Echolocation calls of bats vary in design, and their
structure often reflects the ecological and sensory
requirements of the bats (Neuweiler, 1989; Fenton, 1990).
Individual bats can modify their calls according to
conditions: for instance, in confined spaces, calls may
become shorter and of broader bandwidth (e.g. Kalko and
Schnitzler, 1993). Individual flexibility in echolocation calls
is clearly illustrated during feeding attempts of bats that feed
on aerial prey or on prey on the water surface (e.g. Hartley
et al., 1989; Surlykke et al., 1993). When searching for prey,
the bat emits search-phase calls. On detection of the prey,
pulse repetition rate increases and pulse duration and
interpulse interval (IPI) decrease during the approach phase,
reaching extreme values during the terminal phase of the buzz
immediately prior to capture (Griffin et al., 1960). In many
species, the terminal phase can be subdivided into the phases
‘buzz I’ and ‘buzz II’. An abrupt fall in call frequency
typically marks the onset of buzz II (e.g. Kalko and
Schnitzler, 1989; Surlykke et al., 1993). Echolocation calls
emitted during the search phase are designed for detecting
targets, whereas calls emitted during the approach phase are
modified to provide more information on target location and
type (Simmons et al., 1979). The function of signals emitted
during the terminal phase is to provide information on a prey
item’s position immediately prior to capture (Fenton and
Bell, 1979). The terminal phase of echolocation behaviour is
synonymous with the phrase ‘feeding buzz’ (e.g. Griffin,
1958; Acharya and Fenton, 1992), but here we use the phrase
‘terminal buzz’ since it is often unclear whether the capture
attempt was successful.
The structure of the terminal buzz may vary according to the
nature of prey attacked and whether the capture attempt is
successful. In Lasiurus borealis and L. cinereus, the duration
of the terminal buzz was not related to prey size, but the silent
period after the buzz (which we call the ‘post-buzz pause’) was
longer after successful than after unsuccessful capture attempts
(Acharya and Fenton, 1992). Here, we develop some
hypotheses about how the structure of various components of
the terminal buzz should vary in relation to feeding behaviour,
and test these predictions both in the field and in the laboratory
by studying prey capture from the water surface (trawling) in
Daubenton’s bat Myotis daubentonii.
M. daubentonii is classified in the subgenus Leuconoe on
morphological characteristics. Many species in this subgenus
1793 The Journal of Experimental Biology 202, 1793–1801 (1999)
Printed in Great Britain © The Company of Biologists Limited 1999
JEB1951
During prey-capture attempts, many echolocating bats
emit a ‘terminal buzz’, when pulse repetition rate is
increased and pulse duration and interpulse interval are
shortened. The buzz is followed by a silent interval (the
post-buzz pause). We investigated whether variation in the
structure of the terminal buzz, and the calls and silent
periods following it, may provide information about
whether the capture attempt was successful and about the
size of prey detected – detail that is valuable in studies of
habitat use and energetics. We studied the trawling bat
Myotis daubentonii. The time between the first call of the
approach phase and the end of the terminal phase was not
related to prey size in the laboratory. The last portion of
the terminal buzz (buzz II) was shortened or omitted
during aborted capture attempts. Both in the laboratory
and in the field, the mean interpulse interval immediately
after the terminal buzz (post-buzz interpulse interval) was
longer in successful captures than in unsuccessful attempts.
In the laboratory, the post-buzz pause was longer after
successful captures than for unsuccessful attempts, and the
minimum frequency of the first search-phase call emitted
after the buzz (Fmin
) was higher than that of the last such
call prior to the buzz. These effects were not apparent in
field data. Both in the laboratory (85 %) and in the field
(74 %), significant discrimination between successful and
unsuccessful capture attempts was possible when the
duration of the post-buzz pause, post-buzz interpulse
interval and F
min
were entered into a discriminant analysis.
Thus, variation in the echolocation calls of bats during
prey-capture attempts can reveal substantial information
about capture success and prey size.
Key words: echolocation, bat, Myotis daubentonii, prey capture,
sonar.
Summary
Introduction
ECHOLOCATION BEHAVIOUR AND PREY-CAPTURE SUCCESS IN FORAGING BATS:
LABORATORY AND FIELD EXPERIMENTS ON MYOTIS DAUBENTONII
ADAM R. C. BRITTON AND GARETH JONES*
School of Biological Sciences, University of Bristol, Woodland Road, Bristol BS8 1UG, UK
*Author for correspondence (e-mail: Gareth.Jones@bris.ac.uk)
Accepted 1 April; published on WWW 8 June 1999