Reviews in Fish Biology and Fisheries

, Volume 16, Issue 2, pp 183–200 | Cite as

Vertebrate exploitation of pulsed marine prey: a review and the example of spawning herring

Original Paper


Short-term bursts of prey availability occur in many ecosystems and have potential important consequences for both predator biology and ecosystem function. Examples of prey ‘pulses’ in marine ecosystems include spawning runs of several anadromous and marine fishes, horseshoe crab spawning, and salmonid juvenile outmigrations, which are exploited by numerous species of vertebrate predators. In a few cases, the fitness or demographic consequences of such predator–prey interactions are known or inferred, but too often that information remains unknown. We explored the extent of temporal and spatial variation in one example of a pulsed marine resource: the spawning of Pacific herring (Clupea pallasii). Spawning herring provide a rich, aggregated resource to which dozens of species of vertebrate predators often exhibit strong numerical responses. However, the spawning events are often variable in both time (annual differences of several to many weeks) and space (both regional and more local differences in size and timing of events). Such variability must affect more mobile predators less than area-restricted predators, and thus its effect would vary not only among species but also within species, depending on constraints of the predator life history. Unpredictability of the prey concentrations, whatever their proximate causes, may contribute to maintenance of metapopulations of prey such as herring, if unpredictability lessens the impact of predation.


Clupea pallasii Pacific herring Pulsed resources Short-term prey exploitation Vertebrate predators in marine ecosystems 


Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.


  1. Baker AJ, González PM, Piersma T, Niles LJ, de Lima Serrano do Nascimento I, Atkinson PW, Clark NA, Minton CDT, Peck MK, Aarts G (2004) Rapid population decline in red knots: fitness consequences of decreased refuelling rates and late arrival in Delaware Bay. Proc Roy Soc Lond B 271:875–882Google Scholar
  2. Bayer RD (1980) Birds feeding on herring eggs at the Yaquina estuary, Oregon. Condor 82:193–198Google Scholar
  3. Bishop MA, Green SP (2001) Predation on Pacific herring (Clupea pallasii) spawn by birds in Prince William Sound, Alaska. Fisher Oceanogr 10(Suppl. 1):149–158Google Scholar
  4. Blaxter JHS, Hunter JR (1982) The biology of the clupeoid fishes. Adv Mar Biol 20:1–223Google Scholar
  5. Botton ML, Loveland RE (1993) Predation by herring gulls and great black-backed gulls on horseshoe crabs. Wils Bull 105:518–521Google Scholar
  6. Burgner RL (1991) Life history of sockeye salmon (Oncorhynchus nerka). In: Groot C, Margolis L (eds) Pacific salmon life histories. University of British Columbia, Vancouver, BC, pp 1–117Google Scholar
  7. Burns JG, Ydenberg RC (2002) The effects of wing loading and gender on the escape flights of least sandpipers (Calidris minutilla) and western sandpipers (Calidris mauri). Behav Ecol Sociobiol 52:128–136Google Scholar
  8. Calkins D, Becker E, Spraker T, Loughlin T (1994) Impacts on Steller sea lions. In: Loughlin TR, (eds) Maine mammals and the Exxon Valdez. Academic Press, San Diego, CA, pp 119–139Google Scholar
  9. Carls MG, Marty GD, Hose JE (2002) Synthesis of the toxicological impacts of the Exxon Valdez oil spill on Pacific herring (Clupea pallasii) in Prince William Sound, Alaska, U.S.A. Can J Fish Aquat Sci 59:153–172Google Scholar
