Aspen recovery in northern Yellowstone: A comment on Brice et al. (2021).

Aspen sapling recruitment increased as browsing by elk decreased, following the 1995-96 reintroduction of wolves in Yellowstone National Park. We address claims by Brice et al. (2021) that previous studies exaggerated recent aspen recovery. We conclude that their results actually supported previous work showing a trophic cascade benefiting aspen.

Bison alter the northern Yellowstone ecosystem by breaking aspen saplings.

The American bison (Bison bison) is a species that strongly interacts with its environment, yet the effects of this large herbivore on quaking aspen (Populus tremuloides) have received little study. We documented bison breaking the stems of aspen saplings (young aspen >2 m tall and ≤5 cm in diameter at breast height) and examined the extent of this effect in northern Yellowstone National Park (YNP). Low densities of Rocky Mountain elk (Cervus canadensis) after about 2004 created conditions conducive for new aspen recruitment in YNP's northern ungulate winter range (northern range). We sampled aspen saplings at local and landscape scales, using random sampling plots in 87 randomly selected aspen stands. Across the YNP northern range, we found that 18% of sapling stems had been broken. The causal attribution to bison was supported by multiple lines of evidence: (1) most broken saplings were in areas of high bison and low elk density; (2) saplings were broken in summer when elk were not foraging on them; (3) we directly observed bison breaking aspen saplings; and (4) mixed-effects modeling showed a positive association between scat density of bison and the proportion of saplings broken. In a stand heavily used by bison, most aspen saplings had been broken, and portions of the stand were cleared of saplings that were present in previous sampling in 2012. Bison numbers increased more than fourfold between 2004 and 2015, and their ecosystem effects have similarly increased, limiting and in some places reversing the nascent aspen recovery. This situation is further complicated by political constraints that prevent bison from dispersing to areas outside the park. Thus, one important conservation goal, the preservation of bison, is affecting another long-term conservation goal, the recovery of aspen and other deciduous woody species in northern Yellowstone.

Forum: Climate, Ecological, and Social Costs of Livestock Grazing on Western Public Lands.

Grazing by domestic livestock is the most widespread use of public lands in the American West (USA) and their effects on climate change and ways to mitigate those effects are of interest to land managers, policy makers, and the broader public. Kauffman et al. (2022a) provided a meta-analysis of the ecosystem impacts, greenhouse gas (GHG) emissions, and social costs of carbon (SCC) associated with livestock grazing on public lands in the western USA. They determined that GHG emissions from cattle on public lands equaled 12.4 million t CO2e/year. At the scale of land use planning utilized by federal agencies, GHG emissions associated with allocated livestock numbers will typically exceed US Environmental Protection Agencies' reporting limits (25,000 t) for certain industrial greenhouse gas emitters. As such, these are essentially unreported sources of GHG emissions from public lands. Using the US government's most recent SCC estimate of $51/t, Kauffman et al. (2022a) determined the total SCC of cattle grazing on public lands to be approximately $264-630 million/year. However, recent advances in the determination of SCC reveal this is to be an underestimate. Using the latest science results in an estimated SCC of $1.1-2.4 billion/year for grazing on public lands. Furthermore, the SCC borne by the public exceeds the economic benefits to private livestock permittees by over $926 million/year. Cessation of public lands grazing is an environmentally and economically sound mitigation and adaptation approach to addressing the climate crisis; an approach that will also facilitate restoration of the myriad of ecosystem services provided by intact wildland ecosystems.

Livestock Use on Public Lands in the Western USA Exacerbates Climate Change: Implications for Climate Change Mitigation and Adaptation.

Public lands of the USA can play an important role in addressing the climate crisis. About 85% of public lands in the western USA are grazed by domestic livestock, and they influence climate change in three profound ways: (1) they are significant sources of greenhouse gases through enteric fermentation and manure deposition; (2) they defoliate native plants, trample vegetation and soils, and accelerate the spread of exotic species resulting in a shift in landscape function from carbon sinks to sources of greenhouse gases; and (3) they exacerbate the effects of climate change on ecosystems by creating warmer and drier conditions. On public lands one cow-calf pair grazing for one month (an "animal unit month" or "AUM") produces 875 kg CO2e through enteric fermentation and manure deposition with a social carbon cost of nearly $36 per AUM. Over 14 million AUMs of cattle graze public lands of the western USA each year resulting in greenhouse gas emissions of 12.4 Tg CO2e year-1. The social costs of carbon are > $500 million year-1 or approximately 26 times greater than annual grazing fees collected by managing federal agencies. These emissions and social costs do not include the likely greater ecosystems costs from grazing impacts and associated livestock management activities that reduce biodiversity, carbon stocks and rates of carbon sequestration. Cessation of grazing would decrease greenhouse gas emissions, improve soil and water resources, and would enhance/sustain native species biodiversity thus representing an important and cost-effective adaptive approach to climate change.

