As the only living species within the odobenid lineage of carnivores, walruses (Odobenus rosmarus) have no close relatives from which auditory information can be extrapolated. Sea lions and fur seals in the otariid lineage are the nearest evolutionary outgroup. To advance understanding of odobenid and otariid hearing, we conducted behavioral testing with two walruses and one California sea lion (Zalophus californianus). Detection thresholds for airborne sounds were measured from 0.08 to at least 16 kHz in ambient noise conditions and then re-measured in the presence of octave-band white masking noise. Walruses were more sensitive than the sea lion at lower frequencies and less sensitive at higher frequencies. Critical ratios for the walruses ranged from 20 dB at 0.2 kHz to 32 dB at 10 kHz, while critical ratios for the sea lion ranged from 16 dB at 0.2 kHz to 35 dB at 32 kHz. The masking values for these species are comparable to one another and to those of terrestrial carnivores, increasing by about 3 dB per octave with increasing frequency. Despite apparent differences in hearing range and sensitivity, odobenids and otariids have a similar ability to hear signals in noisy conditions.
REFERENCES
1.
Babushina
,
Y. S.
,
Zaslavski
,
G. L.
, and
Yurkevich
,
L. I.
(1991
). “
Air and underwater hearing characteristics of the northern fur seal: Audiograms, frequency and differential thresholds
,” Psychophysics
36
, 909
–913
.2.
Berta
,
A.
,
Churchill
,
M.
, and
Boessenecker
,
R. W.
(2018
). “
The origin and evolutionary biology of pinnipeds: Seals, sea lions, and walruses
,” Annu. Rev. Earth Planet. Sci.
46
, 203
–228
.3.
Boessenecker
,
R. W.
, and
Churchill
,
M.
(2018
). “
The last of the desmatophocid seals: A new species of Allodesmus from the upper Miocene of Washington, USA, and a revision of the taxonomy of Desmatophocidae
,” Zool. J. Linn. Soc.
184
, 211
–235
.4.
Branstetter
,
B. K.
, and
Sills
,
J. M.
(2022
). “
Mechanisms of auditory masking in marine mammals
,” Anim. Cogn.
25
, 1029
–1047
.5.
Branstetter
,
B. K.
,
Trickey
,
B. J.
,
Bakhtiari
,
K.
,
Black
,
A.
,
Aihara
,
H.
, and
Finneran
,
J. J.
(2013
). “
Auditory masking patterns in bottlenose dolphins (Tursiops truncatus) with natural, anthropogenic, and synthesized noise
,” J. Acoust. Soc. Am.
133
, 1811
–1818
.6.
Charrier
,
I.
(2021
). “
Non-vocal communication in otariids and odobenids
,” in Ethology and Behavorial Ecology of Otariids and the Odobenid,
Ethology and Behavioral Ecology of Marine Mammals, edited by C. Campagna and R. Harcourt (Springer
, Cham, Switzerland
), pp. 265
–289
.7.
Charrier
,
I.
,
Aubin
,
T.
, and
Mathevon
,
N.
(2010
). “
Mother-calf vocal communication in Atlantic walrus: A first field experimental study
,” Anim. Cogn.
13
, 471
–482
.8.
Cunningham
,
K. A.
,
Hayes
,
S. A.
,
Wargo Rub
,
A. M.
, and
Reichmuth
,
C.
(2014a
). “
Auditory detection of ultrasonic coded transmitters by seals and sea lions
,” J. Acoust. Soc. Am.
135
, 1978
–1985
.9.
Cunningham
,
K. A.
, and
Reichmuth
,
C.
(2016
). “
High-frequency hearing in seals and sea lions
,” Hear. Res.
331
, 83
–91
.10.
Cunningham
,
K. A.
,
Southall
,
B. L.
, and
Reichmuth
,
C.
(2014b
). “
Auditory sensitivity of seals and sea lions in complex listening scenarios
,” J. Acoust. Soc. Am.
136
, 3410
–3421
.11.
Dooling
,
R. J.
, and
Blumenrath
,
S. H.
(2013
). “
Avian sound perception in noise
,” in Animal Communication and Noise
, edited by
H.
Brumm
(
Springer
,
New York
), pp. 229
–250
.12.
Erbe
,
C.
,
Reichmuth
,
C.
,
Cunningham
,
K.
,
Lucke
,
K.
, and
Dooling
,
R.
(2016
). “
Communication masking in marine mammals: A review and research strategy
,” Mar. Pollut. Bull.
