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Cited by 774Wildlife Ecology and ConservationRangeland and Wildlife ManagementAnimal Ecology and Behavior Studies

ELK ALTER HABITAT SELECTION AS AN ANTIPREDATOR RESPONSE TO WOLVES

Open access
Creel, Scott, Winnie, John, Maxwell, Bruce, Hamlin, Ken, Creel, Michael·2005·Ecology
DOI 10.1890/05-0032
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Abstract

For elk (Cervus elaphus) in the Gallatin drainage of the Greater Yellowstone Ecosystem, Montana, USA, wolf movements caused local predation risk to vary substantially on a time scale of days. Spatially and temporally fine-scaled data from GPS radio collars show that elk moved into the protective cover of wooded areas when wolves were present, reducing their use of preferred grassland foraging habitats that had high predation risk. By constraining habitat selection, wolves may have greater effects on elk dynamics than would be predicted on the basis of direct predation alone. Based on changes in the woody vegetation following the reintroduction of wolves, it has been suggested that antipredator responses by elk may be driving a trophic cascade in the Yellowstone ecosystem. However, studies to date have been hampered by a lack of direct data on spatial variation in predation risk, and the ways in which elk respond to variation in risk. Our data support a central portion of the hypothesis that elk antipredator behavior could drive a trophic cascade, but changes in elk numbers are also likely to have affected elk–plant interactions.

Integrity signals

Registry

Crossref · 10.1890/05-0032

Cites

54 of 54 references verified0 citation errors · 0 not checkable

Statistics

Nothing to recompute.

Claims the paper makes

2 of 51 cited sources checked · 0 supported · 0 flagged · accuracy not yet measured

  • Premisecitation

    Recent proposals that wolves trigger elk behavioral responses with cascading plant effects are weakened by missing direct data on how predation risk and elk responses vary in space and time.

    “Recent studies have proposed that wolves (Canis lupus, Linneaus) provoke behavioral responses by elk that cause cascading effects on plants, but this conclusion is weakened by a lack of direct data on spatial and temporal variation in predation risk and the responses of elk to this variation (Ripple and Larsen 2000, Laundre et al. 2001, Childress and Lung 2003, Ripple and Bechsta 2003).”
    Quote in the paperSubstantial claim1 of 3 sources checked
    • cites Ripple and Larsen 2000 source
    • cites Laundre et al. 2001 source

      The Discussion (page unavailable) reports that female elk increased vigilance and reduced foraging in wolf areas, with responses changing as wolves spread to other areas over several years. The Methods (page unavailable) describe comparisons between wolf and non-wolf areas but limit seasonal sampling to late May–early June. The paper does not establish cascading effects on plants or state that such a conclusion is weakened by a lack of direct spatial and temporal risk data. (LLM, conf=0.84)

      Experimental — accuracy not yet measured
    • cites Childress and Lung 2003 source
  • Findingown data

    For elk in the Gallatin drainage, wolf movements made local predation risk vary greatly over days.

    “wolf movements caused local predation risk to vary substantially on a time scale of days.”
    Quote in the paperSubstantial claim
  • Findingown data

    Predation risk across the site varied in space and time because of wolf movements, colonization, and pack failure.

    “Predation risk varied spatially and temporally across the site, due to wolf movements, colonization, and pack failure.”
    Quote in the paperSubstantial claim
    • Rests on an assumption: Wolf presence in a drainage indicates elevated predation risk for elk in that drainage on that day.
  • Findingown data

    Early-winter calf recruitment was ≤0.2 calves per cow in six of seven winters after wolf colonization, compared with one of 13 winters before.

    “Similarly, calf recruitment in early winter has been Յ0.2 calves per cow in six of seven winters since colonization by wolves, compared to one of 13 winters prior to wolf colonization (Fisher's exact test, P ϭ 0.003).”
    Quote in the paperSubstantial claim
  • Findingnone given

    During each winter, every drainage repeatedly changed between days with wolves present and days with none.

    “Throughout a winter, each drainage shifted repeatedly from having wolves present to harboring no wolves on that day.”
    Quote in the paperSubstantial claim
  • May combine two claims. 24
  • Classified as probably not a substantial claim. 12
  • The quote only partly states this claim. 3

  • May combine two claims.

    Premisecitation

    Those studies defined high- or low-risk areas either by habitat features or by large ellipses only loosely tied to wolf home range polygons.