  10. Carlson R (1980) Seasonal distribution and environment of Pacific herring near Auke Bay, Lynn Canal, southeastern Alaska. Trans Am Fish Soc 109:71–78Google Scholar
  11. Casas J, Pincebourde S, Mandon N, Vannier F, Poujol R, Giron D (2005) Lifetime nutrient dynamics reveal simultaneous capital and income breeding in a parasitoid. Ecology 86:545–554Google Scholar
  12. Castro G, Myers JP (1993) Shorebird predation on eggs of horseshoe crabs during spring stopover on Delaware Bay. Auk 110:927–930Google Scholar
  13. Clark KE, Niles LJ, Burger J (1993) Abundance and distribution of migrant shorebirds in Delaware Bay. Condor 95:694–705Google Scholar
  14. Cleaver FC, Franett DM (1946) The predation by sea birds upon the eggs of the Pacific herring (Clupea pallasii) at Holmes Harbor during 1945. Department of Fisheries, Division of Scientific Research, State of Washington, Biological Report 46BGoogle Scholar
  15. Corton A (2002) The role of ‘conservatism’ in herring migrations. Rev Fish Biol Fisher 11:339–361Google Scholar
  16. Dickerson BR, Quinn TP, Willson MF (2002) Body size, arrival date, and reproductive success of pink salmon, Oncorhynchus gorbuscha. Ethol Ecol Evol 14:29–44CrossRefGoogle Scholar
  17. Dragoo DE, Byrd GV, Irons DB (2004) Breeding status, population trends and diets of seabirds in Alaska, 2002. U. S. Fish and Wildlife Service Report AMNWR 04/15Google Scholar
  18. Dunn PO, May TA, McCollough MA, Howe MA (1988) Length of stay and fat content of migrant semipalmated sandpipers in eastern Maine. Condor 90:824–835Google Scholar
  19. Frost KJ, Lowry LF, VerHoef JM (1999) Monitoring the trend of harbor seals in Prince William Sound, Alaska after the Exxon Valdez oil spill. Mar Mammal Sci 15:494–506Google Scholar
  20. Furness RW, Tasker ML (1997) Seabird consumption in sand lance MSVPA models for the North Sea, and the impact of industrial fishing on seabird populations dynamics. In: Forage fishes in marine ecosystems. University of Alaska Sea Grant College Program Report No. 97–01, pp 147–169Google Scholar
  21. Gales NJ, Fraser WR, Costa DP, Southwell C (2004) Do crabeater seals forage cooperatively?. Deep-Sea Res II 51:2305–2310Google Scholar
  22. Gende SM, Womble JN, Willson MF, Marston BH (2001) Cooperative foraging by Steller sea lions (Eumetopias jubatus). Can Field-Nat 115:355–356Google Scholar
  23. Gende SM, Quinn TP, Willson MF, Heintz R, Scott TM (2004) Magnitude and fate of salmon-derived nutrients and energy in a coastal stream ecosystem. J Freshwater Ecol 19:149–160Google Scholar
  24. Gentry RL (1970) Social behavior of the Steller sea lion. PhD. Dissertation. University of California, Santa Cruz, 113 ppGoogle Scholar
  25. Gerke BL (2002) Spawning habitat characteristics of Pacific herring (Clupea pallasii) in PrinceWilliam Sound, Alaska. M.S. thesis. University of Alaska FairbanksGoogle Scholar
  26. Grass A (1973) Mew gulls and black turnstones feeding on herring eggs. Murrelet 54:38–39Google Scholar
  27. Guillemette M, Ouellet J-F (2005) Temporary flightlessness in pre-laying common eiders Somateria mollissima: are females constrained by excessive wing-loading or by minimal flight muscle ratio? Ibis 147:293–300Google Scholar
  28. Haegele CW (1993a) Seabird predation of Pacific herring, Clupea pallasii, spawn in British Columbia. Can Field-Nat 107:73–82Google Scholar
