Advertisement

Reviews in Fish Biology and Fisheries

, Volume 6, Issue 4, pp 379–416 | Cite as

Reproductive strategies of Atlantic salmon: ecology and evolution

  • Ian A. Fleming
Papers

Abstract

Atlantic salmon (Salmo solar, Salmonidae) show a diversity of life history, behavioural and morphological adaptations for reproduction which have evolved as an outcome of competition to maximize reproductive success. Reproductive traits of females have been shaped principally by natural selection for offspring production and survival, those of males by sexual selection for access to matings. Female Atlantic salmon invest approximately six times more energy in offspring production (i.e. gonads) than males and face an important trade-off between number and size of eggs to produce that will maximize the number of surviving offspring. Timing of breeding and the construction of nests appear adapted to increase offspring survival. The most important determinant of female breeding success is body size because it affords high fecundity, access to breeding territories and decreased probability of nest destruction. Asynchronous female spawning and the male ability to spawn rapidly and repeatedly results in male-biased operational sex ratios that generate intense male competition for mates. This has likely been responsible for the evolution of elaborate male secondary sexual characters associated with fighting and status signalling. Furthermore, it has given rise, through frequency-dependent selection, to two alternative male breeding phenotypes: (1) large, anadromous males; and (2) small, mature male parr. Anadromous males invest heavily in behavioural activity on the spawning grounds, searching and fighting for mates and courting them, with body size being an important determinant of their breeding success. This behavioural activity carries a heavy cost, as anadromous males have significantly reduced survival relative to females. In contrast, mature male parr invest proportionally more in testes for sperm competition and attempt to ‘sneak’ access to matings. While this behaviour also carries costs in terms of subsequent growth and survival, male parr are more likely to breed again, either prior to or following a migration to sea, than anadromous males. While knowledge about the breeding of Atlantic salmon is detailed, we are only beginning to understand the ultimate causes and/or functional significances of their reproductive strategies. Predictive models of the life history variation are developing, focusing on the need for empirical study and testing of life history and reproductive patterns.

Keywords

Atlantic Salmon Breeding Success Sperm Competition Offspring Production Life History Variation 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.

Preview

Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.