    “In these studies, areas of high or low risk were defined either by habitat features (Ripple andBechsta 2003, 2004) or by large ellipses with a loose relationship to wolf home range polygons (Laundre et al. 2001).”
    Quote in the paperSubstantial claim1 of 3 sources checked
    • cites Ripple and Bechsta 2003

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

    • cites Ripple and Bechsta 2004

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

    • cites Laundre et al. 2001 source

      The Methods section distinguishes areas with wolves from wolf-free areas and says the latter were selected using Park Service data on wolf-pack occurrence. It does not describe the areas as large ellipses or assess how closely they corresponded to wolf home-range polygons; those details attributed to Laundré et al. are unsupported. (LLM, conf=0.81)

      Experimental — accuracy not yet measured

  • Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

    Findingown data

    Elk made up more than 90% of the ungulate community and usually occurred in small herds.

    “Elk comprised Ͼ90% of the ungulate community, usually in small herds (13.9 Ϯ 0.67 elk, mean Ϯ SE, with a maximum size of 253 elk, based on ground counts of 1143 ungulate herds in winter).”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Findingown data

    Elk shifted into wooded cover and decreased use of preferred grassland foraging areas with high predation risk when wolves were present.

    “Spatially and temporally fine-scaled data from GPS radio collars show that elk moved into the protective cover of wooded areas when wolves were present, reducing their use of preferred grassland foraging habitats that had high predation risk.”
    • Also stated in: § RESULTS · plain
      “In contrast, elk showed a marked shift into coniferous forest in the presence of wolves, leaving the grassy meadows that were preferred in the absence of wolves”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Findingown data

    Gallatin elk respond to wolf presence within one day or less and at a spatial scale of about 1 km or less, with strong effects on habitat use.

    “Gallatin elk respond to the presence of wolves on a time scale of one day or less, and a spatial scale of ϳ1 km or less, with strong effects on habitat use.”
    Quote in the paperSubstantial claim

  • The quote only partly states this claim.

    Findingown data

    Across all four drainages, cows were more likely than bulls to use woodland sites, which are safer but less preferred for foraging.

    “In all four drainages, cows showed a stronger tendency than bulls to occupy woodland sites, which were safer, but not preferred for foraging (effect of gender on probability of conifer occurrence at elk locations: b ϭ 0.029 Ϯ 0.013, t ϭ 2.198, P ϭ 0.028; Table 1).”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Findingown data

    Impulse-response forecasts show elk locations shift strongly from grassland to conifers in every drainage and both sexes, with little change in slope, elevation, or aspect.

    “Fig. 4 reveals differences between the sexes and among drainages in the strength with which elk respond to wolf presence, but a basic pattern of response is common to both sexes in all drainages.In all cases, elk locations shift strongly from grassland to conifers, despite very little change in slope, elevation, or aspect.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Findingreasoning

    Elk probably use open grassland foraging sites when wolves are absent and move into coniferous forest when wolves are present.

    “Elk are likely to occupy foraging sites in open grassland in the absence of wolves, but shift into coniferous forest in the presence of wolves”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Findingcitation

    Without wolves, elk heavily used grassland areas, consistent with earlier work showing they selectively forage for grasses.

    “In the absence of wolves, elk used grassland areas heavily (Figs. 1 and 3a), as expected on the basis of previous behavioral studies showing that elk forage selectively for grasses (Houston 1982, Irwin and Peek 1983, McCorquodale 1993, Unsworth et al. 1998).”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites Houston 1982

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

    • cites Irwin and Peek 1983 source
    • cites McCorquodale 1993 source

  • May combine two claims.

    Findingown data

    Every dependent variable was significantly influenced by at least one other dependent variable, meaning each independent variable had indirect effects on all others.

    “All of the dependent variables (elevation, slope, aspect, conifer occurrence, native grass occurrence, and speed of movement) were significantly affected by at least one other dependent variable (Table 1), so each independent variable had indirect effects on all others.”
    Quote in the paperNot a substantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

    Premisecitation

    Since 1927, the Gallatin winter elk herd has ranged from 1214 to 3028 animals, averaging 1653 ± 63.

    “Based upon aerial total counts conducted by Montana Fish, Wildlife and Parks (Hamlin 2004), the Gallatin winter elk herd has numbered 1214-3028 elk since 1927, holding relatively stable around the long-term average of 1653 Ϯ 63 animals (mean Ϯ SE).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Hamlin 2004

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • Classified as probably not a substantial claim.

    Premisecitation

    Many prey species change habitat use when at risk of predation, exchanging lower forage quality or quantity for greater safety.

