Eight years of passive acoustic data (2007–2014) from the Beaufort Sea were used to estimate the mean cue rate (calling rate) of individual bowhead whales (Balaena mysticetus) during their fall migration along the North Slope of Alaska. Calls detected on directional acoustic recorders (DASARs) were triangulated to provide estimates of locations at times of call production, which were then translated into call densities (calls/h/km2). Various assumptions were used to convert call density into animal cue rates, including the time for whales to cross the arrays of acoustic recorders, the population size, the fraction of the migration corridor missed by the localizing array system, and the fraction of the seasonal migration missed because recorders were retrieved before the end of the migration. Taking these uncertainties into account in various combinations yielded up to 351 cue rate estimates, which summarize to a median of 1.3 calls/whale/h and an interquartile range of 0.5–5.4 calls/whale/h.

1.
Alaska Fisheries Science Center
(
2021
). “
AFSC/MML: Marine mammal aerial surveys in the Bering, Chukchi, and Beaufort Seas, and Amundsen Gulf, 1979-2019
,” https://www.fisheries.noaa.gov/inport/item/17338 (Last viewed 17 May 2021).
2.
Ashjian
,
C. J.
,
Braund
,
S. R.
,
Campbell
,
R. G.
,
George
,
J. C.
,
Kruse
,
J.
,
Maslowski
,
W.
,
Moore
,
S. E.
,
Nicolson
,
C. R.
,
Okkonen
,
S. R.
,
Sherr
,
B. F.
,
Sherr
,
E. B.
, and
Spitz
,
Y. H.
(
2010
). “
Climate variability, oceanography, bowhead whale distribution, and Iñupiat subsistence whaling near Barrow, Alaska
,”
Arctic
63
,
179
194
.
3.
Bivand
,
R. S.
,
Keitt
,
T.
, and
Rowlingson
,
B.
(
2019
). “
rgdal: Bindings for the ‘geospatial’ data abstraction library
,” R package version 1.4-8, https://CRAN.R-project.org/package=rgdal (Last viewed 1 March 2021).
4.
Bivand
,
R.
, and
Lewin-Koh
,
N.
(
2019
). “
maptools: Tools for reading and handling spatial objects
,” R package version 0.9-9, https://CRAN.R-project.org/package=maptools (Last viewed 1 March 2021).
5.
Bivand
,
R. S.
,
Pebesma
,
E. J.
, and
Gomez-Rubio
,
V.
(
2013
).
Applied Spatial Data Analysis with R
(
Springer
,
New York
).
6.
Bivand
,
R.
, and
Rundel
,
C.
(
2019
). “
rgeos: Interface to geometry engine–open source (‘GEOS’)
,” R package version 0.5-2, https://CRAN.R-project.org/package=rgeos (Last viewed 1 March 2021).
7.
Blackwell
,
S. B.
,
Conrad
,
A. S.
, and
Thode
,
A. M.
(
2014
). “
Characterizing underwater sounds at six locations in the U.S. Beaufort Sea during winter months
,” Joint Monitoring Program in the Chukchi and Beaufort Seas, 2012, LGL Alaska Final Report P1272-2 for Shell Offshore, Inc., ION Geophysical, Inc., and Other Industry Contributors (National Marine Fisheries Service and U.S. Fish and Wildlife Service, Washington, DC), Appendix C.
8.
Blackwell
,
S. B.
,
Nations
,
C. S.
,
McDonald
,
T. L.
,
Greene
,
C. R.
, Jr.
,
Thode
,
A. M.
,
Guerra
,
M.
, and
Macrander
,
A. M.
(
2013
). “
Effects of airgun sounds on bowhead whale calling rates in the Alaskan Beaufort Sea
,”
Mar. Mamm. Sci
29
,
E342
E365
.
9.
Blackwell
,
S. B.
,
Nations
,
C. S.
,
McDonald
,
T. L.
,
Thode
,
A. M.
,
Mathias
,
D.
,
Kim
,
K. H.
,
Greene
,
C. R.
, Jr.
, and
Macrander
,
A. M.
(
2015
). “
Effects of airgun sounds on bowhead whale calling rates: Evidence for two behavioral thresholds
,”
PLoS One
10
,
e0125720
.
10.
Blackwell
,
S. B.
,
Nations
,
C. S.
,
Thode
,
A. M.
,
Kauffman
,
M. E.
,
Conrad
,
A. S.
,
Norman
,
R. G.
, and
Kim
,
K. H.
(
2017
). “
Effects of tones associated with drilling activities on bowhead whale calling rates
,”
PLoS One
12
,
e0188459
.
