First published online May 1, 2006
Journal of Experimental Biology 209, 1816-1826 (2006)
Published by The Company of Biologists 2006
doi: 10.1242/jeb.02194
Flight and echolocation behaviour of whiskered bats commuting along a hedgerow: range-dependent sonar signal design, Doppler tolerance and evidence for `acoustic focussing'
Marc W. Holderied1,2,*,
Gareth Jones1 and
Otto von Helversen2
1 School of Biological Sciences, University of Bristol, Woodland Road,
Bristol BS8 1UG, UK
2 Department of Zoology II, University of Erlangen, Staudtstr. 5, 91058
Erlangen, Germany

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Fig. 1. Three examples of the effect of call design on ranging errors. (AC)
Spectrograms (FFT size 512, 256 points Hanning window, 240 points overlap).
Two calls (A,C) were emitted at the locations indicated in
Fig. 7B by stars. DOF indicates
the distance at which overall ranging errors were zero. (DF) Ranging
errors calculated for these calls and indicated as lines of identical error.
Labels on lines indicate size of ranging error in centimetres (>0
represents overestimation and <0 represents underestimation of distance).
The thick line labelled `focus' indicates locations where the overall ranging
error is zero. (GI) Ranging errors as in DF but for a smaller
distance range. `Pinheads' indicate perceived positions and `pin tips' the
actual positions of targets.
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Fig. 2. Four examples of the influence of call bandwidth and frequency modulation
curvature on Doppler-related ranging errors at a flight speed of 8 m
s1. Upper row: spectrograms of four call echo pairs. Lower
row: envelope of the cross correlation function (CCF). Arrows indicate the
actual time delay of 8 ms between call and echo. Red lines show the position
of the peak in the CCF.
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Fig. 3. Flight behaviour along the hedge. (A) 3-D surface of the hedge; (B) as in
A, but with all flight paths; (CF) flight corridor in four cross
sections through the hedge and flight paths at the Y-positions
indicated; (G) individual instantaneous distance to the hedge; (H) individual
flight speed during the flight along the hedge. Flight direction from right to
left, i.e. with decreasing Y.
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Fig. 4. Echolocation behaviour along the hedge (means ± s.d.) in segments of
1 m. (A) pulse interval; (B) pulse duration; (C) bandwidth of the first
harmonic; (D) Doppler ranging error; (E) width of the cross correlation
function (CCF) envelope, i.e. inverse measure of Doppler ranging acuity, for a
flight speed of 6 m s1.
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Fig. 5. Signal overlap zone (SOZ) of nine individual bats flying along the hedge.
Flight direction from right to left. The first individual in the upper row was
the only one not using the corridor below the hedge. Hemispheres indicate the
sizes of the SOZ plotted around the position of the bat at the time of each
call.
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Fig. 6. Distance of focus (DOF) of the same nine individual bats flying along the
hedge as in Fig. 5. Flight
direction from right to left. The first individual in the upper row was the
only one not using the corridor below the hedge. Hemispheres indicate the
surface of zero ranging error, i.e. a hemisphere with the DOF as its radius,
plotted around the position of the bat at the time of each call.
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Fig. 7. Distance of focus (DOF) and signal overlap zone (SOZ) in relation to
position to the hedge (shaded area). (A) Three-dimensional representation of
the flight paths of 22 bats and of the hedge they were flying along. The open
rectangle at Y=2 m indicates the size and position of the cross
section shown in B and C. (B) Cross section through the hedge and flight paths
as indicated by the rectangle in A. Symbols show where each individual bat has
passed the cross-sectional plane. Circles around symbols have a radius that
equals the DOF of the call emitted by the particular bat closest to the cross
sectional plane. (C) Same cross-section as in B, with circles indicating the
limit of SOZ.
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© The Company of Biologists Ltd 2006