Cited by 69Wildlife Ecology and ConservationEcology and biodiversity studiesAnimal Ecology and Behavior Studies

Large herbivores and aquatic–terrestrial links in southern boreal forests

Open access
Bump, Joseph K., Tischler, Keren B., Schrank, Amy J., Peterson, Rolf O., Vucetich, John A.·2009·Journal of Animal Ecology

Abstract

1. Concurrent measurement of population dynamics and associated spatio-temporal patterns of resource flow across aquatic-terrestrial boundaries are rare, yet necessary to understand the consequences of cross-habitat resource flux. Long-term study of the moose Alces alces (L.) population in Isle Royale National Park (Lake Superior, USA) provides an opportunity to examine the patterns of resource flux from aquatic to terrestrial habitats over approximately50 years. 2. We analysed the spatio-temporal dynamics of aquatic-derived nitrogen (N) that moose transfer to terrestrial systems by using excretion models, foraging parameters, moose densities, and moose carcass locations (n = 3616) collected from 1958-2005. 3. Results suggest that moose transfer significant amounts of aquatic-derived N to terrestrial systems, which likely increases terrestrial N availability in riparian zones. A seasonal increase in terrestrial N availability when moose are foraging on N-rich aquatic macrophytes would contrast with the depression of soil N mineralization previously attributed indirectly to moose. 4. Aquatic foraging by moose and moose carcass locations are significantly clustered at multiple scales, indicating that grey wolves Canis lupus (L.) and moose can create concentrated areas of resource transfer due to clustered predation and foraging patterns. 5. This study shows that patterns of faunal-mediated resource transfer can depend significantly on predator-prey dynamics, and that large predators in this system influence herbivore-controlled resource transfer between ecosystems. Given the circumpolar extent of moose, they constitute an important, unquantified aquatic-terrestrial resource vector in boreal systems.

Integrity signals

Registry

Crossref · 10.1111/j.1365-2656.2008.01498.x

Cites

51 of 51 references verified0 citation errors · 0 not checkable

Statistics

Nothing to recompute.

Claims the paper makes

0 of 57 cited sources checked · 0 supported · 0 flagged

  • Findingown data

    Moose aquatic foraging and the places where moose carcasses are found show significant clustering at more than one spatial scale.

    “Aquatic foraging by moose and moose carcass locations are significantly clustered at multiple scales, indicating that grey wolves Canis lupus (L.) and moose can create concentrated areas of resource transfer due to clustered predation and foraging patterns.”
    Quote in the paperSubstantial claim
  • Findingown data

    Over the past century, terrestrial N gain via moose fluctuated tenfold before and after wolf establishment.

    “Over the past century, terrestrial N gain via moose has exhibited 10-fold fluctuations, respectively, before and after wolf establishment (Fig. 3).”
    Quote in the paperSubstantial claim
    • Rests on an assumption: The Hobbs excretion models developed for elk diets in Colorado are applicable to moose in southern boreal forests.
  • Findingown data

    N_aquatic is affected more by plant N concentration than by the aquatic diet fraction.

    “N aquatic is more strongly influenced by plant N concentration than the aquatic diet fraction (note greater slope along z-axis compared to x-axis in Fig. 1a and b).”
    Quote in the paperNot a substantial claim
    • Rests on an assumption: The Hobbs excretion models developed for elk diets in Colorado are applicable to moose in southern boreal forests.
  • Findingown data

    Moose carcasses were strongly clustered at the island scale, with less than a 0.1% probability that the clustering of wolf-killed carcass sites arose by chance.

    “The distribution of moose carcasses showed a striking degree of clustering at the island scale, with less than a 0•1% likelihood that the clustering of wolf-killed moose carcass sites resulted from random chance (Fig. 4b; Getis-Ord Gi z-score = 4•13 standard deviations; Getis and Ord 1992).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Getis and Ord 1992

      Not checked in the free run.