  29. Haegele CW (1993b) Epibenthic invertebrate predation of Pacific herring, Clupea pallasii, spawn in British Columbia. Can Field-Nat 107:83–91Google Scholar
  30. Haegele CW, Schweigert JF (1985) Distribution and characteristics of herring spawning grounds and description of spawning behaviour. Can J Fish Aquat Sci 42:39–55Google Scholar
  31. Haegele CW, Schweigert JF (1989) Egg loss from Pacific herring spawns in Barkley Sound in 1988. Can. Manuscript Report Fish Aquat Sci 2037Google Scholar
  32. Hamilton DJ, Barbeau MA, Diamond AW (2003) Shorebirds, mud snails, and Corophium volutator in the upper Bay of Fundy, Canada: predicting bird activity on intertidal mud flats. Can J Zool 81:1358–1366Google Scholar
  33. Hay DE (1985) Reproductive biology of Pacific herring (Clupea harengus). Can J Fish Aquat Sci 42 (Suppl. 1):111–126Google Scholar
  34. Hay DE, Kronlund AR (1987) Factors affecting the distribution, abundance, and measurement of Pacific herring (Clupea harengus) spawn. Can J Fish Aquat Sci 44:1181–1194Google Scholar
  35. Hay DE, McCarter PB, Kronlund R, Roy C (1989) Spawning areas of British Columbia herring: a review, geographical analysis and classification. Volumes 1-VI. Can. Manuscript Report Fish Aquat Sci 2019Google Scholar
  36. Hay DE, Toresen RR, Stephenson R, Thompson M, Claytor R, Funk F, Ivshina E, Jakobsson J, Kobayashi T, McQuinn I, Melvin G, Molloy J, Naumenko N, Oda KT, Parmanne R, Power M, Radchenko V, Schweigert J, Simmonds J, Sjöstrand B, Stevenson DK, Tanasichuk R, Tang R, Q, Watters DL, Wheeler J (2001) Taking stock: an inventory and review of world herring stocks in 2000. In: Funk F, Blackburn J, Day DH, Paul AJ, Stephenson R, Torresen R, Witherell D (eds) Herring expectation for a new millenium. Proceedings of the herring symposium 2000. Anchorage AK, pp 381–454Google Scholar
  37. Heard WR (1991) Life history of pink salmon (Oncorhynchus gorbuscha). In: Groot C, Margolis L (eds) Pacific salmon life histories. University of British Columbia Press, Vancouver, BC, pp 119–230Google Scholar
  38. Henderson RJ, Almatar SM (1989) Seasonal changes in the lipid composition of herring (Clupea harengus) in relation to gonad maturation. J Mar Biol Assoc U.K. 69:323–334CrossRefGoogle Scholar
  39. Heyman WD, Graham RT, Kjerfve B, Johannes RE (2001) Whale sharks Rhincodon typus aggregate to feed on fish spawn in Belize. Mar Ecol-Prog Ser 215:275–282Google Scholar
  40. Hicklin PW (1987) The migration of shorebirds in the Bay of Fundy. Wils Bull 99:540–570Google Scholar
  41. Hicklin PW, Smith PC (1979) The diets of five species of migrant shorebirds in the Bay of Fundy. Proc Nova Scotia Inst Sci 29:483–488Google Scholar
  42. Hicklin PW, Smith PC (1984) Selection of foraging sites and invertebrate prey by migrant semipalmated sandpipers, Calidris pusilla (Pallas) in Minas Basin, Bay of Fundy. Can J Zool 62:2201–2210CrossRefGoogle Scholar
  43. Hicklin PW, Linkletter LE, Peer DL (1980) Distribution and abundance of Corophium volutator (Pallas), Macoma balthica (L.) and Heteromastus filiformis (Clarapede) in the intertidal zone of Cumberland Basin and Shepody Bay, Bay of Fundy. Can Tech Rep Fish Aquat Sci 965:1–59Google Scholar
  44. Hildebrant GV, Schwartz CC, Robins CT, Jacoby ME, Hanley TA, Arthur SM, Servheen C (1999) The importance of meat, particularly salmon, to body size, population productivity, and conservation of North American brown bears. Can J Zool 77:132–138Google Scholar
