Caribbean Journal of Science, Vol. 42, No.

1, 121-128, 2006
Copyright 2006 College of Arts and Sciences
University of Puerto Rico, Mayaguez

The Echolocation Repertoire of Eptesicus fuscus (Chiroptera:

Vespertilionidae) in Cuba
Annia Rodrguez and Emanuel C. Mora*
Department of Animal and Human Biology, Faculty of Biology, Havana University
calle 25 No. 455 entre J e I, Vedado, CP. 10 400, Ciudad de La Habana, Cuba
*Corresponding author: emanuel_mora@yahoo.com
ABSTRACT.The echolocation behavior of Eptesicus fuscus was studied in the field in western Cuba. Bats
leaving their roost produce short calls of around 2.3 ms showing two harmonics with a downward frequency
modulation between 75 and 33 kHz in the first harmonic. During hunting, search calls typically show a single
harmonic with a downward frequency modulation at the beginning of the signal, followed by a quasiconstant frequency component. Search calls are characterized by durations shorter than 8 ms and bandwidths
of approximately 16 kHz. The minimal frequency is maintained rather constant around 33 kHz. Transitions
from the search to the approach and terminal phases are characterized by an increase in duty cycle as a
consequence of the increase in pulse repetition rate and the decrease in duration. Calls emitted in the
laboratory are similar to those emitted in the wild. Compared with the echolocation repertoire of E. fuscus in
North America, shorter calls with constant minimal frequencies are characteristic of this species in Cuba.
KEYWORDS.bats, echolocation, Eptesicus fuscus

INTRODUCTION
Echolocation is tightly linked to the biology of microchiropteran bats and assists
them to navigate the air and to find food
(Griffin 1958). Variations described in the
echolocation repertoire of several bat species were explained as adaptations of call
designs to solve different perceptual tasks
(Simmons and Stein 1980; Mora et al. 2004),
as geographic variations (Thomas et al.
1987; Barclay et al. 1999), and as changes
devoted to transmit information to others
conspecifics (Fenton et al. 2004; Pearl and
Fenton 1996).
The echolocation system of the big
brown bat, Eptesicus fuscus, has been extensively studied in laboratory experiments
(Covey and Casseday 1995; Simmons 1973;
Simmons et al. 1995; Hartley and Suthers
1989). However, data on its echolocation
call repertoire in the field are limited to
Griffins work (1958), scarce detector re-

ms. received December 14, 2004; resubmitted June

18, 2005; accepted December 22, 2005

cordings (Bett 1998) and cursory data in reviews and papers focusing on other aspects
of echolocation (Simmons et al. 1979; Masters et al. 1995; Obrist 1995). Only recently,
Surlykke and Moss (2000) presented an extensive description of the echolocation call
repertoire of the North American E. fuscus
both in the wild and in the laboratory.
Geographic variations in the echolocation repertoire of E. fuscus are known of
populations from the East and West coast
of the United States of America (Fenton
1995). However, the wide distribution of E.
fuscus in North and South America, as well
as in Bahamas and the Greater Antilles,
predicts the existence of a higher degree of
geographic variations in the echolocation
behavior of this species.
In this paper we describe the echolocation behavior of Eptesicus fuscus in Cuba.
Bats were studied under different flying
conditions: 1) during their hunting behavior in the wild; 2) flying out of their natural
roost; and 3) flying in the laboratory. We
then discuss the differences between the
echolocation behavior of E. fuscus in Cuba
and in North America.

121

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ANNIA RODRGUEZ AND EMANUEL C. MORA

