Large herbivores and aquatic–terrestrial links in southern boreal forests
Open accessAbstract
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.
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- 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
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- 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.
- Findingown data
The mean ± SD extent of the seasonal aquatic resource shadow (~1.2 ± km2, range 0.4-3.3 km2) was smaller than the terrestrial distribution area (~100 km2).
“The mean (± SD) extent of seasonal aquatic resource shadow (~1•2 ± km 2 , range = 0•4-3•3 km 2 ) was less than the scope of terrestrial distribution (~100 km 2 ).”
Quote in the paperNot a substantial claim - Findingown data
Long-term carcass patterns show where these food-web connections appear across the landscape.
“The analysis of long-term carcass patterns (Figs 4 and 5) indicates where such food web linkages occur on the landscape.”
Quote in the paperSubstantial claim- Rests on an assumption: Moose carcass locations are a valid proxy for where aquatic-derived resources are transferred to terrestrial systems.
- Findingown data
Carcass locations show that aquatic subsidies carried in moose biomass reach upper trophic levels and detritivores in clusters across time and scales.
“The moose carcass location pattern reveals that the transfer of aquatic subsidies via moose biomass to upper trophic levels and detritivore communities is clustered over time, at multiple scales (Fig. 4).”
Quote in the paperSubstantial claim- Rests on an assumption: Moose carcass locations are a valid proxy for where aquatic-derived resources are transferred to terrestrial systems.
- Premisecitation
The aquatic diet fraction range is similar to an earlier estimate of 0.18.
“The range in aquatic diet fraction is comparable to a previous estimate of 0•18 (Belovsky & Jordan 1978).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Belovsky & Jordan 1978
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- Interpretationcitation
The N flux estimate (mean ± SD = 1.2 ± 0.2 kg N ha−1 year−1) is similar to N fixation inputs by mosses argued to be important in boreal forests elsewhere.
“However, our N flux estimate (mean ± SD = 1•2 ± 0•2 kg N ha -1 year -1 ) is similar to N fixation inputs by mosses that are argued to be important to boreal forests elsewhere (Zackrisson et al. 2004).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Zackrisson et al. 2004
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- Rests on an assumption: The Hobbs excretion models developed for elk diets in Colorado are applicable to moose in southern boreal forests.
- Interpretationreasoning
N flux could therefore make up at least twice the percentage of net annual N mineralization and other N cycle components.
“Hence, N flux could constitute at least double the percentage of net annual N mineralization and other N cycle components (Fig. 2).”
Quote in the paperSubstantial claim - Interpretationcitation
At high moose densities, N flux may be about three times higher.
“At high moose densities, which often occur in riparian habitats relative to inland habitats during spring and summer (Peterson 1977), N flux may be ~3× higher (Fig. 2).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Peterson 1977
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- Rests on an assumption: The Hobbs excretion models developed for elk diets in Colorado are applicable to moose in southern boreal forests.
- Premisecitation
Water depth usually restricts moose foraging in aquatic habitats to near-shore littoral zones.
“Water depth typically limits moose foraging in aquatic habitats to near-shore, littoral zones (Franzmann & Schwartz 1997).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Franzmann & Schwartz 1997
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- Premisecitation
Long-term studies that measure population dynamics together with the spatio-temporal patterns of resource flow across aquatic-terrestrial boundaries are uncommon.
“Long-term, measurement of population dynamics and associated spatio-temporal patterns of resource flow across aquatic-terrestrial boundaries are rare (reviewed in Polis et al . 2004;Ballinger and Lake 2006; but see Ben-David et al . 2005 for an exception).”
Quote in the paperSubstantial claim0 of 3 sources checked- cites Polis et al. 2004
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- cites Ballinger and Lake 2006
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- cites Ben-David et al. 2005
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- Interpretationreasoning
Because moose are circumpolar, they are an important but unquantified aquatic-terrestrial resource vector in boreal systems.
“Given the circumpolar extent of moose, they constitute an important, unquantified aquatic-terrestrial resource vector in boreal systems.”
Quote in the paperSubstantial claim - Premisecitation
Nitrogen is usually the most limiting element in temperate and boreal forests.
“Nitrogen was the focus because this element is usually the most limiting in temperate and boreal forests (Larsen 1980).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Larsen 1980
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- Interpretationcitation
The aquatic diet fraction range used (0.14-0.37) is conservative because the 95% CI reported by Tischler (2004) is 0.09-0.57.
“The aquatic diet fraction range used (i.e., A in equation (2) = 0•14-0•37) in calculating N aquatic is conservative because the 95% CI reported by Tischler (2004) for this range is 0•09-0•57.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Tischler 2004
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- Premisecitation
The maximum dissolved inorganic N runoff reported for boreal forests was 1.6 kg N ha−1 year−1.