11.
Blackwell
,
S. B.
,
Richardson
,
W. J.
,
Greene
,
C. R.
, Jr.
, and
Streever
,
B.
(
2007
). “
Bowhead whale (Balaena mysticetus) migration and calling behaviour in the Alaskan Beaufort Sea, autumn 2001–04: An acoustic localization study
,”
Arctic
60
,
255
270
.
12.
Braham
,
H. W.
,
Fraker
,
M. A.
, and
Krogman
,
B. D.
(
1980
). “
Spring migration of the western Arctic population of bowhead whales
,”
Mar. Fish. Rev.
42
,
36
46
.
13.
Calambokidis
,
J.
,
Fahlbusch
,
J. A.
,
Szesciorka
,
A. R.
,
Southall
,
B. L.
,
Cade
,
D. E.
,
Friedlaender
,
A. S.
, and
Goldbogen
,
J. A.
(
2019
). “
Differential vulnerability to ship strikes between day and night for blue, fin, and humpback whales based on dive and movement data from medium duration archival tags
,”
Front. Mar. Sci.
6
,
543
.
14.
Citta
,
J. J.
,
Quakenbush
,
L. T.
,
Okkonen
,
S. R.
,
Druckenmiller
,
M. L.
,
Maslowski
,
W.
,
Clement-Kinney
,
J.
,
George
,
J. C.
,
Brower
,
H.
,
Small
,
R. J.
,
Ashjian
,
C. J.
,
Harwood
,
L. A.
, and
Heide-Jørgensen
,
M. P.
(
2015
). “
Ecological characteristics of core-use areas used by Bering–Chukchi–Beaufort (BCB) bowhead whales, 2006–2012
,”
Prog. Oceanogr.
136
,
201
222
.
15.
Clarke
,
J. T.
,
Brower
,
A. A.
,
Christman
,
C. L.
, and
Ferguson
,
M. C.
(
2014
). “
Distribution and relative abundance of marine mammals in the Northeastern Chukchi and Western Beaufort seas, 2013: Annual report
,” OCS Study BOEM 2014-018 (U.S. Department of the Interior, Anchorage, AK).
16.
Clarke
,
J. T.
,
Ferguson
,
M. C.
,
Willoughby
,
A. L.
, and
Brower
,
A. A.
(
2018
). “
Bowhead and beluga whale distributions, sighting rates, and habitat associations in the western Beaufort Sea in summer and fall 2009–16, with comparison to 1982–91
,”
Arctic
71
,
115
138
.
17.
Delarue
,
J.
,
Laurinolli
,
M.
, and
Martin
,
B.
(
2009
). “
Bowhead whale (Balaena mysticetus) songs in the Chukchi Sea between October 2007 and May 2008
,”
J. Acoust. Soc. Am.
126
,
3319
3328
.
18.
Dunn
,
P. K.
, and
Smith
,
G. K.
(
2005
). “
Series evaluation of Tweedie exponential dispersion model densities
,”
Stat. Comput.
15
,
267
280
.
19.
Ferguson
,
M. C.
,
Clarke
,
J. T.
,
Brower
,
A. A.
,
Willoughby
,
A. L.
, and
Okkonen
,
S. R.
(
2021
). “
Ecological variation in the Western Beaufort Sea
,” in
The Bowhead Whale Balaena mysticetus: Biology and Human Interactions
, edited by
J. C.
George
and
J. G. M.
Thewissen
(
Elsevier
,
London
), pp.
365
379
.
20.
Givens
,
G. H.
,
Edmondson
,
S. L.
,
George
,
J. C.
,
Suydam
,
R.
,
Charif
,
R. A.
,
Rahaman
,
A.
,
Hawthorne
,
D.
,
Tudor
,
B.
,
DeLong
,
R. A.
, and
Clark
,
C. W.
(
2013
). “
Estimate of 2011 abundance of the Bering-Chukchi-Beaufort Seas bowhead whale population
,” Report SC/65a/BRG01 to the Scientific Committee of the International Whaling Commission (International Whaling Commission, Cambridge, UK).
21.
Goldbogen
,
J. A.
,
Stimpert
,
A. K.
,
DeRuiter
,
S. L.
,
Calambokidis
,
J.
,
Friedlaender
,
A. S.
,
Schorr
,
G. S.
,
Moretti
,
D. J.
,
Tyack
,
P. L.
, and
Southall
,
B. L.
(
2014
). “
Using accelerometers to determine the calling behavior of tagged baleen whales
,”
J. Exp. Biol.
217
,
2449
2455
.
