Some hot topics in animal bioacoustics

Whitlow W. L. Au
Marine Mammal Research Program, Hawaii Institute of Marine Biology, P.O. Box 1106, Kailua,
Hawaii 96734

~Received 16 November 1996; accepted for publication 19 February 1997!

This paper is derived from a Hot Topics in Animal Bioacoustics presentation at the 130th
meeting of the Acoustical Society of America in St. Louis, Missouri. Six bioacoustics studies on a
wide variety of species are discussed. Two of the studies are concerned with insects, the parasitoid
fly, and cotton bollworms. The remaining bioacoustics studies are on aquatic animals including the
West Indian manatee, elephant seals, and dolphins. 1997 Acoustical Society of America.
@S0001-4966~97!07005-7#
PACS numbers: 43.10.Ln, 43.80.Ka, 43.80.Lb @DWM#

INTRODUCTION

The field of animal bioacoustics is diverse, involving a

wide variety of species varying from insects and birds to
terrestrial and aquatic animals. Interest in this area of acoustics has been growing, and the Animal Bioacoustics Special
Technical Group will become a full committee of the Acoustical Society of America following the fall 1996 meeting in
Honolulu. This paper is derived from a Hot Topics in Animal Bioacoustics presentation at the 130th meeting of the
ASA in St. Louis, Missouri, in which interesting research
being performed with different species was highlighted. Results of some of the studies that will be discussed have not
been published so they can be discussed only in general
terms without compromising the publication plans of the
various investigators. The objective of this paper is to
present the reader with a general flavor of the type of animal
bioacoustics research that is currently being conducted. The
seven specific projects highlighted deal with subjects as varied as insects and mammals of aquatic and marine origin.
I. DIRECTIONAL HEARING IN THE PARASITOID FLY

Humans and most large animals generally localize

sounds by using two mechanisms: time of arrival difference
of sounds arriving at the two ears ~which can also be considered as a phase difference cue!, and the difference in the
intensity of sounds arriving at the two ears caused by diffraction of sounds around the head and body. However, the ears
of insects are often fractions of a wavelength apart, making
these two cues relatively ineffective. Yet insects and other
small animals can and do localize sounds using very different mechanisms such as pressure gradient receivers in some
crickets ~Michelsen, 1992!.
The female Parasitoid fly, Ormia ochracea must find
and deposit her parasitic larvae on a live field cricket. It
locates the host cricket at night, apparently using acoustics
cues. However, the flys ears are not physically separated but
are contained within a common air-filled chamber, only
about 450 to 520 mm apart ~Miles et al., 1995!. The small
separation distance between tympanum makes the encoding
of time difference and intensity difference cues impractical.
A group from Cornell University ~Robert et al., 1992, 1994!
has been studying the hearing processes of the parasoitoid fly
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and has discovered a new mechanism of sound localization

involving a mechanical coupling between the tympanal
membranes of the flys hearing organ ~Miles et al., 1995!.
Figure 1 is a schematic showing the external anatomy,
location of the ears, and an electron micrograph of the tympanal membranes. The bulbae acoustica are attached to the
membranes through a stiff cuticular rod that connects to the
tympanum at the tympanal pit ~TP!. The cuticular structure
connecting the tympanum pits to each other and to the pivot
point is known as the intertympanal bridge. Each bulba acustica contains about 7075 auditory reception cells that are
innervated by the auditory nerve.
Miles et al. ~1995! measured the response of the tympanal membranes to sounds arriving at different angles by using a laser vibrometer, as depicted in Fig. 2. The laser light
could be focused to an area of approximately 5 mm in diameter allowing Miles and his colleagues to measure the membrane vibration at a large number of locations. The incident
sound was also measured using a small microphone, as
shown in Fig. 2. The acoustic stimulus was a burst of bandlimited white noise lasting 0.01 s and having a frequency
range of 1 to 30 kHz. The signals from the laser vibrometer
and the microphone were digitized and the transfer function
of the different locations on the tympanum were measured.
Miles and colleagues found that the tympanal pit ~TP!
which was closest to the sound source ~ipsilateral! responded
with as much as 20 dB greater amplitude than the tympanal
pit farthest from the sound source ~contralateral!. The tympanal structure rocks about the central pivot point with the
two ears moving in nearly opposite phase. A mechanical
model of the tympanal structure on a pivot is depicted in Fig.
3. When a sound first arrives at the ipsilateral tympanal pit,
the structure deflects downward and this motion produces
forces on the contralateral ear via the intertympanal bridge,
which tend to cancel the effects of the external acoustic pressure. This mechanical action essentially produces an interaural intensity difference that would normally not exist since
the ears are closer than 1/200th of a wavelength at 5 kHz.
II. ACOUSTIC DETECTION OF PINK BOLLWORMS