  45. Hose JE, McGurk MD, Marty GD, Hinton DE, Brown ED, Baker TT (1996) Sublethal effects of the Exxon Valdez oil spill on herring embryos and larvae: morphological, cytogenetic, and histopathological assessments, 1989–1991. Can J Fish Aquat Sci 53:2355–2365Google Scholar
  46. Janzen DH (1976) Why bamboos wait so long to flower. Annu Rev Ecol Syst 7:347–391Google Scholar
  47. Jehl JR Jr (1988) Biology of the eared grebe and Wilson’s phalarope in the nonbreeding seasons: a study of adaptations to saline lakes. Stud Avian Biol 12:1–74Google Scholar
  48. Jönsson KI (1997) Capital and income breeding as alternative tactics of resource use in reproduction. Oikos 78:57–66Google Scholar
  49. Jurasz C, Jurasz V (1979) Feeding modes of the humpback whale, Megaptera novaengliae, in southeast Alaska. Scientific Rep Whale Res Inst 31:69–83Google Scholar
  50. Kålås JA, Heggberget TG, Bjørn PA, Reitan O (1993) Feeding behaviour and diet of goosanders (Mergus merganser) in relation to salmonid seaward migration. Aquat Living Resour 6:31–38Google Scholar
  51. Klaassen M, Lindström A, Meltofte H, Piersma T (2001) Arctic waders are not capital breeders. Nature 413:794PubMedGoogle Scholar
  52. Kvist A, Lindström A, Green M, Piersma T, Visser GH (2001) Carrying large fuel loads during sustained bird flight is cheaper than expected. Nature 413:730–732PubMedGoogle Scholar
  53. Lack D (1966) Population studies of birds. Clarendon Press, OxfordGoogle Scholar
  54. Lasker R (1985) What limits clupeoid production? Can J Fish Aquat Sci 42 s(Suppl. 1):31–38Google Scholar
  55. Lillegård M, Engen S, Sæther B, Toresen R (2005) Harvesting strategies for Norwegian spring-spawning herring. Oikos 110:567–577Google Scholar
  56. Loughlin TR, Perlov AS, Vlaadimirov VA (1992) Range-wide survey and estimation of total number of Steller sea lions in 1989. Mar Mammal Sci 8:220–239Google Scholar
  57. Loughlin TR, Sterling JT, Merrick RL, Sease JL, York AE (2003) Diving behaviour of immature Steller sea lions (Eumetopias jubatus). Fish Bull 101:566–582Google Scholar
  58. Lowry LF, Frost KJ, VerHoef JM, DeLong RA (2001) Movements of satellite tagged subadult and adult harbor seals in Prince William Sound, Alaska. Mar Mam Sci 17:835–861Google Scholar
  59. Marston BH, Willson MF, Gende SM (2002) Predator aggregations at eulachon (Thaleichthys pacificus) spawning runs in southeast Alaska. Mar Ecol-Prog Ser 231:229–236Google Scholar
  60. Matthews SL, Boates JS, Walde SJ (1992) Shorebird predation may cause discrete generations in an amphipod prey. Ecography 15:393–400Google Scholar
  61. Maurer BA (1990) Extensions of optimal foraging theory for insectivorous birds: implications for community structure. Stud Avian Biol 13:455–461Google Scholar
  62. McClelland BR, Young LS, Shea DS, McClelland PT, Allen HL, Spetttigue EB (1982) The bald eagle concentration in Glacier National Park, Montana: origin, growth, and variation in numbers. Living Bird 19:133–155Google Scholar
  63. McGurk MD, Brown ED (1996) Egg-larval mortality of Pacific herring in Prince William Sound, after the Exxon Valdez oil spill. Can J Fish Aquat Sci 53:2243–2354Google Scholar
  64. McLarney WO (1967) Intrastream movement, feeding habits, and structure of a population of coast range sculpin, Cottus aleuticus, in relation to eggs of the pink salmon, Oncorhynchus gorbuscha, in Alaska. PhD. dissertation, University of Michigan, Ann ArborGoogle Scholar