    “Many prey species alter their use of habitats in response to predation risk, trading a reduction in forage quality or quantity for increased security (Abramsky 1996, Heithaus andDill 2002).”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites Abramsky 1996

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

    • cites Heithaus and Dill 2002 source

  • May combine two claims.

    Premisecitation

    A recent meta-analysis found behaviorally mediated effects on prey demography were usually stronger than direct consumption, but no study among 166 involved a terrestrial vertebrate pair.

    “A recent meta-analysis showed that behaviorally mediated effects on prey demography were generally stronger than the effects of direct consumption, but none of the 166 studies involved a pair of terrestrial vertebrates (Preisser et al. 2005).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Preisser et al. 2005 source

  • May combine two claims.

    Interpretationown data

    Local precipitation changed little after wolf colonization, so abiotic factors do not explain the elk demographic changes well.

    “Much of Montana has experienced drought during the period since wolf colonization, but local precipitation has showed little change, so these changes in elk demography are not well explained by abiotic factors (Fig. 2).”
    Quote in the paperSubstantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    Selection is generally thought to favor individuals that best balance the benefits of reducing risk against its costs.

    “It is generally accepted that selection will favor individuals that optimally balance the benefits of risk reduction against its costs (Lima 1998(Lima , 2002)).”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Lima 1998 source
    • cites Lima 2002 source

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    Ungulate populations are well known to shape plant communities via aggregation and foraging choices.

    “It is well established that ungulate populations can structure plant communities through patterns of aggregation and foraging decisions (McNaughton 1985, Augustine and McNaughton 1998, Bardgett and Wardle 2003).”
    Quote in the paperNot a substantial claim0 of 3 sources checked
    • cites McNaughton 1985 source
    • cites Augustine and McNaughton 1998 source
    • cites Bardgett and Wardle 2003 source

  • May combine two claims.

    Interpretationreasoning

    Habitat-selection changes in response to wolves are consistent with a behaviorally mediated trophic cascade, but they do not exclude the possibility that plant-community changes mainly reflect declining elk numbers.

    “Although changes in habitat selection in response to wolves are compatible with the hypothesis of a behaviorally mediated trophic cascade, they do not rule out the hypothesis that changes in plant communities are primarily a response to declining elk numbers.”
    Quote in the paperSubstantial claim
    • Rests on an assumption: Gallatin elk numbers have declined significantly, mainly because of changes since wolf recolonization.

  • May combine two claims.

    Premisecitation

    During winter, males in this population were in poorer physical condition than females, with significantly lower marrow fat concentrations.

    “Males in this population were in worse physical condition than females throughout the winter (due to loss of mass during the fall rut), as shown by significantly lower marrow fat concentrations (Winnie and Creel, in press).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Winnie and Creel, in press

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    Several other studies of North American ungulates have reported high predation risk in open areas.

    “High predation risk in open areas has been reported by several other studies of North American ungulates (e.g., Kunkel andPletscher 2000, Pierce et al. 2004).”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Kunkel and Pletscher 2000 source
    • cites Pierce et al. 2004 source

  • May combine two claims.

    Premisecitation

    Most antipredator-behavior research has studied ambush predators or simulated attacks, producing a conventional view of predator-prey encounters as brief and small-scale.

    “Most research on antipredator behavior has focused on ambush predators or simulated attacks (Kenward 1978, Elgar 1989, Lima 1995, Caro et al. 2004), yielding a conventional view of predator-prey encounters as events that occur quickly (seconds to minutes), on a small spatial scale (a few meters or body lengths).”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites Kenward 1978 source
    • cites Elgar 1989 source
    • cites Lima 1995 source

  • Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims. The quote only partly states this claim.

    Study design and conductown data

    The plant occurrence regression models were chosen with AICc and validated against 315 independent sampling sites.

    “These regressions were selected using AIC c via single-term deletions from a full model including slope, elevation, sine (E/W) and cosine (N/S) of aspect, and seven bands of Landsat 7 ETM reflectance data.”
    • Also stated in: § Habitat variables · plain
      “We validated the regression models using 315 independent sampling locations stratified by drainage and elevation across the Gallatin study site.”
    Quote in the paperNot a substantial claim

  • Probably not a substantial claim: may not pass S4 (based).

    Interpretationnone given

    The move into coniferous forest when wolves are present is presumably for protective cover.

    “presumably to benefit from protective cover.”
    Quote in the paperNot a substantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

    Study design and conductnone given

    The autoregression system is stable, so responses settle to an equilibrium over time.