22.
Greene
,
C. R.
, Jr.
,
McLennan
,
M. W.
,
Norman
,
R. G.
,
McDonald
,
T. L.
,
Jakubczak
,
R. S.
, and
Richardson
,
W. J.
(
2004
). “
Directional frequency and recording (DIFAR) sensors in seafloor recorders to locate calling bowhead whales during their fall migration
,”
J. Acoust. Soc. Am.
116
,
799
813
.
23.
Guazzo
,
R. A.
,
Weller
,
D. W.
,
Europe
,
H. M.
,
Durban
,
J. W.
,
D'Spain
,
G. L.
, and
Hildebrand
,
J. A.
(
2019
). “
Migrating eastern North Pacific gray whale call and blow rates estimated from acoustic recordings, infrared camera video, and visual sightings
,”
Sci. Rep.
9
,
12517
.
24.
Harris
,
D. V.
,
Miksis-Olds
,
J. L.
,
Vernon
,
J. A.
, and
Thomas
,
L.
(
2018
). “
Fin whale density and distribution estimation using acoustic bearings derived from sparse arrays
,”
J. Acoust. Soc. Am.
143
,
2980
2993
.
25.
Harwood
,
L. A.
,
Quakenbush
,
L. T.
,
Small
,
R. J.
,
George
,
J. C.
,
Pokiak
,
J.
,
Pokiak
,
C.
,
Heide-Jørgensen
,
M. P.
,
Lea
,
E. V.
, and
Brower
,
H.
(
2017
). “
Movements and inferred foraging by bowhead whales in the Canadian Beaufort Sea during August and September, 2006–12
,”
Arctic
70
,
161
176
.
26.
Hijmans
,
R. J.
(
2020
). “
raster: Geographic data analysis and modeling
,” R package version 3.0.2, http://CRAN.R-project.org/package=raster (Last viewed 1 March 2021).
27.
Huntington
,
H. P.
, and
Quakenbush
,
L. T.
(
2009
). “
Traditional knowledge of bowhead whale migratory patterns near Kaktovik and Barrow, Alaska
,” Final report to the Barrow and Kaktovik Whaling Captains Associations, the Alaska Eskimo Whaling Commission, ConocoPhillips, and the Minerals Management Service (Alaska Department of Fish and Game, Juneau, AK).
28.
Johnson
,
H. D.
,
Stafford
,
K. M.
,
George
,
J. C.
,
Ambrose
,
W. G.
, Jr.
, and
Clark
,
C. W.
(
2015
). “
Song sharing and diversity in the Bering-Chukchi-Beaufort population of bowhead whales (Balaena mysticetus), spring 2011
,”
Mar. Mamm. Sci.
31
,
902
922
.
29.
Johnson
,
M.
(
2014
). “
On-animal methods for studying echolocation in free-ranging animals
,” in
Biosonar
, edited by
A.
Surlykke
,
P. E.
Nachtigall
,
R. R.
Fay
, and
A. N.
Popper
(
Springer-Verlag
,
New York
), pp.
195
229
.
30.
Kim
,
K. H.
,
Norman
,
R. G.
,
Burgess
,
W. C.
,
Blackwell
,
S. B.
, and
Greene
,
C. R.
, Jr.
(
2014
). “
Acoustic monitoring of bowhead whale migration, autumn 2013
,” in
Monitoring of Industrial Sounds, Seals, and Bowhead Whales near BP's Northstar Oil Development, Alaskan Beaufort Sea, 2013: Annual Summary Report
, edited by
W. J.
Richardson
and
K. H.
Kim
(
BP Exploration (Alaska) Inc
.,
Anchorage, AK
).
31.
Koski
,
W. R.
, and
Miller
,
G. W.
(
2009
). “
Habitat use by different size classes of bowhead whales in the central Beaufort Sea during late summer and autumn
,”
Arctic
62
,
137
150
.
32.
Marques
,
T. A.
,
Munger
,
L.
,
Thomas
,
L.
,
Wiggins
,
S.
, and
Hildebrand
,
J. A.
(
2011
). “
Estimating North Pacific right whale Eubalaena japonica density using passive acoustic cue counting
,”
Endang. Species Res.
13
,
163
172
.
33.
Marques
,
T. A.
,
Thomas
,
L.
,
Martin
,
S. W.
,
Mellinger
,
D. K.
,
Ward
,
J. A.
,
Moretti
,
D. J.
,
Harris
,
D.