“dissolved inorganic N runoff [1•6 kg N ha -1 year -1 reported maximum for boreal forests (Wetzel 2001)]”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Wetzel 2001
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- Interpretationown data
Aquatic-derived excretory resources moved by moose are likely most dispersed in riparian zones and within about 0.5 km of lake shorelines.
“Terrestrial dispersion of aquatic-derived excretory resources transferred via moose is likely highest in riparian zones and within ~0•5 km of lake shorelines (Fig. 4b).”
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.
- Interpretationcitation
The effects of moose foraging in littoral zones probably reach pelagic zones, depending on lake mixing.
“However, the effects of littoral zone foraging by moose probably extend to pelagic zones, depending on lake mixing dynamics (Wetzel 2001).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Wetzel 2001
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- Premisecitation
Atmospheric deposition reported for Isle Royale was 3 kg N ha−1 year−1.
“atmospheric deposition [3 kg N ha -1 year -1 reported for Isle Royale (Stottlemyer, Travis, & Toczydlowski 1995)]”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Stottlemyer et al. 1995
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- Premisecitation
The fraction of total daily excretory N that came from aquatic sources was 0.14-0.37, based on recent stable carbon and nitrogen isotope estimates of the aquatic portion of summer moose diets.
“The proportion of N t (equation 1) that was of aquatic origin, 0•14 -0•37, was determined using recent estimates of the aquatic fraction of summer moose diets based on stable carbon and nitrogen isotope analyses (Tischler 2004).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Tischler 2004
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- Interpretationreasoning
Aquatic-derived resources can move through moose excretion or moose biomass and then enter consumer, producer, or detritivore communities.
“Consequently, aquatic-derived resources can flow through either moose excretion or biomass pathways and enter consumer, producer, or detritivore communities.”
Quote in the paperSubstantial claim- Rests on an assumption: Excretion occurs daily, while mortality from predation or starvation is relatively pulsed.
- Interpretationcitation
N_aquatic could be much lower or much higher for an individual moose.
“Hence, N aquatic could be substantially lower or higher for individual moose.”
Quote in the paperNot a substantial claim - Premisecitation
Adult moose carcasses are large (270-550 kg), concentrated pools of labile nutrients available to predators, scavengers, and detritivores.
“Adult moose carcasses represent substantial (270 -550 kg), highly concentrated pools of labile nutrients available to predators (e.g., grey wolves), scavengers (Wilmers et al . 2003), and detritivore communities (Carter et al . 2007).”
Quote in the paperSubstantial claim0 of 2 sources checked- cites Wilmers et al. 2003
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- cites Carter et al. 2007
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- Interpretationown data
Grey wolves and moose can create concentrated areas of resource transfer because of clustered predation and foraging patterns.
“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 - Premisecitation
Stottlemyer et al. (1995) found annual net N mineralization rates under dominant boreal tree species on Isle Royale to be less than half the rate used in this analysis (20 kg ha−1 year−1 from Pastor et al. 1993).
“For example, Stottlemyer et al. (1995) found annual net N mineralization rates under dominant boreal tree species on Isle Royale to be below half of the rate used in this analysis (20 kg ha -1 year -1 from Pastor et al. 1993).”
Quote in the paperSubstantial claim0 of 2 sources checked- cites Stottlemyer et al. 1995
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- cites Pastor et al. 1993
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- Premisecitation
The ecological importance of resource flux is thought to increase with asymmetrical productivity between ecosystems.
“This forest-aquatic comparison is important because the ecological significance of resource flux is thought to be positively related to asymmetrical productivity between ecosystems (Polis et al . 1997).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Polis et al. 1997
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- Premisecitation
The soil and plant responses to urine and dung returns are well quantified for large herbivores, including moose on Isle Royale.
“Significant soil and terrestrial plant responses to urine and faecal returns are well quantified for large herbivores (McNaughton 1985;Hobbs 1996;Frank 1998;Olofsson, Stark, & Oksanen 2004), including moose on Isle Royale (Pastor et al . 1993).”
Quote in the paperNot a substantial claim0 of 3 sources checked- cites McNaughton 1985
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- cites Hobbs 1996
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- cites Frank 1998
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- Interpretationreasoning
The relative importance of the moose-mediated N flux depends on net N mineralization in the terrestrial system.
“The relative importance of the moose-mediated N flux is dependent on terrestrial system net N mineralization.”
Quote in the paperSubstantial claim - Premisecitation
In some parts of the landscape, wolves kill moose at rates up to twelve times higher than in other parts.