    “The system of equations in the autoregression is stable, so responses converge to an equilibrium, given time.”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Interpretationcitation

    A large shift in habitat selection probably changes diets or energy budgets, as shown for elk that move from grassland to woodland in response to human hunters.

    “A strong shift in habitat selection is likely to alter diets or energy budgets, as has been shown for elk that move from grassland to woodland in response to human hunters (Morgantini and Hudson 1985).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Morgantini and Hudson 1985 source

  • May combine two claims.

    Study design and conductreasoning

    Because wolves were probably missed on some days they were present, the authors label data as 'wolves present' or 'wolves not detected'; this is conservative.

    “Nonetheless, it is likely that we failed to detect wolves on some days that they were present.Thus, we categorize our data as ''wolves present'' and ''wolves not detected.''This classification is conservative: failure to detect wolf presence might mask responses by elk to wolves, but should not create apparent differences where none exist.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Interpretationreasoning

    Larger habitat patches on the Northern Range than in Gallatin Canyon would tend to raise travel costs and limit habitat selection as an antipredator response.

    “Second, habitat patches are larger on the Northern Range than in the Gallatin Canyon, which would tend to increase travel costs and constrain habitat selection as an antipredator response.”
    Quote in the paperSubstantial claim
    • Rests on an assumption: Habitat patches are larger on the Northern Range than in Gallatin Canyon.

  • May combine two claims.

    Interpretationown data

    Both elk numbers and elk behavior have changed since wolf recovery.

    “Elk numbers and behavior have both changed since wolf recovery.”
    Quote in the paperSubstantial claim

  • Classified as probably not a substantial claim. May combine two claims.

    Premisecitation

    Mao (2003) concluded that Northern Range elk did not spatially separate from wolves weekly in winter, and that elk appear to use habitat selection to avoid wolves in summer but rely on other methods in winter.

    “For Yellowstone's Northern Range, Mao (2003) concluded: ''In winter, elk did not spatially separate themselves from wolves on a weekly timescale. . . .Elk appear to use habitat selection to avoid wolves in summer, but must rely on [other methods] in winter.''”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Mao 2003

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • May combine two claims.

    Interpretationreasoning

    Because elk change habitat types without topographic changes, their habitat selection in response to wolves appears direct rather than correlated.

    “The fact that elk change habitat types without changes in topography suggests that habitat selection in response to wolf presence is direct (i.e., not a correlated response).”
    Quote in the paperSubstantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Interpretationreasoning

    Species and ecosystems vary in how habitat features affect risk, which is unsurprising given differences in predator and prey tactics.

    “Clearly there is variation among species and ecosystems in the way that habitat features affect risk, which is not surprising, given that predators and prey vary in their behavioral tactics (e.g., stalking vs. coursing by predators, defense vs. escape by prey).”
    Quote in the paperNot a substantial claim

  • The quote only partly states this claim.

    Study design and conductreasoning

    Nonlinear vector autoregression was used to handle autocorrelation, simultaneous changes in location attributes, and confounding from other factors.

    “We used nonlinear vector autoregression to deal with three general characteristics of data from GPS collars.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Interpretationreasoning

    The habitat shift appears to be cover-seeking, though the data do not directly show that moving into woodland lowers predation risk.

    “Together, these results suggest that the habitat shift is a form cover-seeking, although the data do not directly demonstrate that shifting into woodland reduces predation risk (see Plate 1).”
    Quote in the paperSubstantial claim
    • Rests on an assumption: Shifting into coniferous woodland reduces predation risk for elk.

  • May combine two claims.

    Interpretationcitation

    Bulls responded less than cows in habitat selection, consistent with other antipredator behaviors.

    “As with other aspects of antipredator behavior (Creel and Winnie 2005;Winnie and Creel, in press), bulls were less responsive than cows with respect to habitat selection.”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Creel and Winnie 2005 source
    • cites Winnie and Creel, in press

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    Elk elsewhere respond similarly to humans during hunting season.

    “Elk in other ecosystems show a similar response to humans during the hunting season (Morgantini andHudson 1985, Skovlin et al. 2002).”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Morgantini and Hudson 1985 source
    • cites Skovlin et al. 2002

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

33 of 33

  • The paper relies on this without support.

    Wolf presence in a drainage indicates elevated predation risk for elk in that drainage on that day.

    “Predation risk varied spatially and temporally across the site, due to wolf movements, colonization, and pack failure.”
    “We scored wolves as present within in a drainage if we located them via radiotelemetry or found fresh tracks, kills, or scats.”

  • The paper relies on this without support.

