After long-term decline, are aspen recovering in northern Yellowstone?
Abstract
In northern Yellowstone National Park, quaking aspen (Populus tremuloides) stands were dying out in the late 20th century following decades of intensive browsing by Rocky Mountain elk (Cervus elaphus). In 1995–1996 gray wolves (Canis lupus) were reintroduced, joining bears (Ursus spp.) and cougars (Puma concolor) to complete the guild of large carnivores that prey on elk. This was followed by a marked decline in elk density and change in elk distribution during the years 1997–2012, due in part to increased predation. We hypothesized that these changes would result in less browsing and an increase in height of young aspen. In 2012, we sampled 87 randomly selected stands in northern Yellowstone, and compared our data to baseline measurements from 1997 and 1998. Browsing rates (the percentage of leaders browsed annually) in 1997–1998 were consistently high, averaging 88%, and only 1% of young aspen in sample plots were taller than 100cm; none were taller than 200cm. In 2012, browsing rates were much lower at 44%, and young aspen were taller on average with 34% >100cm and 5% >200cm. Most (62%) of the variation in height of young aspen in 2012 was explained by browsing intensity. Furthermore, in 2012, 25% of stands had at least five aspen saplings tall enough to escape elk browsing (⩾200cm spring height), a condition that has not occurred for decades and happened despite a recent drought. Sapling recruitment did not increase until browsing decreased, following substantial changes…
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Browsing pressure was reduced and more spatially variable, and subsequent aspen studies reported new recruitment of aspen saplings.
“Browsing pressure was reduced and more spatially variable, and subsequent aspen studies reported new recruitment of aspen saplings (Beschta et al., 2023; Painter et al., 2014, 2015; Ripple & Beschta, 2007, 2012).”
From Bison alter the northern Yellowstone ecosystem by breaking aspen saplings (2023) · cites this paper as Painter et al., 2014
In northern Yellowstone, herbivory has been the primary factor limiting the growth of aspen from sprouts to saplings, and eventually trees.
“Recent trends toward a warmer and drier climate may cause additional stress (Kulakowski et al., 2013; Worrall et al., 2013), but in northern Yellowstone herbivory has been the primary factor limiting the growth of aspen from sprouts to saplings, and eventually trees (Beschta et al., 2016; Kay, 2018; Painter et al., 2014, 2018).”
From Bison alter the northern Yellowstone ecosystem by breaking aspen saplings (2023) · cites this paper as Painter et al., 2014
Willow recovery, like aspen recovery, began in the early 2000s during a time of drought and has continued under a long-term warming and drying trend.
“willow recovery, like aspen recovery, began in the early 2000s during a time of drought (McMenamin et al. 2008, Painter et al. 2014) and has continued under a long-term warming and drying trend.”
From Tall willow thickets return to northern Yellowstone (2020) · cites this paper as Painter et al. 2014
Cites
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BackgroundGray Wolves as Climate Change Buffers in Yellowstone
Wilmers, C. C., & Getz, W. M. (2005) · PLoS Biology · doi:10.1371/journal.pbio.0030092 · Cited by 211 · cited 1× · see citing works
“4) (Wilmers and Getz, 2005; McMenamin et al., 2008; Frank et al., 2013), and in northern YNP every winter from 2000 to 2010 had snow accumulations below average.”
BackgroundHYDROLOGIC REGIME AND HERBIVORY STABILIZE AN ALTERNATIVE STATE IN YELLOWSTONE NATIONAL PARK
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WILLOW ON YELLOWSTONE'S NORTHERN RANGE: EVIDENCE FOR A TROPHIC CASCADE?
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“In some places willow heights are now influenced more by abiotic factors such as water and soil conditions than by browsing (Bilyeu et al., 2008; Tercek et al., 2010; Marshall et al., 2013), evidence of a significant reduction in herbivory (Singer et al.”
