First published online October 7, 2005
Journal of Experimental Biology 208, 3835-3849 (2005)
Published by The Company of Biologists 2005
doi: 10.1242/jeb.01856
Submerged swimming of the great cormorant Phalacrocorax carbo sinensis is a variant of the burst-and-glide gait
Gal Ribak1,
Daniel Weihs2,* and
Zeev Arad1
1 Department of Biology, Technion-Israel Institute of Technology, Haifa
32000, Israel
2 Faculty of Aerospace Engineering, Technion-Israel Institute of Technology,
Haifa 32000, Israel

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Fig. 1. Shape of the cormorant's body in three dimensions. (A) Schematic
illustration (not to scale) of the apparatus used to measure the diameters of
the cormorant carcass along the body's main axis.(B) Diameter of the width
(lateral axis, solid rectangles) and height (dorsoventral axis, empty
circles). The x-axis represents the position along the body's main
axis in % of body length (total body length = 85 cm). The diameters were used
to calculate surface area, volume and added mass coefficients. The broken
vertical lines mark the definition of the body excluding the neck and tail
(trunk).
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Fig. 2. Variation in velocity during one paddling cycle. The plot was obtained from
one video sequence (Bird 5) and smoothed by a stepwise 5-point moving average
for easy viewing. The plot shows the kinematic parameters defined by Weihs
(1974 ) for the burst-and-glide
model (L1, L2, Ui,
Uf), and the additional parameters defined in the present
study (Lp, Lg, Lr,
Ur, tp, tg,
tr); see List of symbols for the description of each
parameter. The bottom bar indicates the division of the paddling cycle into
three stages based on the motion of the feet relative to the body (stroke,
glide and recovery). Average swimming speed for this cycle was 1.72 m
s1. Note that the bird did not swim at a constant speed but
rather accelerated during the power stage and decelerated during the glide and
recovery stages in a burst-and-glide swimming pattern.
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Fig. 3. The average swimming speed observed for each cormorant in the experiment
relative to body length. Each point is the mean swimming speed obtained from
510 sequences taken of the same bird (values are ±
S.E.M.).
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Fig. 4. The duration of each stage in the paddling cycle of the great cormorant (A)
and the proportion of the power (burst) and deceleration (glide + recovery)
stages in the paddling cycle (B) in relation to paddling frequency. Note that
the glide duration decreased with the increase in paddling frequency. At a
paddling frequency of 2.7 Hz, the duration of the glide phase was diminished
to almost zero so that the power and recovery stages each occupy 50% of
the paddling cycle duration.
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Fig. 5. The distribution of the momentary drag along an average paddling cycle of
the great cormorant. Momentary drag is calculated using the
Cd average for each of the paddling stages. To average all
the analyzed sequences despite the differences in cycle duration, the
x-axis is the proportion of the cycle duration that is normalized by
dividing the period of each stage (stroke, glide and recovery) by the mean
duration of that stage. To allocate equal weight to all birds, the sequences
of each bird were first averaged and then means ±
S.E.M. of all birds were calculated.
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© The Company of Biologists Ltd 2005