MATERIALS AND METHODS

Species identification and field locations
We studied the echolocation behavior of
Eptesicus fuscus (Chiroptera: Vespertilionidae) at three different localities of Havana
Province, western Cuba, between November, 2001 and September, 2003. In each locality bats were observed living in caves,
with colonies of animals in cracks and narrow fissures of the cave ceiling. Species
identification follows Silva (1979). Each animal was released immediately after identification, except for 10 of them that we used
for laboratory recordings. Recordings of
the bats hunting behavior were made
within 1 km around colonies. In each case
the bats were hunting for insects while flying at heights up to 5 m above ground level
in an area with scarce bushed vegetation.
Recordings from bats flying out of the roost
were made during their evening exodus.
The study was conducted between 18:00 to
22:00 hours, a period corresponding to the
first peak of E. fuscus nocturnal activity.
Sound recordings
We recorded the echolocation calls using
an U30 Ultrasound Advice ultrasonic detector (London, UK) with a flat response
characteristic ( 2 dB) between 20 and 200
kHz. During the recording sessions the heterodyne output was tuned to 35 kHz, thus
allowing the acoustic monitoring of individuals. The high frequency output of the
bat detector was fed into an analogdigital
input port of a digital signalprocessing
board (model PCM-DAS 16S/330) which
was controlled with the commercial software BatSound 2.1 (Pettersson Elektronik).
Calls were recorded at a sampling frequency of 312 kHz. We analyzed recordings sequences with at least 5 pulses of
good signal noise ratio (peak intensity with
more than 20 dB above noise level measured in the power spectra). A database
with 4 h of recordings (from which 34 sequences and 475 calls were finally selected)
was available for the analysis of echolocation.
As they usually fly in small circles of less

than 10 m in diameter, each E. fuscus was

visually followed during the recordings.
The microphone was always directed to
solitary bats and care was taken to record
only bats approaching the microphone. In
the same area we recorded different bats
one after the other to be confident about the
source of the echolocation passes.
Recordings in an enclosed space were
made from 10 bats, which were liberated
individually in a laboratory room (6 m x 6
m x 3 m). For recording bats flying out of
their roost the microphone was placed at
the entrance of their roosting cave.
Sound analysis
Sequences of echolocation calls were displayed simultaneously as spectrograms,
and temporal digitized recordings (oscillograms) using BatSound 2.1. Spectrograms
were made of consecutive fast Fourier
transforms (FFTs) with a 99% overlap.
Time was measured manually on the
screen. On oscillograms time resolution
was 0.1 ms. To obtain power spectra, FFTs
were calculated using 1024-2048 data
points.
From each call that registered a maximal
intensity of more than 20 dB above noise
level we measured the following parameters: (1) duration (time between start and
end of a call, measured in ms in the oscillogram); (2) peak frequency (frequency in
kHz corresponding with maximal intensity
in the power spectrum); (3) minimal and (4)
maximal frequency, respectively, lower
and higher values of frequency measured
20 dB below maximal intensity in the
power spectrum; (5) bandwidth (calculated
as difference between maximal frequency
and minimal frequency); (6) Q10-dB (calculated as peak frequency divided by bandwidth measured 10 dB below maximal intensity); (7) slope of frequency modulation
(calculated by the following ratio: difference in kHz between the initial and final
frequency of the call divided by the duration of the call). The variables included in
parameter 7 were measured in the spectrogram. In each call sequence we measured
interpulse interval from the beginning of a
call to the start of next call.

ECHOLOCATION OF BIG BROWN BAT IN CUBA

Since our sample included more than one

recording from the same individual, to
minimize pseudoreplication, the parameters of each pass of a bat were averaged
and treated as a single measurement before
further analysis. All the values of the acoustic parameters are given as mean 1 SD.
Since the data sets were not normally distributed (Kolmogorov-Smirnov test), we
used nonparametric statistics (KruskalWallis ANOVA). Statistical differences between several mean values were analyzed
with a nonparametric Dunns post-hoc test
(identical letters mean no statistically significant difference between the means).
Correlation analysis (Spearmans coefficient) was used to explore the strength of
relationship between acoustic parameters.
All analyses were made using the mean
values for each pass and the level of significance was = 0.05.
A multivariate discriminant function
analysis (DFA) was performed. The parameters used in this analysis were: duration,
maximal frequency, minimal frequency,
slope, peak frequency, and Q10-dB. The
cases were each of the situations and conditions of the recordings (e.g., flying from
the roost, flying in the room, search, approach, and terminal phases of the hunting
behavior). The DFA was employed to observe if cases could be separated in independent groups. The statistical partial was
used to determine the contribution of each
parameter to the classification of each case.
RESULTS
Bats flying out of the roost
Bats leaving their roost during the beginning of their evening foraging activity produced short calls (2.3 0.5 ms; N = 8 passes,
71 calls) with two harmonics overlapped in
frequency when measured in the power
spectra 20 dB below peak intensity. Each
call is a downward frequency modulated
sweep with the first most prominent harmonic between 75 and 33 kHz (Fig. 1A,
Table 1). Calls were emitted during the first
second of flight with an average interval of
63.0 24.9 ms.
Bats leaving the roost emitted calls that