  65. Meijer T, Drent R (1999) Re-examination of the capital and income dichotomy in breeding birds. Ibis 141:399–414Google Scholar
  66. Merrick RL, Loughlin TR (1997) Foraging behavior of adult female and young-of-the-year Steller sea lions in Alaskan waters. Can J Zool 75:776–786Google Scholar
  67. Morrison RIG, Hobson KA (2004) Use of body stores in shorebirds after arrival on high-arctic breeding grounds. Auk 121:333–344Google Scholar
  68. Moulton LL (1999) Review of Lynn Canal herring. MJM Research, 1012 Shoreland Drive, Lopez Island, WAGoogle Scholar
  69. Munro JA, Clemens WA (1931) Water fowl in relation to the spawning of herring in British Columbia. Biol Board Canada Bull XVII:1–46Google Scholar
  70. Murdoch MH, Bärlocher F, Laltoo ML (1986) Population dynamics and nutrition of Corophium volutator (Pallas) in the Cumberland Basin (Bay of Fundy). J Exp Mar Biol Ecol 103:235–249Google Scholar
  71. Myers JP (1986) Sex and gluttony on Delaware Bay. Nat Hist 95:68–77Google Scholar
  72. Newton I (1998) Population limitation in birds. Academic Press, San DiegoGoogle Scholar
  73. Norcross BL, Hose JE, Fransden M, Brown ED (1996) Distribution, abundance, morphological condition, and cytogenetic abnormalities of larval Pacific herring in Prince William Sound, Alaska following the Exxon Valdez oil spill. Can J Fish Aquat Sci 53:2376–2387Google Scholar
  74. Norton DW, Senner SE, Gill RE Jr, Martin PD, Wright JM, Fukuyama AK (1990) Shorebirds and herring roe in Prince William Sound, Alaska. Am Birds 44:367–371Google Scholar
  75. Odum WE, Odum EP, Odum HT (1995) Nature’s pulsing paradigm. Estuaries 18:547–555Google Scholar
  76. Olson JM, McNeil WJ (1967) Research on pink salmon at Little Port Walter, Alaska, 1934–1964. U. S. Fish and Wildlife Service Data Report 17, Washington DCGoogle Scholar
  77. Ostfeld RS, Keesing F (2000) Pulsed resources and community dynamics of consumers in terrestrial ecosystems. Trends Ecol Evol 15:232–237PubMedGoogle Scholar
  78. Outram DN (1958) The magnitude of herring spawn losses due to bird predation on the west coast of Vancouver Island. Fish Res Board Canada, Progress Reports of the Pacific Coast Stations No. 111:9–13Google Scholar
  79. Outram DN, Humphreys RD (1974) The Pacific herring in British Columbia waters. Circular 100, Fisheries and Marine Service, Pacific Biological Station, Nanaimo, B. CGoogle Scholar
  80. Palsson WA (1984) Egg mortality upon natural and artificial substrata within Washington State spawning grounds of Pacific herring (Clupea harengus). M. S. thesis, University of Washington, Seattle. 191 ppGoogle Scholar
  81. Paton DC (1986) Honeyeaters and their plants in south-eastern Australia. In: Ford HA, Paton DC (eds) The dynamic partnership: Birds and plants in southern Australia. Woolman, Government Printer, South Australia, pp 9–19Google Scholar
  82. Paul AJ, Paul JM (1999) Energy contents of whole body, ovaries, and ova from pre-spawning Pacific herring. Alaska Fishery Res Bull 6(1):29–34Google Scholar
  83. Paul AJ, Paul JM, Brown ED (1996) Ovarian energy content of Pacific herring from Prince William Sound, Alaska. Alaska Fishery Res Bull 3(2):103–111Google Scholar
  84. Pearce-Higgins JW, Yalden DW (2004) Habitat selection, diet, arthropod availability and growth of a moorland wader: the ecology of European golden plover Pluvialis apricaria chicks. Ibis 146:335–346Google Scholar
