Does restoring apex predators to food webs restore ecosystems? Large carnivores in Y ellowstone as a model system
Open accessAbstract
Modification of food webs is a frequent cause of shifts in ecosystem states that resist reversal when the food web is restored to its original condition. We used the restoration of the large carnivore guild including gray wolves ( Canis lupis ), cougars ( Felis concolor ), and grizzly bears ( Ursus arctos horribilis ) to the northern range of Yellowstone National Park as a model system to understand how ecosystems might resist reconfiguration after the restoration of apex predators to the food web. The absence of wolves, cougars, and grizzly bears for nearly a century from the northern range was the primary cause of dramatic changes in riparian plant communities. Willows ( Salix spp.) were suppressed in height by intense browsing by the dominant herbivore, elk ( Cervus canadensis ). The loss of activity by beavers ( Castor canadensis ) coincided with the loss of tall willows. We hypothesized that intense elk browsing interrupted the mutualism between willow and beavers: ecosystem engineering by beavers was a critical component of willow habitat and tall willows were a critical component of habitat for beavers. This interruption made riparian communities resilient to the disturbance caused by the restoration of apex predators. We hypothesized further that reductions in elk browsing attributable to reductions in elk population size were not sufficient to prevent the suppression of willow growth. To test these hypotheses, we conducted a 20‐year, factorial experiment that crossed…
Integrity signals
Registry
Cites
Statistics
Claims the paper makes
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
Annual browsing intensity in the Hobbs et al. plots was rarely below 0.25 on average, and frequently exceeded 0.50 for individual plants.
“In contrast, annual browsing intensity in the Hobbs et al. plots, defined as the proportion of shoots browsed, was rarely below 0.25 on average, and frequently exceeded 0.50 for individual plants (see Fig.12and 13 inHobbs et al. 2024).”
From Flawed analysis invalidates claim of a strong Yellowstone trophic cascade after wolf reintroduction: A comment on Ripple et al. (2025) (2025) · cites this paper as Hobbs et al. 2024
Hobbs et al. concluded that carnivore recovery produced a relatively weak trophic cascade.
“In contrast, Hobbs et al. concluded that carnivore recovery produced a relatively weak trophic cascade.”
From Flawed analysis invalidates claim of a strong Yellowstone trophic cascade after wolf reintroduction: A comment on Ripple et al. (2025) (2025) · cites this paper as Hobbs et al. 2024
In a landscape where willow recovery is highly variable, omitting context allows isolated successes to stand in for more complex and uneven outcomes.
“In a landscape where willow recovery is highly variable (e.g., Hobbs et al. 2024), omitting such context allows isolated successes to stand in for more complex and uneven outcomes.”
From Flawed analysis invalidates claim of a strong Yellowstone trophic cascade after wolf reintroduction: A comment on Ripple et al. (2025) (2025) · cites this paper as Hobbs et al. 2024
This structural bias, amplified by increasing height but persistent asymmetry, likely contributed to Ripple et al.'s inference of a strong trophic cascade, despite underlying trends in willow height and browsing that align more closely with Hobbs et al.'s conclusion of a relatively weak one.
“This structural bias, amplified by increasing height but persistent asymmetry, likely contributed to Ripple et al.'s inference of a strong trophic cascade, despite underlying trends in willow height and browsing that align more closely with Hobbs et al.'s conclusion of a relatively weak one.”
From Flawed analysis invalidates claim of a strong Yellowstone trophic cascade after wolf reintroduction: A comment on Ripple et al. (2025) (2025) · cites this paper as Hobbs et al. 2024
These shortcomings explain the apparent conflict with Hobbs et al. (2024), who found evidence for a relatively weak trophic cascade based on the same height data and a long-term factorial field experiment.
“These shortcomings explain the apparent conflict with Hobbs et al. (2024), who found evidence for a relatively weak trophic cascade based on the same height data and a long-term factorial field experiment.”
From Flawed analysis invalidates claim of a strong Yellowstone trophic cascade after wolf reintroduction: A comment on Ripple et al. (2025) (2025) · cites this paper as Hobbs et al. 2024
Thirty-eight percent of the 53 willows used to construct the model were Salix lutea, a species absent from the Hobbs et al. dataset, and 11% were sampled outside northern Yellowstone.
“Thirty-eight percent of the 53 willows used to construct the model were Salix lutea, a species absent from the Hobbs et al. dataset, and 11% were sampled outside northern Yellowstone.”
From Flawed analysis invalidates claim of a strong Yellowstone trophic cascade after wolf reintroduction: A comment on Ripple et al. (2025) (2025) · cites this paper as Hobbs et al. 2024
Hobbs et al.'s interpretation was based on the same height data as well as a factorial field experiment that manipulated browsing pressure and stream hydrology.
“Their interpretation was based on the same height data as well as a factorial field experiment that manipulated browsing pressure and stream hydrology.”
