First published online September 9, 2005
Journal of Experimental Biology 208, 3451-3461 (2005)
Published by The Company of Biologists 2005
doi: 10.1242/jeb.01791
Energetic costs of diving and thermal status in European shags (Phalacrocorax aristotelis)
Manfred R. Enstipp1,*,
David Grémillet1 and
Svein-Håkon Lorentsen2
1 Centre d'Ecologie et Physiologie Energétiques, CNRS, 23 Rue
Becquerel, F-67087 Strasbourg Cedex 2, France
2 Norwegian Institute for Nature Research, Tungasletta 2, N-7485 Trondheim,
Norway

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Fig. 1. Side view and dimensions of the dive trench and the set-up of the
respirometry system within the laboratory container. The approximate
underwater routes taken by the birds are indicated by the arrows, with
arrowheads indicating the direction of locomotion.
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Fig. 2. Respirometry trace (A) and stomach temperature (B) of a European shag
during a dive trial (post-absorptive). Arrows indicate when the bird entered
and left the respirometry set-up. The trial lasted 32 min in water of
10.1°C. Note, the trace in A does not represent instantaneous metabolic
rate but gives an indication of metabolic rate during a dive trial. See
Materials and methods for details on how diving metabolic rate was
calculated.
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Fig. 3. Metabolic rate of European shags during various activities. Basal metabolic
rate (BMR) was measured in air temperatures between 10 and 19°C.
All measurements in water were made at water temperatures between 5 and
13°C. Values are grand means ± 1 S.D., which
were established from individual bird means. N=3 birds for all
activities except `preening', where N=1 bird. Asterisks indicate a
significant difference from BMR.
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Fig. 4. Metabolic rate of post-absorptive European shags during diving (triangles)
and when resting on the water surface (circles) at various water temperatures.
There was a significant negative relationship between metabolic rate and water
temperature during diving. The regression line shows the average relationship
for all shags, which takes into account variability between subjects. It is
best described by y=28.461-0.671x
(r2=0.69, N=3 birds, n=25 trials), where
y is metabolic rate during diving and x is water
temperature. During resting, there was a non-significant trend for metabolic
rate to increase with a decrease in water temperature (N=3 birds,
n=12 trials).
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Fig. 5. Stomach temperatures of European shags during rest at night and during the
day and during diving. Values are grand means ± 1
S.D., which were established from individual bird means.
N=3 birds. *Significantly different from day (rest) value.
Significantly different from diving (peak) value.
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© The Company of Biologists Ltd 2005