Transportation of Pacific salmon carcasses from streams to riparian forests by bears
Abstract
Predation on Pacific salmon by bears (genus Ursus L., 1758) can be an important ecosystem process because the spatial distribution of carcasses largely determines whether marine-derived nutrients cycle through aquatic or terrestrial pathways. Direct observations on three streams in southeastern Alaska indicated that 49% of the pink (Oncorhynchus gorbuscha (Walbaum, 1792)) and chum (Oncorhynchus keta (Walbaum in Artedi, 1792)) salmon killed by bears were carried into the forest. The tendency of bears to transport carcasses was independent of the sex and species of salmon, but unspawned fish were more often transported than fish that had completed spawning. Data on tagged sockeye salmon (Oncorhynchus nerka (Walbaum in Artedi, 1792)) in one southwestern Alaska stream indicated that 42.6% of the killed salmon were transported, and that higher percentages were transported in years when salmon densities were greater. At six other streams, on average, 68.1% of the sockeye salmon killed were apparently transported away from the stream into the forest. Combining the data from all sites, the proportion of carcasses transported increased with water depth at the site. These results emphasize the role that bears play in mediating the interactions between nutrients from salmon and the terrestrial and aquatic ecosystems, and the variation in carcass distribution among streams and among years.
Résumé
La prédation des ours (le genre Ursus L., 1758) sur les carcasses de saumons du Pacifique peut être un processus écosystémique important parce que la répartition spatiale des carcasses détermine en grande partie si les nutriments d’origine marine sont recyclés par la voie aquatique ou la voie terrestre. Des observations directes sur trois cours d’eau du sud-est de l’Alaska indiquent que 49 % des saumons roses (Oncorhynchus gorbuscha (Walbaum, 1792)) et kéta (Oncorhynchus keta (Walbaum in Artedi, 1792)) tués par les ours sont transportés dans la forêt. La tendance qu’ont les ours à transporter les carcasses est indépendante du sexe et de l’espèce de saumon, mais les poissons qui n’ont pas frayé sont transportés plus fréquemment que les poissons qui ont terminé leur reproduction. Des données de marquage de saumons rouges (Oncorhynchus nerka (Walbaum in Artedi, 1792)) dans un cours d’eau du sud-ouest de l’Alaska indiquent que 42,6 % des saumons tués ont été transportés et que des pourcentages plus élevés sont transportés les années de plus forte densité de saumons. Dans six autres cours d’eau, en moyenne 68,1 % des saumons rouges tués ont apparemment été transportés du cours d’eau vers la forêt. Si les données sont combinées pour tous les sites, il appert que le pourcentage de carcasses transportées augmente en fonction de la profondeur de l’eau au site. Ces résultats soulignent le rôle joué par les ours comme médiateurs des interactions entre les nutriments des saumons et les écosystèmes terrestres et aquatiques et responsables de la variation de la répartition des carcasses d’un cours d’eau à un autre et d’une année à l’autre.
Get full access to this article
View all available purchase options and get full access to this article.
References
Ben-David, M., Hanley, T.A., Klein, D.R., and Schell, D.M. 1997. Seasonal changes in diets of coastal and riverine mink: the role of spawning Pacific salmon. Can. J. Zool. 75: 803–811.
Ben-David, M., Hanley, T.A., and Schell, D.M. 1998. Fertilization of terrestrial vegetation by spawning Pacific salmon: the role of flooding and predator activity. Oikos, 83: 47–55.
Ben-David, M., Titus, K., and Beier, L.R. 2004. Consumption of salmon by Alaskan brown bears: a trade-off between nutritional requirements and the risk of infanticide? Oecologia 138: 465–474.
Carlson, S.M., and Quinn, T.P. 2007. Ten years of varying lake level and selection on size-at-maturity in sockeye salmon. Ecology, 88: 2620–2629.
Carlson, S.M., Hilborn, R., Hendry, A.P., and Quinn, T.P. 2007. Predation by bears drives senescence in natural populations of salmon. PLoS One, 2: e1286.
Cederholm, C.J., Houston, D.B., Cole, D.L., and Scarlett, W.J. 1989. Fate of coho salmon (Oncorhynchus kisutch) carcasses in spawning streams. Can. J. Fish. Aquat. Sci. 46: 1347–1355.