“Wolves kill moose in some regions of the landscape at up to 12× the rate of other regions (Bump et al. in review).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Bump et al. in review
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- Interpretationreasoning
Moose add a net amount of nitrogen from water to land habitats.
“conferring a net N influx to terrestrial habitats from aquatic habitats.”
Quote in the paperSubstantial claim - Interpretationreasoning
When moose reuse particular places and carcasses cluster over time, resource transfer in the receiving system can become highly concentrated.
“Repeated use of specific areas and clustering of moose carcasses over time can create highly concentrated areas of resource transfer in recipient systems.”
Quote in the paperNot a substantial claim- Rests on an assumption: Moose carcass locations are a valid proxy for where aquatic-derived resources are transferred to terrestrial systems.
- Interpretationreasoning
If moose eat aquatic plants, part of their body mass must come from aquatic sources.
“Even so, where moose consume aquatic vegetation, some fraction of moose biomass is undeniably aquatic in origin.”
Quote in the paperSubstantial claim - Premisecitation
Forage nitrogen concentrations are critical in determining whether nutrient cycling accelerates or decelerates.
“Forage N concentrations are critical in determining nutrient accelerating or decelerating scenarios (sensu Ritchie, Tilman, & Knops 1998).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Ritchie, Tilman, & Knops 1998
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- Interpretationown data
The buffer around interior lakes marks likely average summer core areas where moose feed on aquatic macrophytes.
“Buffer zone around interior lakes indicates probable mean summer core areas for moose foraging on aquatic macrophytes.”
Quote in the paperSubstantial claim - Interpretationreasoning
Through moose, wolves indirectly shape the flow of subsidies between aquatic and terrestrial systems.
“Wolves, therefore, have a cascading impact on aquatic-terrestrial resource subsidies transferred via moose.”
Quote in the paperSubstantial claim - Interpretationreasoning
The findings show that moose act as a major vector moving resources between water and land in boreal systems.
“These results indicate that moose are an important aquaticterrestrial resource vector in boreal systems”
Quote in the paperSubstantial claim- Rests on an assumption: The patterns and processes observed on Isle Royale are representative of boreal systems generally.
- Premisecitation
Potamogeton sp. and Myriophyllum sp. dominate shallow aquatic habitats on Isle Royale.
“dominate shallow aquatic habitats on Isle Royale (Qvarnemark and Sheldon 2004)”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Qvarnemark and Sheldon 2004
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- Interpretationreasoning
Aquatic foraging would cause moose to excrete excess nitrogen primarily as urea in riparian zones and within about 0.5 km of lakeshores.
“Aquatic foraging, therefore, would cause moose to excrete excess N primarily as urea in riparian zones and within ~0•5 km of lakeshores (Fig. 5b).”
Quote in the paperSubstantial claim - Premisecitation
Large mammal herbivory produces two contrasting patterns of nitrogen cycling in terrestrial ecosystems.
“Terrestrial ecosystems exhibit a dichotomous N cycling response to large mammal herbivory (reviewed in Pastor et al. 2006).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Pastor et al. 2006
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- Premisecitation
Winter moose density was on average 415% higher on small islets than on the main island.
“Moose densities in winter, which are important for assessing where aquatic resources incorporated into moose biomass are distributed, averaged 415% greater on small islets than on the main island (Stephens and Peterson 1984).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Stephens and Peterson 1984
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- Premisecitation
Up to nine moose have been seen feeding together in an area smaller than one square kilometre on a single lake.
“as many as nine different moose have been observed simultaneously foraging in an area less than 1 km -2 , on a single lake (Peterson 1995)”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Peterson 1995
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- Interpretationreasoning
Moose connect the seasonal productivity of aquatic systems to terrestrial predators, scavengers, and detritivores.
“Thus, moose link seasonal aquatic system productivity with terrestrial predator, scavenger, and detritivore communities.”
Quote in the paperSubstantial claim - Premisecitation
How much, how long, and how intensively moose forage in water varies, from a few months in this system to year-round use on the Copper River Delta, Alaska.
“It is important to note that the timing, duration, and intensity of aquatic foraging by moose is variable, ranging from the few months observed in our system to the year-round use of aquatic plants reported by Mac-Cracken, Vanballenberghe & Peek (1993) on the Copper River Delta, Alaska.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Mac-Cracken, Vanballenberghe & Peek 1993
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- Interpretationreasoning
Excretory nitrogen moved from aquatic to terrestrial habitats by moose probably raises terrestrial nitrogen availability in moose summer core areas.