Pink bollworm infestation of cotton bolls is a serious

problem for cotton growers. The present methods for exam-

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1997 Acoustical Society of America

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FIG. 1. ~a! External anatomy of the Ormia orchracea showing the location
of the ears. ~b! Frontal view of the ears with the head removed in left panel
and a frontal scanning electron micrograph view of the ears. The prosternal
tympanal membranes ~PTM! show radial corrugations which converge upon
the tympanal pit ~TP! to which the internal sensory organ is attached ~from
Miles et al., 1995!.

ining samples of cotton bolls to determine if a field has been

infested are laborious and prone to error. In order to solve
this problem and devise a more expedient method of examining cotton bolls for bollworms, Robert Hickling and colleagues from the National Center for Physical Acoustics
have turned to acoustic detection of larvae. Larvae constantly
eat and move, generating sounds that can be detected by a
sensitive acoustic sensor.
Boll cutting is the traditional method to determine if boll
infestation has occurred. The cotton boll are pulled from the
plant and are cut open and examined for pink bollworm lar-

FIG. 2. Schematic of the experimental geometry ~adapted from Miles et al.,

1995!.
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J. Acoust. Soc. Am., Vol. 101, No. 5, Pt. 1, May 1997

FIG. 3. Mechanical model of the parasitoid fly ears. The intertympanal

bridge is assumed to be two rigid bars connected at the pivot through a
coupling spring k 3 and dashpot c 3 ~from Miles et al., 1995!.

vae. However, the larvae are usually very small, and so this
technique is not only error prone but also time intensive. In a
second method, cotton bolls are stored and left in boll boxes
at the appropriate temperature and daylength regime for a
time. Larvae within the bolls finish feeding and cut out of the
boll. The number of damaged bolls and larvae/pupae produced are then counted. However, this technique requires up
to 2 weeks and therefore has limited value.
The prototype multiple acoustic sensor developed by
Hickling et al. ~1994! to detect the sounds of pink bollworm
larvae eating and moving in cotton bolls has four major components: the sound isolation box, sensors, amplification and
filtering components, and power supply. The individual sensors are based on low cost fetal monitors consisting of
adapted electret microphones connected to stethoscope headphones that were developed at the National Center of Physical Acoustics. A photograph of the prototype multiple acoustic sensor consisting of 48 sensors is shown in Fig. 4. The
sensor signals are amplified and bandpass-filtered so that
only a narrow band of low-frequency sound is detected.
Bolls from a Pima cotton field in Coolidge, Arizona
were used in a test of the acoustic detection system. Three
hundred bolls were cut using standard procedures, and were
put into boll boxes and held in an insectary. Three hundred
other bolls were used with the multiple acoustic sensor and
then were carefully cut to verify sensing results. Binocular
microscopes were used in several cases to find suspected
larvae for the bolls tested with the acoustic unit. The bolls
were warmed to a temperature of 38 C prior to sensing. This
increased audible larval activity thereby increasing the probability of sensing the larvae.
Results of the test are shown in Fig. 5. The acoustic
sensor found as many or more larvae than did the other
methods. Furthermore, acoustic sensing took less time and
labor. As with the standard boll cutting and storing techniques, the acoustic sensor unit was not error free. Cases in
which a larva was heard, but could not be found ~false positive! occurred as shown in Fig. 5, along with cases in which
the sensors did not detect larvae but one was found upon
Whitlow W. L. Au: Animal topics

2434

FIG. 4. Photograph of the multiple acoustic sensor system ~courtesy of R.

