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Abstract Large mammalian herbivores exert strong top‐down control on plants, which in turn influence most ecological processes. Accordingly, the decline, displacement, or extinction of wild large herbivores in African savannas is expected to alter the physical structure of vegetation, the diversity of plant communities, and downstream ecosystem functions. However, herbivore impacts on vegetation comprise both direct and indirect effects and often depend on herbivore body size and plant type. Understanding how herbivores affect savanna vegetation requires disaggregating the effects of different herbivores and the responses of different plants, as well as accounting for both the structural complexity and composition of plant assemblages. We combined high‐resolution Light Detection and Ranging (LiDAR) with field measurements from size‐selective herbivore exclosures in Kenya to determine how herbivores affect the diversity and physical structure of vegetation, how these impacts vary with body size and plant type, and whether there are predictable associations between plant diversity and structural complexity. Herbivores generally reduced the diversity and abundance of both overstory and understory plants, though the magnitude of these impacts varied substantially as a function of body size and plant type: only megaherbivores (elephants and giraffes) affected tree cover, whereas medium‐ and small‐bodied herbivores had stronger effects on herbaceous diversity and abundance. We also found evidence that herbivores altered the strength and direction of interactions between trees and herbaceous plants, with signatures of facilitation in the presence of herbivores and of competition in their absence. While megaherbivores uniquely affected tree structure, medium‐ and small‐bodied species had stronger (and complementary) effects on metrics of herbaceous vegetation structure. Plant structural responses to herbivore exclusion were species‐specific: of five dominant tree species, just three exhibited significant individual morphological variation across exclosure treatments, and the size class of herbivores responsible for these effects varied across species. Irrespective of exclosure treatment, more species‐rich plant communities were more structurally complex. We conclude that the diversity and architecture of savanna vegetation depend on consumptive and nonconsumptive plant–herbivore interactions; the roles of herbivore diversity, body size, and plant traits in mediating those interactions; and a positive feedback between plant diversity and structural complexity.more » « less
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African savannas are the last stronghold of diverse large-mammal communities, and a major focus of savanna ecology is to understand how these animals affect the relative abundance of trees and grasses. However, savannas support diverse plant life-forms, and human-induced changes in large-herbivore assemblages—declining wildlife populations and their displacement by livestock—may cause unexpected shifts in plant community composition. We investigated how herbivory affects the prevalence of lianas (woody vines) and their impact on trees in an East African savanna. Although scarce (<2% of tree canopy area) and defended by toxic latex, the dominant liana,Cynanchum viminale(Apocynaceae), was eaten by 15 wild large-herbivore species and was consumed in bulk by native browsers during experimental cafeteria trials. In contrast, domesticated ungulates rarely ate lianas. When we experimentally excluded all large herbivores for periods of 8 to 17 y (simulating extirpation), liana abundance increased dramatically, with up to 75% of trees infested. Piecewise exclusion of different-sized herbivores revealed functional complementarity among size classes in suppressing lianas. Liana infestation reduced tree growth and reproduction, but herbivores quickly cleared lianas from trees after the removal of 18-y-old exclosure fences (simulating rewilding). A simple model of liana contagion showed that, without herbivores, the long-term equilibrium could be either endemic (liana–tree coexistence) or an all-liana alternative stable state. We conclude that ongoing declines of wild large-herbivore populations will disrupt the structure and functioning of many African savannas in ways that have received little attention and that may not be mitigated by replacing wildlife with livestock.more » « less
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Michener, William K. (Ed.)Diverse communities of large mammalian herbivores (LMH), once widespread, are now rare. LMH exert strong direct and indirect effects on community structure and ecosystem functions, and measuring these effects is important for testing ecological theory and for understanding past, current, and future environmental change. This in turn requires long-term experimental manipulations, owing to the slow and often nonlinear responses of populations and assemblages to LMH removal. Moreover, the effects of particular species or body-size classes within diverse LMH guilds are difficult to pinpoint, and the magnitude and even direction of these effects often depends on environmental context. Since 2008, we have maintained the Ungulate Herbivory Under Rainfall Uncertainty (UHURU) experiment, a series of size-selective LMH exclosures replicated across a rainfall/productivity gradient in a semi-arid Kenyan savanna. The goals of the UHURU experiment are to measure the effects of removing successively smaller size classes of LMH (mimicking the process of size-biased extirpation) and to establish how these effects are shaped by spatial and temporal variation in rainfall. The UHURU experiment comprises three LMH-exclusion treatments and an unfenced control, applied to 9 randomized blocks of contiguous 1-ha plots (n = 36). The fenced treatments are: “MEGA” (exclusion of megaherbivores, elephant and giraffe); “MESO” (exclusion of herbivores ≥40 kg); and “TOTAL” (exclusion of herbivores ≥5 kg). Each block is replicated three times at three sites across the 20-km rainfall gradient, which has fluctuated over the course of the experiment. The first five years of data were published previously (Ecological Archives E095-064) and have been used in numerous studies. Since that publication, we have (a) continued to collect data following the original protocols, (b) improved the taxonomic resolution and accuracy of plant and small-mammal identifications, and (c) begun collecting several new data sets. Here, we present updated and extended raw data from the first 12 years of the UHURU experiment (2008–2019). Data include daily rainfall data throughout the experiment; annual surveys of understory plant communities; annual censuses of woody-plant communities; annual measurements of individually tagged woody plants; monthly monitoring of flowering and fruiting phenology; every-other-month small-mammal mark-recapture data; and quarterly large-mammal dung surveys. There are no copyright restrictions; notification of when and how data are used is appreciated and users of UHURU data should cite this data paper when using the data.more » « less
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Abstract Differences in the bacterial communities inhabiting mammalian gut microbiomes tend to reflect the phylogenetic relatedness of their hosts, a pattern dubbed phylosymbiosis. Although most research on this pattern has compared the gut microbiomes of host species across biomes, understanding the evolutionary and ecological processes that generate phylosymbiosis requires comparisons across phylogenetic scales and under similar ecological conditions. We analysed the gut microbiomes of 14 sympatric small mammal species in a semi‐arid African savanna, hypothesizing that there would be a strong phylosymbiotic pattern associated with differences in their body sizes and diets. Consistent with phylosymbiosis, microbiome dissimilarity increased with phylogenetic distance among hosts, ranging from congeneric sets of mice and hares that did not differ significantly in microbiome composition to species from different taxonomic orders that had almost no gut bacteria in common. While phylosymbiosis was detected among just the 11 species of rodents, it was substantially weaker at this scale than in comparisons involving all 14 species together. In contrast, microbiome diversity and composition were generally more strongly correlated with body size, dietary breadth, and dietary overlap in comparisons restricted to rodents than in those including all lineages. The starkest divides in microbiome composition thus reflected the broad evolutionary divergence of hosts, regardless of body size or diet, while subtler microbiome differences reflected variation in ecologically important traits of closely related hosts. Strong phylosymbiotic patterns arose deep in the phylogeny, and ecological filters that promote functional differentiation of cooccurring host species may disrupt or obscure this pattern near the tips.more » « less
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