Elsevier

Animal Behaviour

Volume 78, Issue 4, October 2009, Pages 777-789
Animal Behaviour

Reviews
Organized flight in birds

https://doi.org/10.1016/j.anbehav.2009.07.007Get rights and content

The organized flight of birds is one of the most easily observed, yet challenging to study, phenomena in biology. Birds that fly in organized groups generally do so in one of two fashions: Line formations and Cluster formations. The former groups are typical of large birds such as waterfowl, where birds fly arranged in single lines, often joined together. The scientific questions about these groups usually involve potential adaptive functions, such as why geese fly in a V. Cluster formations are typically made up of large numbers of smaller birds such as pigeons or starlings flying in more irregular arrangements that have a strong three-dimensional character. The groups are defined by synchronized and apparently simultaneous rapid changes in direction. Scientific questions about these groups are usually concerned with mechanism such as how synchrony is achieved. Although field observations about the phenomenon date to the origins of natural history, experimental studies did not begin until the 1970s. Early experimenters and theoreticians were primarily biologists, but more recently aeronautical engineers, mathematicians, computer scientists and, currently, physicists have been attracted to the study of organized flight. Computer modelling has recently generated striking visual representations of organized flight and a number of hypotheses about its functions and mechanisms, but the ability to test these hypotheses lags behind the capacity to generate them. We suggest that a multi disciplinary approach to the phenomenon will be necessary to resolve apparently conflicting current hypotheses.

Section snippets

The era of anecdote and speculation

Several ornithologists of the 1930s made field observations that would later be very provocative to experimentalists and theoreticians. Nichols (1931) noted that in turning and wheeling pigeon, Columba livia, flocks, the position of the birds at the head of a turning flock would be exchanged with birds at the side after the completion of a turn; there did not appear to be consistent ‘leadership’ in such flocks. He speculated that this behaviour might be the result of faster birds in the front

Line formations

Line-flying birds typically fly in staggered, or ‘echelon’, formations rather than in straight lines nose-to-tail. If two such formations are joined at an apex at the front of the formation, we have a V or a J, its asymmetric variant. Franzisket, 1951, von Holst, 1952 and Hochbaum (1955) suggested that close formation flight might provide the advantage of a turbulence-free zone behind a bird ahead, but that would seem to apply only if the birds flew immediately behind the bird in front, like

Cluster flocks

There is an extensive literature discussing the biological value of flocking in general (Krebs & Barnard 1980), but very few papers have appeared with specific reference to the highly organized turning and wheeling (‘cluster’) flocks of some small birds. The most commonly offered hypothesis is that the closely spaced cluster flocks offer protection against aerial predators such as hawks, presumably by increasing the risk of collision to the predator (Tinbergen 1953). Examples have been reported

Conclusions

Advances in the understanding of the function and mechanisms of organized flight have been strongly linked to the introduction of new techniques or technologies. Heppner (1997) identified several areas that might be expected to produce such advances, but a decade later, although it has been possible to refine and more closely define these needs, much still needs to be done.

(1) Three-dimensional simulations. Some of the existing simulations (Vicsek et al., 1995, Lebar Bajec et al., 2005, Moškon

Acknowledgments

We sincerely thank Maja Lebar Bajec, Michael Byrne, Andrea Cavagna, Marjorie Heppner, Jim Kennedy, Craig Reynolds and Timothy Williams for reading the manuscript. This work was funded in part by the Slovenian Research Agency (ARRS) through the Pervasive Computing research programme (P2-0395).

References (129)

  • G. Grégoire et al.

    Moving and staying together without a leader

    Physica D

    (2003)
  • C. Huepe et al.

    New tools for characterizing swarming systems: a comparison of minimal models

    Physica A

    (2008)
  • D. Hummel

    Aerodynamic aspects of formation flight in birds

    Journal of Theoretical Biology

    (1983)
  • I. Lebar Bajec et al.

    Simulating flocks on the wing: the fuzzy approach

    Journal of Theoretical Biology

    (2005)
  • A. Okubo

    Dynamical aspects of animal grouping: swarms, schools, flocks, and herds

    Advances in Biophysics

    (1986)
  • H. Pomeroy et al.