, and
Tyack
,
P. L.
(
2013
). “
Estimating animal population density using passive acoustics
,”
Biol. Rev.
88
,
287
309
.
34.
Martin
,
S. W.
,
Marques
,
T. A.
,
Thomas
,
L.
,
Morrissey
,
R. P.
,
Jarvis
,
S.
,
DiMarzio
,
N.
,
Moretti
,
D.
, and
Mellinger
,
D. K.
(
2013
). “
Estimating minke whale (Balaenoptera acutorostrata) boing sound density using passive acoustic sensors
,”
Mar. Mamm. Sci.
29
,
142
158
.
35.
Mate
,
B. R.
,
Krutzikowsky
,
G. K.
, and
Winsor
,
M. H.
(
2000
). “
Satellite-monitored movements of radio-tagged bowhead whales in the Beaufort and Chukchi seas during the late-summer feeding season and fall migration
,”
Can. J. Zool.
78
,
1168
1181
.
36.
May-Collado
,
L. J.
, and
Quiñones-Lebrón
,
S. G.
(
2014
). “
Dolphin changes in whistle structure with watercraft activity depends on their behavioral state
,”
J. Acoust. Soc. Am.
135
,
EL193
.
37.
Miller
,
G. W.
,
Elliott
,
R. E.
, and
Richardson
,
W. J.
(
1996
). “
Marine mammal distribution, numbers and movements
,” in
Northstar Marine Mammal Monitoring Program, 1995: Baseline Surveys and Retrospective Analyses of Marine Mammal and Ambient Noise Data from the Central Alaskan Beaufort Sea
(
BP Exploration (Alaska) Inc.
,
Anchorage, AK
), pp.
3
72
.
38.
Moore
,
S. E.
,
DeMaster
,
D. P.
, and
Dayton
,
P. K.
(
2000
). “
Cetacean habitat selection in the Alaskan Arctic during summer and autumn
,”
Arctic
53
,
341
480
.
39.
Moore
,
S. E.
, and
Reeves
,
R. R.
(
1993
). “
Distribution and movement
,” in
The Bowhead Whale
, edited by
J. J.
Burns
,
J. J.
Montague
, and
C. J.
Cowles
(
Allen Press
,
Lawrence, KS
), pp.
313
386
.
40.
National Snow and Ice Data Center
(
2021
). https://nsidc.org/ (Last viewed 17 May 2021).
41.
Noad
,
M. J.
,
Dunlop
,
R. A.
, and
Mack
,
A. K.
(
2017
). “
Changes in humpback whale singing behavior with abundance: Implications for the development of acoustic surveys of cetaceans
,”
J. Acoust. Soc. Am.
142
,
1611
1618
.
42.
Okkonen
,
S. R.
,
Clarke
,
J. T.
, and
Potter
,
R. A.
(
2018
). “
Relationships among high river discharges, upwelling events, and bowhead whale (Balaena mysticetus) occurrence in the central Alaskan Beaufort Sea
,”
Deep-Sea Res. Pt. II
152
,
195
202
.
43.
Olnes
,
J.
,
Citta
,
J. J.
,
Quakenbush
,
L. T.
,
George
,
J. C.
,
Harwood
,
L. A.
,
Lea
,
E. V.
, and
Heide-Jørgensen
,
M. P.
(
2020
). “
Use of the Alaskan Beaufort Sea by bowhead whales (Balaena mysticetus) tagged with satellite transmitters, 2006–18
,”
Arctic
73
,
278
291
.
44.
Pebesma
,
E. J.
, and
Bivand
,
R. S.
(
2005
). “
Classes and methods for spatial data in R
,”
R News
, http://cran.r-project.org/doc/Rnews/ (Last viewed 1 March 2021).
45.
Quakenbush
,
L.
,
Citta
,
J.
,
George
,
J. C.
,
Heide-Jørgensen
,
M. P.
,
Small
,
R.
,
Brower
,
H.
,
Harwood
,
L.
,
Adams
,
B.
,
Brower
,
L.
,
Tagarook
,
G.
,
Pokiak
,
C.
, and
Pokiak
,
J.
(
2012
). “
Seasonal movements of the Bering-Chukchi-Beaufort stock of bowhead whales: 2006–2011 satellite telemetry results
,” Report SC/64/BRG1 to the scientific committee of the International Whaling Commission (International Whaling Commission, Cambridge, UK).
46.
R Core Team
(
2019
). “
R: A language and environment for statistical computing
,” R Foundation for Statistical Computing, Vienna, Austria, http://www.R-project.org/ (Last viewed 1 March 2021).