  85. Peer DL, Linkletter LE, Hicklin PW (1986) Life history and reproductive biology of Corophium volutator (Crustacea: Amphipoda) and the influence of shorebird predation on population structure in Chignecto Bay, Bay of Fundy, Canada. Netherlands J Sea Res 20:359–373Google Scholar
  86. Pfister C, Kasprzyk MJ, Harrington BA (1998) Body-fat levels and annual return in migrating semipalmated sandpipers. Auk 115:904–915Google Scholar
  87. Piatt JF (1990) The aggregative response of common murres and Atlantic puffins to schools of capelin. Stud Avian Biol 14:36–51Google Scholar
  88. Pierotti R (1988) Associations between marine birds and mammals in the northwest Atlantic Ocean. In: Seabirds and other marine vertebrates: competition, predation, and other interactions. Columbia University Press, New York, pp 31–58Google Scholar
  89. Piersma T (1998) Phenotypic flexibility during migration: optimization of organ size contingent on the risks and rewards of fueling and flight? J Avian Biol 29:511–520Google Scholar
  90. Pitcher KW (1980) Food of the harbor seal (Phoca vitulina richardsi) in the Gulf of Alaska. Fishery Bull 78:544–549Google Scholar
  91. Pitcher KW (1981) Prey of the Steller sea lion, Eumetopias jubatus, in the Gulf of Alaska. Fishery Bull 79(3):467–472Google Scholar
  92. Raum-Suryan KL, Rehberg MJ, Pendleton GW, Pitcher KW, Gelatt TG (2004) Dispersal, movement patterns, and haulout use of pup and juvenile Steller sea lions (Eumetopias jubatus) in Alaska. Mar Mamm Sci 20:823–850Google Scholar
  93. Restani M, Harmata AR, Madden EM (2000) Numerical and functional responses of migrant bald eagles exploiting a seasonally concentrated food source. Condor 102:561–568Google Scholar
  94. Rodway MS, Cooke F (2001) Effect of food availability on arrival and departure decisions of harlequin ducks at diurnal feeding grounds. Condor 103:870–874Google Scholar
  95. Rodway MS, Cooke F (2002) Use of fecal analysis to determine seasonal changes in the diet of wintering harlequin ducks at a herring spawning site. J Field Ornith 73:363–371Google Scholar
  96. Rodway MS, Regehr HM, Ashley J, Clarkson PV, Goudie RI, Hay DE, Smith CM, Wright KG (2003) Aggregative response of harlequin ducks to herring spawning in the Strait of Georgia, British Columbia. Can J Zool 81:1–11Google Scholar
  97. Rooper CN (1996) Physical and biological factors affecting Pacific herring egg loss in Prince William Sound, Alaska. M.S. thesis, University of Alaska-Fairbanks, 195 ppGoogle Scholar
  98. Rosen DAS, Trites AW (2000) Digestive efficiency and dry-matter digestibility in Steller sea lions fed herring, pollock, squid, and salmon. Can J Zool 78:234–239Google Scholar
  99. Rosen DAS, Trites AW (2002) Cost of transport in Steller sea lions, Eumetopias jubatus. Mar Mam Sci 18:513–524Google Scholar
  100. Rounsefell GA, Dahlgren EH (1935) Races of herring, Clupea pallasii, in southeastern Alaska. Bull Bureau Fisher 58:119–141Google Scholar
  101. Sandegreen FE (1970) Breeding and maternal behavior of the Steller sea lion (Eumetopias jubatus) in Alaska. M.S. thesis, University of Alaska Fairbanks, 138 ppGoogle Scholar
  102. Sigler MF, Womble, J.N, Vollenweider JJ (2004) Availability to Steller sea lions (Eumetopias jubatus) of a seasonal prey resource: a prespawning aggregation of eulachon (Thaleichthys pacificus). Can J Fish Aquat Sci 61:1475–1484Google Scholar
  103. Similä T, Ugarte F (1993) Surface and underwater observations of cooperatively feeding killer whales in northern Norway. Can J Zool 71:1494–1499CrossRefGoogle Scholar