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“In some places willow heights are now influenced more by abiotic factors such as water and soil conditions than by browsing (Bilyeu et al., 2008; Tercek et al., 2010; Marshall et al., 2013), evidence of a significant reduction in herbivory (Singer et al., 2003; Hebblewhite and Smith, 2010).”
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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
“Although a broadening release of aspen in both riparian and upland aspen stands currently appears to be underway (Painter et al., 2014, 2015), there is considerable variability as to where and when aspen recovery has begun occur.”
BackgroundMethodResultInfluentialSelf-citationResolving the value of the dingo in ecological restoration
Newsome, T. M., Ballard, G., Crowther, M. S., Dellinger, J. A., Fleming, P. J. S., Glen, A. S., Greenville, A. C., Johnson, C. N., Letnic, M., Moseby, K. E., et al. (2015) · Restoration Ecology · doi:10.1111/rec.12186 · Cited by 81 · see citing works
“…there is debate about the exact mechanisms by which wolves affect their main prey, elk (Cervus elaphus) (Kauffman et al. 2007; Middleton et al. 2013), and how widespread the cascading effects on vegetation have been (Kauffman et al. 2010, but see Painter et al. 2014 for the most recent analyses).”
BackgroundSelf-citationRecovering aspen follow changing elk dynamics in Yellowstone: evidence of a trophic cascade?
Painter, L. E., Beschta, R. L., Larsen, E. J., & Ripple, W. J. (2015) · Ecology · doi:10.1890/14-0712.1 · Cited by 76 · see citing works
“…in aspen recruitment represents a fundamental shift in aspen dynamics, because recruitment of new aspen trees declined and virtually ceased by the mid-20th century due to intensive herbivory by elk (Ripple and Larsen 2000, Kauffman et al. 2010, Beschta and Ripple 2013, Painter et al. 2014).”
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“Browsing effects depend on ungulate behavior and population size, which varies over time; a substantial decline in Yellowstone elk numbers between 1997 and 2014 (Smith et al., 2015) has been linked to increased aspen-ramet heights in mature stands (Painter et al., 2014).”
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Sampling bias exaggerates a textbook example of a trophic cascade
(2021) · doi:10.1111/ele.13915 · Cited by 56 · see citing works
Self-citationBison limit ecosystem recovery in northern Yellowstone
Beschta, R. L., Ripple, W. J., Kauffman, J. B., & Painter, L. E. (2020) · Food Webs · doi:10.1016/j.fooweb.2020.e00142 · Cited by 41 · see citing works
Self-citationCan large carnivores change streams via a trophic cascade?
Beschta, R. L., & Ripple, W. J. (2019) · Ecohydrology · doi:10.1002/eco.2048 · Cited by 46 · see citing works
Self-citationRoles for the Canidae in food webs reviewed: Where do they fit?
Fleming, P. J. S., Nolan, H., Jackson, S., Ballard, G., Bengsen, A. J., Brown, W. Y., Meek, P. D., Mifsud, G., Pal, S., & Sparkes, J. (2017) · Food Webs · doi:10.1016/j.fooweb.2017.03.001 · Cited by 52 · see citing works
“Where this occurs, indirect effects on stream morphology (Beschta & Ripple 2012), woody plants (Painter et al., 2014) and berry production (Ripple et al. 2014b) have been observed as well as for a suite of animals including medium-sized ungulates (e.g. White-tailed Deer, Odocoileus virginianus, and…”
BackgroundFlagel, D. G., Belovsky, G. E., & Beyer, D. E. (2015) · Oecologia · doi:10.1007/s00442-015-3515-z · Cited by 48 · see citing works
Newsome, T. M., Fleming, P. J. S., Dickman, C. R., Doherty, T. S., Ripple, W. J., Ritchie, E. G., & Wirsing, A. J. (2017) · BioScience · doi:10.1093/biosci/bix022 · Cited by 39 · see citing works
Self-citationTrophic cascades at multiple spatial scales shape recovery of young aspen in Yellowstone
Beschta, R. L., Painter, L. E., & Ripple, W. J. (2018) · Forest Ecology and Management · doi:10.1016/j.foreco.2018.01.055 · Cited by 40 · see citing works
“…young aspen (< 200 cm in height) and overstory trees (typically> 20 cm in diameter at breast height) was characteristic of aspen stands across the northern range due to the historical suppression of young plants by elk (Larsen and Ripple, 2005; Ripple and Beschta, 2007; Painter et al., 2014).”
