Sperm whales produce codas for communication that can be grouped into different types according to their temporal patterns. Codas have led researchers to propose that sperm whales belong to distinct cultural clans, but it is presently unclear if they also convey individual information. Coda clicks comprise a series of pulses and the delay between pulses is a function of organ size, and therefore body size, and so is one potential source of individual information. Another potential individual-specific parameter could be the inter-click intervals within codas. To test whether these parameters provide reliable individual cues, stereo-hydrophone acoustic tags (Dtags) were attached to five sperm whales of the Azores, recording a total of 802 codas. A discriminant function analysis was used to distinguish 288 5 Regular codas from four of the sperm whales and 183 3 Regular codas from two sperm whales. The results suggest that codas have consistent individual features in their inter-click intervals and inter-pulse intervals which may contribute to individual identification. Additionally, two whales produced different coda types in distinct foraging dive phases. Codas may therefore be used by sperm whales to convey information of identity as well as activity within a social group to a larger extent than previously assumed.

1.
Antunes
,
R.
(
2000
). “
Study of the sperm whale (Physeter macrocephalus) coda repertoire at the Azores
,” Lic thesis,
Universidade do Algarve, Faro
,
Portugal
,
43
pp.
2.
Antunes
,
R.
(
2009
). “
Variation in sperm whale (Physeter macrocephalus) coda vocalizations and social structure in the North Atlantic Ocean
,” Ph.D. thesis.
University of St. Andrews
,
Scotland
,
123
pp.
3.
Antunes
,
R.
,
Schulz
,
T.
,
Gero
,
S.
,
Whitehead
,
H.
,
Gordon
,
J.
, and
Rendell
,
L.
(
2011
). “
Individually distinctive acoustic features in sperm whale codas
,”
Anim. Behav.
81
,
723
730
.
4.
Caldwell
,
M. C.
, and
Caldwell
,
D. K.
(
1965
). “
Individualized whistle contours in bottlenosed dolphins (Tursiops truncatus)
,”
Nature
207
,
434
435
.
5.
Christal
,
J.
,
Whitehead
,
H.
, and
Lettevall
,
E.
(
1998
). “
Sperm whale social units: Variation and change
,”
Can. J. Zool.
76
,
1431
1440
.
6.
Fischer
,
J.
,
Hammerschmidt
,
K.
,
Cheney
,
D. L.
, and
Seyfarth
,
R. M.
(
2001
). “
Acoustic features of female chacma baboon barks
,”
Ethology
107
,
33
54
.
7.
Frantzis
,
A.
, and
Alexiadou
,
P.
(
2008
). “
Male sperm whale (Physeter macrocephalus) coda production and coda-type usage depend on the presence of conspecifics and the behavioural context
,”
Can. J. Zool.
86
,
62
75
.
8.
Gannon
,
W. L.
,
Sikes
,
R. S.
, and The Animal Care and Use Committee of the American Society of Mammalogists. (
2007
). “
Guidelines of the American Society of Mammalogists for the use of wild mammals in research
,”
J. Mammal.
88
,
809
823
.
9.
Gero
,
S.
,
Engelhaupt
,
D.
, and
Whitehead
,
H.
(
2008
). “
Heterogenous social associations within a sperm whale, Physeter macrocephalus, unit reflect pairwise relatedness
,”
Behav. Ecol. Sociobiol.
63
,
143
151
.
10.
Gero
,
S.
,
Gordon
,
J.
, and
Whitehead
,
H.
(
2013
). “
Calves as social hubs: Dynamics of the social network within sperm whale units
,”
Proc. R. Soc. B
280
,
1
9
.
11.
Gero
,
S.
,
Whitehead
,
H.
, and
Rendell
,
L.
(
2016
). “
Individual, unit and vocal clan level identity cues in sperm whale codas
,”
R. Soc. Open Sci.
3
,
150372
.
12.
Gillespie
,
D.
(
1997
). “
An acoustic survey for sperm whales in the Southern Ocean sanctuary conducted from the RSV Aurora Australis
,”
Rep. Int. Whal. Commn.
47
,
897
907
.
13.
Gordon
,
J. C.
(
1991
). “
Evaluation of a method for determining the length of sperm whales (Physeter catodon) from their vocalizations
,”
J. Zool.
224
(
2
),
301
314
.
14.
Hammerschmidt
,
K.
, and
Todt
,
D.
(
1995
). “
Individual differences in vocalisations of young Barbary macaques (Macaca sylvanus): A multi-parametric analysis to identify critical cues in acoustic signaling
,”
Behaviour
132
(
5/6
),
381
399
.
15.
Jaquet
,
N.
,
Dawson
,
S.
, and
Douglas
,
L.
(
2001
). “
Vocal behavior of male sperm whales: Why do they click?
,”
J. Acoust. Soc. Am.
109
(
5
),
2254
2259
.
16.
Johnson
,
M.
,
Aguilar de Soto
,
N.
, and
Madsen
,
P. T.
(
2009
). “
Studying the behaviour and sensory ecology of marine mammals using acoustic recording tags: A review
,”
Mar. Ecol. Prog. Ser.
395
,
55
73
.
17.
