First published online June 16, 2004
Journal of Experimental Biology 207, 2649-2662 (2004)
Published by The Company of Biologists 2004
doi: 10.1242/jeb.01067
Antennae on transmitters on penguins: balancing energy budgets on the high wire
Rory P. Wilson1,*,
Jan M. Kreye1,
Klaus Lucke2 and
Heather Urquhart3
1 Institut für Meereskunde, Düsternbrooker Weg 20, D-24105 Kiel,
Germany
2 Forschungs- und Technologiezentrum Westküste, Hafentörn, D-25761
Büsum, Germany
3 New England Aquarium, Central Wharf, Boston, MA 02110, USA

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Fig. 1. Schematic diagram of the system used for measuring antenna drag showing
details of the relationship between antenna and pressure transducer and the
attachment of the measuring system to the penguin model (inset).
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Fig. 2. Relationship between pressure measured by the drag measurement system shown
in Fig. 1 and the torque
calculated by hanging known weights at specific distances from the fulcrum
(see text).
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Fig. 3. (A) Relationship between recorded pressure and swim speed for essentially
rigid antennae of various dimensions and set at various angles to the
direction of water flow mounted on a model penguin. (B) Relationship between
recorded pressure and swim speed for highly flexible antennae of various
dimensions and set at various angles to the direction of water flow mounted on
a model penguin. White symbols, length=200 mm; grey symbols, length=150 mm;
black symbols, length=100 mm.
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Fig. 4. Relationship between recorded pressure and swim speed for rigid antennae
(length 200 mm and set at an angle of 90° to water flow) with differing
diameters mounted on a model penguin.
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Fig. 5. (A) Frequency distribution of swim speeds used by nine Magellanic penguins
swimming from a colony at Punta Norte (N=8302). (B) Swim speed and
dive depth over three consecutive dives made by a Magellanic penguin foraging
from Punta Norte, Argentina. Note that the first and last dives in the series
show gradually changing speeds during the dives whereas the second dive shows
an abrupt change in speed (marked by an arrow) associated with a similarly
abrupt change in depth, which we assume is due to prey capture (see
Simeone and Wilson, 2003 ).
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Fig. 6. Frequency distribution of the number of consecutive dives (black bars;
N=302) where Magellanic penguins (N=25) foraging from Punta
Norte, Argentina, were considered to be exploiting a prey patch (this being
defined by higher swim speeds; see text) and the frequency distribution of the
amounts of estimated food ingested per patch for Magellanic penguins (grey
bars; N=4 birds for 65 patches) foraging from Punta Norte, Argentina.
Note that these two data sets were not derived from the same birds (see
text).
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Fig. 7. Frequency distribution (N=60) of the time between prey patches for
four Magellanic penguins foraging from Punta Norte, Argentina. The inset shows
all data for periods up to, and including, 6 h, to highlight the bimodality of
the data, whereas the main graph shows only those data for up to and including
120 min, to show more detail. Note that these data do not include periods
where birds presumably rested overnight.
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Fig. 8. (A) Drag (for calculations, see text) induced by antennae of various
dimensions on a model penguin as a function of swim speed. (B) Power output
needed to drive antennae of different dimensions attached to the body of a
penguin through water at various speeds.
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Fig. 9. Power input (energy expended per second) for a swimming Magellanic penguin
as a function of drag. This was derived by using a polynomial fit for the
mass-specific power requirements as a function of swim speed for Humboldt
penguins given by Luna-Jorquera and Culik
(2000 ) (Equation 9 in text)
and then regressing these power-requirements against the drag experienced by
the penguins swimming at the corresponding speed. The dragspeed
relationship was determined from the standard formula (Equation 10 in text),
which incorporates a drag coefficient of 0.03 (Bannasch, 1995) and a
cross-sectional area of 0.018m2 (see text).
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Fig. 10. (A) Relationship between energy expended per second and speed for a
Magellanic penguin swimming unequipped (bottom line) and equipped with
antennae (flexible or stiff) of various dimensions. (B) Relationship between
the cost of transport and speed for a Magellanic penguin swimming unequipped
(bottom line) and equipped with antennae (flexible or stiff) of various
dimensions. The arrows show the speeds at which costs of transport are
minimized for the various scenarios.
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Fig. 11. Relationship between foraging efficiency (dimensionless) and prey capture
speed for a Magellanic penguin foraging according to the conditions set out in
the text. The upper line (closed circles) shows the efficiency for an
unequipped bird while the lines delineated by squares and diamonds show the
efficiency of birds transporting external antennae (200 mmx3 mm) at
cruising speeds of 1 m s1 and 1.77 m s1,
respectively. The formula used for the antenna-derived drag was
Fd=0.913v20.91v1.5+
0.183v0.5+0.014 and is the best-fit curve
(r2=0.99997, F=10946, P<0.0001) from
the data corresponding to the relevant antenna (see
Fig. 8A). Note that the model
assumes that birds encounter a prey patch once every 36.3 min, travelling at a
mean speed of 1.77 m s1, which corresponds to a patch
separation of 3.86 km. Thus, swimming at 1 m s1, patches
with the same spatial distribution are encountered less often (only once every
64.25 min), although the overall foraging efficiency rises. Arrows show the
approximate scenarios expected for Adélie and Magellanic penguins due
to their different prey capture speeds (see text).
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© The Company of Biologists Ltd 2004