    Shifting into coniferous woodland reduces predation risk for elk.

    “Together, these results suggest that the habitat shift is a form cover-seeking, although the data do not directly demonstrate that shifting into woodland reduces predation risk (see Plate 1).”

  • The paper relies on this without support.

    Habitat patches are larger on the Northern Range than in Gallatin Canyon.

    “habitat patches are larger on the Northern Range than in the Gallatin Canyon”

  • The paper relies on this without support.

    Wolves in this population select prey in favor of males.

    “wolves in this population bias their prey selection in favor of males”

  • The paper relies on this without support.

    Gallatin elk numbers have declined significantly, mainly because of changes since wolf recolonization.

    “In the Gallatin, elk numbers have shown a significant decline, driven mainly by changes since wolf recolonization.”

  • The paper relies on this without support.

    Montana hunting quotas were liberalized in 2004-2005 for much of the state but reduced in the Montana portion of the Greater Yellowstone Ecosystem.

    “Hunting quotas were liberalized for much of Montana in 2004-2005, but were reduced in elk management units in the Montana portion of the Greater Yellowstone Ecosystem.”
6 of 6

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?

  • After wolf reintroduction, strong behavioral and nutritional responses to predation risk from wolves were detected, together with declines in survival and fecundity too large to be explained by wolf predation alone.

    “After wolf reintroduction, strong behavioral and nutritional responses to predation risk from wolves were detected [60]–[63], together with declines in survival and fecundity too large to be explained by wolf predation alone [16], [43], [64], [65].”

    From Ecosystem Scale Declines in Elk Recruitment and Population Growth with Wolf Colonization: A Before-After-Control-Impact Approach (2014) · cites this paper as [60]

    The Results report that elk shifted from preferred grassland forage into coniferous forest when wolves were present, supporting the behavioral-response portion of the claim. The Discussion says this shift is likely to alter diets or energy budgets; it does not report a detected nutritional response. The survival and fecundity declines are attached to different citations in the citing sentence, so they are not assessed here. (LLM, conf=0.86)

    Experimental — accuracy not yet measured

Self-check

Percentages, sums and reported differences re-computed from the paper’s own numbers.

Each contradiction between two passages listed here was confirmed by an independent check.

  • Wolf effect on conifer use differs between the results and Table 1

    Major

    The results report a wolf-presence coefficient of 0.477 for conifer occurrence, but Table 1 reports 0.079 for the same effect. The associated standard errors also differ, so this is not a rounding difference.

  • The aerial photograph is assigned to different drainages

    Minor

    The methods identify the photograph in Fig. 1c as showing Daly drainage, while the figure caption identifies it as Porcupine drainage.

2 of 2

Cites

Flagged first, then by citations

2 of the 56 references couldn’t be matched to a known paper.

Cited by

Sorted by citations

Related

Authors
Creel, S., Winnie, J., Maxwell, B., Hamlin, K., & Creel, M.
Affiliations
  • Montana State University (US)
  • Montana Fish, Wildlife & Parks (US)
  • Universitat Autònoma de Barcelona (ES)
Corresponding author
Creel, Scott
ORCID
Venue
Ecology
Publisher
Wiley
Published
2005-12
Volume
86
Issue
12
Pages
3387-3397
ISSN
0012-9658 (ISSN-L)
DOI
10.1890/05-0032
OpenAlex
W2133681478
Cited by
774
References
56
Citations by year
2026: 14 · 2025: 35 · 2024: 43 · 2023: 27 · 2022: 51 · 2021: 36 · 2020: 41 · 2019: 50 · 2018: 45 · 2017: 49 · 2016: 44 · 2015: 44 · 2014: 40 · 2013: 39 · 2012: 33
FWCI
23.16
Type
journal article
Language
en
Open access
Licence
doi.wiley.com/10.1002/tdm_license_1.1 (tdm, 3561 d)
Locations
Ecology (publishedVersion)
Funders
  • National Science Foundation (0238169)
  • Office of Experimental Program to Stimulate Competitive Research
Topics
  • Physical Sciences › Environmental Science › Ecology › Wildlife Ecology and Conservation
  • Physical Sciences › Environmental Science › Ecology › Rangeland and Wildlife Management
  • Physical Sciences › Environmental Science › Ecology › Animal Ecology and Behavior Studies
Keywords
elk, red deer, wolf reintroduction, antipredator behavior, habitat selection, predation risk, trophic cascades, Greater Yellowstone Ecosystem, woody vegetation, foraging habitat, predator-prey interactions

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