Johnson
,
M.
,
Hickmott
,
L. S.
,
Aguilar Soto
,
N.
, and
Madsen
,
P. T.
(
2008
). “
Echolocation behaviour adapted to prey in foraging Blainville's beaked whale (Mesoplodon densirostris)
,”
Proc. R. Soc. B
275
,
133
139
.
18.
Johnson
,
M.
,
Madsen
,
P. T.
,
Zimmer
,
W. M. X.
,
Aguilar de Soto
,
N.
, and
Tyack
,
P. L.
(
2006
). “
Foraging Blainville's beaked whales (Mesoplodon densirostris) produce distinct click types matched to different phases of echolocation
,”
J. Exp. Biol.
209
,
5038
5050
.
19.
Johnson
,
M. P.
, and
Tyack
,
P. L.
(
2003
). “
A digital acoustic recording tag for measuring the response of wild marine mammals to sound
,”
IEEE J. Ocean. Eng.
28
(
1
),
3
12
.
20.
Kiefte
,
M.
(
1999
). “
Discriminant analysis toolbox (version 0.3)
,” http://www.mathworks.com/matlabcentral/fileexchange/loadFile.do?objectId=189 (Last viewed October 8, 2013).
21.
Madsen
,
P. T.
,
Carder
,
D. A.
,
Au
,
W. W.
,
Nachtigall
,
P. E.
,
Møhl
,
B.
, and
Ridgway
,
S. H.
(
2003
). “
Sound production in neonate sperm whales
,”
J. Acoust. Soc. Am.
113
(
6
),
2988
2991
.
22.
Madsen
,
P. T.
,
Payne
,
R.
,
Kristiansen
,
N. U.
,
Wahlberg
,
M.
,
Kerr
,
I.
, and
Møhl
,
B.
(
2002a
). “
Sperm whale sound production studied with ultrasound time/depth recording tags
,”
J. Exp. Biol.
205
,
1899
1906
.
23.
Madsen
,
P. T.
,
Wahlberg
,
M.
, and
Møhl
,
B.
(
2002b
). “
Male sperm whale (Physeter macrocephalus) acoustics in a high-latitude habitat: Implications for echolocation and communication
,”
Behav. Ecol. Sociobiol.
53
,
31
41
.
24.
Madsen
,
P. T.
,
Wilson
,
M.
,
Johnson
,
M.
,
Hanlon
,
R. T.
,
Bocconcelli
,
A.
,
Aguilar Soto
,
N.
, and
Tyack
,
P. L.
(
2007
). “
Clicking for calamari: Toothed whales can echolocate squid Loligo pealeii
,”
Aquat. Biol.
1
,
141
150
.
25.
Mate
,
B.
,
Mesecar
,
R.
, and
Lagerquist
,
B.
(
2007
). “
The evolution of satellite-monitored radio tags for large whales: One laboratory's experience
,”
Deep Sea Res. Part 2
54
,
224
247
.
26.
Matthews
,
J. N.
,
Steiner
,
L.
, and
Gordon
,
J.
(
2001
). “
Mark-recapture analysis of sperm whale (Physeter macrocephalus) photo-id data from the Azores (1987–1995)
,”
J. Cetacean Res. Manag.
3
(
3
),
219
226
.
27.
McComb
,
K.
,
Reby
,
D.
,
Baker
,
L.
,
Moss
,
C.
, and
Sayialel
,
S.
(
2003
). “
Long-distance communication of acoustic cues to social identity in African elephants
,”
Anim. Behav.
65
,
317
329
.
28.
Melendez
,
K. V.
, and
Feng
,
A. S.
(
2010
). “
Communication calls of little brown bats display individual-specific characteristics
,”
J. Acoust. Soc. Am.
128
(
2
),
919
923
.
29.
Miller
,
P. J. O.
,
Johnson
,
M. P.
, and
Tyack
,
P. L.
(
2004
). “
Sperm whale behaviour indicates the use of echolocation click buzzes ‘creaks’ in prey capture
,”
Proc. R. Soc. B
271
,
2239
2247
.
62.
Møhl
,
B.
(
2001
). “
Sound transmission in the nose of the sperm whale Physeter catodon. A post mortem study
,”
J. Comp. Physiol. A.
187
,
335
340
.
31.
Møhl
,
B.
,
Wahlberg
,
M.
, and
Madsen
,
P. T.
(
2003
). “
The monopulsed nature of sperm whale clicks
,”
J. Acoust. Soc. Am.
114
(
2
),
1143
1154
.
30.
Møhl
,
B.
,
Wahlberg
,
M.
,
Madsen
,
P. T.
,
Miller
,
L. A.
, and
Surlykke
,
A.
(
2000
). “
Sperm whale clicks: Directionality and source level revisited
,”
J. Acoust. Soc. Am.
107
,
638
648
.
32.
Moore
,
K. E.
,
Watkins
,
W. A.
, and
Tyack
,
P. L.
(
1993
). “
Pattern similarity in shared codas from sperm whales (Physeter catodon)
,”
Mar. Mamm. Sci.
9
(
1
),
1
9
.
33.
Mullins
,
J.
,
Whitehead
,
H.