103
, 15
–38
.13.
Fay
,
F. H.
(1982
). “
Ecology and biology of the Pacific walrus, Odobenus rosmarus divergens
,” North Am. Fauna
74
, 1
–279
.14.
Fay
,
R. R.
(1988
). “
Comparative psychoacoustics
,” Hear. Res.
34
, 295
–306
.15.
Finneran
,
J. J.
(2003
). “
An Integrated Computer-Controlled System for Marine Mammal Auditory Testing
,” Technical Report
(
Space and Naval Warfare Systems Center
,
San Diego, CA
).16.
Finneran
,
J. J.
(2015
). “
Noise-induced hearing loss in marine mammals: A review of temporary threshold shift studies from 1996 to 2015
,” J. Acoust. Soc. Am.
138
, 1702
–1726
.17.
Fletcher
,
H.
(1940
). “
Auditory patterns
,” Rev. Mod. Phys.
12
, 47
–65
.18.
Hanke
,
F. D.
, and
Reichmuth
,
C.
(2022
). “
Phocid sensory systems and cognition
,” in Ethology and Behavorial Ecology of Marine Mammals
, edited by
D. P.
Costa
and
E.
McHuron
(
Springer
,
Cham, Switzerland
), pp. 31
–68
.19.
Hanke
,
F. D.
,
Reichmuth
,
C.
, and
Cook
,
P.
(2021
). “
The sensory world of otariids
,” in Ethology and Behavorial Ecology of Otariids and the Odobenid
, edited by
C.
Campagna
and
R. G.
Harcourt
(
Springer
,
Cham, Switzerland
), pp. 305
–321
.20.
Hawkins
,
T. E.
, and
Stevens
,
S. S.
(1950
). “
The masking of pure tones and of speech by white noise
,” J. Acoust. Soc. Am.
22
, 6
–13
.21.
Heffner
,
H. E.
, and
Heffner
,
R. S.
(2008
). “
High-frequency hearing
,” in Handbook of the Senses: Audition
, edited by
P.
Dallos
,
D.
Oertel
, and
R.
Hoy
(
Elsevier
,
New York
), pp. 55
–60
.22.
Hienz
,
R. D.
, and
Sachs
,
M. B.
(1987
). “
Effects of noise on pure-tone thresholds in blackbirds (Agelaius phoeniceus and Molothrus ater) and pigeons (Columba livia)
,” J. Comp. Psychol.
101
, 16
–24
.23.
Holt
,
M. M.
, and
Schusterman
,
R. J.
(2007
). “
Spatial release from masking of aerial tones in pinnipeds
,” J. Acoust. Soc. Am.
121
, 1219
–1225
.24.
Houtsma
,
A. J. M.
(2005
). “
A note on pure-tone masking by broadband noise under free-field and insert-phone conditions
,” J. Acoust. Soc. Am.
117
, 490
–491
.25.
Kastak
,
D.
, and
Schusterman
,
R. J.
(1998
). “
Low-frequency amphibious hearing in pinnipeds: Methods, measurements, noise, and ecology
,” J. Acoust. Soc. Am.
103
, 2216
–2228
.26.
Kastak
,
D.
, and
Schusterman
,
R. J.
(2002
). “
Changes in auditory sensitivity with depth in a free-diving California sea lion (Zalophus californianus)
,” J. Acoust. Soc. Am.
112
, 329
–333
.27.
Kastelein
,
R. A.
,
Dubbeldam
,
J. L.
,
de Bakker
,
M. A. G.
, and
Gerrits
,
N. M.
(1996b
). “
The anatomy of the walrus head (Odobenus rosmarus). Part 4: The ears and their function in aerial and underwater hearing
,” Aquat. Mamm.
22
(2
), 95
–125
.28.
Kastelein
,
R. A.
,
Mosterd
,
P.
,
van Ligtenberg
,
C. L.
, and
Verboom
,
W. C.
(1996a
). “
Aerial hearing sensitivity tests with a male Pacific walrus (Odobenus rosmarus divergens), in the free field and with headphones
,” Aquat. Mamm.
22
, 81
–93
.29.
Kastelein
,
R. A.
,
Mosterd
,
P.
,
van Santen
,
B.
,
Hagedoorn
,
M.
, and
de Haan
,
D.
(2002
). “
Underwater audiogram of a Pacific walrus (Odobenus rosmarus divergens) measured with narrow-band frequency-modulated signals
,” J. Acoust. Soc. Am.
