Wilson, Rory P., Kierspel, Mandy, Scolaro, J. A., Upton, J., Gallelli, H., Frere, E. and Gandini, P. (1999) To think or swim: Does it really cost penguins more to waddle?. Journal of Avian Biology, 30 . pp. 221-224. DOI 10.2307/3677132.

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Abstract

We looked at the routes taken by Magellanic Penguins up the beach while moving between the sea and the colony at a breeding site in San Julian, Argentina. Birds swam parallel to the shore for a period before leaving the water to cross the beach but trajectories over the beach were not perpendicular to the water's edge but at an angle of 39°. We examined the premise that birds might be optimizing for a trade off between time or energy by adopting this procedure and found that birds can gain little or no time by walking obliquely but can, under particular circumstances, save energy. These circumstances require that the previously-calculated costs of transport for walking have been over-estimated by a factor of two

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