References

  1. Aksnes, A., Gjerde, B. and Roald, S.O. (1986) Biological, chemical and organoleptic changes during maturation of farmed Atlantic salmon, Salmo salar. Aquaculture 53, 7–20.Google Scholar
  2. Alm, G. (1959) Connection between maturity, size, and age in fishes. Rep. Inst. Freshwat. Res., Drottningholm 40, 5–145.Google Scholar
  3. Andersson, M. (1994) Sexual selection. Princeton: Princeton University Press. 599 pp.Google Scholar
  4. Anonymous (1980) Investigation into the alleged decline in the migratory salmonids: final report. Wessex Wat. Authority Rep. 40 pp.Google Scholar
  5. Anonymous (1990) Annual Report. Farran Laboratory, Newport, Mayo, Ireland: The Salmon Research Trust of Ireland Incorporated. No. 34. 71 pp.Google Scholar
  6. Armstrong, J.D. and West, C.L. (1994) Relative ventricular weight of wild Atlantic salmon parr in relation to sex, gonad maturation and migratory activity. J. Fish Biol. 44, 453–7.Google Scholar
  7. Armstrong, R.H. (1986) A review of Arctic grayling in Alaska, 1952–1982. Biol. Pap. Univ. Alaska No. 23. 17 pp.Google Scholar
  8. Bagenal, T.B. (1969) Relationship between egg size and fry survival in brown trout, Salmo trutta L. J. Fish Biol. 1, 349–53.Google Scholar
  9. Baglinière, J.L., Maisse, G. and Nihouarn, A. (1990) Migratory and reproductive behaviour of female adult Atlantic salmon, Salmo salar L., in a spawning stream. J. Fish Biol. 36, 511–20.Google Scholar
  10. Baglinière, J.L., Maisse, G. and Nihouarn, A. (1991) Radio-tracking of male adult Atlantic salmon, Salmo salar L., during the last phase of spawning migration in a spawning stream (Brittany, France). Aquat. Living Resour. 4, 161–7.Google Scholar
  11. Barlaup, B.T., Lura, H., Sáegrov, H. and Sundt, R.C. (1994) Inter- and intra-specific variability in female salmonid spawning behaviour. Can. J. Zool. 72, 636–42.Google Scholar
  12. Baum, E.T. and Meister, A.L. (1971) Fecundity of Atlantic salmon (Salmo salar) from two marine rivers. J. Fish. Res. Board Can. 28, 764–7.Google Scholar
  13. Beacham, T.D. and Murray, C.B. (1987) Adaptive variation in body size, age, morphology, egg size, and developmental biology of chum salmon (Oncorhynchus keta) in British Columbia. Can. J. Fish. Aquat. Sci. 44, 244–61.Google Scholar
  14. Beacham, T.D. and Murray, C.B. (1993) Fecundity and egg size variation in North American Pacific salmon (Oncorhynchus). J. Fish Biol. 42, 485–508.Google Scholar
  15. Beacham, T.D., Wither, E.C. and Morley, R.B. (1985) Effect of egg size on incubation and alevin and fry size in chum salmon (Oncorhynchus keta) and coho salmon (O. kisutch). Can. J. Zool. 63, 847–50.Google Scholar
  16. Beacham, T.D., Withler, R.E., Murray, C.B. and Barner, L.W. (1988) Variation in body size, morphology, egg size, and biochemical genetics of pink salmon in British Columbia. Trans. Am. Fish. Soc. 117, 109–26.Google Scholar
  17. Beall, E. and Marty, C. (1983) Reproduction du saumon Atlantique, Salmo salar L. en milieu seminaturel controlé. Bull. Fr. Piscic. 289, 77–93.Google Scholar
  18. Begon, M. and Parker, G.A. (1986) Should egg size and clutch size decrease with age? Oikos 47, 293–302.Google Scholar
  19. Beland, K.F., Jordan, R.M. and Meister, A.L. (1982) Water depth and velocity preferences of spawning Atlantic salmon in Maine rivers. N. Am. J. Fish. Manage. 2, 11–3.Google Scholar
  20. Belding, D.L. (1934a) The spawning habits of the Atlantic salmon. Trans. Am. Fish. Soc. 64, 211–8.Google Scholar
  21. Belding, D.L. (1934b) The cause of high mortality in the Atlantic salmon after spawning. Trans. Am. Fish. Soc. 64, 219–24.Google Scholar
  22. Berg, O.K. (1985) The formation of non-anadromous populations of Atlantic salmon, Salmo salar L., in Europe. J. Fish Biol. 27 805–15.Google Scholar
  23. van den Berghe, E.P. and Gross, M.R. (1989) Natural selection resulting from female breeding competition in a Pacific salmon (coho: Oncorhynchus kisutch). Evolution 43, 125–40.Google Scholar
  24. van den Berghe, E.P., Wernerus, F. and Warner, R.R. (1989) Female choice and mating cost of peripheral males. Anim. Behav. 38, 875–84.Google Scholar
  25. Berglund, I. (1992) Growth and early sexual maturation in Baltic salmon (Salmo salar) parr. Can. J. Zool 70, 205–11.Google Scholar
  26. Berglund, I., Schmitz, M. and Lundqvist, H. (1992) Seawater adaptability in Baltic salmon (Salmo salar): a bimodal smoltification pattern in previously mature males. Can. J. Fish. Aquat. Sci. 49, 1097–106.Google Scholar
  27. Billard, R. (1974) La production spermatogénétique de la truite arc-en-ciel au cours du premier cycle reproducteur. Bull. Fr. Piscic. 253, 139–49.Google Scholar
  28. Bilton, H.T. (1970) Comparison of the fecundity of sockeye salmon (Oncorhynchus nerka) in the Skeena River catch with the fecundity of those in the escapement. Fish. Res. Board Can. MS Rep. Ser. No. 1096. 8 pp.Google Scholar
  29. Bilton, H.T. and Smith, H.D. (1973) Relationship between egg size and fish size in pink salmon (Oncorhynchus gorbuscha) from the lower Babine River in 1966 and 1967. Fish. Res. Board Can. MS Rep. Ser. No. 1235. 5 pp.Google Scholar
  30. Birkhead, T.R. (1991) Sperm depletion in the Bengalese finch Lochura striata. Behav. Ecol. 2, 267–75.Google Scholar
  31. Blair, G.R., Rogers, D.E. and Quinn, T.P. (1993) Variation in life history characteristics and morphology of sockeye salmon in the Kvichak River system, Bristol Bay, Alaska. Trans. Am. Fish. Soc. 122, 550–9.Google Scholar
  32. Bohlin, T., Dellefors, C. and Faremo, U. (1986) Early sexual maturation of male sea trout and salmon — an evolutionary model and some practical implications. Rep. Inst. Freshwat. Res., Drottningholm 63, 17–25.Google Scholar
  33. Bohlin, T., Dellefors, C. and Faremo, U. (1990) Large or small at maturity — theories on the choice of alternative male strategies in anadromous salmonids. Ann. Zool. Fennici 27, 139–47.Google Scholar
  34. Boyce, N.P.J. (1974) Biology of Eubothrium salvelini (Cestoda: Pseudophyllidea), a parasite of juvenile sockeye salmon (Oncorhynchus nerka) of Babine Lake, British Columbia. J. Fish. Res. Board Can. 31, 1735–42.Google Scholar
  35. Brännäs, E. (1995) First access to territorial space and exposure to strong predation pressure: a conflict in early emerging Atlantic salmon (Salmo salar L.) fry. Evol. Ecol. 9, 411–20.Google Scholar
  36. Brännäs, E., Brännäs, K. and Eriksson, L.-O. (1985) Egg characteristics and hatchery survival in a Baltic salmon, Salmo salar L., population. Rep. Inst. Freshwat. Res., Drottningholm 62, 4–11.Google Scholar
  37. Brannon, E.L. (1987) Mechanisms stabilizing salmonid fry emergence timing. Can. Spec. Publ. Fish. Aquat. Sci. 96, 120–4.Google Scholar
  38. Brown, J.S. and Parman, A. (1993) Consequences of size-selective harvesting as an evolutionary game. In Stokes, T.K., McGlade, J.M. and Law, R., eds. The Exploitation of Evolving Resources. Berlin: Springer-Verlag, pp. 248–61.Google Scholar