Hickling!.

cutting ~false negative!. The results indicate that use of a

multiple acoustic sensor may be a good way to examine
infestation of bollworms.
III. HEARING IN MANATEES

Manatees were once very plentiful in the warm waters of

Florida but were placed on the endangered species list in
1973. A variety of causes could account for the decrease in
population: destruction of habitat, loss of sea grasses, and
collisions with motorboats. Boating accidents involving
manatees have been particularly disturbing to Florida officials as the number of boats using Floridas waterways continues to increase. It seems strange that manatees do not

FIG. 5. Results of test one using Pima cotton ~from Hickling et al., 1994!.
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J. Acoust. Soc. Am., Vol. 101, No. 5, Pt. 1, May 1997

FIG. 6. ~a! Picture of the experimental apparatus showing the J-9 sound
projector, H-56 measuring hydrophone, a hoop station and a manatee approaching the hoop. ~b! Manatee in the hoop station awaiting an acoustic
signal ~courtesy of Edmund Gerstein!.

detect the approach of on-coming boats and avoid collision

by swimming away. Lack of knowledge on the hearing capabilities of manatees has made it difficult to solve the boat
collision problem without placing heavy constraints on the
use of Floridas waterways by boaters.
A team of scientists led by Edmund Gerstein has been
performing auditory experiments with the West Indian manatee ~Trichecus manatus! in order to address the boat collision
problem. They have performed experiments to measure the
underwater hearing sensitivity as a function of frequency
~audiogram! for two 7-year-old captive born males ~Gerstein
et al., 1993!, and their masked hearing threshold and sound
localization capabilities ~Gerstein et al., 1995!. The hearing
studies are being conducted at the Lowry Park Zoo in
Tampa, Florida.
The hearing sensitivity of the manatees was determined
by a controlled psychoacoustic experiment in which they
were trained to place their head into a hoop facing a J-9
transducer. The J-9 sound projector, H-56 measuring hydrophone, a hoop station, and a manatee approaching the hoop
are shown in Fig. 6~a!; and the manatee in the hoop station is
depicted in Fig. 6~b!. A pure-tone signal was then played to
the animal. If it could hear the tone, the animal would respond by touching the signal-present paddle, and if a signal
could not be heard, the animal would respond by touching
the signal-absent paddle. The signal level was systematically
decreased after correct detection trials and increased after the
animal missed or failed to respond to a sound. Thus, the
Whitlow W. L. Au: Animal topics

2435

FIG. 7. ~top panel! Audiogram of a manatee compared with a California sea

lion and a harbor seal. ~bottom panel! Manatee audiogram compared with
that of a bottlenose dolphin and a killer whale ~from Gerstein et al., 1997!.

signal level was varied in an updown staircase fashion depending on the response of the subject.
The audiogram of one of the subjects ~Gerstein et al.,
1997! is shown in Fig. 7 along with the audiograms for a
California sea lion and for a harbor seal in the top panel; and
for a killer whale and a bottlenose dolphin in the bottom
panel. The manatee audiogram has a typical u shape in the
hearing range from 500 Hz to 38 kHz. The data also indicated that a manatee can hear low-frequency sounds ~below
16 kHz! as well or better than other marine mammals.
Manatees are agile enough to avoid oncoming boats
~Gerstein, 1994, 1995!, if they are aware of impending collision, but yet they do not. However, the low-frequency nature of boat noise may preclude detection at sufficient range
because of elevated hearing sensitivity at low frequencies
~below 1 kHz!. It is also possible that a manatee may hear an
oncoming boat but not be able to localize the direction of the
sound. Finally, their hearing in noise may also not be good.
Gerstein and colleagues are continuing to study the hearing
of manatees by conducting a masked hearing threshold study
and a sound localization study.

have recently expanded their research to consider the effects

of low-frequency sounds, including the ATOC signal on
these animals ~Fletcher et al., 1996!.
Special electroacoustics instrument packages have been
developed that can be attached to elephant seals to study the
acoustic environment of these animals when they are out at
sea. Adult female elephant seals spend 83% of the year at
sea, and 90% of the time they are submerged ~Le Boeuf
et al., 1988!. They are deep divers, averaging 500-m depth
on each dive with maximum dives to 1500-m depth. They
are one of few marine mammals that can dive down to the
axis of the deep sound channel in temperate waters. Therefore, the kinds of sounds that these animals encounter, especially when they are in the deep sound channel are of much
interest.
The objectives of Le Beoufs project are to determine ~a!
if diving seals make active sounds, ~b! the frequencies and
levels of sounds diving seals encounter in their environment,
and ~c! the best package design and attachment process for
minimizing flow noise. Instrument packages were attached to
o Nuevo Island from
juvenile seals, on their return to An
Monterey Bay. The first model of the attachment package
contained a Sony digital audio tape recorder with a hydrophone, along with time depth recorder and swim speed detectors and the appropriate datalogger. Later models use a
Tattletale data logger which also performs the analog-todigital conversion of the incoming acoustic signals and
stores the data on a hard disk. A picture of the latest electroacoustic package is shown in Fig. 8. The hydrophone is
mounted on the back of the package to minimize the effects
of flow noise. The top panel of Fig. 9 shows a seal being
released from a boat with an electroacoustics package attached to its back. The bottom panel shows the seal on the
o Nuevo Island.
beach at An
An example of the acoustic signal received by a seal is
shown in Fig. 10, with the depth of dive shown above the
color sonogram. Most of the received signals had frequencies
in the range of 20200 Hz. Snapping shrimp, cetacean
sounds, boat noise, and seal swim strokes and heart beats are
clearly audible in some of the data. Flow noise correlated
with swim speed, suggests that optimal time for acoustic

