97 research outputs found
Revisiting Trophic Cascades and Aspen Recovery in Northern Yellowstone
We revisit the nature and extent of trophic cascades and quaking aspen (Populus tremuloides) recovery in the northern range of Yellowstone National Park (YNP), where studies have reported on Rocky Mountain elk (Cervus canadensis) browsing and young aspen heights following the St. John, 1995-96 reintroduction of gray wolves (Canis lupus). A recent study by Brice et al. (2021) expressed concerns about methodologies employed in earlier aspen studies and that results from those studies exaggerated the extent to which a trophic cascade has benefitted aspen, concerns such as: (a) the selection of aspen stands, (b) young aspen sampling and measurements within stands, (c) the upper browse level of elk, (d) cause of increased young aspen height growth, (e) interpretation of browsing and height data, and others. We review these concerns but conclude that earlier aspen studies have provided important insights regarding the recovery of aspen that is underway in northern Yellowstone. We also found that Brice et al. (2021) misinterpreted or misrepresented various aspects of those earlier studies, while failing to address potential biases and shortcomings of their own 2007-2017 study, including: (1) sampling aspen stands from only a portion of the park\u27s northern range, (2) not randomly selecting aspen stands across their study area, but only within identified treatments, (3) varying sampling density (stands/km2) by more than an order of magnitude between treatments, and (4) not sampling all stands in most years. Regardless of the advantages or disadvantages of the sampling designs and research methodologies employed in various aspen studies, they have consistently shown that decreased browsing has resulted in greater young plant heights in YNP\u27s northern range, results supportive of an ongoing trophic cascade
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 \u3e 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\u27s 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
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Dynamic modelling of suspended sediment transport in streams as a function of sediment supply
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Wolves and the Ecology of Fear: Can Predation Risk Structure Ecosystems?
We investigated how large carnivores, herbivores, and plants may be linked to the maintenance of native species biodiversity through trophic
cascades. The extirpation of wolves (Canis lupus) from Yellowstone National Park in the mid-1920s and their reintroduction in 1995 provided the
opportunity to examine the cascading effects of carnivore–herbivore interactions on woody browse species, as well as ecological responses involving
riparian functions, beaver (Castor canadensis) populations, and general food webs. Our results indicate that predation risk may have profound
effects on the structure of ecosystems and is an important constituent of native biodiversity. Our conclusions are based on theory involving trophic
cascades, predation risk, and optimal foraging; on the research literature; and on our own recent studies in Yellowstone National Park. Additional
research is needed to understand how the lethal effects of predation interact with its nonlethal effects to structure ecosystems.Keywords: Wolves, Ungulates, Trophic cascades, Predation risk, Woody browse specie
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Trophic cascades in Yellowstone: the first 15 years after wolf reintroduction
The 1995/96 reintroduction of gray wolves (Canis lupus) into Yellowstone National Park after a 70 year absence has allowed for studies of tri-trophic cascades involving wolves, elk (Cervus elaphus), and plant species such as aspen (Populus tremuloides), cottonwoods (Populus spp.), and willows (Salix spp.). To investigate the status of this cascade, in September of 2010 we repeated an earlier survey of aspen and measured browsing and heights of young aspen in 97 stands along four streams in the Lamar River catchment of the park’s northern winter range. We found that browsing on the five tallest young aspen in each stand decreased from 100% of all measured leaders in 1998 to means of <25 % in the uplands and <20 % in riparian areas by 2010. Correspondingly, aspen recruitment (i.e., growth of seedlings/sprouts above the browse level of ungulates) increased as browsing decreased over time in these same stands. We repeated earlier inventories of cottonwoods and found that recruitment had also increased in recent years. We also synthesized studies on trophic cascades published during the first 15 years after wolf reintroduction. Synthesis results generally indicate that the reintroduction of wolves restored a trophic cascade with woody browse species growing taller and canopy cover increasing in some, but not all places. After wolf reintroduction, elk populations decreased, but both beaver (Caster canadensis) and bison (Bison bison) numbers increased, possibly due to the increase in available woody plants and herbaceous forage resulting from less competition with elk. Trophic cascades research during the first 15 years after wolf reintroduction indicated substantial initial effects on both plants and animals, but northern Yellowstone still appears to be in the early stages of ecosystem recovery. In ecosystems where wolves have been displaced or locally extirpated, their reintroduction may represent a particularly effective approach for passive restoration.The 97 images (supplementary data) that are part of this study were originally submitted to ScholarsArchive@OSU in April 2011, eight months before the article was deposited. The original url for the images is http://hdl.handle.net/1957/2084