  • Findingown data

    Mean ± SD annual N gain for riparian habitats from 1958 to 2005 was 1.06 ± 0.4 kg N ha−1 year−1.

    “Mean (± SD) annual N gain for riparian habitats for 1958 -2005 was 1•06 ± 0•4 kg N ha -1 year -1 .”
    Quote in the paperSubstantial claim
    • Rests on an assumption: The Hobbs excretion models developed for elk diets in Colorado are applicable to moose in southern boreal forests.
  • May combine two claims. 29
  • Classified as probably not a substantial claim. 7
  • May combine two claims.

    Findingown data

    The summer daily N of aquatic origin excreted by moose rises with aquatic diet fraction and forage N concentration, averaging 78 ± 12 g N day−1 moose−1 on Isle Royale.

    “Similarly, the summer daily N of aquatic origin (N aquatic ) excreted by moose increases as the aquatic diet fraction and forage N concentration rise, with a mean (± SD) on Isle Royale of 78 ± 12 g N day -1 moose -1 (average of open plot section in Fig. 1b).”
    Quote in the paperSubstantial claim
    • Rests on an assumption: The Hobbs excretion models developed for elk diets in Colorado are applicable to moose in southern boreal forests.
  • May combine two claims.

    Findingown data

    The total daily N excreted by Isle Royale moose in summer rises with intake rate and forage N concentration, averaging 283 ± 37 g N day−1 moose−1.

    “The summer total daily N (N t ) excreted by Isle Royale moose increases as intake rate and forage N concentration rise, with a mean ( ± 1SD) on Isle Royale of 283 ± 37 g N day -1 moose -1 (Fig. 1a).”
    Quote in the paperSubstantial claim
    • Rests on an assumption: The Hobbs excretion models developed for elk diets in Colorado are applicable to moose in southern boreal forests.
  • May combine two claims.

    Findingown data

    The Getis-Ord Gi* and Local Moran's I z-scores together indicated significant clustering of high-density and low-density carcass areas, and that sites of similar density were usually adjacent at local scales.

    “Together, the Getis-Ord Gi* and Local Moran's I z-scores indicated that there was significant clustering of high-density and low-density carcass areas (Fig. 5a) and that sites of similar density were most often located adjacent to each other at local scales (Fig. 5b).”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Findingown data

    Variation in moose density is the most important determinant of total N flux and largely absorbs the uncertainty from excretion and foraging parameters.

    “Variation in moose density is the most important determinant of total N flux (Fig. 3) and largely subsumes the uncertainty associated with excretion and foraging parameters.”
    Quote in the paperSubstantial claim
    • Rests on an assumption: The Hobbs excretion models developed for elk diets in Colorado are applicable to moose in southern boreal forests.
  • May combine two claims.

    Findingreasoning

    The total summer excretory N of aquatic origin transferred to land by moose on Isle Royale averaged 1.2 ± 0.2 kg N ha−1 year−1, which was 6%, 75%, and 40% of net annual N mineralization, annual N runoff, and annual atmospheric N deposition, respectively.

    “The total summer excretory N of aquatic origin (N flux ) transferred to terrestrial systems via moose on Isle Royale was (mean ± SD) 1•2 ± 0•2 kg N ha -1 year -1 , constituting 6, 75%, and 40% of net annual N mineralization, annual N runoff, and annual atmospheric N deposition, respectively (Fig. 2).”
    Quote in the paperSubstantial claim
    • The authors state this as an assumption: Moose excretory returns directly to aquatic systems are negligible.
    • Rests on an assumption: The Hobbs excretion models developed for elk diets in Colorado are applicable to moose in southern boreal forests.
  • May combine two claims.

    Findingown data

    The positively skewed z-score distributions indicated that high-density carcass sites were more strongly associated with one another than with low-density sites, and that similar carcass density values were more strongly associated.