Chi, D.K. 1999. The effects of salmon availability, social dynamics, and people on black bear (Ursus americanus) fishing behavior on an Alaskan salmon stream. Ph.D. dissertation, University of Utah, Logan.
Frame, G.W. 1974. Black bear predation on salmon at Olsen Creek, Alaska. Z. Tierpsychol. 35: 23–38.
Gard, R. 1971. Brown bear predation on sockeye salmon at Karluk Lake, Alaska. J. Wildl. Manage. 35: 193–204.
Gende, S.M., and Quinn, T.P. 2004. The relative importance of prey density and social dominance in determining energy intake by bears feeding on Pacific salmon. Can. J. Zool. 82: 75–85.
Gende, S.M., Quinn, T.P., and Willson, M.F. 2001. Consumption choice by bears feeding on salmon. Oecologia (Berl.), 127: 372–382.
Gende, S.M., Edwards, R.T., Willson, M.F., and Wipfli, M.S. 2002. Pacific salmon in aquatic and terrestrial ecosystems. Bioscience, 52: 917–928.
Gende, S.M., Quinn, T.P., Hilborn, R., Hendry, A.P., and Dickerson, B. 2004a. Brown bears selectively kill salmon with higher energy content but only in habitats that facilitate choice. Oikos, 104: 518–528.
Gende, S.M., Quinn, T.P., Willson, M.F., Heintz, R., and Scott, T.M. 2004b. Magnitude and fate of salmon-derived nutrients and energy in a coastal stream ecosystem. J. Freshwat. Ecol. 19: 149–160.
Gende, S.M., Miller, A.E., and Hood, E. 2007. The effects of salmon carcasses on soil nitrogen pools in a riparian forest of southeastern Alaska. Can. J. For. Res. 37: 1194–1202.
Hanson, R. 1992. Brown bear (Ursus arctos) predation on sockeye salmon (Oncorhynchus nerka) spawners in two tributaries of the Wood River Lake system, Bristol Bay, Alaska. M.S. thesis, University of Washington, Seattle.
Helfield, J.M., and Naiman, R.J. 2002. Salmon and alder as nitrogen sources to riparian forests in a boreal Alaskan watershed. Oecologia (Berl.), 133: 573–582.
Helfield, J.M., and Naiman, R.J. 2006. Keystone interactions: salmon and bear in riparian forests of Alaska. Ecosystems, 9: 167–180.
Hendry, A.P., Berg, O.K., and Quinn, T.P. 1999. Condition dependence and adaptation-by-time: breeding date, life history, and energy allocation within a population of salmon. Oikos, 85: 499–514.
Hilderbrand, G.V., Hanley, T.A., Robbins, C.T., and Schwartz, C.C. 1999. Role of brown bears (Ursus arctos) in the flow of marine nitrogen into a terrestrial ecosystem. Oecologia (Berl.), 121: 546–550.
Hocking, M.D., and Reimchen, T.E. 2006. Consumption and distribution of salmon (Oncorhynchus spp.) nutrients and energy by terrestrial flies. Can. J. Fish. Aquat. Sci. 63: 2076–2086.
Klinka, D.R., and Reimchen, T.E. 2002. Nocturnal and diurnal foraging behaviour of brown bears (Ursus arctos) on a salmon stream in coastal British Columbia. Can. J. Zool. 80: 1317–1322.
Marriott, R.A. 1964. Stream catalog of the Wood River lake system, Bristol Bay, Alaska. U.S. Fish Wildl. Serv. Spec. Sci. Rep. Fish. No. 494.
Meehan, E.P., Seminet-Reneau, E.E., and Quinn, T.P. 2005. Bear predation on Pacific salmon facilitates colonization of carcasses by fly maggots. Am. Midl. Nat. 153: 142–151.
Merrell, T.R.J. 1964. Ecological studies of sockeye salmon and related limnological and climatological investigations, Brooks Lake, Alaska, 1957. U.S. Fish Wildl. Serv. Biol. Rep. No. 466.
Naiman, R.J., Bilby, R.E., Schindler, D.E., and Helfield, J.M. 2002. Pacific salmon, nutrients, and the dynamics of freshwater and riparian ecosystems. Ecosystems, 5: 399–417.