Predation Risk, Elk, and Aspen: Comment
With the exception of humans, gray wolves (Canis lupus) are perhaps the most significant predator of cervids in the northern hemisphere, mainly due to the group-hunting, year-round activity, and widespread geographic distribution (Peterson et al. 2003). Thus, interactions between wolves and large herbivore prey, such as elk (Cervus elaphus) and moose (Alces alces), have long been of interest to biologists (Peterson 1995, Jęodrzejewska et al. 2000, Mech and Boitani 2003). The potential ecological role this apex predator may have, via trophic cascades, has also received attention in recent years by researchers (e.g., Callan et al. 2013, Kuijper et al. 2013, 2014), wildlife management agencies (e.g., state wolf management plans), as well as the general public. Perhaps nowhere in the western United States has a heightened examination of this large predator been more focused than in Yellowstone National Park (YNP; Laundré et al. 2001, Smith et al. 2003, 2013, Fortin et al. 2005). Here, wolves were reintroduced in the mid-1990s, again completing the park\u27s large predator guild after approximately seven decades of absence, thus providing a long-term, landscape-scale, natural experiment (Diamond 1983).
The Gallatin winter range is one of two that occur along the northern portion of YNP, the other is the northern ungulate winter range, or “northern range,” located some 25 km or more to the east. Of these, the Gallatin has been less studied. Nevertheless, the Gallatin winter range, like the northern range, experienced high levels of elk herbivory following the extirpation of wolves in the early 1900s. Over a period of approximately seven decades, intensive herbivory by elk led to the long-term decline in aspen (Populus tremuloides) and willow (Salix spp.) recruitment (i.e., growth of young plants above the browse level of elk) in the Gallatin winter range, leaving these plant communities in an impoverished condition (Lovaas 1967, Patten 1968, Kay 2001, Ripple and Beschta 2004, Halofsky and Ripple 2008). Accelerated soil and channel erosion also occurred (Lovaas 1967, Beschta and Ripple 2006). Thus, when wolves were reintroduced into Yellowstone in the mid-1990s, aspen recruitment within the Gallatin elk winter range, had been largely absent for several decades (Kay 2001, Halofsky and Ripple 2008).
In 2010, Winnie (2012) sampled 65 aspen stands in the northwestern corner of YNP, within the Gallatin elk winter range, to determine if a behaviorally mediated trophic cascade (BMTC) was occurring. As background information Winnie (2012:2600) included only a single sentence about wolves in the Greater Yellowstone Ecosystem and the remainder of the paragraph briefly discussed elk numbers, with most of the emphasis on elk in YNP\u27s northern range where there has been a pronounced redistribution of elk since the reintroduction of wolves (White et al. 2012). A more complete summary regarding the status and dynamics of wolves and elk over the last 15 years (i.e., 1995–2010) in the Gallatin elk winter range, as well as in the Daly Creek sub-drainage where Winnie\u27s study occurred, would have helped readers better understand the context of his study. Furthermore, information regarding human harvest of elk in the Gallatin winter range since the return of wolves, or whether such hunting has been affecting elk numbers or distribution in recent years was not provided.
As part of his 2010 field study, Winnie (2012) characterized the presence or absence of several hypothesized risk factors (independent variables) for each aspen stand, including escape impediments, visual impediments, distance to conifer forest edge, and presence of deadfall trees. For dependent variables, Winnie (2012) recorded the presence or absence of browsing on aspen suckers (ramets \u3c2 m in height) and the number of aspen juveniles (plants \u3e2 m in height but \u3c6 cm in diameter at breast height). A height of 2 m generally represents the upper browse level of elk, and young aspen exceeding this height are considered to have successfully recruited. Such recruitment would represent a major departure from the browsing suppression that occurred in his study area over recent decades (Kay 2001, Halofsky and Ripple 2008) and an indication that a tri-trophic cascade involving wolves, elk, and aspen may be underway.