    Laboratory determination of startle reaction time in the European starling (Sturnus vulgaris)

    Animal Behaviour

    (1977)
  • AMD

    ATI Radeon HD 4800 Series Real-time Demos

    (2008)
  • M. Andersson et al.

    Kin selection and reciprocity in flight formation?

    Behavioral Ecology

    (2004)
  • J.P. Badgerow

    An analysis of function in the formation flight of Canada geese

    Auk

    (1988)
  • M. Ballerini et al.

    Interaction ruling animal collective behavior depends on topological rather than metric distance: evidence from a field study

    Proceedings of the National Academy of Sciences, U.S.A

    (2008)
  • J.R. Beer

    The composition of bird flocks

    Flicker

    (1958)
  • D.C. Brogan et al.

    Dynamically simulated characters in virtual environments

    Computer Graphics and Applications

    (1998)
  • H. Chaté et al.

    Collective motion of self-propelled particles interacting without cohesion

    Physical Review E

    (2008)
  • E.A. Codling et al.

    Group navigation and the ‘many-wrongs principle’ in models of animal movement

    Ecology

    (2007)
  • C.R., Cone

    The Aerodynamics of Flapping Birdflight

    (1968)
  • I.D. Couzin et al.

    Effective leadership and decision-making in animal groups on the move

    Nature

    (2005)
  • J.H. Crook

    Adaptive significance of avian social organization

    Symposia of the Zoological Society of London

    (1965)
  • C.J. Cutts et al.

    Energy savings in formation flight of pink-footed geese

    Journal of Experimental Biology

    (1994)
  • A. Czirók et al.

    Collective motion

  • A. Czirók et al.

    Spontaneously ordered motion of self-propelled particles

    Journal of Physics A: Mathematical and General

    (1997)
  • L.M. Dill et al.

    Predicting the three-dimensional structure of animal aggregations from functional considerations: the role of information

  • Dimock, G. A. & Selig, M. S. 2003. The aerodynamic benefits of self-organization in bird flocks. In: Proceedings of the...
  • P.M. Driver et al.

    Protean displays as inducers of conflict

    Nature

    (1970)
  • J.T. Emlen

    Flocking behavior in birds

    Auk

    (1952)
  • A.P. Engelbrecht

    Fundamentals of Computational Swarm Intelligence

    (2006)
  • I. Farkas et al.

    Social behaviour: Mexican waves in an excitable medium

    Nature

    (2002)
  • T. Feder

    Statistical physics is for the birds

    Physics Today

    (2007)
  • G.W. Flake

    The Computational Beauty of Nature: Computer Explorations of Fractals, Chaos, Complex Systems, and Adaptation

    (2000)
  • L. Franzisket

    Über die Ursachen des Formationsfluges

    Vogelwarte

    (1951)
  • M. Gardner

    Mathematical games: the fantastic combinations of John Conway's new solitaire game ‘life’

    Scientific American

    (1970)
  • R.W. Gerard

    Synchrony in flock wheeling

    Science

    (1943)
  • B. Gönci et al.

    Phase transition in the scalar noise model of collective motion in three dimensions

    European Physical Journal Special Topics

    (2008)
  • Gould, L.L. 1972. Formation flight in the Canada goose (Branta c. canadensis). M.Sc. thesis, University of Rhode...
  • L.L. Gould et al.

    The Vee formation of Canada Geese

    Auk

    (1974)
  • G. Grégoire et al.

    Onset of collective and cohesive motion

    Physical Review Letters

    (2004)
  • M. Gschwind et al.

    Synergistic processing in Cell's multicore architecture

    IEEE Micro

    (2006)
  • Haffner, J. 1977. A flapping wing model for avian formation flight. M.Sc. thesis, University of Rhode Island,...
  • F.R. Hainsworth

    Precision and dynamics of positioning by Canada geese flying in formation

    Journal of Experimental Biology

    (1987)
  • F.R. Hainsworth

    Induced drag savings from ground effect and formation flight in brown pelicans

    Journal of Experimental Biology

    (1988)
  • W.J. Hamilton

    Social aspects of bird orientation mechanisms

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    F. H. Heppner is at the Department of Biological Sciences, University of Rhode Island, 102 Morrill Hall, Kingston, RI 02881-0816, U.S.A.

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