  39. Brown, M.E. (1946) The growth of brown trout (Salmo trutta Linn.). I. Factors influencing the growth of trout fry. J. Exp. Biol. 22, 118–29.Google Scholar
  40. Calderwood, W. (1900) Appendix II. Note by the Inspector of Salmon Fisheries on the range of the salmon spawning season in Scotland. Ann. Rep. Fish. Board Scotland 19, 53–6.Google Scholar
  41. Campbell, J.S. (1977) Spawning characteristics of brown trout and sea trout, Salmo trutta L., in Kirk Burn, River Tweed, Scotland. J. Fish Biol. 11, 217–29.Google Scholar
  42. Carss, D.N., Kruuk, H. and Conroy, J.W.H. (1990) Predation on adult Atlantic salmon, Salmo salar L., by otters Lutra lutra (L.), within the River Dee system, Aberdeenshire, Scotland. J. Fish Biol. 37, 935–44.Google Scholar
  43. Chadwick, E.M.P., Porter, T.R. and Downton, P. (1978) Analysis of growth of Atlantic salmon (Salmo salar) in a small Newfoundland river. J. Fish. Res. Board Can. 35, 60–8.Google Scholar
  44. Chapman, (1962).Google Scholar
  45. Chapman, D.W. (1988) Critical review of variables used to define effects of fines in redds of large salmonids. Trans. Am. Fish. Soc. 117, 1–21.Google Scholar
  46. Chebanov, N.A., Varnavskaya, N.V. and Varnavskiy, V.S. (1984) Effectiveness of spawning of male sockeye salmon, Oncorhynchus nerka (Salmonidae), of differing hierarchical rank by means of genetic-biochemical markers. J. Ichthyol. 23(5), 51–5.Google Scholar
  47. Clutton-Brock, T.H. and Parker, G.A. (1995) Punishment in animal societies. Nature 373, 209–16.Google Scholar
  48. Congdon, J.D. and Gibbons, J.W. (1987) Morphological constraint on egg size: a challenge of optimal egg size theory? Proc. Natn. Acad. Sci., USA 84, 4145–7.Google Scholar
  49. Cooper, A.C. (1965) The effect of transported stream sediments on survival of sockeye and pink salmon eggs and alevin. Int. Pac. Salmon Fish. Comm. Bull. 18, 1–71.Google Scholar
  50. Crisp, D.T. (1981) A desk study of the relationship between temperature and hatching time for the eggs of five species of salmonid fishes. Freshwat. Biol. 11, 361–8.Google Scholar
  51. Crisp, D.T. and Carling, P.A. (1989) Observations on siting, dimensions and structure of salmonid redds. J. Fish Biol. 34, 119–34.Google Scholar
  52. Cuinal, R. (1988) Atlantic salmon in an extensive French river-system; the Loire-Allier. In Mills, D. and Piggins, D., eds. Atlantic Salmon: Planning for the Future. London: Croom Helm, pp. 389–9.Google Scholar
  53. Dalley, E.L., Andrews, C.W. and Green, J.M. (1983) Precocious male Atlantic salmon parr (Salmo salar) in insular Newfoundland. Can. J. Fish. Aquat. Sci. 40, 647–52.Google Scholar
  54. Darwin, C. (1871) The Descent of Man, and Selection in Relation to Sex, 1st edn. London: J. Murray. 611 pp.Google Scholar
  55. Daye, P.G. and Glebe, B.D. (1984) Fertilization success and sperm motility of Atlantic salmon (Salmo salar L.) in acidified water. Aquaculture 43, 307–12.Google Scholar
  56. Davidson, F.A. (1935) The development of the secondary sexual characters of the pink salmon (Oncorhynchus gorbuscha). J. Morph. 57, 169–83.Google Scholar
  57. von der Decken, A. (1992) Physiological changes in skeletal muscle by maturation-spawning of non-migrating female Atlantic salmon, Salmo salar. Comp. Biochem. Physiol. 101B, 299–301.Google Scholar
  58. Dirin, D.K. (1983) Biological considerations relating to minimum commercial size for the lake salmon, Salmo salar, morpha Sebago (Salmonidae). J. Ichthyol. 23, 37–52.Google Scholar
  59. Ducharme, L.J.A. (1969) Atlantic salmon returning for their fifth and sixth consecutive spawning trips. J. Fish. Res. Board Can. 26, 1661–4.Google Scholar
  60. Elson, P.F. (1973) Genetic polymorphism in Northwest Miramichi salmon, in relation to season of river ascent and age at maturation and its implications for management of stocks. Int. Comm. Northw. Atlantic Fish., Res. Doc. 73/76. 6 pp.Google Scholar
  61. Endler, J.A. (1986) Natural Selection in the Wild. Princeton: Princeton University Press. 336 pp.Google Scholar
  62. Eriksson, T., Eriksson, L.-O. and Lundqvist, H. (1987) Adaptive flexibility in life history tactics of mature male Baltic salmon parr in relation to body size and environment. Am. Fish. Soc. Symp. 1, 236–43.Google Scholar
  63. Fabricius, E. and Gustafson, K.-J. (1953) Further aquarium observations on the spawning behaviour of the char, Salvelinus alpinus L. Rep. Inst. Freshwat. Res. Drottningholm 35, 58–104.Google Scholar
  64. Ferguson, A., Taggart, J.B., Prodöhl, P.A., McMeel, O., Thompson, C., Stone, C., McGinnity, P. and Hynes, R.A. (1995) The application of molecular markers to the study and conservation of fish populations, with special reference to Salmo. J. Fish Biol. 47(Suppl. A), 103–26.Google Scholar
  65. Fleming, I.A. and Gross, M.R. (1989) Evolution of adult female life history and morphology in a Pacific salmon (coho: Oncorhynchus kisutch). Evolution 43, 141–57.Google Scholar
  66. Fleming, I.A. and Gross, M.R. (1990) Latitudinal clines: a trade-off between egg number and size in Pacific salmon. Ecology 71, 1–11.Google Scholar
  67. Fleming, I.A. and Gross, M.R. (1992) Reproductive behavior of hatchery and wild coho salmon (Oncorhynchus kisutch): does it differ? Aquaculture 103, 101–21.Google Scholar
  68. Fleming, I.A. and Gross, M.R. (1993) Breeding success of hatchery and wild coho salmon (Oncorhynchus kisutch) in competition. Ecol. Appl. 3, 230–45.Google Scholar
  69. Fleming, I.A. and Gross, M.R. (1994) Breeding competition in a Pacific salmon (coho: Oncorhynchus kisutch): measures of natural and sexual selection. Evolution 48, 637–57.Google Scholar
  70. Fleming, I.A. and Ng, S. (1987) Evaluation of techniques for fixing, preserving, and measuring salmon eggs. Can. J. Fish. Aquat. Sci. 44, 1957–62.Google Scholar
  71. Fleming, I.A., Jonsson, B. and Gross, M.R. (1994) Phenotypic divergence of sea-ranched, farmed and wild salmon. Can J. Fish. Aquat. Sci. 51, 2808–24.Google Scholar
  72. Fleming, I.A., Jonsson, B., Gross, M.R. and Lamberg, A. An experimental study of the reproductive behaviour and success of farmed and wild Atlantic salmon (Salmo salar). J. Appl. Ecol. (in press).Google Scholar
  73. Foote, C.J. (1988) Male mate choice dependent on male size in salmon. Behaviour 106, 63–80.Google Scholar
  74. Foote, C.J. (1989) Female mate preference in Pacific salmon. Anim. Behav. 38, 721–3.Google Scholar
  75. Foote, C.J. (1990) An experimental comparison of male and female spawning territoriality in a Pacific salmon. Behaviour 115, 283–314.Google Scholar
  76. Foote, C.J. and Larkin, P.A. (1988) The role of male choice in the assortative mating of anadromous and non-anadromous sockeye salmon (Oncorhynchus nerka). Behaviour 106, 43–62.Google Scholar