“Accordingly, the excretory N transferred from aquatic to terrestrial habitats via moose (Figs 1b, 2 and 4) likely increases terrestrial N availability within moose summer core areas.”
Quote in the paperSubstantial claim - Interpretationcitation
When moose disturb lake sediments while feeding on aquatic plants, phosphorus is released from interstitial sediment water, where concentrations are far higher than in the water above.
“turbation of lake sediments by moose while foraging for aquatic plants certainly releases P from interstitial sediment waters, where concentrations are orders of magnitude greater than in overlying water (Carpenter & Lodge 1993).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Carpenter & Lodge 1993
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- Interpretationreasoning
Moose could seasonally increase terrestrial nitrogen cycling when they forage on nitrogen-rich aquatic macrophytes.
“Moose could seasonally increase terrestrial N cycling when foraging on N-rich aquatic macrophytes.”
Quote in the paperSubstantial claim - Interpretationnone given
Mechanisms influencing carcass hot and cold spots include island-scale habitat features and smaller-scale predation patterns.
“Mechanisms influencing carcass 'hot' and 'cold' spots include island-scale habitat features and smaller-scale predation patterns.”
Quote in the paperSubstantial claim- Rests on an assumption: Moose carcass locations are a valid proxy for where aquatic-derived resources are transferred to terrestrial systems.
- Premisecitation
Potamogeton sp. and Myriophyllum sp. are readily consumed by moose.
“two genera, Potamogeton sp. and Myriophyllum sp., are readily consumed by moose”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Qvarnemark and Sheldon 2004
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- May combine two claims. 29
- Classified as probably not a substantial claim. 7
May combine two claims.
Findingown dataThe 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 dataThe 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 dataThe 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 claimMay combine two claims.
Findingown dataVariation 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.
FindingreasoningThe 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 dataThe 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 claimMay combine two claims.
FindingcitationTotal 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
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- cites Hobbs 2006
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- 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.
FindingreasoningUnder 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 claimMay combine two claims.
Findingown dataLocal 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 claimMay combine two claims.
PremisecitationLarge 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
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- cites Danell et al. 2002
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- cites Carter et al. 2007
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PremisecitationMore 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
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PremisecitationMost 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
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- cites Vucetich & Peterson 2004
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Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Study design and conductcitationMoose 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
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May combine two claims.
InterpretationreasoningA 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 claimProbably not a substantial claim: may not pass S3 (load-bearing).
InterpretationreasoningN 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.
PremisecitationMoose 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
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- cites Peterson 1977
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- cites Ballinger and Lake 2006
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May combine two claims.
Interpretationown dataMoose 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.
PremisecitationDuring 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
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PremisecitationTerrestrial 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
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Probably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationTheory 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
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- cites Loreau & Holt 2004
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May combine two claims.
PremisecitationAt 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 givenPre-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 claimMay combine two claims.
PremisecitationFaecal 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.
InterpretationreasoningWolves 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 claimMay combine two claims.
PremisecitationCalf 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 givenLater 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 dataThis 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 claimMay combine two claims.
InterpretationreasoningResources 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 claimMay combine two claims.
PremisecitationIn 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.
PremisecitationThe 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.
PremisecitationUrea-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.
PremisecitationMoose 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.
PremisecitationThe 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.
InterpretationreasoningWolves 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
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.”
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
MinorThe 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.
“Z-scores less than -1•96 or greater than 1•96”
“(i.e., one standard deviation)”
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Self-citationEffects of invasive aquatic carrion on soil chemistry and terrestrial microbial communities
Novais, A., Pascoal, C., & Sousa, R. (2017) · Biological Invasions · doi:10.1007/s10530-017-1459-4 · Cited by 14 · see citing works
Fertilizing riparian forests: nutrient repletion across ecotones with trophic rewilding
Bump, J. K. (2018) · Philosophical Transactions of the Royal Society B Biological Sciences · doi:10.1098/rstb.2017.0439 · Cited by 14 · see citing works
Self-citationFresh Waters, Climate Change and UK Nature Conservation
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White‐tailed deer consumption of emergent macrophytes mediates aquatic‐to‐terrestrial nutrient flows
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The sixth R: Revitalizing the natural phosphorus pump
Abraham, A. J., Roman, J., & Doughty, C. E. (2022) · The Science of The Total Environment · doi:10.1016/j.scitotenv.2022.155023 · Cited by 9 · see citing works
Towards a mechanistic understanding of animal–ecosystem interactions
Meyer, G. A., & Leroux, S. (2023) · Journal of Animal Ecology · doi:10.1111/1365-2656.14023 · Cited by 8 · see citing works
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- 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
- bronze
- doi.org
- 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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