123

FIG. 1. A. Oscillogram (above) and spectrogram (below) of the echolocation calls emitted by Eptesicus fuscus when leaving the roost during the beginning of
their evening foraging activity. B. Above: Acoustic parameters characteristics of the calls shown in (A).

increased in duration as the animals flew

further away from the colony entrance and
decrease in frequency slope modulation.
The minimal frequency, however, remained constant at about 33 kHz (Fig. 1B).
Bats hunting in the field
Eptesicus fuscus, hunting in the wild in
Cuba, emitted search calls of a single harmonic above which occasionally appeared
a very faint (more than 20 dB difference in
intensity) second harmonic (Fig. 2). At the
beginning of the signal, the calls show a
downward frequency modulation, followed by a quasi-constant frequency component (Fig. 2). Search calls are characterized by durations shorter than 8 ms and
bandwidths of approximately 16 kHz. As
evidenced by a variation coefficient of 2.5,
the smallest among the acoustic parameters, the minimal frequency is maintained
rather constant around 33 kHz. Search calls
are emitted at intervals of 87.7 19.2 ms.
Calls emitted during the approach phase

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ANNIA RODRGUEZ AND EMANUEL C. MORA

TABLE 1. Results of Kruskal-Wallis test used in the comparison of the echolocation calls emitted by Eptesicus
fuscus when leaving the roost, during the foraging activity (represented by the search calls) and flying in an
enclosed space. Number of calls analyzed in each behavioral situation is in parenthesis. The values of H show
significant differences (p < 0.05, one asterisk) and very significant differences (p < 0.01, two asterisks). The letters
in minuscule represent the results of the SNK test.

Parameters
Duration (ms)
Slope (kHZ/ms)
Peak frequency (kHz)
Minimal frequency (kHz)
Maximal frequency (kHz)
Q10-dB
Bandwidth (kHz)

Leaving the roost

(N = 8 passes;
71 calls)

Foraging
(N = 11 passes;
138 calls)

Enclosed space
(N = 15 passes;
166 calls)

2.3 0.3 c
15.7 1.8 a
46.6 3.6 a
33.2 1.1
67.1 3.1 a
2.2 0.5 c
33.3 4.6 a

4.6 1.0 a
5.9 2.2 c
36.5 2.0 c
32.8 0.8
47.3 2.8 c
3.7 1.0 a
16.4 6.1 c

3.0 0.7 b
11.1 1.5 b
41.9 2.6 b
33.0 1.1
61.9 4.4 b
2.3 0.8 c
29.3 6.9 b

22.80**
28.97**
26.55**
1.23 n.s.
29.10**
25.49**
28.76**

show a similar design to search calls (Fig.

2B). The start of the approach phase was
defined as the first signal in the sequence
with a monotonic change in the parameter
duty cycle that increased as a consequence
of the increase in pulse repetition rate (inverse of PI) and a decrease in duration; this
indicated a bat reacting to a potential prey
(Fig. 3). Of four approaching sequences
which grouped 53 calls, the shortest contained 6 and the longest 17 approach calls.
In the terminal phase (N = 42 calls distributed in 3 sequences), E. fuscus emitted
steep downward modulated frequency signals (Fig. 2). The beginning of the final buzz
is characterized by an abrupt increase in
pulse repetition rate and a decrease in call
duration (Fig. 3).
While hunting, E. fuscus operates at duty
cycles below 20%. In addition, it keeps the
minimal frequency of its calls relatively
constant at around 33 kHz during the
search and approach phases and only during the terminal phase drops below 25 kHz
(Fig. 3; Table 2). As a consequence, changes
in bandwidth are correlated (R = 0.98; N =
42 passes) with the maximal frequency (Fig.
4), which implies that broader signals are
achieved by increasing this parameter.
Call duration is also correlated with
bandwidth and thus with maximal frequency (R = 0.68) but not with minimal
frequency (R = -0.09; N = 42; p = 0.55). In
addition, call duration is correlated with
the interpulse interval. The minimum values of interpulse intervals correspond to

FIG. 2. A. Oscillograms (above) and spectrograms

(below) of the echolocation calls emitted by Eptesicus
fuscus during their foraging behavior. The three panels
correspond to a unique and continuous sequence in
which the three phases of the foraging behavior can be
recognized: search (s), approach (a) and final buzz (b).
B. Above: Enlarged spectrograms of echolocation calls
or groups of them, representative of each foraging
phase, marked with an asterisk in (A). Below: Power
spectra. The power spectrum of the final buzz call
corresponds with the marked call with asterisk.

