*Article* **Gradients in the Diversity of Plants and Large Herbivores Revealed with DNA Barcoding in a Semi-Arid African Savanna**

**Patrick T. Freeman 1,2, Robert O. Ang'ila 3,4, Duncan Kimuyu 3,4, Paul M. Musili 5, David Kenfack 6, Peter Lokeny Etelej 4, Molly Magid 1,2, Brian A. Gill 1,2 and Tyler R. Kartzinel 1,2,4,\***


**Abstract:** 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-km<sup>2</sup> 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 sugges<sup>t</sup> 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.

**Keywords:** behavioral ecology; DNA barcoding; elephant (*Loxodonta africana*); ForestGEO; geographic mosaic of species interactions; phylogenetic community ecology; landscape ecology; megaherbivores; phylogenetic signal; plant–herbivore interactions
