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.
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Gradients in the Diversity of Plants and Large Herbivores Revealed with DNA Barcoding in a Semi-Arid African Savanna
Do hotspots of plant biodiversity translate into hotspots in the abundance and diversity of large mammalian herbivores? A common expectation in community ecology is that the diversity of plants and animals should be positively correlated in space, as with the latitudinal diversity gradient and the geographic mosaic of biodiversity. Whether this pattern ‘scales down’ to landscape-level linkages between the diversity of plants or the activities of highly mobile megafauna has received less attention. We investigated spatial associations between plants and large herbivores by integrating data from a plant-DNA-barcode phylogeny, camera traps, and a comprehensive map of woody plants across the 1.2-km2 Mpala Forest Global Earth Observatory (ForestGEO) plot, Kenya. Plant and large herbivore communities were strongly associated with an underlying soil gradient, but the richness of large herbivore species was negatively correlated with the richness of woody plants. Results suggest thickets and steep terrain create associational refuges for plants by deterring megaherbivores from browsing on otherwise palatable species. Recent work using dietary DNA metabarcoding has demonstrated that large herbivores often directly control populations of the plant species they prefer to eat, and our results reinforce the important role of megaherbivores in shaping vegetation across landscapes.
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- Award ID(s):
- 1930820
- PAR ID:
- 10388597
- Date Published:
- Journal Name:
- Diversity
- Volume:
- 14
- Issue:
- 3
- ISSN:
- 1424-2818
- Page Range / eLocation ID:
- 219
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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Introduction Plants and their insect herbivores represent a large fraction of the species in Amazonian forests and are often directly implicated in the origin and maintenance of biodiversity at local and regional scales. How these interactions may change over geographic distance is unknown because very few studies have investigated the herbivore fauna and defense chemicals of any host plant species at multiple sites in tropical forests. One hypothesis, the Geographic Mosaic Theory of Coevolution, predicts that if herbivore assemblages turn over in different parts of a plant’s range, then plant defense chemicals should also change, reflecting local selection pressures. Methods We tested this theory by studying 12 species of Protium (Burseraceae) trees that occur in both Iquitos, Peru, and Manaus, Brazil, in rainforests separated by 1500 km. We surveyed all insects observed directly feeding on the plants in both locations for 48 weeks in Manaus and 64 weeks in Iquitos. We analyzed the secondary metabolites in the leaves of all species in both locations using GC/MS and HPLC. Results and Discussion Although in both locations we found that Protium herbivores were dominated by insects from the orders Hemiptera, Coleoptera and Lepidoptera, we found almost complete turnover in the herbivore species composition in the two sites, and each host plant species had a different assemblage of herbivores in each location. Comparing the phylogenetic beta-diversity, we found low similarity in herbivore phylogenetic relatedness between host plant species in the two locations. However, the secondary metabolites found within a Protium species were similar across the two locations. We found no strong evidence that individuals from a host plant species in Iquitos or Manaus expressed locally-adapted defense chemicals, as individuals from geographic locations did not form clusters when looking at patterns of chemical similarity. These results are not consistent with the Geographic Mosaic Theory of Coevolution. The most intriguing pattern we found was a strong correlation between the diversity of herbivores per host plant species in both locations. We also found that plants with high chemical richness had lower numbers of herbivore species and numbers of total herbivores in both locations. We conclude that high chemical diversity is the most effective strategy for Protium trees to reduce insect herbivore attacks. We speculate that each secondary metabolite is effective at repelling only a few insect herbivores, and that different chemicals are likely effective in different parts of a plants’ geographic range. Future studies should investigate additional locations and additional natural enemies (i.e., fungal pathogens) to test the hypothesis that chemical diversity reduces attack from natural enemies and may explain the ecological and evolutionary success of rainforest trees over time and space.more » « less
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