47.
Rugh
,
D. J.
, and
Cubbage
,
J. C.
(
1980
). “
Migration of bowhead whales past Cape Lisburne, Alaska
,”
Mar. Fish. Rev.
42
(9–10),
46
51
.
48.
Simon
,
M.
,
Johnson
,
M.
,
Tyack
,
P.
, and
Madsen
,
P. T.
(
2009
). “
Behaviour and kinematics of continuous ram filtration in bowhead whales (Balaena mysticetus
),”
Proc. R. Soc. B
276
,
3819
3828
.
49.
Stafford
,
K. M.
,
Lydersen
,
C.
,
Wiig
,
Ø.
, and
Kovacs
,
K. M.
(
2018
). “
Extreme diversity in the songs of Spitsbergen's bowhead whales
,”
Biol. Lett.
14
,
20180056
.
50.
Thode
,
A. M.
,
Blackwell
,
S. B.
,
Conrad
,
A. S.
,
Kim
,
K. H.
,
Marques
,
T.
,
Thomas
,
L.
,
Oedekoven
,
C. S.
,
Harris
,
D.
, and
Bröker
,
K.
(
2020
). “
Roaring and repetition: How bowhead whales adjust their call density and source level (Lombard effect) in the presence of natural and seismic airgun survey noise
,”
J. Acoust. Soc. Am.
147
,
2061
2080
.
51.
Thode
,
A. M.
,
Blackwell
,
S. B.
,
Seger
,
K. D.
,
Conrad
,
A. S.
,
Kim
,
K. H.
, and
Macrander
,
A. M.
(
2016
). “
Source level and calling depth distributions of migrating bowhead whale calls in the shallow Beaufort Sea
,”
J. Acoust. Soc. Am.
140
,
4288
4297
.
52.
Thode
,
A. M.
,
Kim
,
K. H.
,
Blackwell
,
S. B.
,
Greene
,
C. R.
, Jr.
,
Nations
,
C. S.
,
McDonald
,
T. L.
, and
Macrander
,
A. M.
(
2012
). “
Automated detection and localization of bowhead whale sounds in the presence of seismic airgun surveys
,”
J. Acoust. Soc. Am.
131
,
3726
3747
.
53.
Tweedie
,
M. C. K.
(
1984
). “
An index which distinguishes between some important exponential families
,” in
Statistics: Applications and New Directions: Proceedings of the Indian Statistical Institute Golden Jubilee International Conference
, edited by
J. K.
Ghosh
and
J.
Roy
(
Indian Statistical Institute
,
Calcutta, India
), pp.
579
604
.
54.
Ver Hoef
,
J. M.
, and
Boveng
,
P. L.
(
2007
). “
Quasi-Poisson vs. negative binomial regression: How should we model overdispersed count data?
,”
Ecology
88
,
2766
2772
.
55.
Warren
,
V. E.
,
Marques
,
T. A.
,
Harris
,
D.
,
Thomas
,
L.
,
Tyack
,
P. L.
,
Aguilar de Soto
,
N.
,
Hickmott
,
L. S.
, and
Johnson
,
M. P.
(
2017
). “
Spatio-temporal variation in click production rates of beaked whales: Implications for passive acoustic density estimation
,”
J. Acoust. Soc. Am.
141
,
1962
1974
.
56.
Warren
,
V. E.
,
Miller
,
P. J. O.
, and
Tyack
,
P. L.
(
2020
). “
Short-term responses of sperm whales Physeter macrocephalus to the attachment of suction cup tags
,”
Mar. Ecol. Prog. Ser.
645
,
219
234
.
57.
Wood
,
S. N.
(
2017
).
Generalized Additive Models: An Introduction with R
(
CRC
,
Boca Raton, FL
).
58.
Würsig
,
B.
, and
Clark
,
C.
(
1993
). “
Behavior
,” in
The Bowhead Whale
, edited by
J. J.
Burns
,
J. J.
Montague
, and
C. J.
Cowles
(
Allen Press
,
Lawrence, KS
), pp.
157
199
.
59.
Zeh
,
J. E.
,
Clark
,
C. W.
,
George
,
J. C.
,
Withrow
,
D.
,
Carroll
,
G. M.
, and
Koski
,
W. R.
(
1993
). “
Current population size and dynamics
,” in
The Bowhead Whale
, edited by
J. J.
Burns
,
J. J.
Montague
, and
C. J.
Cowles
(
Allen Press
,
Lawrence, KS
), pp.
409
489
.

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