First published online October 7, 2004
Journal of Experimental Biology 207, 3917-3926 (2004)
Published by The Company of Biologists 2004
doi: 10.1242/jeb.01232
Heart rate and energetics of free-ranging king penguins (Aptenodytes patagonicus)
G. Froget1,2,
P. J. Butler1,*,
A. J. Woakes1,
A. Fahlman1,
G. Kuntz2,
Y. Le Maho2 and
Y. Handrich2
1 School of Biosciences, University of Birmingham, Edgbaston, Birmingham,
B15 2TT, UK
2 Centre d'Écologie et Physiologie Énergétiques, Centre
National de la Recherche Scientifique, 23 rue Becquerel, 67087 Strasbourg
cedex, France

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Fig. 1. Recording of depth (blue trace; top) and heart rate (red trace; bottom) of
a king penguin (bird B00, mass=11.65 kg) over a 6 day period. Black arrow
indicates the time when the bird went to sea.
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Fig. 2. Percentage frequency distribution of dive durations from 10 king penguins
(only dives longer than 1 min and inter-dive interval shorter than 10 min were
considered). Dive durations are divided into 20 s intervals.
N=5230.
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Fig. 3. Percentage frequency distribution of 30 min averaged heart rate from king
penguins ashore (solid line) and while at sea (dotted line);
N=10.
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Fig. 4. Mean heart rates at half-hourly intervals (continuous line), and mean
maximum diving depth (broken line) of foraging king penguins (N=10)
during the incubation and chick-guarding period. Each point is the mean
± S.E.M. from all the birds while they were at sea.
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Fig. 5. Temporal changes in heart rate (beats min1; bottom), and
depth (m; top) occurring during diving relative to the point of submergence
and the points of surfacing, for long dives (240250 s; red traces) and
short dives (150140 s; black traces). Each point is the mean ±
S.E.M. of the mean values from 7 birds. The blue horizontal lines
are mean ± S.E.M. of heart rate when ashore.
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© The Company of Biologists Ltd 2004