  77. Forbes, L.S. and Peterman, R.M. (1994) Simple size-structured models of recruitment and harvest in Pacific salmon (Oncorhynchus spp.). Can. J. Fish. Aquat. Sci. 51, 603–16.Google Scholar
  78. Forseth, T. and Jonsson, B. (1994) The growth and food ration of piscivorous brown trout (Salmo trutta). Funct. Ecol. 8, 171–7.Google Scholar
  79. Fowler, L.G. (1972) Growth and mortality of fingerling chinook salmon as affected by egg size. Progve Fish-Cult. 34, 66–9.Google Scholar
  80. Gage, M.J.G., Stockley, P. and Parker, G.A. (1995) Effects of alternative male mating strategies on characteristics of sperm production in the Atlantic salmon (Salmo salar): theoretical and empirical investigations. Phil. Trans. R. Soc. Lond. 350B, 391–9.Google Scholar
  81. Galkina, Z.I. (1970) Dependence of egg size on the size and age of female salmon (Salmo salar (L.)) and rainbow trout (Salmo irideus (Gib.)). J. Ichthyol. 10, 625–33.Google Scholar
  82. Garcia de Leániz, C., Hawkins, A.D., Hay, D.W. and Martinez, J.J. (1987) The Atlantic salmon of Spain. Pitlochry: The Atlantic Salmon Trust. 28 pp.Google Scholar
  83. Gjerde, B. (1984) Variation in semen production of farmed Atlantic salmon and rainbow trout. Aquaculture 40, 109–14.Google Scholar
  84. Glebe, B.D. and Saunders, R.L. (1986) Genetic factors in sexual maturity of cultured Atlantic salmon (Salmo salar) parr and adults reared in sea cages. Can. Spec. Publ. Fish. Aquat. Sci. 89, 24–9.Google Scholar
  85. Glebe, B.D., Appy, T.D. and Saunders, R.L. (1979) Variation in Atlantic salmon (Salmo salar) reproductive traits and their implications in breeding programs. Int. Counc. Explor. Sea C.M. 1979/M:23. 11 pp.Google Scholar
  86. Greenstreet, S.P.R., Morgan, R.I.G. and Thorpe, J.E. (1992) Effect of stripping on the length, condition and gonadal state of mature male Atlantic salmon, Salmo salar L., parr during autumn and winter. J. Fish Biol. 41, 191–200.Google Scholar
  87. Groot, C. and Margolis, L. (eds) (1991) Pacific Salmon Life Histories. Vancouver: University of British Columbia Press. 564 pp.Google Scholar
  88. Gross, M.R. (1984) Sunfish, salmon, and the evolution of alternative reproductive strategies and tactics in fishes. In Potts, G. and Wootton, R.J., eds. Fish Reproduction: Strategies and Tactics. London: Academic Press, pp. 55–75.Google Scholar
  89. Gross, M.R. (1985) Alternative breeding strategies in male salmon. Nature 313, 47–8.Google Scholar
  90. Gross, M.R. (1991) Salmon breeding behavior and life history evolution in changing environments. Ecology 72, 1180–6.Google Scholar
  91. Gross, M.R. (1996) Alternative reproductive strategies and tactics: diversity within sexes. Trends Ecol. Evol. 11, 92–8.Google Scholar
  92. Gustafson-Greenwood, K.I. and Moring, J.R. (1991) Gravel compaction and permeabilities in redds of Atlantic salmon, Salmo salar L. Aquacult. Fish. Manage. 22, 537–40.Google Scholar
  93. Hansen, L.P. (1980) Tagging and release of Atlantic salmon (Salmo salar) smolts in the river Glomma. Fauna 33, 89–97. (In Norwegian with English abstract).Google Scholar
  94. Hansen, L.P., H»stein, T., Nævdal, G., Saunders, R.L. and Thorpe, J.E. (eds) (1991) Interactions between cultured and wild Atlantic salmon. Aquaculture 98, 1–324.Google Scholar
  95. Hanson, A.J. and Smith, H.D. (1967) Mate selection in a population of sockeye salmon (Oncorhynchus nerka) of mixed age groups. J. Fish. Res. Board Can. 24, 1955–77.Google Scholar
  96. Harper, D.G.C. (1991) Communication. In Krebs, J.R. and Davies, N.B., eds. Behavioural Ecology: an Evolutionary Approach. Oxford: Blackwell, pp. 374–97.Google Scholar
  97. Hasler, A.D. and Scholz, A.T. (1983) Olfactory Imprinting and Homing in Salmon. Berlin: Springer-Verlag. 134 pp.Google Scholar
  98. Hawkins, A.D. (1989) Factors affecting the timing of entry and upstream movement of Atlantic salmon in the Aberdeenshire Dee. In Brannon, E. and Jonsson, B., eds. Proceedings of the Salmonid Migration and Distribution Symposium, Trondheim, June 1987. Seattle: School of Fisheries, University of Washington, pp. 101–5.Google Scholar
  99. Hawkins, A.D. and Smith, G.W. (1986) Radio-tracking observations on Atlantic salmon ascending the Aberdeenshire Dee. Scott. Fish. Res. Rep. No. 36. 24 pp.Google Scholar
  100. Hay, D.W. (1984) The relationship between redd counts and the number of spawning salmon in the Girnock Burn (Scotland). Int. Counc. Explor. Sea CM 1984/M:22.Google Scholar
  101. Healey, M.C. (1986) Optimum size and age at maturity in Pacific salmon and effects of size-selective fisheries. Can. Spec. Publ. Fish. Aquat. Sci. 89, 39–52.Google Scholar
  102. Healey, M.C. and Heard, W.R. (1984) Inter- and intra-population variation in the fecundity of chinook salmon (Oncorhynchus tshawytscha) and its relevance to life history theory. Can. J. Fish. Aquat. Sci. 41, 476–83.Google Scholar
  103. Heggberget, T.G. (1988) Timing of spawning in Norwegian Atlantic salmon (Salmo salar). Can. J. Fish. Aquat. Sci. 45, 845–9.Google Scholar
  104. Heggberget, T.G., Lund, R.A., Ryman, N. and St»hl, G. (1986) Growth and genetic variation of Atlantic salmon (Salmo salar) from different section of the River Alta, North Norway. Can. J. Fish. Aquat. Sci. 43, 1828–35.Google Scholar
  105. Heggberget, T.G., Hansen, L.P. and Næsje, T.F. (1988a) Within-river spawning migration of Atlantic salmon (Salmo salar). Can. J. Fish. Aquat. Sci. 10, 1691–8.Google Scholar
  106. Heggberget, T.G., Haukebø, T., Mork, J. and Stahl, G. (1988b) Temporal and spatial segregation of spawning in sympatric populations of Atlantic salmon, Salmo salar L., and brown trout, Salmo trutta L. J. Fish Biol. 33, 347–56.Google Scholar
  107. Hindar, K. (1994) Alternative life histories and genetic conservation. In Loeschcke, V, Tomiuk, J. and Jain, S.K., eds. Conservation Genetics. Basel: Birkhäuser Verlag, pp. 323–36.Google Scholar
  108. Hindar, K., Ryman, N. and Utter, F. (1991) Genetic effects of cultured fish on natural fish populations. Can. J. Fish. Aquat. Sci. 48, 945–57.Google Scholar
  109. Huntingford, F.A., Metcalfe, N.B., Thorpe, J.E., Graham, W.D. and Adams, C.E. (1990) Social dominance and body size in Atlantic salmon parr, Salmo salar. J. Fish. Biol. 36, 877–81.Google Scholar
  110. Hutchings, J.A. (1986) Lakeward migrations by juvenile Atlantic salmon, Salmo Salar. Can. J. Fish. Aquat. Sci. 43, 732–41.Google Scholar
  111. Hutchings, J.A. (1991) Fitness consequences of variation in egg size and food abundance in brook trout, Salvelinus fontinalis. Evolution 45, 1162–8.Google Scholar
  112. Hutchings, J.A. (1993) Adaptive life histories effected by age-specific survival and growth rate. Ecology 74, 673–84.Google Scholar
  113. Hutchings, J.A. and Myers, R.A. (1985) Mating between anadromous and nonanadromous Atlantic salmon, Salmo salar. Can. J. Zool. 63, 2219–21.Google Scholar