From the results of his analyses, Winnie (2012:2600) concluded that “aspen were not responding to hypothesized fine-scale risk factors in ways consistent with the current BMTC hypothesis.” We respectfully submit that the design and analysis used to support such a conclusion may be deficient for two reasons, the first based on conceptual concerns and the second on statistical concerns. (1) Unfortunately, some aspen stands Winnie (2012) sampled contained juveniles associated with “physical barriers,” barriers that could prevent elk from browsing young aspen. To be scientifically valid, a risk assessment using young aspen as the dependent variable must inherently assure that all evaluated plants were accessible to elk browsing. (2) The inclusion of 10 aspen stands containing some physically protected aspen likely confounded results from his predation risk analyses (i.e., Figs. 5, 6, and 7 in Winnie 2012). While the inclusion of stands with protected aspen may increase the variance associated with his dependent variables (i.e., browsing rate, number of juveniles), the fallacy of doing so is revealed by inspecting these variables for the 85% of his stands (n = 55 stands) that did not have physically protected aspen. Here, a browsing rate of ∼99% and an average of \u3c1 juvenile per stand occurred (back-transformed means from Fig. 8b and a, respectively [Winnie 2012:2609]), indicating a general lack of variance in the dependent variables associated with these stands and little likelihood of a statistically significant outcome. Thus, we suspect that the “statistically significant” results Winnie (2012) found in Figs. 5, 6, and 7, whether contrary to or in support of a BMTC hypothesis, are primarily influenced by the occurrence of risk factors associated with those stands where some of the young aspen were physically protected. A reanalysis by Winnie of browsing rate and number of juveniles vs. his risk factors, using just the 55 stands accessible to elk, could clarify this issue.
Because of the above concerns, we would offer that results of Winnie\u27s (2012) analyses of “proportion of sprouts browsed” or “number of juveniles per stand” relative to his hypothesized risk factors may well be spurious. If so, any discussions and conclusions based on those results are in question.
A 2004 field study of aspen stands in the Gallatin winter range found aspen recruitment had declined precipitously following the extirpation of wolves in the 1920s and remained essentially absent through the late 1990s (Halofsky and Ripple 2008). Thus, when Winnie (2012) undertook his field study in 2010, a wolf–elk–aspen trophic cascade had not yet been confirmed. While the occurrence of juvenile aspen would be important to the long-term survival of aspen stands, the data for elk-accessible stands continue to show exceptionally high browse rates and little or no recruitment (Winnie 2012). This situation contrasts with YNP\u27s northern range where decreased browsing and increased heights of young aspen in portions of that range have been observed some 6–10 years after the occurrence of increased willow growth, although this recruitment has been spatially patchy (e.g., Ripple and Beschta 2012, Painter 2013; also see northern range photos of aspen recruitment available online).5 It should be noted that decreased browsing and increased heights of willows in the Gallatin winter range (at the base of the Daly Creek watershed) following the return of wolves, and consistent with the occurrence of a trophic cascade, were documented as early as 1999–2000 (Ripple and Beschta 2004), with heights continuing to increase in more recent years (Beschta and Ripple 2010). Also consistent with a trophic cascade, various northern range studies have found increased willow growth/canopy cover, sometimes interacting with climatic fluctuations, following wolf reintroduction (e.g., Groshong 2004, Beschta and Ripple 2007, Beyer et al. 2007, Baril 2009, Tercek et al. 2010, Marshall 2012).