  104. Similä T, Holst JC, Christensen I (1996) Occurrence and diet of killer whales in northern Norway: seasonal patterns relative to the distribution and abundance of Norwegian spring-spawning herring. Can J Fish Aquat Sci 53:769–779Google Scholar
  105. Smith RJ (2003) Resources and arrival of landbird migrants at northerly breeding grounds: linking en route with breeding season events. PhD thesis, University of Southern MississippiGoogle Scholar
  106. Soto KH, Trites AW, Arias-Schreiber M (2004) The effects of prey availability on pup mortality and the timing of birth of South American sea lions (Otaria flavescens) in Peru). J Zool Lond 264:419–428Google Scholar
  107. Soto KH, Trites AW, Arias-Schreiber M (2006) Changes in diet and maternal attendance of South American sea lions indicate changes in the marine environment and prey abundance. Mar Ecol Prog Ser 312:277–290Google Scholar
  108. Stephens DW (1990) Foraging theory: up, down, and sideways. Stud Avian Biol 13:444–454Google Scholar
  109. Straley JM (1990) Fall and winter occurrence of humpback whales (Megaptera novaeangliae) in southeastern Alaska. Rep Int Whaling Commission, Special Issue 12:319–323Google Scholar
  110. Sullivan TM, Butler RW, Boyd WS (2002) Seasonal distribution of waterbirds in relation to spawning Pacific herring, Clupea pallasii, in the Strait of Georgia, British Columbia. Can Field-Nat 116:366–370Google Scholar
  111. Taylor SG, Lum JL (2005) Auke Creek Weir 2004. Annual report, operations, fish counts, and historical summaries. National Marine Fisheries Service, Auke Bay Laboratory, 22305 Glacier Hwy, Juneau AK 99801. Unpublished report 26 ppGoogle Scholar
  112. Thomas GL, Thorne RE (2001) Night-time predation by Steller sea lions. Nature 411:1013PubMedGoogle Scholar
  113. Thompson JN, Willson MF (1979) Evolution of temperate fruit/bird interactions: phenological strategies. Evolution 33:973–982Google Scholar
  114. Thorne RE (2005) Monitoring Pacific herring abundance with combined acoustic and optical technologies. Proceedings Oceans 05. Washington, D.C. Sept. 20–23Google Scholar
  115. Trillmich F, Ono KA (1991) Pinnipeds and El Niño: responses to environmental stress. Springer-Verlag, BerlinGoogle Scholar
  116. Tsipoura N, Burger J (1999) Shorebird diet during spring migration stopover on Delaware Bay. Condor 101:635–644Google Scholar
  117. Van Opzeeland IC, Corkeron PJ, Leyssen T, Similiä T, Van Parijs SM (2005) Acoustic behaviour of Norwegian killer whales, Orcinus orca, during carousel and seiner foraging on spring-spawning herring. Aquat Mammals 31:110–119Google Scholar
  118. VerHoef JM, Frost KJ (2003) Bayesian hierarchical model for monitoring harbor seal changes in Prince William Sound, Alaska. Environ Ecol Stat 10:201–209Google Scholar
  119. Vermeer K (1981) Food and populations of surf scoters in British Columbia. Wildfowl 32:107–116Google Scholar
  120. Vermeer K (1983) Marine bird populations in the Strait of Georgia: comparison with the west coast of Vancouver Island. Can Tech Rep Hydrography Ocean Sci 19:1–18Google Scholar
  121. Vermeer K (1992) The diet of birds as a tool for monitoring the biological environment. Occas. Paper, Can Wildlife Serv 75:41–50Google Scholar