  114. Hutchings, J.A. and Myers, R.A. (1987) Escalation of an asymmetric contest: mortality resulting from mate competition in Atlantic salmon, Salmo salar. Can. J. Zool. 65, 766–8.Google Scholar
  115. Hutchings, J.A. and Myers, R.A. (1988) Mating success of alternative maturation phenotypes in male Atlantic salmon, Salmo salar. Oecologia 75, 169–74.Google Scholar
  116. Hutchings, J.A. and Myers, R.A. (1994) The evolution of alternative mating strategies in variable environments. Evol. Ecol. 8, 256–68.Google Scholar
  117. Järvi, T. (1990) The effects of male dominance, secondary sexual characteristics and female mate choice on the mating success of male Atlantic salmon Salmo salar. Ethology 84, 123–32.Google Scholar
  118. Jessop, B.M. (1986) Atlantic salmon (Salmo salar) of the Big Salmon River, New Brunswick. Can. Tech. Rep. Fish. Aquat. Sci. No. 1415, 1–50.Google Scholar
  119. Johnstone, A.D.F., Lucas, M.C., Boylan, P. and Carter, T.J. (1992) Telemetry of tail-beat frequency of Atlantic salmon (Salmo salar L.) during spawning. In Priede, I.G. and Swift, S.M., eds. Wildlife Telemetry: Remote Monitoring and Tracking of Animals. New York: Ellis Horwood, pp. 456–65.Google Scholar
  120. Jones, J.W. (1959) The Salmon. London: Collins. 192 pp.Google Scholar
  121. Jones, J.W. and Ball, J.N. (1954) The spawning behaviour of brown trout and salmon. Brit. J. Anim. Behav. 2, 103–14.Google Scholar
  122. Jones, J.W. and King, G.M. (1949) Experimental observations on the spawning behaviour of the Atlantic salmon (Salmo salar Linn.). Proc. Zool. Soc. Lond. 119, 33–48.Google Scholar
  123. Jones, J.W. and King, G.M. (1950) Further experimental observations on the spawning behaviour of the Atlantic salmon (Salmo salar Linn.). Proc. Zool. Soc. Lond. 120, 317–23.Google Scholar
  124. Jones, J.W. and King, G.M. (1952) The spawning of the male salmon parr (Salmo salar Linn. Juv.). Proc. Zool. Soc. Lond. 122, 615–9.Google Scholar
  125. Jonsson, B. (1985).Google Scholar
  126. Jonsson, B. and Fleming, I.A. (1993) Enhancement of wild salmon populations. In Sundnes, G., ed. Human Impact on Self-recruiting Populations. Trondheim: Tapir Press, pp. 209–42.Google Scholar
  127. Jonsson, B. and Hindar, K. (1982) Reproductive strategy of dwarf and normal Arctic charr (Salvelinus alpinus) from Vangsvatnet Lake, western Norway. Can. J. Fish. Aquat. Sci. 39, 1404–13.Google Scholar
  128. Jonsson, B., Jonsson, N. and Hansen, L.P. (1990) Does juvenile experience affect migration and spawning of adult Atlantic salmon. Behav. Ecol. Sociobiol. 26, 225–30.Google Scholar
  129. Jonsson, B., Jonsson, N., and Hansen, L.P. (1991) Differences in life history and migratory behaviour between wild and hatchery-reared Atlantic salmon in nature. Aquaculture 98, 69–78.Google Scholar
  130. Jonsson, N., Jonsson, B. and Hansen, L.P. (1990) Partial segregation in the timing of migration of Atlantic salmon of different ages. Anim. Behav. 40, 313–21.Google Scholar
  131. Jonsson, N., Hansen, L.P. and Jonsson, B. (1991a) Variation in age, size and repeat spawning of adult Atlantic salmon in relation to river discharge. J. Anim. Ecol. 60, 937–47.Google Scholar
  132. Jonsson, N., Jonsson, B. and Hansen, L.P. (1991b) Energetic cost of spawning in male and female Atlantic salmon (Salmo salar L.). J. Fish Biol. 39, 739–44.Google Scholar
  133. Jonsson, N., Jonsson, B. and Fleming, I.A. (1996) Phenotypically plastic response of egg production to early growth in Atlantic salmon. Funct. Ecol. 10, 89–96.Google Scholar
  134. Jonsson, N., Jonsson, B. and Hansen, L.P. (1997) Changes in proximate composition and estimates of energetic costs during upstream migration and spawning in Atlantic salmon, Salmo salar. J. Anim. Ecol. (in press).Google Scholar
  135. Jordan, W.C. and Youngson, A.F. (1992) The use of genetic marking to assess the reproductive success of mature male Atlantic salmon parr (Salmo salar, L.) under natural spawning conditions. J. Fish Biol. 41, 613–8.Google Scholar
  136. Judson, O.P. (1994) The rise of the individual-based model in ecology. Trends Ecol. Evol. 9, 9–14.Google Scholar
  137. Kadri, S., Metcalfe, N.B., Huntingford, F.A. and Thorpe, J.E. (1995) What controls the onset of anorexia in maturing adult female Atlantic salmon? Funct. Ecol. 9, 790–7.Google Scholar
  138. Kazakov, R.V. (1981a) The effect of the size of Atlantic salmon, Salmo salar L., eggs on embryos and alevins. J. Fish Biol. 19, 353–60.Google Scholar
  139. Kazakov, R.V. (1981b) Peculiarities of sperm production by anadromous and parr Atlantic salmon (Salmo salar L.) and fish cultural characteristics of such sperm. J. Fish Biol. 18, 1–8.Google Scholar
  140. Keenleyside, M.H.A. and Dupuis, H.M.C. (1988a) Comparison of digging behavior by female pink salmon (Oncorhynchus gorbuscha) before and after spawning. Copeia 1988, 1092–5.Google Scholar
  141. Keenleyside, M.H.A. and Dupuis, H.M.C. (1988b) Courtship and spawning competition in pink salmon (Oncorhynchus gorbuscha). Can. J. Zool. 66, 262–5.Google Scholar
  142. Kitano, S. (1996) Size-related factors causing individual variation in seasonal reproductive success of fluvial male Dolly Varden (Salvelinus malma). Ecol. Freshw. Fish 5, 59–67.Google Scholar
  143. L'Abée-Lund, J.H. (1991) Variation within and between rivers in adult size and sea age at maturity of anadromous brown trout, Salmo trutta. Can. J. Fish. Aquat. Sci. 48, 1015–21.Google Scholar
  144. L'Abée-Lund, J.H. and Hindar, K. (1990) Interpopulation variation in reproductive traits of anadromous female brown trout, Salmo trutta L. J. Fish Biol. 37, 755–63.Google Scholar
  145. Larsson, P.-O. and Pickova, J. (1978) Egg size of salmon (Salmo salar L.) in correlation to female age and weight in three river stocks. Laxforskningsinstitutt Meddelande 2, 1–6.Google Scholar
  146. Laughton, R. and Smith, G.W. (1992) The relationship between date of river entry and the estimated spawning position of adult Atlantic salmon (Salmo salar L.) in two major Scottish east coast rivers. In Priede, I.G. and Swift, S.M., eds. Wildlife Telemetry: Remote Monitoring and Tracking of Animals. New York: Ellis Horwood, pp. 423–33.Google Scholar
  147. Lee, R.L.G. and Power, G. (1976) Atlantic salmon (Salmo salar) of the Leaf River, Ungava Bay. J. Fish. Res. Board Can. 33, 2616–21.Google Scholar
  148. Leggett, W.C. and Power, G. (1969) Differences between two populations of landlocked Atlantic salmon (Salmo salar) in Newfoundland. J. Fish. Res. Board Can. 26, 1585–96.Google Scholar
  149. Leider, S.A., Hulett, P.L., Loch, J.J. and Chilcote, M.W. (1990) Electrophoretic comparison of the reproductive success of naturally spawning transplanted and wild steelhead trout through the returning adult stage. Aquaculture 88, 239–52.Google Scholar
  150. Leonardsson, K. and Lundberg, P. (1986) The choice of reproductive tactics as a mixed evolutionary stable strategy: the case of male Atlantic salmon (Salmo salar L.). Rep. Inst. Freshwat. Res. Drottningholm 63, 69–76.Google Scholar
