Reviews in Fish Biology and Fisheries

, Volume 25, Issue 1, pp 21–37 | Cite as

Towards of a firmer explanation of large shoal formation, maintenance and collective reactions in marine fish

  • Guillaume Rieucau
  • Anders Fernö
  • Christos C. Ioannou
  • Nils Olav Handegard


Avoiding predation is generally seen as the most common explanation for why animals aggregate. However, it remains questionable whether the existing theory provides a complete explanation of the functions of large shoals formation in marine fishes. Here, we consider how well the mechanisms commonly proposed to explain enhanced safety of group living prey explain fish shoals reaching very large sizes. By conceptually re-examining these mechanisms for large marine shoals, we find little support from either empirical studies or classical models. We address first the importance of reassessing the functional theory with predator-dependent models and the need to consider factors other than predation to explain massive fish shoals. Second, we argue that taking into account the interplay between ultimate benefits and proximate perspectives is a key step in understanding large fish shoals in marine ecosystems. Third, we present the growing body of evidence from field studies that identify shoal internal structure as an important feature for how large shoals can form, maintain and react as a coordinated unit to external stimuli. In particular, we consider a mechanistic basis of local rules of interaction for group formation and collective dynamic properties that can account for groups reaching very large sizes. Recent research in collective animal behaviour has shifted focus from the importance of global properties (group size) to local properties (local density and information transfer). In contrast to studies of fish shoals in the laboratory, the difficulty in measuring behaviour in large shoals in marine systems remains a major constraint to further work. Advances in acoustical observation have shown the greatest potential to provide data that can link proximate mechanisms in, and ultimate functions of, large marine fish shoals.


Large marine fish aggregations Shoaling behaviour Collective behaviour functional explanations Local properties Shoal internal structure 



This work was financed by the Norwegian Research Council (Grant 204229/F20). CCI was supported by a Leverhulme Trust Early Career Fellowship and a NERC Independent Research Fellowship. We thank Lise Doksæter, Olav Rune Godø, Egil Ona, Espen Johnsen for providing us with helpful comments on this manuscript. We are also grateful to Graeme Ruxton and Iain Couzin for comments while we were developing the original idea of this paper.


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Copyright information

© Springer International Publishing Switzerland 2014

Authors and Affiliations

  • Guillaume Rieucau
    • 1
  • Anders Fernö
    • 2
  • Christos C. Ioannou
    • 3
  • Nils Olav Handegard
    • 1
  1. 1.Institute of Marine ResearchNordnes, BergenNorway
  2. 2.Department of BiologyUniversity of BergenBergenNorway
  3. 3.School of Biological SciencesUniversity of BristolBristolUK

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