Skip to main content

Thank you for visiting You are using a browser version with limited support for CSS. To obtain the best experience, we recommend you use a more up to date browser (or turn off compatibility mode in Internet Explorer). In the meantime, to ensure continued support, we are displaying the site without styles and JavaScript.

  • Letter
  • Published:

Survival variability and population density in fish populations


To understand the processes that regulate the abundance and persistence of wild populations is a fundamental goal of ecology and a prerequisite for the management of living resources. Variable abundance data, however, make the demonstration of regulation processes challenging1,2,3. A previously overlooked aspect in understanding how populations are regulated4,5,6 is the possibility that the pattern of variability—its strength as a function of population size—may be more than ‘noise’, thus revealing much about the characteristics of population regulation. Here we show that patterns in survival variability do provide evidence of regulation through density. Using a large, global compilation of marine, anadromous and freshwater fisheries data, we examine the relationship between the variability of survival and population abundance. The interannual variability in progeny survival increases at low adult abundance in an inversely density-dependent fashion. This pattern is consistent with models in which density dependence enters after the larval stage. The findings are compatible with very simple forms of density dependence: even a linear increase of juvenile mortality with adult density adequately explains the results. The model predictions explain why populations with strong regulation may experience large increases in variability at low densities7. Furthermore, the inverse relationship between survival variability and the strength of density dependence has important consequences for fisheries management and recovery, and population persistence or extinction8,9,10.

This is a preview of subscription content, access via your institution

Access options

Buy this article

Prices may be subject to local taxes which are calculated during checkout

Figure 1: Example relationships between the survival index and adult abundance.
Figure 2: The predicted relationships between adult abundance and recruitment, survival and survival variability.
Figure 3: Estimates of the heteroscedastic coefficient η 1 in survival across available fish species.

Similar content being viewed by others


  1. Murdoch, W. W. Population regulation in theory and practice. Ecology 75, 271–287 (1994)

    Article  Google Scholar 

  2. den Boer, P. J. & Reddingius, J. Regulation and Stabilization Paradigms in Population Ecology (Chapman & Hall, London, 1996)

    Google Scholar 

  3. Shenk, T. M., White, G. C. & Burnham, K. P. Sampling-variance effects on detecting density dependence from temporal trends in natural populations. Ecol. Monogr. 68, 445–463 (1998)

    Article  Google Scholar 

  4. Murray, B. G. Can the population regulation controversy be buried and forgotten? Oikos 84, 148–152 (1999)

    Article  Google Scholar 

  5. Turchin, P. Population regulation: a synthetic view. Oikos 84, 153–159 (1999)

    Article  Google Scholar 

  6. Berryman, A. A., Lima Arce, M. & Hawkins, B. A. Population regulation, emergent properties, and a requiem for density dependence. Oikos 99, 600–606 (2002)

    Article  Google Scholar 

  7. Hsiesh, C.-h. et al. Fishing elevates the variability in the abundance of exploited species. Nature 443, 859–862 (2006)

    Article  ADS  Google Scholar 

  8. Sæther, B.-E., Engen, S., Islam, A., McCleery, R. & Perrins, C. Environmental stochasticity and extinction risk in a population of a small songbird, the great tit. Am. Nat. 151, 441–450 (1998)

    Article  Google Scholar 

  9. Lande, R., Engen, S. & Sæther, B.-E. Stochastic Population Dynamics in Ecology and Conservation (Oxford Univ. Press, Oxford, 2003)

    Book  Google Scholar 

  10. Drake, J. M. Density-dependent demographic variation determines extinction rate of experimental populations. PLoS Biol. 3, 1300–1304 (2005)

    Article  CAS  Google Scholar 

  11. Wolda, H. & Dennis, B. Density dependence tests, are they? Oecologia 95, 581–591 (1993)

    Article  ADS  Google Scholar 

  12. Godfray, H. C. L. & Hassell, M. P. Long time series reveal density dependence. Nature 359, 673–674 (1992)

    Article  ADS  Google Scholar 

  13. Myers, R. A. Stock and recruitment: generalizations about maximum reproductive rate, density dependence, and variability using meta-analytic approaches. ICES J. Mar. Sci. 58, 937–951 (2001)

    Article  Google Scholar 

  14. Sale, P. F. & Tolimieri, N. Density dependence at some time and place? Oecologia 124, 166–171 (2000)

    Article  ADS  CAS  Google Scholar 

  15. Myers, R. A. & Cadigan, N. G. Density-dependent juvenile mortality in marine demersal fish. Can. J. Fish. Aquat. Sci. 50, 1576–1590 (1993)

    Article  Google Scholar 

  16. Deriso, R. B. Harvesting strategies and parameter estimation for an age-structured model. Can. J. Fish. Aquat. Sci. 37, 268–282 (1980)