BackgroundInfluentialSelf-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-citationTrophic cascades from wolves to alders in Yellowstone
Ripple, W. J., Beschta, R. L., & Painter, L. E. (2015) · Forest Ecology and Management · doi:10.1016/j.foreco.2015.06.007 · Cited by 31 · see citing works
“However, the annual SWE was below average most years after 1997, consistent with the fact that the region was in a drought (Painter et al., 2014).”
BackgroundResultInfluentialSelf-citationLong-term aspen dynamics, trophic cascades, and climate in northern Yellowstone National Park
Beschta, R. L., Painter, L. E., Levi, T., & Ripple, W. J. (2016) · Canadian Journal of Forest Research · doi:10.1139/cjfr-2015-0301 · Cited by 26 · see citing works
“An inverse relationship between browsing intensity and height of young woody plants is central to the trophic cascade hypotheses, as it clearly identifies the mechanism via which top-down pressure ensues (Beyer et al. 2007; Painter et al. 2014).”
BackgroundMethodResultInfluentialSelf-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-citationThe primacy of density‐mediated indirect effects in a community of wolves, elk, and aspen
Brice, E. M., Larsen, E. J., Stahler, D. R., & MacNulty, D. R. (2024) · Ecological Monographs · doi:10.1002/ecm.1627 · Cited by 17 · see citing works
Self-citationPotter, C. S. (2020) · Landscape Ecology · doi:10.1007/s10980-019-00951-3 · Cited by 18 · see citing works
Tall willow thickets return to northern Yellowstone
Painter, L. E., & Tercek, M. T. (2020) · Ecosphere · doi:10.1002/ecs2.3115 · Cited by 11 · see citing works
“In fact, willow recovery, like aspen recovery, began in the early 2000s during a time of drought (McMenamin et al. 2008, Painter et al. 2014) and has continued under a long-term warming and drying trend.”
BackgroundSelf-citationThe Condition and Trend of Aspen, Willows, and Associated Species on the Northern Yellowstone Range
Kay, C. E. (2018) · Rangelands · doi:10.1016/j.rala.2018.10.005 · Cited by 10 · see citing works
Large carnivore extirpation linked to loss of overstory aspen in Yellowstone
Beschta, R. L., & Ripple, W. J. (2019) · Food Webs · doi:10.1016/j.fooweb.2019.e00140 · Cited by 7 · see citing works
“This ongoing recovery started slowly but has nevertheless increased in strength over time (Painter et al., 2014; Beschta et al., 2018).”
BackgroundSelf-citationForest growth trends in Canada
Loehle, C., & Solarik, K. A. (2019) · The Forestry Chronicle · doi:10.5558/tfc2019-027 · Cited by 14 · see citing works
“Painter et al. (2014) and Rooney (2001) came to this same conclusion.”