    “The positively skewed z-score distributions (Fig. 5a) indicated that high-density ('hot spots') carcass sites were more strongly associated with one another than with low-density carcass sites ('cold spots'), and that similar carcass density values were more strongly associated (Fig. 5b).”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Findingcitation

    Total daily N excretion is affected more by plant N concentration than by intake rate, because urinary N excretion rises quadratically with forage N concentration.

    “N t is more strongly influenced by plant N concentration than intake rate because urinary N excretion increases quadratically with forage N concentrations (Hobbs 1996;2006).”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites Hobbs 1996

      Not checked in the free run.

    • cites Hobbs 2006

      Not checked in the free run.

    • Rests on an assumption: The Hobbs excretion models developed for elk diets in Colorado are applicable to moose in southern boreal forests.
  • May combine two claims.

    Findingreasoning

    Under such concentrated conditions, nitrogen moved from aquatic to terrestrial systems through moose might reach roughly 4.5 kg N per hectare per year in those places, which would be a large share of nitrogen cycling components.

    “Under such conditions, N transferred to terrestrial systems via moose could approach ~4•5 kg N ha -1 year -1 in those areas, constituting much higher proportions of N cycling components (Fig. 2).”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Findingown data

    Local Moran's I z-scores show that 4% of carcass sites have densities unlike neighboring areas, while about 52% have densities similar to neighbors.

    “Local Moran's I z-scores indicate 4% of carcass sites exhibit carcass densities dissimilar to adjacent areas, while about half (52%) of carcass sites are located in areas of similar carcass density.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Premisecitation

    Large herbivores are usually viewed as a resource for secondary consumers, but they can also contribute directly to detritivore and producer communities.

    “Large herbivores are typically considered a resource for secondary consumers (e.g., Wilmers et al . 2003), but can also directly contribute to detritivore and producer communities (Danell, Berteaux, & Brathen 2002;Carter, Yellowlees, & Tibbett 2007).”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites Wilmers et al. 2003

      Not checked in the free run.

    • cites Danell et al. 2002

      Not checked in the free run.

    • cites Carter et al. 2007

      Not checked in the free run.

  • May combine two claims.

    Premisecitation

    More than three moose per hectare have been recorded visiting mineral licks on Isle Royale, and all these licks are close to inland lakes.

    “Over three individual moose per hectare have been recorded visiting Isle Royale mineral licks, all of which are near inland lakes (Risenhoover & Peterson 1986)”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Risenhoover & Peterson 1986

      Not checked in the free run.

  • May combine two claims.

    Premisecitation

    Most North American moose population densities exceed 1 moose km−2, and Isle Royale densities generally range from about 1 to 6 individuals km−2.

    “Most North American moose population densities are > 1 moose km -2 (Franzmann & Schwartz 1997) and Isle Royale densities generally range ~1-6 individuals km -2 (Vucetich & Peterson 2004).”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Franzmann & Schwartz 1997

      Not checked in the free run.

    • cites Vucetich & Peterson 2004

      Not checked in the free run.

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

    Study design and conductcitation

    Moose population estimates before 1958 are less reliable and are used only to show general trends.

    “Moose population estimates before 1958 are less reliable and are included to illustrate general trends only (Peterson 1977).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Peterson 1977

      Not checked in the free run.

  • May combine two claims.

    Interpretationreasoning

    A seasonal rise in terrestrial N availability while moose feed on N-rich aquatic macrophytes would contrast with the previously reported indirect depression of soil N mineralization by moose.

    “A seasonal increase in terrestrial N availability when moose are foraging on N-rich aquatic macrophytes would contrast with the depression of soil N mineralization previously attributed indirectly to moose.”
    Quote in the paperSubstantial claim
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Interpretationreasoning

    N flux could be substantially higher where animals concentrate in preferred foraging and resting sites.

    “However, N flux could be substantially higher in areas where animals are concentrated in preferred foraging and resting sites.”
    Quote in the paperNot a substantial claim
    • Rests on an assumption: The Hobbs excretion models developed for elk diets in Colorado are applicable to moose in southern boreal forests.
  • May combine two claims.