IV. ACOUSTIC ENVIRONMENT OF DIVING ELEPHANT

SEALS

Many marine animals rely on acoustics to capture prey,

avoid predators, reproduce, and navigate. Yet the ocean is a
very noisy environment, especially at low frequencies.
Buney Le Boeuf and his colleagues at the University of California at Santa Cruz have spent many years studying the
natural behavior and physiology of the northern elephant seal
o Nuevo rookery off the coast of California. They
on the An
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FIG. 8. Picture of the electroacoustic package showing the hydrophone

located on the back of the package to minimize flow noise ~courtesy of W.
Burgess!.
Whitlow W. L. Au: Animal topics

2436

rine mammals ~Burgess et al., 1996!. This capability has

been a important bonus to the project. Overall, this project
demonstrates that it is feasible to monitor the acoustic environment of free swimming seals, and, that such electroacoustics packages could provide important data on the seals environment.
V. AUDITORY-EVOKED POTENTIAL FROM TRAINED
DOLPHINS

FIG. 9. The top panel shows a seal with an instrumentation package being
o
released at sea and the bottom panel shows a seal on the beach at An
Nuevo Island ~courtesy of B. LeBeouf!.

sampling would be when the seals are swimming slowly.

Results of several deployments have indicated that it is also
feasible to obtain long-term, reliable, quantitative, and noninvasive cardiac monitoring of elephant seals and other ma-

Auditory-evoked potentials on dolphins and on small

whales have been measured with the animals housed in a
small container where there is a considerable amount of
acoustic reflection from the container walls ~Ridgway et al.,
1981; Supin and Popov, 1985!. The length of these containers is generally just larger than the animals length making it
difficult to position a transducer directly on the beam axis of
the animals auditory reception system. Furthermore, brainstem-evoked potentials have been measured with either a
click or short tone-burst stimuli. With the click signal, it is
impossible to specify the amplitude that would relate to a
threshold obtained in a behavioral study in which the rms
sound-pressure level at threshold is used. A click signal is
also broadband so that it is difficult to determine what part of
the spectrum the animals auditory system is responding to.
Tone bursts also have some frequency and amplitude ambiguities associated with them.
Auditory-evoked potentials recorded from the human
scalp have been shown to follow the periodicity of pure-tone
sinusoidal signals for frequencies up to about 1 kHz ~Moushegian et al., 1983! and have been termed the frequency following response. Scalp-recorded responses have also been
found to follow the envelope of continuous sinusoidally amplitude modulated stimuli in humans ~Richards and Clark,
1984! and in gerbils ~Dolphin and Mountain, 1993! using
carrier frequencies considerably higher than the pure-tone
phase locking capability of the frequency following re-

FIG. 10. Example of the acoustic signal received by the instrumentation package on a diving elephant seal ~courtesy of W. Burgess!.
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sponse. Continuous sinusoidal amplitude-modulated ~SAM!

signals produced by amplitude modulating a high-frequency
sinusoidal carrier with a low-frequency sine wave can be
expressed as
s ~ t ! 5sin~ 2 p f c t !@ 11m sin~ 2 p f modt # ,

~1!

where t is time, m is the modulation depth, f mod is the modulation frequency, and f c is the carrier frequency. Such a signal will have energy only at f c and f c 6 f mod . However when
such a signal is passed through a nonlinear detection system,
such as a mammalian auditory system, the signal is demodulated and energy at f mod will appear. The evoked response to
such a signal contains significant energy at the modulation,
or envelope frequency of the stimulus ~Dolphin and Mountain, 1993!. Hence this phenomenon has been termed the
envelope flowing response ~EFR!. With such a continuous
signal, the rms value of the acoustic pressure can be readily
measured and associated with the behavioral measure of
hearing sensitivity measured at the carrier frequency.
Amplitude-modulated signals can also be produced by
summing two sinusoidal or tonal signals of different frequencies so that
s ~ t ! 5sin~ 2 p f 1 t ! 1sin~ 2 p f 2 t ! ,

~2!

where f 1 and f 2 are the frequencies of sinusoid 1 and 2,

respectively. A two-tone signal ~TT! contains energy only at
the frequencies of the individual tones. However, the amplitude of the signal will be modulated with a period corresponding to the absolute difference in the frequency between
the two tones, or f 2.1 5 u f 2 2 f 1 u .