  151. Lloyd, D.G. (1987) Selection of offspring size at independence and other size-versus-number strategies. Am. Nat. 129, 800–17.Google Scholar
  152. Lura, H. and Sægrov, H. (1991) Documentation of successful spawning of escaped farmed female Atlantic salmon, Salmo salar, in Norwegian rivers. Aquaculture 98, 151–9.Google Scholar
  153. Lura, H., Barlaup, B.T. and Sægrov, H. (1993) Spawning behaviour of a farmed escaped female Atlantic salmon (Salmo salar). J. Fish Biol. 42, 311–3.Google Scholar
  154. McCart, P. (1969) Digging behaviour of Oncorhynchus nerka spawning in streams at Babine Lake, British Columbia. In Northcote, T.D., ed. Symposium on Salmon and Trout in Streams (H.R. MacMillan Lectures in Fisheries). Vancouver: University of British Columbia, pp. 39–51.Google Scholar
  155. MacCrimmon, H.R. and Gots, B.L. (1979) World distribution of Atlantic salmon, Salmo salar. Can. J. Fish. Aquat. Sci. 36, 422–57.Google Scholar
  156. McNeil, W.J. (1969) Survival of pink salmon eggs and alevins. In Northcote, T.G., ed. Symposium on Salmon and Trout in Streams (H.R. MacMillan Lectures in Fisheries). Vancouver: University of British Columbia, pp. 101–19.Google Scholar
  157. Maekawa, K. and Onozato, H. (1986) Reproductive tactics and fertilization success of mature male Miyabe charr, Salvelinus malma miyabei. Env. Biol. Fishes 15, 119–29.Google Scholar
  158. Maekawa, K., Nakano, S. and Yamamoto, S. (1994) Spawning behaviour and size-assortative mating of Japanese charr in an artificial lake-inlet stream system. Env. Biol. Fishes 39, 109–17.Google Scholar
  159. Maynard Smith, J. (1982) Evolution and the Theory of Games. Cambridge: Cambridge University Press. 224 pp.Google Scholar
  160. Metcalfe, N.B. and Thorpe, J.E. (1992) Early predictors of life-history events: the link between first feeding date, dominance and seaward migration in Atlantic salmon, Salmo salar L. J. Fish Biol. 41(Suppl. B), 93–9.Google Scholar
  161. Metcalfe, N.B., Taylor, A.C. and Thorpe, J.E. (1995) Metabolic rate, social status and life-history strategies in Atlantic salmon. Anim. Behav. 49, 431–6.Google Scholar
  162. Mills, D. (1989) Ecology and Management of Atlantic Salmon. London: Chapman and Hall. 351 pp.Google Scholar
  163. Moore, A. and Scott, A.P. (1991) Testosterone is a potent odorant in precocious male Atlantic salmon (Salmo salar L.) parr. Phil. Trans. R. Soc. Lond. 332B, 241–4.Google Scholar
  164. Moore, A. and Scott, A.P. (1992) 17α,20β-dihydroxy-4-pregnen-3-one 20-sulphate is a potent odorant in precocious male Atlantic salmon (Salmo salar L.) parr which have been pre-exposed to the urine of ovulating females. Proc. R. Soc. Lond. 249B, 205–9.Google Scholar
  165. Munkittrick, K.R. and Moccia, R.D. (1987) Seasonal changes in the quality of rainbow trout (Salmo gairdneri) semen: effect of a delay in stripping on spermatocrit, motility, volume and seminal plasma constituents. Aquaculture 64, 147–56.Google Scholar
  166. Myers, R.A. (1984) Demographic consequences of precocious maturation of Atlantic salmon (Salmo salar). Can. J. Fish. Aquat. Sci. 41, 1349–53.Google Scholar
  167. Myers, R.A. (1986) Game theory and the evolution of Atlantic salmon (Salmo salar) age at maturation. Can. Spec. Publ. Fish. Aquat. Sci. 89, 53–61.Google Scholar
  168. Myers, R.A. and Hutchings, J.A. (1986) Selection against parr maturation in Atlantic salmon. Aquaculture 53, 313–20.Google Scholar
  169. Myers, R.A. and Hutchings, J.A. (1987a) A spurious correlation in an interpopulation comparison of Atlantic salmon life histories. Ecology 68, 1839–43.Google Scholar
  170. Myers, R.A. and Hutchings, J.A. (1987b) Mating of anadromous Atlantic salmon, Salmo salar L., with mature male parr. J. Fish Biol. 31, 143–6.Google Scholar
  171. Myers, R.A., Hutchings, J.A. and Gibson, R.J. (1986) Variation in male parr maturation within and among populations of Atlantic salmon, Salmo salar. Can. J. Fish. Aquat. Sci. 43, 1242–8.Google Scholar
  172. Næsje, T.F., Hansen, L.P. and Järvi, T. (1988) Sexual dimorphism in the adipose fin of Atlantic salmon, Salmo salar L. J. Fish Biol 33, 955–6.Google Scholar
  173. Nævdal, G., Holm, M., Møller, D. and østhus, O.D. (1976) Variation in growth rate and age at sexual maturity in Atlantic salmon. Int. Counc. Explor. Sea CM 1976/E:40. 10 pp.Google Scholar
  174. Nakatsuru, K. and Kramer, D.L. (1982) Is sperm cheap? Limited male fertility and female choice in the lemon tetra (Pisces, Characidae). Science 216, 753–5.Google Scholar
  175. Nielsen, J.L. (ed.) (1995) Evolution and the Aquatic Ecosystem: Defining Unique Units in Population Conservation (American Fisheries Society Symposium No. 17). Bethesda, MD: American Fisheries Society. 435 pp.Google Scholar
  176. Nordqvist, O. (1924) Times of entering of the Atlantic salmon (Salmo salar L.) in the rivers. Rapp. P.-v. Réun. Cons. Int. Explor. Mer 33, 1–58.Google Scholar
  177. økland, F., Heggberget, T.G. and Jonsson, B. (1995) Migratory behaviour of wild and farmed Atlantic salmon (Salmo salar) during spawning. J. Fish Biol. 46, 1–7.Google Scholar
  178. Ottaway, E.M., Carling, P.A., Clarke, A. and Reader, N.A. (1981) Observations on the structure of brown trout, Salmo trutta Linnaeus, redds. J. Fish Biol. 19, 593–607.Google Scholar
  179. Parker, G.A. (1984) Evolutionary stable strategies. In Krebs, J.R. and Davies, N.B., eds. Behavioural Ecology: An Evolutionary Approach. Sunderland: Sinauer, pp. 30–61.Google Scholar
  180. Parker, G.A. (1993) Sperm competition games: sperm size and number under adult control. Proc. R. Soc. Lond. 253B, 245–54.Google Scholar
  181. Parker, R.R. (1971) Size selective predation among juvenile salmonid fishes in a British Columbia inlet. J. Fish. Res. Board Can. 28, 1503–10.Google Scholar
  182. Patten, B.G. (1977) Body size and learned avoidance as factors affecting predation on coho salmon, Oncorhynchus kisutch, fry by torrent sculpin, Cottus rhotheus. Fish. Bull. US 75, 457–9.Google Scholar
  183. Perrin, N. (1988) Why are offspring born larger when it is colder? Phenotypic plasticity for offspring size in the cladoceran Simocephalus vetulus (Müller). Funct. Ecol. 2, 283–8.Google Scholar
  184. Perrin, N. (1989) Population density and offspring size in the cladoceran Simocephalus vetulus (Müller). Funct. Ecol. 3, 29–36.Google Scholar
  185. Peterson, N.P. and Quinn, T.P. (1996) Persistence of egg pocket architecture in redds of chum salmon, Oncorhynchus keta. Env. Biol. Fishes 46, 243–53.Google Scholar
  186. Peterson, R.H. (1978) Physical characteristics of Atlantic salmon spawning gravel in some New Brunswick streams. Fish. Mar. Serv. Tech. Rep. No. 785, 1–28.Google Scholar