    Premisecitation

    Moose specialize at the aquatic-terrestrial interface, feed heavily on aquatic macrophytes in spring and summer, and most often excrete and die on land.

    “Moose are aquatic-terrestrial interface specialists ( sensu Ballinger and Lake 2006), foraging extensively on aquatic macrophytes during spring and summer (Franzmann & Schwartz 1997), and excreting and dieing most frequently in terrestrial habitats (Franzmann & Schwartz 1997;Peterson 1977).”
    Quote in the paperSubstantial claim0 of 3 sources checked
    • cites Franzmann & Schwartz 1997

      Not checked in the free run.

    • cites Peterson 1977

      Not checked in the free run.

    • cites Ballinger and Lake 2006

      Not checked in the free run.

  • May combine two claims.

    Interpretationown data

    Moose move substantial amounts of aquatic-derived N to land, which probably raises terrestrial N availability in riparian zones.

    “Results suggest that moose transfer significant amounts of aquatic-derived N to terrestrial systems, which likely increases terrestrial N availability in riparian zones.”
    Quote in the paperSubstantial claim
    • Rests on an assumption: The Hobbs excretion models developed for elk diets in Colorado are applicable to moose in southern boreal forests.
  • May combine two claims.

    Premisecitation

    During 65 h of watching moose forage in water shallow enough for reliable detection, no defecation was seen, whereas defecation was often seen during 48 h of land observation.

    “In 65 h of viewing moose foraging in water shallow enough to permit proper detection, defecation was never observed, while in 48 h of terrestrial observation, defecation was frequently observed (Belovsky & Jordan 1978).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Belovsky & Jordan 1978

      Not checked in the free run.

  • May combine two claims.

    Premisecitation

    Terrestrial nutrient fluxes are informative and comparable when expressed per unit area per year.

    “Terrestrial nutrient fluxes are informative and comparable when carried out on a per unit area per year basis (Hobbs 2006).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Hobbs 2006

      Not checked in the free run.

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

    Premisecitation

    Theory indicates that spatial subsidies can strongly affect community structure by changing local competition among producers and consumers.

    “Theory indicates that spatial subsidies can substantially influence community structure by affecting rates of local competition for producers and consumers (Holt 2004;Loreau & Holt 2004).”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Holt 2004

      Not checked in the free run.

    • cites Loreau & Holt 2004

      Not checked in the free run.

  • May combine two claims.

    Premisecitation

    At Isle Royale kill sites, moose carcasses raise soil macronutrients and microbial biomass, alter microbial composition, and increase leaf nitrogen in plants for at least two to three years, even though wolves and scavengers use them heavily.

    “Isle Royale moose carcasses, despite being very well utilized by wolves and scavengers, increase soil macronutrients and microbial biomass, shift soil microbial composition, and elevate leaf nitrogen for at least 2 -3 years in plants at kill sites (Bump et al. in review).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Bump et al. in review

      Not checked in the free run.

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

    Study design and conductnone given

    Pre-1958 data are only illustrative of a broad trend.

    “Data before 1958 should be considered illustrative of a general trend only.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Premisecitation

    Faecal nitrogen excretion rises linearly and urinary nitrogen excretion rises quadratically as forage nitrogen concentrations increase.

    “faecal N excretion increases linearly and urinary N excretion quadratically with forage N concentrations (Hobbs 1996;2006).”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites Hobbs 1996

      Not checked in the free run.

    • cites Hobbs 2006

      Not checked in the free run.

  • May combine two claims.

    Interpretationreasoning

    Wolves clearly shape where carcasses are distributed and thus where moose-mediated resources flow.

    “The direct influence of wolves on the spatial distribution of carcasses, and thereby patterns in the moose-mediated resource flux, is readily observable.”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Premisecitation

    Calf kills on Isle Royale tend to occur near shorelines in dense cover, whereas adult kills are more randomly placed.