    Article  Google Scholar 

  17. Schnute, J. A general theory for analysis of catch and effort data. Can. J. Fish. Aquat. Sci. 42, 414–429 (1985)

    Article  ADS  Google Scholar 

  18. Harvey, A. C. Estimating regression models with multiplicative heteroscedasticity. Econometrica 44, 461–465 (1976)

    Article  MathSciNet  Google Scholar 

  19. Mertz, G. & Myers, R. A. Estimating the predictability of recruitment. Fish. Bull. 93, 657–665 (1995)

    Google Scholar 

  20. Myers, R. A. & Cadigan, N. G. Is juvenile natural mortality in marine demersal fish variable? Can. J. Fish. Aquat. Sci. 50, 1591–1598 (1993)

    Article  Google Scholar 

  21. Mertz, G. & Myers, R. A. Match/mismatch predictions of spawning duration versus recruitment variability. Fish. Oceanogr. 3, 236–245 (1994)

    Article  Google Scholar 

  22. May, R. M. C. Stability and Complexity in Model Ecosystems Ch. 2 (Princeton Univ. Press, Princeton, 1973)

    Google Scholar 

  23. Myers, R. A., Barrowman, N. J., Hutchings, J. A. & Rosenberg, A. A. Population dynamics of exploited fish stocks at low population levels. Science 269, 1106–1108 (1995)

    Article  ADS  CAS  Google Scholar 

  24. Walters, C. & Kitchell, J. F. Cultivation/depensation effects on juvenile survival and recruitment: implications for the theory of fishing. Can. J. Fish. Aquat. Sci. 58, 39–50 (2001)

    Article  Google Scholar 

  25. Leigh, E. G. The average lifetime of a population in a varying environment. J. Theor. Biol. 90, 213–239 (1981)

    Article  MathSciNet  Google Scholar 

  26. Rosenberg, A. A. et al. The history of ocean resources: modeling cod biomass using historical records. Front. Ecol. Environ. 3, 78–84 (2005)

    Article  Google Scholar 

  27. Hutchings, J. A. & Myers, R. A. The effect of age on the seasonality of maturation and spawning of Atlantic cod, Gadus morhua. Can. J. Fish. Aquat. Sci. 50, 2468–2474 (1993)

    Article  Google Scholar 

  28. Peterman, R. M. Form of random variation in salmon smolt-to-adult relations and its influence on production estimates. Can. J. Fish. Aquat. Sci. 38, 1113–1119 (1981)

    Article  Google Scholar 

  29. Myers, R. A., Bridson, J. & Barrowman, N. J. Summary of worldwide stock and recruitment data. Can. Tech. Rep. Fish. Aquat. Sci. 2024, 1–327 (1995)

    Google Scholar 

  30. Hassell, M. P., Latto, J. & May, R. M. Seeing the wood for the tree: detecting density dependence from existing life-table studies. J. Anim. Ecol. 58, 883–892 (1989)

    Article  Google Scholar 

  31. Mertz, G. & Myers, R. A. Influence of fecundity on recruitment variability of marine fish. Can. J. Fish. Aquat. Sci. 53, 1618–1625 (1996)

    Article  Google Scholar 

  32. Manly, B. F. Stage-structured Populations: Sampling, Analysis and Simulation (Chapman and Hall, London, 1990)

    Book  Google Scholar 

  33. Stuart, A. & Ord, J. K. Kendall’s Advanced Theory of Statistics Vol. 1 Distribution Theory (Oxford Univ. Press, New York, 1987)

    MATH  Google Scholar 

Download references


We thank D. Tittensor for comments on the variance derivations and A. Edwards, B. Worm, D. Ricard and C. Barber for comments on the manuscript. This work was funded partly by the Irish government and partly by the European Union, under the National Development Plan 2000–2006, through “Supporting Measures in the Fisheries Sector” (to C.M.).

Author Contributions The original idea for this study was conceived by R.A.M. R.A.M., C.M. and W.B. developed the theoretical models for the variance in survival. C.M. conducted the empirical analyses. All authors contributed to the writing of the manuscript.

Author information

Authors and Affiliations


Corresponding author

Correspondence to Coilín Minto.

Additional information

All data used are available at the stock-recruitment database

Supplementary information

Supplementary Information

The file contains Supplementary Methods, Supplementary Figures 1-8 with Legends and Supplementary Table 1. The Supplementary Methods include analytical models for survival variability; delta approximations to survival variability; sensitivity analysis and meta-analytical methods. (PDF 927 kb)

Rights and permissions

Reprints and permissions

About this article

Cite this article

Minto, C., Myers, R. & Blanchard, W. Survival variability and population density in fish populations. Nature 452, 344–347 (2008).

Download citation

  • Received:

  • Accepted:

  • Issue Date:

  • DOI:

This article is cited by


By submitting a comment you agree to abide by our Terms and Community Guidelines. If you find something abusive or that does not comply with our terms or guidelines please flag it as inappropriate.


Quick links

Nature Briefing

Sign up for the Nature Briefing newsletter — what matters in science, free to your inbox daily.

Get the most important science stories of the day, free in your inbox. Sign up for Nature Briefing