Pedestrian locomotion energetics and gait characteristics of a diving bird, the great cormorant, Phalacrocorax carbo

  • Craig R. White
  • Graham R. Martin
  • Patrick J. Butler
Original Paper


Great cormorants Phalacrocorax carbo are foot propelled diving birds that seem poorly suited to locomotion on land. They have relatively short legs, which are presumably adapted for the generation of high forces during the power stroke of aquatic locomotion, and walk with a pronounced “clumsy waddle”. We hypothesise (1) that the speed, independent minimum cost of locomotion (C min, ml O2 m−1) will be high for cormorants during treadmill exercise, and (2) that cormorants will have a relatively limited speed range in comparison to more cursorial birds. We measured the rate of oxygen consumption \( (\dot{V}_{{\rm O}_{2}}) \) of cormorants during pedestrian locomotion on a treadmill, and filmed them to determine duty factor (the fraction of stride period that the foot is in contact with the ground), foot contact time (t c), stride frequency (f), swing phase duration and stride length. C min was 2.1-fold higher than that predicted by their body mass and phylogenetic position, but was not significantly different from the C min of runners (Galliformes and Struthioniformes). The extrapolated y-intercept of the relationship between \( \dot{V}_{{\rm O}_{2}} \) and speed was 1.9-fold higher than that predicted by allometry. Again, cormorants were not significantly different from runners. Contrary to our hypothesis, we therefore conclude that cormorants do not have high pedestrian transport costs. Cormorants were observed to use a grounded gait with two double support phases at all speeds measured, and showed an apparent gait transition between 0.17 and 0.25 m s−1. This transition occurs at a Froude number between 0.016 and 0.037, which is lower than the value of ~0.5 observed for many other species. However, despite the use of a limited speed range, and a gait transition at relatively low speed, we conclude that the pedestrian locomotion of these foot propelled diving birds is otherwise generally similar to that of cursorial birds at comparable relative velocities.


Froude Number Treadmill Exercise Stride Length Duty Factor Stride Frequency 
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We thank Lewis Halsey, Alex Kabat, and Susannah Thorpe for many helpful discussions, and two anonymous referees for their constructive comments. This research was supported by Natural Environment Research Council Grant NER/A/2003/00542 to G.R.M., P.J.B. and A. J. Woakes. All experiments were conducted under licence in accordance with the Animals (Scientific Procedures) Act 1986.


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Copyright information

© Springer-Verlag 2008

Authors and Affiliations

  • Craig R. White
    • 1
    • 2
  • Graham R. Martin
    • 1
  • Patrick J. Butler
    • 1
  1. 1.Centre for Ornithology, School of BiosciencesThe University of BirminghamEdgbaston, BirminghamUK
  2. 2.School of Integrative BiologyThe University of QueenslandSt LuciaAustralia

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