Riparian vegetation composition and diversity shows resilience following cessation of livestock grazing in northeastern Oregon, USA.

Riparian ecosystem restoration has been accomplished through exclusion of livestock using corridor fencing along hundreds of kilometers of streams in the western United States, for the benefit of riparian-obligate wildlife and endangered fishes. Yet, there are limited scientific studies that have evaluated more than short-term shifts in vegetation composition and diversity at a single location or handful of locations following grazing. We sampled riparian vegetation composition along 11-paired grazed and ungrazed (exclosed) stream reaches in northeastern Oregon. Exclosure ages ranged from 2 to >30 years and grazing treatments varied from light grazing every one out of three years to heavy season-long grazing. Species richness and diversity was higher in the ungrazed reaches (p = 0.002). The abundance of native sedges (Carex spp.) and broad-leaved forbs were also significantly (p ≤ 0.05) greater in ungrazed areas. In contrast, exotic species adapted to grazing such as Poa pratensis and Trifolium repens were more abundant in grazed stream reaches. The prevalence of hydrophytic species significantly increased (p ≤ 0.01) in ungrazed reaches, (based on wetland species indicator scores), indicating that wetland-dominated communities within the ungrazed stream reaches were replacing ones adapted to drier environments. The increased abundance of facultative and wetland-obligate species in ungrazed reaches compared to grazed reaches suggests that livestock grazing exacerbates those climate change effects also leading to warmer and drier conditions. Further, riparian-obligate shrub cover along the streambank was higher in 7 of 8 exclosures that were older than 5 years (p = 0.05). As a restoration approach, the inherent resilience of riparian ecosystems exhibited in ungrazed riparian zones suggest positive feedbacks to other beneficial ecosystem processes such as increased species and habitat diversity, increased carbon sequestration, enhanced allochthonous inputs and greater sediment retention, that would affect the aquatic and terrestrial biota, water quality, and stream morphology.

Recovering aspen follow changing elk dynamics in Yellowstone: evidence of a trophic cascade?

To investigate the extent and causes of recent quaking aspen (Populus tremuloides) recruitment in northern Yellowstone National Park, we measured browsing intensity and height of young aspen in 87 randomly selected aspen stands in 2012, and compared our results to similar data collected in 1997-1998. We also examined the relationship between aspen recovery and the distribution of Rocky Mountain elk (Cervus elaphus) and bison (Bison bison) on the Yellowstone northern ungulate winter range, using ungulate fecal pile densities and annual elk count data. In 1998, 90% of young aspen were browsed and none were taller-than 200 cm, the height at which aspen begin to escape from elk browsing. In 2012, only 37% in the east and 63% in the west portions of the winter range were browsed, and 65% of stands in the east had young aspen taller than 200 cm. Heights of young aspen were inversely related to browsing intensity, with the least browsing and greatest heights in the eastern portion of the range, corresponding with recent changes in elk density and distribution. In contrast with historical elk distribution (1930s-1990s), the greatest densities of elk recently (2006-2012) have been north of the park boundary (approximately 5 elk/km2), and in the western part of the range (2-4 elk/km2), with relatively few elk in the eastern portion of the range (<2 elk/km2), even in mild winters. This redistribution of elk and decrease in density inside the park, and overall reduction in elk numbers, explain why many aspen stands have begun to recover. Increased predation pressure following the reintroduction of gray wolves (Canis lupius) in 1995-1996 played a role in these changing elk population dynamics, interacting with other influences including increased predation by bears (Ursus spp.), competition with an expanding bison population, and shifting patterns of human land use and hunting outside the park. The resulting new aspen recruitment is evidence of a landscape-scale trophic cascade in which a resurgent large carnivore community, combined with other ecological changes, has benefited aspen through effects on ungulate prey.

Wolves trigger a trophic cascade to berries as alternative food for grizzly bears.

This is a Forum article in response to: Barber-Meyer, S. (2015) Trophic cascades from wolves to grizzly bears or changing abundance of bears and alternate foods? Journal of Animal Ecology, 83, doi: 10.1111/1365-2656.12338. We used multiple data sets and study areas as well as several lines of evidence to investigate potential trophic linkages in Yellowstone National Park. Our results suggest that a trophic cascade from wolves to elk to berry production to berry consumption by grizzly bears may now be underway in the Park.

Reducing livestock effects on public lands in the western United States as the climate changes: a reply to Svejcar et al.

