Migrating bison engineer the green wave.

Newly emerging plants provide the best forage for herbivores. To exploit this fleeting resource, migrating herbivores align their movements to surf the wave of spring green-up. With new technology to track migrating animals, the Green Wave Hypothesis has steadily gained empirical support across a diversity of migratory taxa. This hypothesis assumes the green wave is controlled by variation in climate, weather, and topography, and its progression dictates the timing, pace, and extent of migrations. However, aggregate grazers that are also capable of engineering grassland ecosystems make some of the world's most impressive migrations, and it is unclear how the green wave determines their movements. Here we show that Yellowstone's bison (Bison bison) do not choreograph their migratory movements to the wave of spring green-up. Instead, bison modify the green wave as they migrate and graze. While most bison surfed during early spring, they eventually slowed and let the green wave pass them by. However, small-scale experiments indicated that feedback from grazing sustained forage quality. Most importantly, a 6-fold decadal shift in bison density revealed that intense grazing caused grasslands to green up faster, more intensely, and for a longer duration. Our finding broadens our understanding of the ways in which animal movements underpin the foraging benefit of migration. The widely accepted Green Wave Hypothesis needs to be revised to include large aggregate grazers that not only move to find forage, but also engineer plant phenology through grazing, thereby shaping their own migratory movements.

The effects of herbivore aggregations at water sources on savanna plants differ across soil and climate gradients.

Water sources in arid and semiarid ecosystems support humans, wildlife, and domestic animals, forming nodes of activity that sculpt surrounding plant communities and impact critical grazing and soil systems. However, global aridification and changing surface water supply threaten to disrupt these water resources, with strong implications for conservation and management of these ecosystems. To understand how effects of herbivore aggregation at water impact plant communities across contexts, we measured herbivore activity, plant height, cover (trees, grasses, forbs, and bare ground), diversity, and composition at 17 paired water sources and matrix sites across a range of abiotic factors in a semiarid savanna in Kenya. The effects of proximity to surface water and herbivore aggregation on plant communities varied substantially depending on soil and rainfall. In arid areas with nutrient-poor sandy soils, forb and tree cover were 50% lower at water sources compared to neighboring matrix sites, bare ground was 20% higher, species richness was 15% lower, and a single globally important grazing grass (Cynodon dactylon) dominated 60% of transects. However, in mesic areas with nutrient-rich finely textured soils, species richness was 25% higher, despite a 40% increase in bare ground, concurrent with the decline of a dominant tall grass (Themeda triandra) and increase in C. dactylon and other grass species near water sources. Recent rainfall was important for grasses; cover was higher relative to matrix sites only during wet periods, a potential indication of compensatory grazing. These findings suggest that effects of herbivore aggregation on vegetation diversity and composition will vary in magnitude, and in some cases direction, depending on other factors at the site. Where moisture and nutrient resources are high and promote the dominance of few plant species, herbivore aggregations may maintain diversity by promoting grazing lawns and increasing nondominant species cover. However, in arid conditions and sites with low nutrient availability, diversity can be substantially reduced by these aggregations. Our results highlight the importance of considering abiotic conditions when managing for effects of herbivore aggregations near water. This will be particularly important for future managers in light of growing global aridification and surface water changes.

Diverse effects of the common hippopotamus on plant communities and soil chemistry.

The ecological importance of the common hippopotamus (Hippopotamus amphibius) in aquatic ecosystems is becoming increasingly well known. These unique megaherbivores are also likely to have a formative influence on the terrestrial ecosystems in which they forage. In this study, we employed a novel exclosure design to exclude H. amphibius from experimental plots on near-river grasslands. Our three-year implementation of this experiment revealed a substantial influence of H. amphibius removal on both plant communities and soil chemistry. H. amphibius significantly reduced grassland canopy height, increased the leafiness of common grasses, reduced woody plant abundance and size, and increased the concentrations of several soil elements. Many of the soil chemistry changes that we experimentally induced by exclusion of H. amphibius were mirrored in the soil chemistry differences between naturally occurring habitats of frequent (grazing lawns) and infrequent (shrub forest) use by H. amphibius and other grazing herbivores. In contrast to existing hypotheses regarding grazing species, we found that H. amphibius had little effect on local plant species richness. Simultaneous observations of exclosures designed to remove all large herbivores revealed that H. amphibius removal had ecologically significant impacts, but that the removal of all species of large herbivores generated more pronounced impacts than the removal of H. amphibius alone. In aggregate, our results suggest that H. amphibius have myriad effects on their terrestrial habitats that likely improve the quality of forage available for other herbivores. We suggest that ongoing losses of this vulnerable megaherbivore are likely to cause significant ecological change.

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.