Where the buffalo roam: Ungulate influences on quaking aspen and willow communities in the Greater Yellowstone Ecosystem
Open accessAbstract
Quaking aspen (Populus tremuloides) and willows (Salix spp.) are keystone species of montane and shrub‐steppe landscapes of the Western United States. Intact communities dominated by these species provide a wide range of ecosystem services, harboring an exceptional proportion of landscape biodiversity. Land use, especially overgrazing by large ungulates, is among the greatest threats to these ecosystems. To examine the effects of wild ungulates and levels of grazing at Yellowstone National Park (YNP) and the adjacent Gallatin National Forest, we sampled plant community composition and vegetation structure of aspen and willow communities both inside and outside of exclosures. Within the Park, we found that current grazing pressures from large ungulates, principally American Bison (Bison bison), have a dramatic effect on community composition and structure, resulting in a shift from a structurally diverse forest or tall shrub dominance to that of a grassland. On heavily grazed sites, shrubs common to semiarid uplands are now relatively abundant, as are exotic grass species, in contrast to an abundance of berry‐producing shrubs within exclosures. Finally, large herbivores at unnaturally high densities are resulting in the simplification of landscape diversity outside of exclosures through a decline in the patch diversity of the site (i.e., homogenization of the landscape). Increases in exotic species and those adapted to semiarid environments suggest that current levels of bison use at YNP are amplifying the effects of climate change as well as resulting in a loss of biodiversity values. Inside exclosures, the inherent resilience of these keystone ecosystems was demonstrated by the recovery of plant diversity following decades of large ungulate utilization exceeding natural carrying capacity. This suggests that reductions in current grazing pressures within YNP would have positive feedbacks to beneficial ecosystem processes such as increased species and habitat diversity, increased carbon sequestration, and a greater adaptive capacity to climate change.
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Heavy ungulate use shifts the composition and structure of forest and shrubland plant communities to simplified grass-dominated systems, often with an increasing prevalence of exotic plant species.
“Shifting the composition and structure of forest and shrubland plant communities to simplified grass-dominated systems, often with an increasing prevalence of exotic plant species (Kauffman et al. 2023, 2025).”
From A Century of Overabundant Ungulates in Yellowstone’s Lamar Valley: First Elk and Now Bison (2026) · cites this paper as Kauffman et al. 2025
There is a lack of recent aspen and willow recruitment outside of long-term ungulate exclosures in areas of high bison use.
“there is a lack of recent aspen and willow recruitment outside of long-term ungulate exclosures, such as the Lamar West (Fig. 11) and Lamar East exclosures (Kauffman et al. 2025).”
From A Century of Overabundant Ungulates in Yellowstone’s Lamar Valley: First Elk and Now Bison (2026) · cites this paper as Kauffman et al. 2025
Aspen stands on adjacent hillslopes will continue to be replaced by exotic-dominated grassland species.
“Similarly, aspen stands on adjacent hillslopes will continue to be replaced by exotic-dominated grassland species (Kauffman et al. 2025).”