From Flawed analysis invalidates claim of a strong Yellowstone trophic cascade after wolf reintroduction: A comment on Ripple et al. (2025) (2025) · cites this paper as Hobbs et al. 2024
Although browsing intensity declined through 2010, it increased again in later years, even as some willows grew taller.
“Although browsing intensity declined through 2010, it increased again in later years, even as some willows grew taller.”
From Flawed analysis invalidates claim of a strong Yellowstone trophic cascade after wolf reintroduction: A comment on Ripple et al. (2025) (2025) · cites this paper as Hobbs et al. 2024
In 2001, willows in the Hobbs plots were short, heavily browsed, and structurally deformed, conditions inconsistent with the half-ellipsoidal crown architecture assumed by the model.
“In 2001, willows in the Hobbs plots were short, heavily browsed, and structurally deformed-conditions clearly inconsistent with the half-ellipsoidal crown architecture assumed by the model.”
From Flawed analysis invalidates claim of a strong Yellowstone trophic cascade after wolf reintroduction: A comment on Ripple et al. (2025) (2025) · cites this paper as Hobbs et al. 2024
The strong cascade inferred by Ripple et al. does not reflect different empirical patterns, but rather a circular reinterpretation of the same height data that underpinned Hobbs et al.'s finding of a relatively weak trophic cascade.
“This helps explain the conflicting conclusions: the strong cascade inferred by Ripple et al. does not reflect different empirical patterns, but rather a circular reinterpretation of the same height data that underpinned Hobbs et al.'s finding of a relatively weak trophic cascade.”
From Flawed analysis invalidates claim of a strong Yellowstone trophic cascade after wolf reintroduction: A comment on Ripple et al. (2025) (2025) · cites this paper as Hobbs et al. 2024
Cites
Status and Ecological Effects of the World’s Largest Carnivores
Ripple, W. J., Estes, J. A., Beschta, R. L., Wilmers, C. C., Ritchie, E. G., Hebblewhite, M., Berger, J., Elmhagen, B., Letnic, M., Nelson, M. P., et al. (2014) · Science · doi:10.1126/science.1241484 · Cited by 3,667 · see citing works
Trophic cascades in Yellowstone: The first 15years after wolf reintroduction
Ripple, W. J., & Beschta, R. L. (2012) · Biological Conservation · doi:10.1016/j.biocon.2011.11.005 · Cited by 884 · see citing works
Restoring Yellowstone’s aspen with wolves
Ripple, W. J., & Beschta, R. L. (2007) · Biological Conservation · doi:10.1016/j.biocon.2007.05.006 · Cited by 319 · see citing works
Kauffman, M. J., Brodie, J. F., & Jules, E. S. (2010) · Ecology · doi:10.1890/09-1949.1 · Cited by 275 · see citing works
Influence of harvest, climate and wolf predation on Yellowstone elk, 1961-2004
Vucetich, J. A., Smith, D. W., & Stahler, D. R. (2005) · Oikos · doi:10.1111/j.0030-1299.2005.14180.x · Cited by 222 · see citing works
HYDROLOGIC 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
Self-citationRiparian 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
Trophic Cascades in a Multicausal World: Isle Royale and Yellowstone
Peterson, R. O., Vucetich, J. A., Bump, J. M., & Smith, D. W. (2014) · Annual Review of Ecology, Evolution, and Systematics · doi:10.1146/annurev-ecolsys-120213-091634 · Cited by 150 · see citing works
Resolving Ecosystem Complexity (MPB-47)
Schmitz, O. J. (2010) · doi:10.1515/9781400834174 · Cited by 145 · see citing works
WILLOW ON YELLOWSTONE'S NORTHERN RANGE: EVIDENCE FOR A TROPHIC CASCADE?