  122. Vermeer K, Bourne N (1983) The white-winged scoter diet in British Columbia waters: resource partitioning with other scoters. In: Nettleship DN, Sanger GA, Springer PF (eds) Marine birds: their feeding ecology and commercial fishing relationships, Can. Wildlife Serv. Report for the Pacific Seabird Group, pp. 30–38Google Scholar
  123. Vermeer K, Morgan KH (1992) Marine bird populations and habitat use in a fjord on the west coast of Vancouver Island. In Vermeer K, Butler RW and Morgan KH (eds) The ecology, status, and conservation of marine and shoreline birds on the west coast of Vancouver Island. Occas. Paper No. 75, Can. Wildlife Ser, pp 86–95Google Scholar
  124. Vermeer, K., Bentley M, Morgan KH, Smith GEJ (1997) Association of feeding flocks of brant and sea ducks with herring spawn at Skidegate Inlet. In: Vermeer K, Morgan KH (eds) The ecology, status, and conservation of marine and shoreline birds of the Queen Charlotte Islands. Occas Paper Can Wildlife Serv. 93:102–107Google Scholar
  125. Vermeer K, Morgan KH, Dorst A, Whittington B (1992) Bird populations of estuaries on the southwest coast of Vancouver Island. In: Vermeer K, Butler RW, Morgan KH (eds) The ecology, status, and conservation of marine and shoreline birds in the west coast of Vancouver Island. Occas Pap Can. Wildlife Serv 75:97–108Google Scholar
  126. Ware DM (1985) Life history characteristics, reproductive value, and resilience of Pacific herring (Clupea harengus). Can J Fish Aquat Sci 42 (Suppl. 1):127–137Google Scholar
  127. Weidensaul S (1999) Living on the wind. North Point Press, New York, 340 ppGoogle Scholar
  128. Wespestad VG, Barton LH (1979) Distribution, migration, and status of Pacific herring. Nat. Mar. Fish. Serv., Northwest and Alaska Fisheries Center, Seattle WA. pp 509–525Google Scholar
  129. Willson MF, Halupka KC (1995) Anadromous fish as “keystone” resources in vertebrate communities. Cons Biol 9:489–497Google Scholar
  130. Willson MF, Marston BH (2002) Fishing success of gulls at a Southeast Alaska smelt run. J Field Ornith 73:91–96Google Scholar
  131. Willson MF, Gende SM (2004) Seed dispersal by brown bears, Ursus arctos, in southeastern Alaska. Can Field-Nat 118:499–503Google Scholar
  132. Willson MF, Gende SM, Marston BH (1998) Fishes and the forest. BioScience 48:455–462Google Scholar
  133. Wilson WH Jr (1990) Relationship between prey abundance and foraging site selection by semipalmated sandpipers on a Bay of Fundy mudflat. J Field Ornith 61:9–19Google Scholar
  134. Womble JN, Sigler MF (in press) Seasonal availability of abundant, energy-rich prey influences the abundance and diet of a marine predator, the Steller sea lion Eumetopias jubatus. Mar Ecol-Prog SerGoogle Scholar
  135. Womble JN, Willson MF, Sigler MF, Kelly BP, VanBlaricom GR (2005) Distribution of Steller sea lions (Eumetopias jubatus) in relation to spring-spawning prey species in Southeastern Alaska. Mar Ecol-Prog Ser 294:271–282Google Scholar
  136. Wood CC (1987) Predation of juvenile Pacific Salmon by the Common Merganser (Mergus merganser) on eastern Vancouver Island. I: predation during the seaward migration. Can J Fish Aquat Sci 44:941–949Google Scholar
  137. Yang LH (2004) Periodical cicadas as resource pulses in North American forests. Science 306:1565–1567PubMedGoogle Scholar

Copyright information

© Springer Science+Business Media B.V. 2006

Authors and Affiliations

  1. 1.School of Fisheries and Ocean SciencesUniversity of Alaska-FairbanksJuneauUSA
  2. 2.National Park ServiceGlacier Bay Field StationJuneauUSA

Personalised recommendations