Naiman, R.J., Helfield, J.M., Bartz, K.K., Drake, D.C., and Honea, J.M. 2009. Pacific salmon, marine-derived nutrients and the dynamics of aquatic ecosystems. Am. Fish. Soc. Symp. In press.
O’Keefe, T.C., and Edwards, R.T. 2002. Evidence for hyporheic transfer and removal of marine-derived nutrients in a sockeye stream in southwest Alaska. Am. Fish. Soc. Symp. 33: 99–107.
Quinn, T.P., and Buck, G.B. 2000. Scavenging by brown bears, Ursus arctos, and glaucous-winged gulls, Larus glaucescens, on adult sockeye salmon, Oncorhynchus nerka. Can. Field-Nat. 114: 217–223.
Quinn, T.P., and Buck, G.B. 2001. Size and sex selective mortality on adult Pacific salmon: bears, gulls and fish out of water. Trans. Am. Fish. Soc. 130: 995–1005.
Quinn, T.P., and Kinnison, M.T. 1999. Size-selective and sex-selective predation by brown bears on sockeye salmon. Oecologia (Berl.), 121: 273–282.
Quinn, T.P., Hendry, A.P., and Buck, G.B. 2001. Balancing natural and sexual selection in sockeye salmon: interactions between body size, reproductive opportunity and vulnerability to predation by bears. Evol. Ecol. Res. 3: 917–937.
Quinn, T.P., Gende, S.M., Ruggerone, G.T., and Rogers, D.E. 2003. Density dependent predation by brown bears (Ursus arctos) on sockeye salmon (Oncorhynchus nerka). Can. J. Fish. Aquat. Sci. 60: 553–562.
Reimchen, T.E. 2000. Some ecological and evolutionary aspects of bear–salmon interactions in coastal British Columbia. Can. J. Zool. 78: 448–458.
Ruggerone, G.T., Hanson, R., and Rogers, D.E. 2000. Selective predation by brown bears (Ursus arctos) foraging on spawning sockeye salmon (Oncorhynchus nerka). Can. J. Zool. 78: 974–981.
Schindler, D.E., Scheuerell, M.D., Moore, J.W., Gende, S.M., Francis, T.B., and Palen, W.J. 2003. Pacific salmon and the ecology of coastal ecosystems. Front. Ecol. Environ, 1: 31–37.
Shuman, R.F. 1950. Bear depredations on red salmon spawning populations in the Karluk River system, 1947. J. Wildl. Manage. 14: 1–9.
Thomas, S.C., Halpern, C.B., Falk, D.A., Liguori, D.A., and Austin, K.A. 1999. Plant diversity in managed forests: understory responses to thinning and fertilization. Ecol. Appl. 9: 864–879.
Wilkinson, C.E., Hocking, M.D., and Reimchen, T.E. 2005. Uptake of salmon-derived nitrogen by mosses and liverworts in coastal British Columbia. Oikos, 108: 85–98.
Willson, M.F., and Halupka, K.C. 1995. Anadromous fish as keystone species in vertebrate communities. Conserv. Biol. 9: 489–497.
Willson, M.F., Gende, S.M., and Marston, B.H. 1998. Fishes and the forest: expanding perspectives on fish–wildlife interactions. Bioscience, 48: 455–462.
Willson, M.F., Gende, S.M., and Bisson, P.A. 2004. Anadromous fishes as ecological links between ocean, fresh water, and land. In Food webs at the landscape level. Edited by G.A. Polis, M.E. Power, and G.R. Huxel. University of Chicago Press, Chicago. pp. 284–300.
Winder, M., Schindler, D.E., Moore, J.W., Johnson, S.P., and Palen, W.J. 2005. Do bears facilitate transfer of salmon resources to aquatic macroinvertebrates? Can. J. Fish. Aquat. Sci. 62: 2285–2293.
Information & Authors
Information
Published In
Canadian Journal of Zoology
Volume 87 • Number 3 • March 2009
Pages: 195 - 203
History
Received: 13 October 2008
Accepted: 5 January 2009
Version of record online: 13 February 2009
Authors
Metrics & Citations
Metrics
Other Metrics
Citations
Cite As
Thomas P.QuinnT.P. Quinn, Stephanie M.CarlsonS.M. Carlson, Scott M.GendeS.M. Gende, and Harry B.Rich, Jr.H.B. Rich, Jr. 2009. Transportation of Pacific salmon carcasses from streams to riparian forests by bears. Canadian Journal of Zoology.