  187. Policansky, D. (1993) Fishing as a cause of evolution in fishes. In Stokes, T.K., McGlade, J.M. and Law, R., eds. The Exploitation of Evolving Resources. Berlin: Springer-Verlag, pp. 2–18.Google Scholar
  188. Pope, J.A., Mills, D.H. and Shearer, W.M. (1961) The fecundity of Atlantic salmon (Salmo salar L.). Freshwat. Salm. Fish. Res., Scotland 26, 1–12.Google Scholar
  189. Power, G. (1958) The evolution of freshwater races of the Atlantic salmon (Salmo salar L.). Arctic 11, 86–92.Google Scholar
  190. Power, G. (1969) The salmon of Ungava Bay. Arctic Inst. N. Am. Tech. Pap. No. 22, 1–72.Google Scholar
  191. Power, G. (1981) Stock characteristics and catches of Atlantic salmon (Salmo salar) in Quebec, and Newfoundland and Labrador in relation to environmental variables. Can J. Fish. Aquat. Sci. 38, 1601–11.Google Scholar
  192. Power, G., Power, M.V., Dumas, R. and Gordon, A. (1987) Marine migrations of Atlantic salmon from rivers in Ungava Bay, Quebec. Am. Fish. Soc. Symp. 1, 364–76.Google Scholar
  193. Prouzet, P. and Gaignon, J.L. (1982) Fécondité des saumons Atlantiques adultes capturés sur le bassin versant de L'Elorn (rivière de Bretagne Nord) et caractéristiques de leurs pontes. Bull. Fr. Piscic. 283, 233–43.Google Scholar
  194. Prouzet, P., LeBail, P.Y. and Heydorff, M. (1984) Sex ratio and potential fecundity of Atlantic salmon (Salmo salar L.) caught by anglers on the Elorn River (northern Brittany, France) during 1979 and 1980. Fish. Manage. 15, 123–30.Google Scholar
  195. Quinn, T.P. and Bloomberg, S. (1992) Fecundity of chinook salmon (Oncorhynchus tshawytscha) from the Waitaki and Rakaia Rivers, New Zealand. N.Z. J. Mar. Freshwat. Res. 26, 429–34.Google Scholar
  196. Quinn, T.P. and Foote, C.J. (1994) The effects of body size and sexual dimorphism on the reproductive behaviour of sockeye salmon, Oncorhynchus nerka. Anim. Behav. 48, 751–61.Google Scholar
  197. Quinn, T.P., Hendry, A.P. and Wetzel, L.A. (1995) The influence of life history trade-offs and the size of incubation gravels on egg size variation in sockeye salmon (Oncorhynchus nerka). Oikos 74, 425–38.Google Scholar
  198. Quinn, T.P., Adkison, M.D. and Ward, M.B. (1996) Behavioral tactics of male sockeye salmon (Oncorhynchus nerka) under varying operational sex ratio. Ethology 102, 304–322.Google Scholar
  199. Randall, R.G. (1989) Effect of sea-age on the reproductive potential of Atlantic salmon (Salmo salar) in eastern Canada. Can. J. Fish. Aquat. Sci. 46, 2210–8.Google Scholar
  200. Reynolds, J.D. and Gross, M.R. (1990) Costs and benefits of female mate choice: is there a lek paradox? Am. Nat. 136, 230–43.Google Scholar
  201. Riddell, B.E. (1986) Assessment of selective fishing on the age at maturity in Atlantic salmon (Salmo salar): a genetic perspective. Can. Spec. Publ. Fish. Aquat. Sci. 89, 102–9.Google Scholar
  202. Robitaille, J.A., Coté, Y., Shooner, G. and Hayeur, G. (1986) Growth and maturation patterns of Atlantic salmon, Salmo salar, in the Koksoak River, Ungava, Quebec. Can. Spec. Publ. Fish. Aquat. Sci. 89, 62–9.Google Scholar
  203. Roff, D.A. (1992) The Evolution of Life Histories. New York: Chapman & Hall. 535 pp.Google Scholar
  204. Rogers, D.E. (1987) The regulation of age at maturity in Wood River sockeye salmon (Oncorhynchus nerka). Can. Spec. Publ. Fish. Aquat. Sci. 96, 78–89.Google Scholar
  205. Roni, P. and Quinn, T.P. (1995) Geographic variation in size and age of North American chinook salmon. N. Am. J. Fish. Manage. 15, 325–45.Google Scholar
  206. Rowe, D.K. and Thorpe, J.E. (1990) Differences in growth between maturing and non-maturing male Atlantic salmon, Salmo salar L., parr. J. Fish Biol. 26, 643–58.Google Scholar
  207. Rowe, D.K., Thorpe, J.E. and Shanks, A.M. (1991) Role of fat stores in maturation of male Atlantic salmon (Salmo salar) parr. Can. J. Fish. Aquat. Sci. 48, 405–13.Google Scholar
  208. Royce, W.F. (1951) Breeding habits of lake trout in New York. Fish. Bull. US 52, 59–76.Google Scholar
  209. Rubin, J.-F. and Glimsäter, C. (1996) Egg-to-fry survival of the sea trout in some streams of Gotland. J. Fish Biol. 48, 585–606.Google Scholar
  210. Ryman, N. (1991) Conservation genetics considerations in fishery management. J. Fish Biol. 39(Suppl. A), 211–24.Google Scholar
  211. Sargent, R.C., Gross, M.R. and van den Berghe, E.P. (1986) Male mate choice in fishes. Anim. Behav. 34, 545–50.Google Scholar
  212. Saunders, R.L. (1967) Seasonal pattern of return of Atlantic salmon in the Northwest Miramichi River, New Brunswick. J. Fish. Res. Board Can. 24, 21–32.Google Scholar
  213. Saunders, R.L. (1981) Atlantic salmon (Salmo salar) stocks and management implications in the Canadian Atlantic Provinces and New England, USA. Can. J. Fish. Aquat. Sci. 38, 1612–25.Google Scholar
  214. Saunders, R.L., Henderson, E.B. and Glebe, B.D. (1982) Precocious sexual maturation and smoltification in male Atlantic salmon (Salmo salar). Aquaculture 28, 211–29.Google Scholar
  215. Scarnecchia, D.L. (1983) Age at sexual maturity in Icelandic stocks of Atlantic salmon (Salmo salar). Can. J. Fish. Aquat. Sci. 40, 1456–68.Google Scholar
  216. Schaffer, W.M. and Elson, P.F. (1975) The adaptive significance of variations in life history among local populations of Atlantic salmon in North America. Ecology 56, 577–90.Google Scholar
  217. Schluter, D. and McPhail, J.D. (1993) Character displacement and replicate adaptive radiation. Trends Ecol. Evol. 8, 197–200.Google Scholar
  218. Schroder, S.L. (1981) The role of sexual selection in determining overall mating patterns and mate choice in chum salmon. PhD thesis, Univ. Washington. 274 pp.Google Scholar
  219. Schroder, S.L. (1982) The influence of intrasexual competition on the distribution of chum salmon in an experimental stream. In Brannon, E.L. and Salo, E.O., eds. Salmon and Trout Migratory Symposium. Seattle: University of Washington Press, pp. 275–85.Google Scholar
  220. Scott, W.B. and Crossman, E.J. (1973) Freshwater fishes of Canada. Fish. Res. Board Can. Bull. No. 184. 966 pp.Google Scholar
  221. Sharov, A.F. and Zubchenko, A.V. (1993) Influence of human activity on properties of Atlantic salmon populations. In Stokes, T.K., McGlade, J.M. and Law, R., eds. The Exploitation of Evolving Resources. Berlin: Springer-Verlag, pp. 248–61.Google Scholar
  222. Shearer, W.M. (1992) The Atlantic Salmon: Natural History, Exploitation and Future Management. Oxford: Fishing News Books. 244 pp.Google Scholar
  223. Sibly, R.M. and Calow, P. (1983) An integrated approach to life-cycle evolution using selective landscapes. J. Theor. Biol. 102, 527–47.Google Scholar
  224. Sibly, R.M. and Smith, R.H. (eds) (1985) Behavioural Ecology: Ecological Consequences of Adaptive Behaviour. Oxford: Blackwell. 620 pp.Google Scholar
  225. Skúlason, S. and Smith, T.B. (1995) Resource polymorphisms in vertebrates. Trends Ecol. Evol. 10, 366–70.Google Scholar