ECHOLOCATION OF BIG BROWN BAT IN CUBA

125

monics. The design is similar to those emitted by the animals in the surroundings of
their roost or during the approach phase of
the foraging behavior. The bandwidth,
slope and call duration show intermediate
values with regard to the rest of the calls in
the echolocation call repertoire (Table 1).
However, the minimal frequency remained
around 33 kHz, showing no statistical differences with the other call types.
Overall variability

FIG. 3. Acoustic parameters that characterize the

echolocation calls emitted by Eptesicus fuscus during
its foraging behavior. The variations occur without
sudden changes from search (s) to approach (a) and
final buzz (b). PRR: pulse repetition rate; Initial: initial
frequency; Final: final frequency. Top: open circles:
duration; closed circles: PRR.

the shortest calls (R = 0.89; N = 22 passes;

p = 0.00).

Each type of call in the echolocation repertoire of E. fuscus was included in a discriminant function analysis to look for similarities in their spectral and temporal
design. A MANOVA showed that the
model was significant (Wilks = 0.00367;
F = 13.769; p < 0.01). The first two discriminant functions accounted for 95.88% of the
variation in the data. The values of partial
illustrated the following increasing discrimination power for the six parameters
used: peak frequency > Q10 > minimal frequency > slope > duration > maximal frequency. Calls emitted during the final buzz
were 100% of the time correctly identified
and grouped separately of the rest of the
calls (Fig. 5; Table 3). However, other types
of calls evidenced overlapping characteristics in their design thus difficult to determine their true group/nature.

Bats flying in enclosed space

DISCUSSION

Eptesicus fuscus flying in an enclosed

space emitted short calls (<4 ms; N = 15
passes, 166 calls) with two overlapping har-

Call design
In Cuba, the echolocation calls emitted
by E. fuscus show the downward frequency

TABLE 2. Mean DS values of acoustic parameters that characterize the echolocation calls emitted by Eptesicus
fuscus during each one of the phases of their foraging activity. The number of calls analyzed in each phase is in
parenthesis.

Parameters
Duration (ms)
Slope (kHz/ms)
Peak frequency (kHz)
Minimal frequency (kHz)
Maximal frequency (kHz)
Q10-dB
Bandwidth (kHz)

Search
(N = 11 passes;
138 calls)

Approach
(N = 4 passes;
53 calls)

Buzz
(N = 3 passes;
42 calls)

4.6 1.0
5.9 2.2
36.5 2.0
32.8 0.8
47.3 2.8
3.7 0.4
14.4 2.2

2.8 1.0
9.8 5.1
36.3 3.3
29.4 4.0
50.8 6.3
2.9 0.8
21.3 6.8

0.9 0.1
10.2 3.9
24.7 4.2
20.0 0.4
37.2 4.2
2.4 0.5
17.3 4.0

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ANNIA RODRGUEZ AND EMANUEL C. MORA

FIG. 4. Positive correlation between bandwidth and

maximal frequency (R = 0.98; N = 42 passes; p = 0.00).
Means per passes of the maximal and minimal frequencies are represented by open and closed symbols,
respectively. There is no correlation between the minimal frequency and bandwidth (R = 0.13; N = 42
passes; p = 0.38). Circles: search calls, approach calls.
Triangles: calls emitted when leaving the roost. Stars:
calls emitted at the enclosed space.

modulated sweep design characteristic of

vespertilionid bats (Fenton and Bell 1981;
Kalko and Schnitzler 1993; O Farrell et al.
1999; Parson and Jones 2000). The combination in the same signal of a frequency
modulated (FM) and a quasi-constant frequency (QCF) component allows bats to
modify independently both of them; this
guaranties the adequate acoustic call characteristic to fulfill a specific task. For example, during the search phase insectivorous bats decrease the duration of the FM
component and increase the QCF of the signal. As a consequence, the search calls are
longer and narrower which causes the signal energy to concentrate in a narrow band
of frequencies, thus facilitating the detection of distant prey (Simmons and Stein
1980; Schnitzler and Kalko 1998). On the
other hand, the FM component is well
suited for a precise localization and characterization of target (Simmons et al. 1979).
When compared to the search calls, the approach and final buzz calls design in E. fuscus is dominated by the initial FM component that determines an increase in the

FIG. 5. Discriminant functions analysis applied to

the echolocation calls emitted by Eptesicus fuscus during the search phase (open circles), approach phase
(open triangles) and final buzz (open rhombus) as well
as calls emitted close to the roost (grays triangles) and
flying in enclosed space (closed circles).

bandwidth and a decrease in call duration.