87(3): 195-203. https://doi.org/10.1139/Z09-004
Export Citations
If you have the appropriate software installed, you can download article citation data to the citation manager of your choice. Simply select your manager software from the list below and click Download.
Cited by
1. Spider waste enhances soil nutrient content, soil respiration, and plant growth
2. Carrion ecology in inland aquatic ecosystems: a systematic review
3. Brown bear (Ursus arctos) foraging in a mosaic of spatially discrete and variable habitats over 25 years of shifting Pacific salmon densities
4. Establishment of terrestrial mammals on former reservoir beds following large dam removal on the Elwha River, Washington, USA
5. Looking forward; A synthesis of stream research undertaken in Glacier Bay
6. Estimates of Fall-run Chinook Salmon Escapement in Two San Joaquin River Tributaries from Device-based and Survey-based Methods
7. The influence of human disturbance on Pacific salmon (Oncorhynchus spp.) in the diet of American black bears (Ursus americanus) in two areas of coastal British Columbia, Canada
8. Amphibian Biomass Export from Geographically Isolated Wetlands: Temporal Variability, Species Composition, and Potential Implications for Terrestrial Ecosystems
9. Examining the Role of Marine Mammals and Seabirds in Southeast Alaska’s Marine Ecosystem Dynamics
10. Salmon subsidies predict territory size and habitat selection of an avian insectivore
11. Local Values and Data Empower Culturally Guided Ecosystem‐Based Fisheries Management of the Wuikinuxv Bear–Salmon–Human System
12. A methodological roadmap to quantify animal‐vectored spatial ecosystem subsidies
13. Terrestrial and semi-aquatic scavengers on invasive Pacific pink salmon (Oncorhynchus gorbuscha) carcasses in a riparian ecosystem in northern Norway
14. Automated analysis of lateral river connectivity and fish stranding risks—Part 1: Review, theory and algorithm
15. Simulating the relative effects of movement and sociality on the distribution of animal-transported subsidies
16. Effects of subsidies from small anadromous Pacific salmon populations on stream and riparian food webs are mediated by channel gradient
17. Prevalence and patterns of scavenging by brown bears (Ursus arctos) on salmon (Oncorhynchus spp.) carcasses
18. Community Ecology and Conservation of Bear-Salmon Ecosystems
19. Do brown bears Ursus arctos avoid barbed wires deployed to obtain hair samples? A videographic assessment
20. Significance of anecdotes for historical perspective: black bear predation on sea turtle eggs
21. Relationships between Pacific salmon and aquatic and terrestrial ecosystems: implications for ecosystem‐based management