112
, 2173
–2182
.30.
Kastelein
,
R. A.
,
Van Ligtenberg
,
C. L.
,
Gjertz
,
I.
, and
Verboom
,
W. C.
(1993
). “
Free field hearing tests on wild Atlantic walruses (Odobenus rosmarus rosmarus) in air
,” Aquat. Mamm.
19
, 143
–148
.31.
Kastelein
,
R. A.
,
van Schie
,
R.
,
Verboom
,
W. C.
, and
de Haan
,
D.
(2005
). “
Underwater hearing sensitivity of a male and a female Steller sea lion (Eumetopias jubatus)
,” J. Acoust. Soc. Am.
118
, 1820
–1829
.32.
Langemann
,
U.
,
Gauger
,
B.
, and
Klump
,
G. M.
(1998
). “
Auditory sensitivity in the great tit: Perception of signals in the presence and absence of noise
,” Anim. Behav.
56
, 763
–769
.33.
MacCracken
,
J. G.
,
Beatty
,
W. S.
,
Garlich-Miller
,
J. L.
,
Kissling
,
M. L.
, and
Snyder
,
J. A.
(2017
). “
Final species status assessment for the Pacific walrus (Odobenus romarus divergins), May 2017 (version 1.0)
,” (
US Fish and Wildlife Service Marine Mammals Management
,
Anchorage, AK
).34.
Miller
,
E. H.
(1985
). “
Airborne acoustic communication in the walrus Odobenus rosmarus
,” Natl. Geogr. Res.
1
, 124
–145
.35.
Miller
,
E. H.
, and
Kochnev
,
A. A.
(2021
). “
Ethology and behavioral ecology of the walrus (Odobenus rosmarus), with emphasis on communication and social behavior
,” in Ethology and Behavioral Ecology of Otariids and the Odobenid,
Ethology and Behavioral Ecology of Marine Mammals, edited by C. Campagna and R. Harcourt (Springer, Cham, Switzerland), pp. 437
–488
.36.
Mooney
,
T. A.
,
Yamato
,
M.
, and
Branstetter
,
B. K.
(2012
). “
Hearing in cetaceans: From natural history to experimental biology
,” Adv. Mar. Biol.
63
, 197
–246
.37.
Moore
,
B. C. J.
(1993
). “
Frequency analysis and pitch perception
,” in Human Psychophysics
, edited by
W.
Yost
,
A.
Popper
, and
R. R.
Fay
(
Springer-Verlag
,
New York
), pp. 56
–115
.38.
Moore
,
P. W. B.
, and
Schusterman
,
R. J.
(1987
). “
Audiometric assessment of northern fur seals, Callorhinus ursinus
,” Mar. Mamm. Sci.
3
, 31
–53
.39.
Mouy
,
X.
, and
Hannay
,
D.
(2012
). “
Tracking of Pacific walruses in the Chukchi Sea using a single hydrophone
,” J. Acoust. Soc. Am.
131
, 1349
–1358
.40.
Mulsow
,
J.
,
Finneran
,
J. J.
, and
Houser
,
D. S.
(2011
). “
California sea lion (Zalophus californianus) aerial hearing sensitivity measured using auditory steady-state response and psychophysical methods
,” J. Acoust. Soc. Am.
129
, 2298
–2306
.41.
Mulsow
,
J.
,
Houser
,
D. S.
, and
Finneran
,
J. J.
(2012
). “
Underwater psychophysical audiogram of a young male California sea lion (Zalophus californianus)
,” J. Acoust. Soc. Am.
131
, 4182
–4187
.42.
Mulsow
,
J.
, and
Reichmuth
,
C.
(2010
). “
Psychophysical and electrophysiological aerial audiograms of a Steller sea lion (Eumetopias jubatus)
,” J. Acoust. Soc. Am.
127
, 2692
–2701
.43.
Nummela
,
S.
(2008
). “
Hearing in aquatic mammals
,” in Sensory Evolution on the Threshold
, edited by
J. G.
Thewissen
and
S.
Nummella
(
University of California
,
Berkeley
), pp. 211
–232
.44.
Reichmuth
,
C.
(2012
). “
Psychophysical studies of auditory masking in marine mammals: Key concepts and new directions
,” Adv. Exp. Med. Biol.
730
, 23
–27
.45.
Reichmuth
,
C.