    “On Isle Royale, kills of calves are usually near shorelines, in heavy cover, while adult kills appear more randomly distributed (Peterson 1977;Stephens and Peterson 1984).”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Peterson 1977

      Not checked in the free run.

    • cites Stephens and Peterson 1984

      Not checked in the free run.

  • Probably not a substantial claim: may not pass S2 (checkable), S3 (load-bearing).

    Interpretationnone given

    Later improvements to these initial estimates may change model parameters and flux values, but they probably will not change the qualitative conclusions.

    “Future refinement of the first approximations made in this study may result in different modelling parameters and flux estimates, but such improvements will not likely alter the qualitative importance of our conclusions.”
    Quote in the paperNot a substantial claim
    • Rests on an assumption: The patterns and processes observed on Isle Royale are representative of boreal systems generally.
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Study design and conductown data

    This study did not measure how much of the aquatic forage eaten by moose becomes moose biomass.

    “In this study, we did not quantify the fraction of aquatic forage incorporated into moose biomass.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Interpretationreasoning

    Resources usually spread more widely in the receiving system than in the donor system, because moose range over the landscape while their littoral foraging is limited by depth.

    “Resource dispersion in the recipient system will typically be more extensive than the donor system resource shadow due to the landscape-level terrestrial range of moose and depth-restricted extent of their littoral zone foraging.”
    Quote in the paperNot a substantial claim
  • May combine two claims.

    Premisecitation

    In summer, cow moose and calves are significantly associated with human camp grounds (about twice as likely as bulls), which are almost all on shorelines and have no wolves.

    “In summer, cow moose and calves are significantly associated (~2× more likely than bulls) with human camp grounds (and no wolves), which are almost all situated on shorelines (Stephens and Peterson 1984).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Stephens and Peterson 1984

      Not checked in the free run.

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

    Premisecitation

    The significant influence consumers have on internal phosphorus cycling has been recognized, but such analyses have not included moose.

    “The significant influence consumers have on internal P cycling has been recognized (reviewed by Vanni 2002), but such analyses have not included moose.”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Vanni 2002

      Not checked in the free run.

  • May combine two claims.

    Premisecitation

    Urea-derived nitrogen is readily available to plants, whereas faecal nitrogen is less available and can lead to slower nitrogen mineralization than in soil humus alone.

    “Urea-derived N is readily available to plants, but faecal N is less so and can lead to slower N mineralization rates than found in soil humus alone (Pastor et al. 1993).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Pastor et al. 1993

      Not checked in the free run.

  • May combine two claims.

    Premisecitation

    Moose density is highest at the east end (about 5.4 per km^2), lowest in mid-island 1936 burn areas (about 0.8 per km^2), and moderate at the west end (about 1.8-3.4 per km^2).

    “The highest moose densities (~5•4 individuals km -2 ) are at the east end, low densities mid-island in major 1936 burn areas (~0•8 individuals km -2 ), and moderate densities (~1•8-3•4 individuals km -2 ) at the west end (Vucetich and Peterson 2004).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Vucetich and Peterson 2004

      Not checked in the free run.

  • May combine two claims.

    Premisecitation

    The critical nitrogen concentration in forage is about 1.5% dry mass; above it mammalian herbivores excrete nitrogen mainly as urea to remove excess, and below it they excrete mainly in faeces to maximize retention.

    “The resulting critical N concentration in forage is ~1•5% dry mass, above which mammalian herbivores excrete N primarily as urea to remove excesses, below which N is excreted primarily in faeces to maximize N retention (Pastor et al 2006).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Pastor et al 2006

      Not checked in the free run.

  • May combine two claims.

    Interpretationreasoning

    Wolves affect moose numbers, where moose live, and where their carcasses end up.

    “Wolves influence moose populations, habitat use, and carcass distribution.”
    Quote in the paperSubstantial claim
34 of 34
  • The authors state this as an assumption.

    Moose excretory returns directly to aquatic systems are negligible.