FIG. 11. A false killer whale in a hoop station with suction cup electrodes
on its head, facing a J-9 sound projector.

Researchers from Boston University and the University

of Hawaii have been performing auditory-evoked potential
measurement, using continuous amplitude-modulated
stimuli, with three species of trained dolphins ~an Atlantic
bottlenose dolphin, a Rissos dolphin, and a false killer
whale! in an open water environment ~Dolphin et al., 1995!.
The animals are trained to wear suction cup electrodes and
station themselves within a hoop which has its center at a
depth of 1 m. The electrodes were silver discs ~2.4-cm diameter! imbedded in custom designed latex suction cups.
Evoked responses were recorded differentially from the scalp

FIG. 12. ~a! Example of the two-tone amplitude modulated acoustic stimulus in the time domain. ~b! Frequency domain representation of the two-tone signal.
~c! The evoked potential response in the time domain. ~d! The evoked potential response in the frequency domain ~from Dolphin et al., 1995!.
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source between the parietal ~noninverting! just posterior to

the blowhole and the mastoid ~inverting!. A ground electrode
was placed either on the melon or trunk region. A J-9 sound
projector is located 1 m from the animal as shown in Fig. 11.
The white suction cup electrodes on the animals head can
also be seen in the figure.
An example of the signal is depicted in the time domain
@Fig. 12~a!#, showing the amplitude modulation, and in the
frequency domain @Fig. 12~b!#, showing the energies at f 1
and f 2 . The evoked potential response averaged over 350
presentations of the stimulus in 164-ms blocks are shown in
Fig. 12~c! and ~d!. The nonlinear property of the auditory
system demodulates the amplitude modulated signal so that
energy at the modulated or envelope frequency is present in
the evoked response. The question of what would be the
most ideal relationship between the frequencies of the carrier
and modulated signals can be addressed by measuring the
modulation rate transfer function. This was determined for
dolphins by modulating the primary or carrier tone at frequencies between 184019 Hz and measuring the response
magnitude and phase at f mod or f 2.1 . Examples of the modulation rate transfer function obtained from Pseudorca using
both TT and SAM signals of different carrier and primary
frequencies are shown in Fig. 13. From the figure one can
see that there are fairly broad areas in which the magnitude
of the evoked potential responses does not vary much with
the modulation or difference frequency.
The hearing sensitivity of the three dolphin species to
low-frequency sounds ~below 1.6 kHz! has been determined
behaviorally and by measuring evoked potentials. The results
of both techniques agree very well, suggesting that determining hearing sensitivity with continuous amplitude modulated
sound may be a good method to measure the hearing sensitivity of large whales that are beached or trapped in nets.

the bite plate. Hearing thresholds were determined by presenting a sound stimulus at random intervals. Every time the
whale heard a sound, it would produce a whistle, which was
detected by another hydrophone and sent topside for analysis. The amplitude of the test tone was varied in a staircase
fashion. Echolocation was tested by presenting a target in
line with the whales longitudinal axis, 1 m away. Each time
a door hiding the target on the platform opened, the whale
was trained to echolocate. The animal responded by producing a whistle if it perceived the target or remained silent
when the target was absent. The echolocation signals were
detected by a hydrophone and the information sent topside
for recording.
Preliminary results indicate that the whales hearing sensitivity is not affected by the high pressure at depth. Their
thresholds are as good or slightly better at depth than at the
surface. However, preliminary results indicate that the ani-