22. Factors affecting the fate of Pacific lamprey carcasses and resource transport to riparian and stream macrohabitats
23. Matching habitat choice: it's not for everyone
24. Managing salmon for wildlife: Do fisheries limit salmon consumption by bears in small Alaskan streams?
25. Beyond “Donors and Recipients”: Impacts of Species Gains and Losses Reverberate Among Ecosystems Due to Changes in Resource Subsidies
26. Reverberating effects of resource exchanges in stream–riparian food webs
27. Examining the relative influence of animal movement patterns and mortality models on the distribution of animal transported subsidies
28. Landscape Structure and Species Interactions Drive the Distribution of Salmon Carcasses in Coastal Watersheds
29. Optimal foraging or surplus killing: selective consumption and discarding of salmon by brown bears
30. Efficiency and composition of vertebrate scavengers at the land-water interface in the Chernobyl Exclusion Zone
31. Carrion Availability in Space and Time
32. A multidecade experiment shows that fertilization by salmon carcasses enhanced tree growth in the riparian zone
33. Alaskan brown bears (
Ursus arctos
) aggregate and display fidelity to foraging neighborhoods while preying on Pacific salmon along small streams
34. Habitat features mediate selective consumption of salmon by bears
35. Watershed influences on the structure and function of riparian wetlands associated with headwater streams – Kenai Peninsula, Alaska
36. Soil biogeochemical responses to the deposition of anadromous fish carcasses in inland riparian forests of the Pacific Northwest, USA
37. Advancing research on animal‐transported subsidies by integrating animal movement and ecosystem modelling
38. Intrapopulation diversity in isotopic niche over landscapes: Spatial patterns inform conservation of bear–salmon systems
39. Diverse foraging opportunities drive the functional response of local and landscape-scale bear predation on Pacific salmon
40. Importance of Riparian Zone: Effects of Resource Availability at Land-water Interface
41. Exploitation of marine resources by wolves in southwestern Alaska
42. Aerial insect responses to non-native Chinook salmon spawning in a Great Lakes tributary
43. Effects of experimentally added salmon subsidies on resident fishes via direct and indirect pathways
44. Modeling terrestrial carbon sources for juvenile Chinook salmon in the Merced River, California
45. Salmon nutrients are associated with the phylogenetic dispersion of riparian flowering‐plant assemblages
46. Exploratory behavior of dispersers within a metapopulation of sockeye salmon
47. Novel species interactions: American black bears respond to Pacific herring spawn
48. Modeling the influence of salmon spawning on hyporheic exchange of marine-derived nutrients in gravel stream beds
49. Temperate Rain Forest
50. Tundra
51. Heterogeneity of riparian habitats mediates responses of terrestrial arthropods to a subsidy of Pacific salmon carcasses
52. Complementary use of motion-activated cameras and unbaited wire snares for DNA sampling reveals diel and seasonal activity patterns of brown bears (Ursusarctos) foraging on adult sockeye salmon (Oncorhynchusnerka)
53. Can intense predation by bears exert a depensatory effect on recruitment in a Pacific salmon population?
54. Using stable isotopes to understand the feeding ecology of the Hokkaido brown bear (
Ursus arctos
) in Japan
55. Effects of animal movement strategies and costs on the distribution of active subsidies across simple landscapes
56. Terrestrial Salmon Carcass Decomposition: Nutrient and Isotopic Dynamics in Central Idaho
57. Selecting for the phenotypic optimum: size‐related trade‐offs between mortality risk and reproductive output in female sockeye salmon
58. Seasonal persistence of marine‐derived nutrients in south‐central Alaskan salmon streams
59. Stable isotope evidence indicates the incorporation into Japanese catchments of marine‐derived nutrients transported by spawning Pacific Salmon
60. Hyporheic Restoration in Streams and Rivers
61. Quantifying associations of large vertebrates with salmon in riparian areas of British Columbia streams by means of camera-traps, bait stations, and hair samples
62. Salmon consumption by Kodiak brown bears (Ursus arctos middendorffi) with ecosystem management implications
63. Spatio-temporal variation in river otter (
Lontra canadensis
) diet and latrine site activity
64. Salmon subsidize an escape from a size spectrum
65. Reciprocal subsidies in ponds: does leaf input increase frog biomass export?
66. Alternative foraging strategies among bears fishing for salmon: a test of the dominance hypothesis
67. Nitrogen uptake by plants subsidized by Pacific salmon carcasses: a hierarchical experiment
68. Ecological flexibility of brown bears on Kodiak Island, Alaska
69. Population dynamics and asynchrony at fine spatial scales: a case history of sockeye salmon (Oncorhynchus nerka) population structure in Alaska, USA
70. Holocene soil-geomorphic surfaces influence the role of salmon-derived nutrients in the coastal temperate rainforest of Southeast Alaska
71. Eco-evolutionary dynamics in Pacific salmon
72. Effects of hydromorphological integrity on biodiversity and functioning of river ecosystems
73. Salmon for terrestrial protected areas
74. The Benefits of Harvesting Wetland Invaders for Cellulosic Biofuel: An Ecosystem Services Perspective
75. Predator control of ecosystem nutrient dynamics
76. Effects of hydromorphological integrity on biodiversity and functioning of river ecosystems
77. Spatial and Temporal Patterns of Upriver Migration by Sockeye Salmon Populations in the Wood River System, Bristol Bay, Alaska
78. Diet, Movement, and Growth of Dolly Varden in Response to Sockeye Salmon Subsidies
View Options
Login options
Check if you access through your login credentials or your institution to get full access on this article.