    “This estimate assumes moose excretory returns directly to aquatic systems are negligible, which is initially reasonable given that aquatic feeding bouts average 42•8 min day -1 on Isle Royale, digestive retention times typically exceed 12 h, and moose in summer spend about 5 × more time per day on land than foraging in aquatic habitats (Belovsky & Jordan 1978).”
  • The paper relies on this without support.

    Physical features do not generally limit where moose forage on land, and mortality can occur anywhere on the landscape.

    “In contrast, physical features do not generally limit where moose forage in terrestrial habitats and mortality can occur anywhere on the landscape.”
  • The paper relies on this without support.

    Excretion occurs daily, while mortality from predation or starvation is relatively pulsed.

    “Excretion occurs daily whereas mortality due to predation or starvation is relatively pulsed.”
  • The paper relies on this without support.

    The Hobbs excretion models developed for elk diets in Colorado are applicable to moose in southern boreal forests.

    “These excretory models were empirically developed for elk diets in Colorado, USA (Hobbs 2006).”
  • The paper relies on this without support.

    The patterns and processes observed on Isle Royale are representative of boreal systems generally.

    “These results indicate that moose are an important aquaticterrestrial resource vector in boreal systems”
  • The paper relies on this without support.

    Moose carcass locations are a valid proxy for where aquatic-derived resources are transferred to terrestrial systems.

    “The analysis of long-term carcass patterns (Figs 4 and 5) indicates where such food web linkages occur on the landscape.”
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?

  • Animals play an important role in transporting nutrients.

    “Animals play an important role in the transport of nutrients, but this role has diminished because many of the largest animals have gone extinct or experienced massive population declines.”

    From Global nutrient transport in a world of giants (2016) · cites this paper as [10]

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.

  • A 1.96 z-score is mislabeled as one standard deviation

    Minor

    The caption equates a z-score magnitude of 1.96 with one standard deviation. A z-score of 1.96 is 1.96 standard deviations from the mean.

1 of 1

Cites

Flagged first, then by citations

Research works printed but not linked: 1

  1. Tischler K.D., 2004, Aquatic Plant Nutritional Quality and Contribution to Moose Diet at Isle Royale National ParK

Cited by

Sorted by citations

Authors
Bump, J. K., Tischler, K. B., Schrank, A. J., Peterson, R. O., & Vucetich, J. A.
Affiliations
  • Michigan Technological University (US)
  • University of Michigan Biological Station (US)
Corresponding author
Bump, Joseph K.
Venue
Journal of Animal Ecology
Publisher
Wiley
Published
2009-02-09
Volume
78
Issue
2
Pages
338-345
ISSN
  • 0021-8790 (ISSN-L)
  • 1365-2656
OpenAlex
W1975795445
Cited by
69
References
51
Citations by year
2025: 2 · 2024: 5 · 2023: 13 · 2022: 10 · 2021: 1 · 2020: 2 · 2019: 2 · 2018: 5 · 2017: 4 · 2016: 4 · 2015: 8 · 2014: 5 · 2013: 4
FWCI
1.15
Type
journal article
Language
en
Open access
Locations
  • Journal of Animal Ecology (publishedVersion)
  • PubMed (publishedVersion)
  • Digital Commons - Michigan Tech (Michigan Technological University) (submittedVersion)
Funders
  • National Science Foundation (9972803, 0424562)
  • U.S. Environmental Protection Agency
Topics
  • Physical Sciences › Environmental Science › Ecology › Wildlife Ecology and Conservation
  • Physical Sciences › Environmental Science › Ecology › Ecology and biodiversity studies
  • Physical Sciences › Environmental Science › Ecology › Animal Ecology and Behavior Studies
Keywords
moose, nitrogen fluxes, riparian zones, aquatic macrophytes, foraging patterns, predator-prey dynamics, grey wolf, wolf, large herbivores, boreal forest, Isle Royale National Park, soil nitrogen mineralization
MeSH terms
Animals, Deer, Fresh Water, Michigan, Nitrogen, Trees, Wolves

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