A wider bandwidth signal offers more information about the physical characteristic
of the reflective surface (Schnitzler and
Henson 1980; Simmons and Stein 1980).
This change in call design is also characteristic of other vespertilionid bats and is
present in the North American E. fuscus
(Surlykke and Moss 2000). In addition, the
high frequency repetition rate of the calls in
the final buzz increase the quantity of information obtained per unit of time and allows the bat to prepare for the capture.
Adaptation of echolocation calls to different
flight conditions
A change in the structure of echolocation
calls to fit the requirements of each hunting
phase (search, approach, and final buzz) is
a common behavior in insectivorous bats,
also found in E. fuscus (Griffin 1958; this
study). Calls emitted under other circumstances are often intermediate in acoustic
features to hunting calls. That is the case of
calls emitted in the laboratory and leaving
the roost in both, the North American and
the Cuban E. fuscus (Surlykke and Moss
2000; this study).
FM bats adapt the characteristics of their
calls to the degree of clutter present in their
foraging areas (Simmons et al. 1979). The

ECHOLOCATION OF BIG BROWN BAT IN CUBA

127

TABLE 3. Correct identification rates from a discriminant function analysis (DFA) including each type of call
emitted by Eptesicus fuscus in Cuba.
True group
Classified as:

Search

Approach

Buzz

Enclosed space

Roost

Search
Approach
Buzz
Enclose space
Roost
N total
% correct

196
5
0
8
2
211
92.89

26
13
3
8
3
53
24.52

0
0
42
0
0
42
100

13
7
0
138
8
166
83.13

1
0
0
19
51
71
71.83

adaptation of call duration to flight conditions, for example, is interpreted as a clutter

rejection strategy (Kalko and Schnitzler
1993). According to this strategy, E. fuscus
flying close to the roost and in enclosed
spaces decreases the duration of its calls to
avoid call-echo overlap.
Geographical variations in echolocation calls
of E. fuscus
The results gathered on the echolocation
of E. fuscus in Cuba and North America
show differences in acoustic parameters
like call duration and minimal frequency.
According to Surlykke and Moss (2000), E.
fuscus in North America emits long signals
(14-19 ms) while hunting in the field, similar to what is found in cruising bats after
flying out of the roost. Likewise, these authors described a negative correlation between the minimal frequency and call duration for calls from E. fuscus in North
America. Cuban E. fuscus appears to keep
the minimal frequency constant during
search and approach phases while hunting;
furthermore, this frequency does not differ
from the minimum frequency of echolocation calls recorded in the laboratory. This
contrasts with a difference of almost 15 kHz
among calls found in North American E.
fuscus (see Fig. 3, Surlykke and Moss 2000).
Part of this difference can be explained by
the fact, that the sample from Cuban E. fuscus only included calls with durations up to
7 ms, whereas Surlykke and Moss (2000)
report call durations up to 20 ms. Differences in call duration may well be explained by differences in hunting heights

since E. fuscus in Cuba searches for prey

flying close to ground and clutter, while in
the continent it can search for insects more
than 10 m above the ground (Surlykke and
Moss 2000). In echolocating bats, the longest and lowest frequency calls are emitted
by high flying bats (Jensen and Miller
1999). It is then, too soon, to propose the
existence of fundamental differences in the
acoustic behavior of E. fuscus in North
American and the Caribbean. As demonstrated previously in Pipistrellus (Russo and
Jones 2000; Mayer and von Helversen
2001): acoustic differences may point to the
existence of sibling species; therefore new
studies should further test this hypothesis.
Acknowledgments.The authors want to
express their gratitude to Elier and Fernando, for field assistance. Special thanks
to Silvio Macas, Adianez Garca, Ileana
Quinez and Danny Rojas for their constant support to the group enthusiasm. The
equipment used in this work was provided
by a grant from the VW Foundation project
(I / 77-306).
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