From A Century of Overabundant Ungulates in Yellowstone’s Lamar Valley: First Elk and Now Bison (2026) · cites this paper as Kauffman et al. 2025
Cites
Historic aspen recruitment, elk, and wolves in northern Yellowstone National Park, USA
Ripple, W. J., & Larsen, E. J. (2000) · Biological Conservation · doi:10.1016/s0006-3207(00)00014-8 · Cited by 308 · see citing works
Self-citationHYDROLOGIC REGIME AND HERBIVORY STABILIZE AN ALTERNATIVE STATE IN YELLOWSTONE NATIONAL PARK
Wolf, E. C., Cooper, D. J., & Hobbs, N. T. (2007) · Ecological Applications · doi:10.1890/06-2042.1 · Cited by 167 · see citing works
Riparian vegetation recovery in Yellowstone: The first two decades after wolf reintroduction
Beschta, R. L., & Ripple, W. J. (2016) · Biological Conservation · doi:10.1016/j.biocon.2016.03.031 · Cited by 160 · see citing works
Self-citationWolves, elk, willows, and trophic cascades in the upper Gallatin Range of Southwestern Montana, USA
Ripple, W. J., & Beschta, R. L. (2004) · Forest Ecology and Management · doi:10.1016/j.foreco.2004.06.017 · Cited by 146 · see citing works
Self-citationPainter, L. E., Beschta, R. L., Larsen, E. J., & Ripple, W. J. (2018) · Ecosphere · doi:10.1002/ecs2.2376 · Cited by 29 · see citing works
Self-citationBison alter the northern Yellowstone ecosystem by breaking aspen saplings
Painter, L. E., Beschta, R. L., & Ripple, W. J. (2023) · Ecology and Evolution · doi:10.1002/ece3.10369 · Cited by 17 · see citing works
Self-citationKauffman, J. B., Cummings, D. L., Kauffman, C., Beschta, R. L., Brooks, J., MacNeill, K., & Ripple, W. J. (2023) · Ecosphere · doi:10.1002/ecs2.4406 · Cited by 15 · see citing works
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MacNulty, D. R., Cooper, D., Procko, M., & Clark-Wolf, T. J. (2025) · Global Ecology and Conservation · doi:10.1016/j.gecco.2025.e03899 · Cited by 1 · see citing works
“…since been applied to at least one other study of northern Yellowstone willow, yielding claims of extraordinarily large differences between browsed and unbrowsed plants (e.g., mean crown volume reported as 56m³ inside vs. 0.001m³ outside exclosures — a 55,000-fold contrast; (Kauffman et al., 2025).”
A Century of Overabundant Ungulates in Yellowstone’s Lamar Valley: First Elk and Now Bison
Beschta, R. L., Ripple, W. J., & Kauffman, J. B. (2026) · Environmental Management · doi:10.1007/s00267-026-02505-1
“By 2020, overstory trees are completely gone while height growth and crown expansion of aspen and willows, in the absence of ungulate herbivory, has continued inside the exclosure (Kauffman et al. 2025).”
InfluentialSelf-citation
- Authors
- Kauffman, J. B., Cummings, D. L., Beschta, R. L., & Ripple, W. J.
- Affiliations
- Oregon State University (US)
- Illahee Sciences International Corvallis Oregon USA
- Corresponding author
- Kauffman, J. Boone
- Venue
- Ecosphere
- Publisher
- Wiley
- Published
- 2025-09
- Volume
- 16
- Issue
- 9
- ISSN
- 2150-8925 (ISSN-L)
- OpenAlex
- W4414519506
- Semantic Scholar
- 772a18ff1ed5a1e347551830e5bb4b6a6e0fafc5
- Cited by
- 5
- References
- 7
- Citations by year
- 2026: 2 · 2025: 3
- FWCI
- 2.66
- Type
- journal article
- Language
- en
- Open access
- gold
- doi.org
- Licence
- Locations
- Ecosphere (publishedVersion, cc-by)
- Digital Commons - USU (Utah State University) (submittedVersion, cc-by)
- DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion, cc-by-sa)
- Topics
- Physical Sciences › Environmental Science › Ecology › Rangeland and Wildlife Management
- Physical Sciences › Environmental Science › Nature and Landscape Conservation › Ecology and Vegetation Dynamics Studies
- Physical Sciences › Environmental Science › Ecology › Wildlife Ecology and Conservation
- Keywords
- Populus tremuloides, willow, ungulate grazing, American bison, overgrazing, exclosures, floristic composition, vegetation structure, shrub-steppe, landscape homogenization, exotic grasses, keystone species, carrying capacity, climate change, biodiversity loss, carbon sequestration, Greater Yellowstone Ecosystem, Yellowstone National Park, Gallatin National Forest
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