The occurrence of 192 juvenile aspen within Winnie\u27s (2012) study area would seem to indicate the beginnings of a tri-trophic cascade, particularly when compared to the lack of juvenile production in the decades immediately before wolf reintroduction (Halofsky and Ripple 2008). However, most of the 192 juveniles were associated with aspen stands characterized as having some degree of physical protection from elk (Fig. 8a in Winnie 2012), making it difficult to confirm if they represent a wolf–elk–aspen trophic cascade involving density and/or behavioral mediation. A trophic cascade involving aspen can be complex and context dependent (e.g., linked to bottom-up factors such as fire [Eisenberg et al. 2013]). Furthermore, undertaking risk assessments associated with large mammalian predators and ungulates in mountainous terrain, where human hunting is also occurring across part of the landscape, can be especially challenging. While we commend Winnie (2012) for attempting such an assessment, without a reanalysis of only those young aspen accessible to elk it would appear that his evaluation may not have been sufficiently rigorous to evaluate the presence or absence of a potential BMTC in the Gallatin winter range
Bison Influences on Composition and Diversity of Riparian Plant Communities in Yellowstone National Park
Riparian zones are among the most biologically diverse ecosystems in the Intermountain West, USA, and provide valuable ecosystem services, including high rates of biotic productivity, nutrient processing, and carbon storage. Thus, their sustainability is a high priority for land managers. Large ungulates affect composition and structure of riparian/stream ecosystems through herbivory and physical effects, via trailing and trampling. Bison (Bison bison) in Yellowstone National Park (YNP) have been characterized as ecosystem engineers because of their demonstrated effects on phenology, aboveground productivity of grasses, and woody vegetation structure. Bison have greatly increased in numbers during the last two decades and spend large periods of time in the broad open floodplains of the Northern Range of the Park, where they are hypothesized to have multiple effects on plant species composition and diversity. We sampled indicators of bison use as well as riparian vegetation composition, diversity, and structure along eight headwater streams within YNP\u27s Northern Range. Total fecal density ranged from 333 to 1833 fecal chips and/or piles/ha, stubble heights ranged from 7 to 49 cm, and streambank disturbance ranged from 9% to 62%. High levels of bison use were positively correlated with exotic species dominance and negatively correlated with species richness, native species diversity, willow (Salix spp.) cover, and wetland species dominance. At three sites, the intensity of bison use exceeded recommended utilization thresholds to avoid degradation of streams and riparian zones on public lands. The influences of large herbivores, principally bison, on vegetation composition and structure suggest the cumulative effects of the current densities on the Northern Range are contributing to biotic impoverishment, representing the loss of ecosystem services that are provided by native riparian plant communities. In addition, contemporary levels of bison use may be exacerbating climate change effects as observed through ungulate-related shifts in composition toward plant assemblages adapted to warmer and drier conditions. However, the resilience of native riparian vegetation suggests that sites currently heavily utilized by bison would have the potential for recovery with a reduction in pressure by this herbivore
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Sediment and organic matter transport in Oregon Coast range streams
Bison influences on composition and diversity of riparian plant communities in Yellowstone National Park
Riparian zones are among the most biologically diverse ecosystems in the Intermountain West, USA, and provide valuable ecosystem services, including high rates of biotic productivity, nutrient processing, and carbon storage. Thus, their sustainability is a high priority for land managers. Large ungulates affect composition and structure of riparian/stream ecosystems through herbivory and physical effects, via trailing and trampling. Bison (Bison bison) in Yellowstone National Park (YNP) have been characterized as “ecosystem engineers” because of their demonstrated effects on phenology, aboveground productivity of grasses, and woody vegetation structure. Bison have greatly increased in numbers during the last two decades and spend large periods of time in the broad open floodplains of the Northern Range of the Park, where they are hypothesized to have multiple effects on plant species composition and diversity. We sampled indicators of bison use as well as riparian vegetation composition, diversity, and structure along eight headwater streams within YNP’s Northern Range. Total fecal density ranged from 333 to 1833 fecal chips and/or piles/ha, stubble heights ranged from 7 to 49 cm, and streambank disturbance ranged from 9% to 62%. High levels of bison use were positively correlated with exotic species dominance and negatively correlated with species richness, native species diversity, willow (Salix spp.) cover, and wetland species dominance. At three sites, the intensity of bison use exceeded recommended utilization thresholds to avoid degradation of streams and riparian zones on public lands. The influences of large herbivores, principally bison, on vegetation composition and structure suggest the cumulative effects of the current densities on the Northern Range are contributing to biotic impoverishment, representing the loss of ecosystem services that are provided by native riparian plant communities. In addition, contemporary levels of bison use may be exacerbating climate change effects as observed through ungulate-related shifts in composition toward plant assemblages adapted to warmer and drier conditions. However, the resilience of native riparian vegetation suggests that sites currently heavily utilized by bison would have the potential for recovery with a reduction in pressure by this herbivore
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