BackgroundRevisiting trophic cascades and aspen recovery in northern Yellowstone
Beschta, R. L., Painter, L. E., & Ripple, W. J. (2023) · Food Webs · doi:10.1016/j.fooweb.2023.e00276 · Cited by 6 · see citing works
“…from across a stand will always be greater than the average height of young aspen measured in a small 20 m 2 plot, such as that used by Brice et al. Painter et al. (2014, 2015) subsequently addressed this issue by measuring, in conjunction with 5T sampling, young aspen heights in random plots for…”
BackgroundMethodInfluentialSelf-citationPotter, C. S. (2019) · Remote Sensing in Earth Systems Sciences · doi:10.1007/s41976-019-00019-5 · Cited by 7 · see citing works
Changing aspen stand structure following large carnivore restoration in Yellowstone
Painter, L. E., Beschta, R. L., & Ripple, W. J. (2025) · Forest Ecology and Management · doi:10.1016/j.foreco.2025.122941 · Cited by 4 · see citing works
“…trend of aspen recovery, we randomly sampled aspen stands across northern YNP and compared results with two previous surveys of aspen in these same stands: (1) in 1997 – 98 (Larsen and Ripple, 2003) shortly after the 1995 – 96 introduction of wolves, and (2) in 2012 (Painter et al., 2014, 2015).”
InfluentialSelf-citationAspen recovery in northern Yellowstone: A comment on Brice et al. (2021)
Painter, L. E., Beschta, R. L., & Ripple, W. J. (2024) · Ecology Letters · doi:10.1111/ele.14353 · Cited by 2 · see citing works
Self-citationMacNulty, D. R., Brice, E. M., Bergeron, N., & Larsen, E. J. (2025) · EcoEvoRxiv · doi:10.1016/j.foreco.2025.123448
Self-citationAn ecologically significant trophic cascade in Yellowstone: Response to MacNulty, et al. (2026)
Painter, L. E., Beschta, R. L., & Ripple, W. J. (2025) · Forest Ecology and Management · doi:10.1016/j.foreco.2025.123449 · Cited by 1 · see citing works
Self-citationBeschta, R. L., & Ripple, W. J. (2017) · Earth Surface Processes and Landforms · doi:10.1002/esp.4289
“…to establish and grow above the upper browse level of elk in the northern winter range of Yellowstone National Park (Beschta and Ripple, 2010, 2016; Painter et al. 2014), as well as in Banff National Park (Hebblewhite and Smith, 2010; White et al., 2015), Jasper National Park (Beschta and Ripple,…”
BackgroundSelf-citationTrophic cascades and Yellowstone’s aspen: A reply to Fleming (2019)
Beschta, R. L., Painter, L. E., & Ripple, W. J. (2019) · Forest Ecology and Management · doi:10.1016/j.foreco.2019.05.014
Self-citationBrice, E. M., & MacNulty, D. R. (2018) · The UW National Parks Service Research Station Annual Reports · doi:10.13001/uwnpsrc.2018.5647
- Authors
- Painter, L. E., Beschta, R. L., Larsen, E. J., & Ripple, W. J.
- Affiliations
- Oregon State University (US)
- University of Wisconsin–Stevens Point (US)
- Corresponding author
- Painter, Luke E.
- Venue
- Forest Ecology and Management
- Publisher
- Elsevier BV
- Published
- 2014-10
- Volume
- 329
- Pages
- 108-117
- ISSN
- 0378-1127 (ISSN-L)
- OpenAlex
- W2085438120
- Cited by
- 31
- References
- 0
- Citations by year
- 2025: 3 · 2024: 2 · 2023: 3 · 2021: 1 · 2020: 3 · 2019: 4 · 2018: 5 · 2017: 3 · 2016: 2 · 2015: 4 · 2014: 1
- FWCI
- 2.89
- Type
- journal article
- Language
- en
- Open access
- closed
- Locations
- Forest Ecology and Management (publishedVersion)
- Digital Commons - USU (Utah State University) (submittedVersion)
- Funders
- University of Wyoming – National Park Service Research Station
- Topics
- Physical Sciences › Environmental Science › Ecology › Wildlife Ecology and Conservation
- Physical Sciences › Environmental Science › Ecology › Rangeland and Wildlife Management
- Physical Sciences › Environmental Science › Ecology › Ecology and biodiversity studies
- Keywords
- Populus tremuloides, elk browsing, Rocky Mountain elk, gray wolf reintroduction, sapling recruitment, browsing intensity
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