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We experimentally examined the influence of grass competition, grass species identity (taxa) and water availability on the seedling growth and survival of two dominant tree species (Vachellia(formerlyAcacia)robustaandV. tortilis) of the Serengeti National Park in Tanzania. Despite being widely distributed, the species have an opposing overstorey dominance across a rainfall and productivity gradient, withV. robustadominating the more productive mesic sites andV. tortilisin the lower productivity, drier sites. We investigated the role of different grass species, which vary in distribution and abundance across the rainfall gradient, in influencing the growth and survival ofVachelliaseedlings. We found a significant effect of grass competition but no effect of grass species identity on the growth or survival of seedlings. Seedling survival was highest in the absence of grass competition, intermediate when grasses were defoliated to simulate grazing and lowest when grown with ungrazed grasses. Grass competition had a more negative effect on the stem diameter ofV. tortilisthanV. robusta.AllV. tortilisseedlings grown under a combination of drought conditions and unclipped grasses died by the end of the experiment. However, reduced grass competition by simulated grazing improvedV. tortilisseedling survival to comparable levels achieved byV. robustaspecies. Our study advances our understanding of tree and grass competition across environmental gradients and suggests that the presence of grass and soil moisture have species‐specific effects on tree seedling growth and survival in African savannas.

 

Understanding the role of species interactions within communities is a central focus of ecology. A key challenge is to understand variation in species interactions along environmental gradients. The stress gradient hypothesis posits that positive interactions increase and competitive interactions decrease with increasing consumer pressure or environmental stress. This hypothesis has received extensive attention in plant community ecology, but only a handful of tests in animals. Furthermore, few empirical studies have examined multiple co‐occurring stressors. Here we test predictions of the stress gradient hypothesis using the occurrence of mixed‐species groups in six common grazing ungulate species within the Serengeti‐Mara ecosystem. We use mixed‐species groups as a proxy for potential positive interactions because they may enhance protection from predators or increase access to high‐quality forage. Alternatively, competition for resources may limit the formation of mixed‐species groups. Using more than 115,000 camera trap observations collected over 5 yr, we found that mixed‐species groups were more likely to occur in risky areas (i.e., areas closer to lion vantage points and in woodland habitat where lions hunt preferentially) and during time periods when resource levels were high. These results are consistent with the interpretation that stress from high predation risk may contribute to the formation of mixed‐species groups, but that competition for resources may prevent their formation when food availability is low. Our results are consistent with support for the stress gradient hypothesis in animals along a consumer pressure gradient while identifying the potential influence of a co‐occurring stressor, thus providing a link between research in plant community ecology on the stress gradient hypothesis, and research in animal ecology on trade‐offs between foraging and risk in landscapes of fear. 

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. 

Although there is a well‐known association between tree cover and soil texture in savannahs, the hydrological drivers of tree cover variation have not been systematically explored, particularly in parallel with factors such as fire, herbivory, and tree–grass interactions. The relationship between hydrological factors and tree cover is important for resolving the relative contribution of bottom‐up versus top‐down factors in structuring savannah vegetation.

We quantified soil moisture dynamics across eight 1‐km transects spanning tree cover gradients from open to woody savannah in Serengeti National Park in Tanzania using soil moisture sensors coupled with dataloggers. We mapped tree cover at two spatial scales through supervised classification of high‐resolution satellite imagery. We simultaneously produced water retention curves in open and woody habitats within each transect to compare soil hydrological properties and to convert volumetric water content (θ) from dataloggers to plant‐available water over the course of an annual cycle. We also quantified grass biomass at 100 locations per transect, estimated fire frequency from MODIS satellite data, and quantified herbivore occupancy with paired camera traps situated in open and woody habitats within each transect.

We found a positive relationship between tree cover and soil moisture drainage rate, and found that open habitats had more negative water potentials than woody habitats for a given value ofθ. In contrast, we found no evidence for a consistent relationship between grass biomass or fire frequency and tree cover. We found evidence for higher browser occupancy in woody than open habitats, but no habitat effects on herbivores as a group (browsers plus grazers), suggesting that herbivory is unlikely to be the dominant factor explaining variation in tree cover.

Synthesis. Our results suggest that variation in tree cover is partly driven by hydrological (edaphic) factors unrelated to fire, herbivory, tree–grass interactions or mean annual precipitation at these spatial scales in Serengeti. We contrast our findings with previous work attributing tree cover shifts in Serengeti to precipitation gradients.