,
Holt
,
M. M.
,
Mulsow
,
J.
,
Sills
,
J. M.
, and
Southall
,
B. L.
(2013
). “
Comparative assessment of amphibious hearing in pinnipeds
,” J. Comp. Physiol. A Neuroethol. Sens. Neural Behav. Physiol.
199
, 491
–507
.46.
Reichmuth
,
C.
,
Sills
,
J. M.
,
Brewer
,
A.
,
Triggs
,
L.
,
Ferguson
,
R.
,
Ashe
,
E.
, and
Williams
,
R.
(2020
). “
Behavioral assessment of in-air hearing range for the Pacific walrus (Odobenus rosmarus divergens)
,” Polar Biol.
43
, 767
–772
.47.
Reichmuth
,
C.
,
Sills
,
J. M.
, and
Ghoul
,
A.
(2017
). “
Psychophysical audiogram of a California sea lion listening for airborne tonal sounds in an acoustic chamber
,” Proc. Mtgs. Acoust.
30
, 010001
.48.
Reichmuth
,
C.
, and
Southall
,
B. L.
(2012
). “
Underwater hearing in California sea lions (Zalophus californianus): Expansion and interpretation of existing data
,” Mar. Mamm. Sci.
28
, 358
–363
.49.
Renouf
,
D.
(1980
). “
Masked hearing thresholds of harbour seals (Phoca vitulina) in air
,” J. Aud. Res.
20
, 263
–269
.50.
Repenning
,
C. A.
(1972
). “
Underwater hearing in seals: Functional morphology
,” in Functional Anatomy of Marine Mammals
, edited by
R. J.
Harrison
(
Academic
,
London
), pp. 307
–331
.51.
Schusterman
,
R. J.
(1974
). “
Auditory sensitivity of a California sea lion to airborne sound
,” J. Acoust. Soc. Am.
56
, 1248
–1251
.52.
Schusterman
,
R. J.
, and
Balliet
,
R. F.
(1969
). “
Underwater barking by male sea lions (Zalophus californianus)
,” Nature
222
, 1179
–1181
.53.
Schusterman
,
R. J.
,
Balliet
,
R. F.
, and
Nixon
,
J.
(1972
). “
Underwater audiogram of the California sea lion by the conditioned vocalization technique
,” J. Exp. Anal. Behav.
17
, 339
–350
.54.
Sills
,
J. M.
,
Southall
,
B. L.
, and
Reichmuth
,
C.
(2014
). “
Amphibious hearing in spotted seals (Phoca largha): Underwater audiograms, aerial audiograms and critical ratio measurements
,” J. Exp. Biol.
217
, 726
–734
.55.
Southall
,
B. L.
,
Finneran
,
J. J.
,
Reichmuth
,
C.
,
Nachtigall
,
P. E.
,
Ketten
,
D. R.
,
Bowles
,
A. E.
,
Ellison
,
W. T.
,
Nowacek
,
D. P.
, and
Tyack
,
P. L.
(2019
). “
Marine mammal noise exposure criteria: Updated scientific recommendations for residual hearing effects
,” Aquat. Mamm.
45
, 125
–232
.56.
Southall
,
B. L.
,
Schusterman
,
R. J.
, and
Kastak
,
D.
(2000
). “
Masking in three pinnipeds: Underwater, low-frequency critical ratios
,” J. Acoust. Soc. Am.
108
, 1322
–1326
.57.
Southall
,
B. L.
,
Schusterman
,
R. J.
, and
Kastak
,
D.
(2003
). “
Auditory masking in three pinnipeds: Aerial critical ratios and direct critical bandwidth measurements
,” J. Acoust. Soc. Am.
114
, 1660
–1666
.58.
Stebbins
,
W. C.
(1970
). “
Principles of animal psychophysics
,” in Animal Psychophysics: The Design and Conduct of Sensory Experiments
, edited by
W. C.
Stebbins
(
Springer
,
Boston
), pp. 1
–19
.59.
Turnbull
,
S. D.
, and
Terhune
,
J. M.
(1990
). “
White noise and pure tone masking of pure tone thresholds of a harbour seal listening in air and underwater
,” Can. J. Zool.
68
, 2090
–2097
.60.
Yost
,
W. A.
(2000
). Fundementals of Hearing: An Introduction
, 4th ed
. (
Academic
,
San Diego, CA
).© 2023 Acoustical Society of America.
2023
Acoustical Society of America
You do not currently have access to this content.