Svejcar et al. (Environ Manage, 2014) offered several perspectives regarding Beschta et al. (Environ Manage 51:474-491, 2013)--a publication that addressed the interacting ecological effects of climate change and domestic, wild, and feral ungulates on public lands in the western United States (US)--by largely focusing on three livestock grazing issues: (1) legacy versus current day impacts; (2) grazing as a fire reduction tool; and (3) the complexity of grazing. Regarding these issues, we indicate that (1) legacy effects to western ecosystems were indeed significant and contemporary livestock use on public lands generally maintains or exacerbates many of those effects; (2) livestock grazing has been a major factor affecting fire frequency, fire severity, and ecosystem trajectories in the western US for over a century; and (3) the removal or reduction of grazing impacts in these altered ecosystems is the most effective means of initiating ecological recovery. Svejcar et al. (Environ Manage, 2014) offer no evidence that livestock use is consistent with the timely recovery of grazing-degraded uplands, riparian areas, or stream systems. We thus conclude that public-land ecosystems can best persist or cope with a changing climate by significantly reducing ungulate grazing and related impacts.

Status and ecological effects of the world's largest carnivores.

Large carnivores face serious threats and are experiencing massive declines in their populations and geographic ranges around the world. We highlight how these threats have affected the conservation status and ecological functioning of the 31 largest mammalian carnivores on Earth. Consistent with theory, empirical studies increasingly show that large carnivores have substantial effects on the structure and function of diverse ecosystems. Significant cascading trophic interactions, mediated by their prey or sympatric mesopredators, arise when some of these carnivores are extirpated from or repatriated to ecosystems. Unexpected effects of trophic cascades on various taxa and processes include changes to bird, mammal, invertebrate, and herpetofauna abundance or richness; subsidies to scavengers; altered disease dynamics; carbon sequestration; modified stream morphology; and crop damage. Promoting tolerance and coexistence with large carnivores is a crucial societal challenge that will ultimately determine the fate of Earth's largest carnivores and all that depends upon them, including humans.

Trophic cascades from wolves to grizzly bears in Yellowstone.

We explored multiple linkages among grey wolves (Canis lupus), elk (Cervus elaphus), berry-producing shrubs and grizzly bears (Ursus arctos) in Yellowstone National Park. We hypothesized competition between elk and grizzly bears whereby, in the absence of wolves, increases in elk numbers would increase browsing on berry-producing shrubs and decrease fruit availability to grizzly bears. After wolves were reintroduced and with a reduced elk population, we hypothesized there would be an increase in the establishment of berry-producing shrubs, such as serviceberry (Amelanchier alnifolia), which is a major berry-producing plant. We also hypothesized that the percentage fruit in the grizzly bear diet would be greater after than before wolf reintroduction. We compared the frequency of fruit in grizzly bear scats to elk densities prior to wolf reintroduction during a time of increasing elk densities (1968-1987). For a period after wolf reintroduction, we calculated the percentage fruit in grizzly bear scat by month based on scats collected in 2007-2009 (n = 778 scats) and compared these results to scat data collected before wolf reintroduction. Additionally, we developed an age structure for serviceberry showing the origination year of stems in a northern range study area. We found that over a 19-year period, the percentage frequency of fruit in the grizzly diet (6231 scats) was inversely correlated (P < 0·001) with elk population size. The average percentage fruit in grizzly bear scats was higher after wolf reintroduction in July (0·3% vs. 5·9%) and August (7·8% vs. 14·6%) than before. All measured serviceberry stems accessible to ungulates originated since wolf reintroduction, while protected serviceberry growing in a nearby ungulate exclosure originated both before and after wolf reintroduction. Moreover, in recent years, browsing of serviceberry outside of the exclosure decreased while their heights increased. Overall, these results are consistent with a trophic cascade involving increased predation by wolves and other large carnivores on elk, a reduced and redistributed elk population, decreased herbivory and increased production of plant-based foods that may aid threatened grizzly bears.

Adapting to climate change on Western public lands: addressing the ecological effects of domestic, wild, and feral ungulates.

Climate change affects public land ecosystems and services throughout the American West and these effects are projected to intensify. Even if greenhouse gas emissions are reduced, adaptation strategies for public lands are needed to reduce anthropogenic stressors of terrestrial and aquatic ecosystems and to help native species and ecosystems survive in an altered environment. Historical and contemporary livestock production-the most widespread and long-running commercial use of public lands-can alter vegetation, soils, hydrology, and wildlife species composition and abundances in ways that exacerbate the effects of climate change on these resources. Excess abundance of native ungulates (e.g., deer or elk) and feral horses and burros add to these impacts. Although many of these consequences have been studied for decades, the ongoing and impending effects of ungulates in a changing climate require new management strategies for limiting their threats to the long-term supply of ecosystem services on public lands. Removing or reducing livestock across large areas of public land would alleviate a widely recognized and long-term stressor and make these lands less susceptible to the effects of climate change. Where livestock use continues, or where significant densities of wild or feral ungulates occur, management should carefully document the ecological, social, and economic consequences (both costs and benefits) to better ensure management that minimizes ungulate impacts to plant and animal communities, soils, and water resources. Reestablishing apex predators in large, contiguous areas of public land may help mitigate any adverse ecological effects of wild ungulates.