regions. Range-restricted birds were most increased by pyrodiversity, suggesting the diversity of fire regimes increases the availability of rare niches. Our findings are significant because they explain the conflicting results found in previous studies of savannas. We argue that managing savanna landscapes to increase pyrodiversity is especially important in wet savannas.

Invasion trajectory of alien trees: the role of introduction pathway and planting history.

Global change is driving a massive rearrangement of the world's biota. Trajectories of distributional shifts are shaped by species traits, the recipient environment and driving forces with many of the driving forces directly due to human activities. The relative importance of each in determining the distributions of introduced species is poorly understood. We consider 11 Australian Acacia species introduced to South Africa for different reasons (commercial forestry, dune stabilization and ornamentation) to determine how features of the introduction pathway have shaped their invasion history. Projections from species distribution models (SDMs) were developed to assess how the reason for introduction influences the similarity between climatic envelopes in native and alien ranges. A lattice model for an idealized invasion was developed to assess the relative contribution of intrinsic traits and introduction dynamics on the abundance and extent over the course of simulated invasions. SDMs show that alien populations of ornamental species in South Africa occupy substantially different climate space from their native ranges, whereas species introduced for forestry occupy a similar climate space in native and introduced ranges. This may partly explain the slow spread rates observed for some alien ornamental plants. Such mismatches are likely to become less pronounced with the current drive towards 'eco gardens' resulting in more introductions of ornamental species with a close climate match between native and newly introduced regions. The results from the lattice model showed that the conditions associated with the introduction pathway (especially introduction pressure) dominate early invasion dynamics. The placement of introduction foci in urban areas limited the extent and abundance of invasive populations. Features of introduction events appear to initially mask the influence of intrinsic species traits on invasions and help to explain the relative success of species introduced for different purposes. Introduction dynamics therefore can have long-lasting influences on the outcomes of species redistributions, and must be explicitly considered in management plans.