Beyer, H. L., Merrill, E. H., Varley, N., & Boyce, M. S. (2007) · Ecological Applications · doi:10.1890/06-1254.1 · Cited by 142 · see citing works
Ungulate Herbivory of Willows on Yellowstone's Northern Winter Range
Singer, F. J., Mark, L. C., & Cates, R. C. (1994) · Journal of Range Management · doi:10.2307/4002993 · Cited by 118 · see citing works
Stream hydrology limits recovery of riparian ecosystems after wolf reintroduction
Marshall, K. N., Hobbs, N. T., & Cooper, D. J. (2013) · Proceedings of the Royal Society B: Biological Sciences · doi:10.1098/rspb.2012.2977 · Cited by 108 · see citing works
Self-citationINTERACTION OF BEAVER AND ELK HERBIVORY REDUCES STANDING CROP OF WILLOW
Baker, B. W., Ducharme, H. C., Mitchell, D. C. S., Stanley, T. R., & Peinetti, H. R. (2005) · Ecological Applications · doi:10.1890/03-5237 · Cited by 89 · see citing works
Wolf presence and increased willow consumption by Yellowstone elk: implications for trophic cascades
Creel, S., & Christianson, D. (2009) · Ecology · doi:10.1890/08-2017.1 · Cited by 84 · see citing works
Holocene beaver damming, fluvial geomorphology, and climate in Yellowstone National Park, Wyoming
Persico, L., & Meyer, G. (2009) · Quaternary Research · doi:10.1016/j.yqres.2008.09.007 · Cited by 77 · see citing works
WATER TABLES CONSTRAIN HEIGHT RECOVERY OF WILLOW ON YELLOWSTONE'S NORTHERN RANGE
Bilyeu, D. M., Cooper, D. J., & Hobbs, N. T. (2008) · Ecological Applications · doi:10.1890/07-0212.1 · Cited by 75 · see citing works
Self-citationWinnie, J. A. (2012) · Ecology · doi:10.1890/11-1990.1 · Cited by 73 · see citing works
Recovering Riparian Plant Communities with Wolves in Northern Yellowstone, U.S.A.
Beschta, R. L., & Ripple, W. J. (2008) · Restoration Ecology · doi:10.1111/j.1526-100x.2008.00450.x · Cited by 60 · see citing works
Elk browsing increases aboveground growth of water-stressed willows by modifying plant architecture
Johnston, D. B., Cooper, D. J., & Hobbs, N. T. (2007) · Oecologia · doi:10.1007/s00442-007-0854-4 · Cited by 34 · see citing works
Self-citationBeschta, R. L., & Ripple, W. J. (2007) · Western North American Naturalist · doi:10.3398/1527-0904(2007)67[613:iwhany]2.0.co;2 · Cited by 33 · see citing works
After long-term decline, are aspen recovering in northern Yellowstone?
Painter, L. E., Beschta, R. L., Larsen, E. J., & Ripple, W. J. (2014) · Forest Ecology and Management · doi:10.1016/j.foreco.2014.05.055 · Cited by 31 · see citing works
The History and Current Status and Distribution of Beavers in Yellowstone National Park
Smith, D. W., & Tyers, D. B. (2012) · Northwest Science · doi:10.3955/046.086.0404 · Cited by 28 · see citing works
Rose, J. R., & Cooper, D. J. (2017) · Ecohydrology · doi:10.1002/eco.1768 · Cited by 10 · see citing works
Self-citation
Flagged first, then by citations
Cited by
Sorted by citations
The strength of the Yellowstone trophic cascade after wolf reintroduction
Ripple, W. J., Beschta, R. L., Wolf, C., Painter, L. E., & Wirsing, A. J. (2025) · Global Ecology and Conservation · doi:10.1016/j.gecco.2025.e03428 · Cited by 25 · see citing works
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
“…’ s willow recovery is ongoing, nonlinear, and shaped by complex biotic interactions such as persistent browsing from a reorganizing ungulate community, as well as abiotic factors including altered hydrological regimes due to lost beaver activity and stream incision (Hobbs et al., 2024).”
InfluentialSelf-citationA 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
MacNulty, D., Cooper, D., Procko, M., & Clark-Wolf, T. (2025) · doi:10.32942/x2qq0c
Self-citation
- Authors
- Hobbs, N. T., Johnston, D. B., Marshall, K. N., Wolf, E. C., & Cooper, D. J.
- Affiliations
- Colorado State University (US)
- Corresponding author
- Hobbs, N. Thompson
- Venue
- Ecological Monographs
- Publisher
- Wiley
- Published
- 2024-01-30
- Volume
- 94
- Issue
- 2
- ISSN
- 0012-9615 (ISSN-L)
- 1557-7015
- OpenAlex
- W4391399227
- Cited by
- 52
- References
- 23
- Citations by year
- 2026: 15 · 2025: 26 · 2024: 11
- FWCI
- 14.27
- Type
- journal article
- Language
- en
- Update notices
- erratum (2024-08-29)
- Open access
- hybrid
- doi.org
- Licence
- Locations
- Ecological Monographs (publishedVersion, cc-by-nc-nd)
- Funders
- National Science Foundation (0717367, 1147369, 1655035)
- Topics
- Physical Sciences › Environmental Science › Ecology › Ecology and biodiversity studies
- Physical Sciences › Environmental Science › Ecology › Wildlife Ecology and Conservation
- Physical Sciences › Environmental Science › Ecology › Animal Ecology and Behavior Studies
- Keywords
- large carnivores, gray wolves, Puma concolor, grizzly bear, elk browsing, beaver dams, ecosystem engineering, riparian plant communities, alternative stable states, ecosystem resilience, Yellowstone National Park, herbivore guilds
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
What counts as a claim ›How we measure confidence ›How we check this ›
Are you an author, or do you see a problem with this page? Request a review