VI. HEARING AND ECHOLOCATION BY DOLPHINS

AT DEPTH

Hearing and echolocation research on dolphins has been

conducted in shallow waters with the animals usually within
2 m of the surface. However, dolphins can dive to depths
greater than 300 m ~Ridgway and Kanwisher, 1969! and the
effects of a high-pressure environment on hearing and
echolocation is not known. Sam Ridgway and colleagues at
the Naval Command Control and Ocean Surveillance Center
in San Diego have been studying the effects of depth on the
hearing sensitivity and the echolocation signals of beluga
whales, Delphinapterus leucas ~Ridgway and Carder, 1995!.
Two beluga whales were trained to dive in the open
ocean and station on a test platform at depths of 6 ~surface!,
100, 200, and 300 m. The test platform with a whale approaching on a bite plate is shown in Fig. 14. On the test
platform are video cameras to monitor the animals behavior,
a sound projecting transducer for audiometric test, a sonar
target and a hydrophone to measure sonar signals. Ridgway
and his team have been working in the deep waters off San
Clemente Island. The whales are trained to boat follow an
inflatable boat to travel to the test site about 1 mile off shore.
A trial consisted of a whale diving to the test platform
suspended below the data collection boat and stationing on
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FIG. 13. Examples of the modulation rate transfer function obtained with a
false killer whale ~from Dolphin et al., 1995!.
Whitlow W. L. Au: Animal topics

2439

mals whistles at depth have larger high-frequency harmonics, as can be seen in Fig. 15. The lower frequency components of the whistles are present at depth and close to the
surface, however, the whistles at depth have larger high frequency harmonics than at the surface. The increased pressure
causes various air sacs and chambers in the nasal system to
change in shape and volume, affecting their resonance frequencies. This increase of whistle frequency may have important implications concerning the mechanisms that produce whistles. Specific sound generation mechanisms in
cetaceans have not yet been pin pointed. However many
theories exist, based mainly on anatomical considerations.
VII. DISCUSSION

FIG. 14. The test platform used to assess the hearing sensitivity and echolocation signals of beluga whales at depth ~courtesy of S. Ridgway!.

There has been an increasing interest on how man is

affecting the environment in which we live, work, and recreate. Animals are an important part of this environment, and
we are slowly becoming aware that our use of natural resources can have important consequences to many different
life forms. Therefore, interest in animal bioacoustics has
slowly been building over recent years. The U.S. military
~Air Force, Army, and Navy! all have some type of bioacoustics programs to study the effects of military exercises,
practices and operations on the environment of animals. The
Air Force has an extensive program to determine the effects

FIG. 15. Example of the beluga whale whistle at the surface and at depth ~courtesy of S. Ridgway!.
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of aircraft noise on a wide variety of animals, from Musk Ox

and Arctic foxes in the high northern latitudes to tortoise and
reptiles in desert locations, and birds such as the Peregrine
falcons in mountainous regions. The Army also has an extensive program studying their effects on the habitat of various endangered birds and animals. The Navy has a program
to determine the effects of low-frequency underwater sounds
on marine mammals. In these programs, various types of
new, small, portable, and state-of-the-art devices are being
developed to monitor location, physiological parameters, and
environmental noise in these programs ~Carter, 1996!. In order to monitor and study the effects of man-made noise on
different animal species, it is important to understand their
auditory capabilities. Therefore, within the militarys acoustic environmental assessment program, the basic hearing capabilities of different species are being determined. The militarys programs are too extensive and broad to be discussed
here.
Biologists are increasingly becoming aware of the usefulness of monitoring the vocalizations of animals as a good
way to study them in their natural habitat with a minimum of
disturbance. New electronic instrumentation that can be connected to laptop computers has been a boon to field researchers. The continual development of sophisticated signal processing and analysis software has made these tools accessible
to many researchers who would not even have dreamed of
using these techniques a decade ago. The maturing of satellite tracking and monitoring instrumentation has also contributed significantly in studying the life cycle and the use of
acoustics by animals in the wild. The use of sophisticated
laser techniques and other measurement techniques has increased the capabilities of many researchers to study the
acoustic functions of different species in new and innovative
ways. In short, we have entered into an exciting and interesting period for people interested in animal bioacoustics.
ACKNOWLEDGMENTS

The author acknowledges and publicly expresses his appreciation to several people who provided various written
materials, 35-mm slides, and data used in this paper. These
individuals are Dr. William Burgess of Stanford Research
Institute, Dr. Edmund Gerstein of Florida Atlantic University, Dr. Robert Hickling of the National Center for Physical
Acoustics, Dr. Burney LeBeouf of the University of California at Santa Cruz, Dr. Sam Ridgway of the Naval Command,
Control and Ocean Surveillance Center, and Dr. Daniel Robert of Cornell University. This is Hawaii Institute of Marine
Biology Contribution 1021.
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