  226. Smith, C.C. and Fretwell, S.D. (1974) The optimal balance between size and number of offspring. Am. Nat. 108, 499–506.Google Scholar
  227. Smith, G.W., Smith, I.P. and Armstrong, S.M. (1994) The relationship between river flow and entry to the Aberdeenshire Dee by returning adult Atlantic salmon. J. Fish Biol. 45, 953–60.Google Scholar
  228. Springate, J.R.C., Bromage, N.R., Elliott, J.A.K. and Hudson, D.L. (1984) The timing of ovulation and stripping and their effects on the rates of fertilization and survival to eyeing, hatch and swim-up in the rainbow trout (Salmo gairdneri R.). Aquaculture 43, 313–22.Google Scholar
  229. Stabell, O.B. (1984) Homing and olfaction in salmonids: a critical review with special reference to the Atlantic salmon. Biol. Rev. 59, 333–88.Google Scholar
  230. Sutterlin, A.M. and MacLean, D. (1984) Age at first maturity and the early expression of oocyte recruitment processes in two forms of Atlantic salmon (Salmo salar) and their hybrids. Can. J. Fish. Aquat. Sci. 41, 1139–49.Google Scholar
  231. Svärdson, G. (1949) Natural selection and egg number in fish. Rep. Inst. Freshwat. Res., Drottningholm 26, 115–22.Google Scholar
  232. Svärdson, G. (1957) Laxen och klimatet. Rep. Inst. Freshwat. Res., Drottningholm 38, 357–84. (In Swedish with an English summary)Google Scholar
  233. Taggart, J.B. and Ferguson, A. (1990) Hypervariable minisatellite DNA single locus probes for the Atlantic salmon, Salmo salar L. J. Fish Biol. 37, 991–3.Google Scholar
  234. Taggart, J.B., Prodöhl, P.A. and Ferguson, A. (1995) Genetic markers for Atlantic salmon (Salmo salar L.): single locus inheritance and joint segregation analyses of minisatellite (VNTR) DNA loci. Animal Genetics 26, 13–20.Google Scholar
  235. Tautz, A.F. and Groot, C. (1975) Spawning behavior of chum salmon (Oncorhynchus keta) and rainbow trout (Salmo gairdneri). J. Fish. Res. Board Can. 32, 633–42.Google Scholar
  236. Taylor, E.B. (1991) A review of local adaptation in Salmonidae, with particular reference to Pacific and Atlantic salmon. Aquaculture 98, 185–207.Google Scholar
  237. Taylor, E.B. and McPhail, J.D. (1985) Burst swimming and size-related predation of newly emerged coho salmon Oncorhynchus kisutch. Trans. Am. Fish. Soc. 114, 546–51.Google Scholar
  238. Tchernavin, V. (1938) Changes in the salmon skull. Trans. Zool. Soc., Lond. 24, 103–84.Google Scholar
  239. Tchernavin, V. (1944) The breeding characters of salmon in relation to their size. Proc. Zool. Soc., Lond. 113B, 206–32.Google Scholar
  240. Thomaz, D. (1995) Alternative life-history strategies in male Atlantic salmon (Salmo salar L.). PhD thesis, Univ. Leicester, 000 pp.Google Scholar
  241. Thorpe, J.E. (1986) Age at first maturity in Atlantic salmon, Salmo salar: Freshwater period influences and conflicts with smolting. Can. Spec. Publ. Fish. Aquat. Sci. 89, 7–14.Google Scholar
  242. Thorpe, J.E. (1994) Reproductive strategies in Atlantic salmon, Salmo salar L. Aquacult. Fish. Manage. 25, 77–87.Google Scholar
  243. Thorpe, J.E. and Mitchell, K.A. (1981) Stocks of Atlantic salmon (Salmo salar) in Britain and Ireland: discreteness and current management. Can. J. Fish. Aquat. Sci. 38, 1576–90.Google Scholar
  244. Thorpe, J.E., Miles, M.S. and Keay, D.S. (1984) Developmental rate, fecundity and egg size in Atlantic salmon, Salmo salar L. Aquaculture 43, 289–305.Google Scholar
  245. Trepanier, S., Rodriguez, M.A. and Magnan, P. (1996) Spawning migrations in landlocked Atlantic salmon: time series modelling of river discharge and water temperature effects. J. Fish Biol. 48, 925–36.Google Scholar
  246. Vaux, W.G. (1968) Interchange of stream and intragravel water in a spawning riffle. US Fish Wildl. Serv., Spec. Sci. Rep. — Fish. No. 405, 1–11.Google Scholar
  247. Verspoor, E. and Cole, L.J. (1989) Genetically distinct sympatric populations of resident and anadromous Atlantic salmon, Salmo salar. Can. J. Zool. 67, 1453–61.Google Scholar
  248. Verspoor, E., Fraser, N.H.C. and Youngson, A.F. (1991) Protein polymorphism in Atlantic salmon within a Scottish river: evidence for selection and estimates of gene flow between tributaries. Aquaculture 98, 217–30.Google Scholar
  249. Vladykov, V.D. (1954) Taxonomic characters of the eastern North America chars (Salvelinus and Cristivomer). J. Fish. Res. Board Can. 11, 904–32.Google Scholar
  250. Vladykov, V.D. (1962) Osteological studies on Pacific salmon of the genus Oncorhynchus. Fish. Res. Board Can. Bull. No. 136. 172 pp.Google Scholar
  251. Wankowski, J.W.J. and Thorpe, J.E. (1979) Spatial distribution and feeding in Atlantic salmon, Salmo salar L. juveniles. J. Fish Biol. 14, 239–47.Google Scholar
  252. Ware, D.M. (1978) Bioenergetics of pelagic fish: theoretical changes in swimming speed and ration with body size. J. Fish. Res. Board Can. 35, 220–8.Google Scholar
  253. Warner, K. (1962) The landlocked salmon spawning at Cross Lake Thoroughfare, Maine. Copeia 1962, 131–6.Google Scholar
  254. Webb, J.H. and Hawkins, A.D. (1989) The movements and spawning behaviour of adult salmon in the Girnock Burn, a tributary of the Aberdeenshire Dee, 1986. Scott. Fish. Res. Rep. No. 40. 42 pp.Google Scholar
  255. Webb, J.H.and McLay}, H.A. (1997) Variation in the timing of spawning of Atlantic salmon, Salmo salar, and its relationship to temperature in the Aberdeenshire Dee. Can. J. Fish. Aquat. Sci. (in press).Google Scholar
  256. Webb, J.H., Hay, D.W., Cunningham, P.D. and Youngson, A.F. (1991) The spawning behaviour of escaped farmed and wild adult Atlantic salmon (Salmo salar L.) in a northern Scottish river. Aquaculture 98, 97–110.Google Scholar
  257. Webb, P.W. (1978) Fast-start performance and body form in seven species of teleost fish. J. Exp. Biol. 74, 211–26.Google Scholar
  258. Webb, P.W. (1984) Body form, locomotion and foraging in aquatic vertebrates. Am. Zool. 22, 329–42.Google Scholar
  259. White, H.C. (1942) Atlantic salmon redds and artificial spawning beds. J. Fish. Res. Board Can. 6, 37–44.Google Scholar
  260. White, H.C. and Medcof, J.C. (1968) Atlantic salmon scales as records of spawning history. J. Fish. Res. Board Can. 25, 2439–41.Google Scholar
  261. Williams, G.C. (1966) Adaptation and Natural Selection. Princeton: Princeton University Press. 307 pp.Google Scholar
  262. Wood, C.C. and Foote, C.J. (1996) Evidence for sympatric genetic divergence of anadromous and nonanadromous morphs of sockeye salmon (Oncorhynchus nerka). Evolution 50, 1265–79.Google Scholar
  263. Youngson, A.F., Jordan, W.C. and Hay, D.W. (1994) Homing of Atlantic salmon (Salmo salar L.) to a tributary spawning stream in a major river catchment. Aquaculture 121, 259–67.Google Scholar

Copyright information

© Chapman & Hall 1996

Authors and Affiliations

  • Ian A. Fleming
    • 1
  1. 1.Norwegian Institute for Nature ResearchTrondheimNorway

Personalised recommendations