, and
Weilgart
,
L. S.
(
1988
). “
Behaviour and vocalizations of two single sperm whales, Physeter macrocephalus, off Nova Scotia
,”
Can. J. Fish. Aquat. Sci.
45
,
1736
1743
.
34.
Norris
,
K. S.
, and
Harvey
,
G. W.
(
1972
). “
A theory for the function of the spermaceti organ of the sperm whale (Physeter catodon L.)
,” in
Animal Orientation and Navigation
, edited by
S. R.
Galler
,
K.
Schmidt-Koenig
,
G. J.
Jacobs
, and
R. E.
Belleville
(
NASA
,
Washington D.C.
), pp.
397
417
.
35.
Oliveira
,
C.
,
Wahlberg
,
M.
,
Johnson
,
M.
,
Miller
,
P. J. O.
, and
Madsen
,
P. T.
(
2013
). “
The function of male sperm whale slow clicks in a high latitude habitat: Communication, echolocation or prey debilitation?
,”
J. Acoust. Soc. Am.
133
(
5
),
3135
3144
.
36.
Pavan
,
G.
,
Hayward
,
T.
,
Borsani
,
J. F.
,
Priano
,
M.
,
Fossati
,
C.
, and
Gordon
,
J.
(
2000
). “
Time pattern of sperm whale codas recorded in the Mediterranean Sea 1985–1996
,”
J. Acoust. Soc. Am.
107
(
6
),
3487
3495
.
37.
Pinela
,
A. M.
,
Quérouil
,
S.
,
Magalhães
,
S.
,
Silva
,
M. A.
,
Prieto
,
R.
,
Matos
,
J. A.
, and
Santos
,
R. S.
(
2009
). “
Population genetics and social organization of the sperm whale (Physeter macrocephalus) in the Azores inferred by microsatellite analyses
,”
Can. J. Zool.
87
,
802
813
.
38.
Rendell
,
L.
, and
Whitehead
,
H.
(
2003a
). “
Comparing repertoires of sperm whale codas: A multiple methods approach,
Bioacoustics.
14
(
1
),
61
81
.
39.
Rendell
,
L.
, and
Whitehead
,
H.
(
2003b
). “
Vocal clans in sperm whales (Physeter macrocephalus)
,”
Proc. R. Soc. B.
270
,
225
231
.
40.
Rendell
,
L.
, and
Whitehead
,
H.
(
2004
). “
Do sperm whales share coda vocalizations? Insights into coda usage from acoustic size measurement
,”
Anim. Behav.
67
,
865
874
.
41.
Rendell
,
L.
, and
Whitehead
,
H.
(
2005
). “
Spatial and temporal variation in sperm whale coda vocalizations: Stable usage and local dialects
,”
Anim. Behav.
70
,
191
198
.
42.
Sayigh
,
L. S.
,
Tyack
,
P. L.
,
Wells
,
R. S.
, and
Scott
,
M. D.
(
1990
). “
Signature whistles of free-ranging bottlenose dolphins Tursiops truncatus: Stability and mother-offspring comparisons
,”
Behav. Ecol. Sociobiol.
26
,
247
260
.
43.
Schulz
,
T.
,
Whitehead
,
H.
,
Gero
,
S.
, and
Rendell
,
L.
(
2011
). “
Individual vocal production in a sperm whale (Physeter macrocephalus) social unit
,”
Mar. Mamm. Sci.
27
(
1
),
149
166
.
44.
Silva
,
M. A.
,
Prieto
,
R.
,
Cascão
,
I.
,
Seabra
,
M. I.
,
Machete
,
M.
,
Baumgartner
,
M. F.
, and
Santos
,
R. S.
(
2014
). “
Spatial and temporal distribution of cetaceans in the mid-Atlantic waters around the Azores
,”
Mar. Biol. Res.
10
(
2
),
123
137
.
45.
Silva
,
M. A.
,
Prieto
,
R.
,
Magalhães
,
S.
,
Cabecinhas
,
R.
,
Cruz
,
A.
,
Gonçalves
,
J. M.
, and
Santos
,
R. S.
(
2003
). “
Occurrence and distribution of cetaceans in waters around Azores (Portugal), Summer and Autumn 1999–2000
,”
Aquat. Mamm.
29
(
1
),
77
83
.
46.
Soltis
,
J.
,
Leong
,
K.
, and
Savage
,
A.
(
2005
). “
African elephant vocal communication II: Rumble variations reflects the individual identity and emotional state of callers
,”
Anim. Behav.
70
,
589
599
.
47.
Teloni
,
V.
(
2005
). “
Patterns of sound production in diving sperm whales in the Northwestern Mediterranean
,”
Mar. Mamm. Sci.
21
(
3
),
446
457
.
48.
Teloni
,
V.
,
Johnson
,
M. P.
,
Miller
,
P. J. O.
, and
Madsen
,
P. T.
(
2008
). “
Shallow food for deep divers: Dynamic foraging behaviour of male sperm whales in a high latitude habitat
,”
J. Exp. Mar. Biol. Ecol.
354
,
119
131
.
49.
Tibbetts
,
E. A.
, and
Dale
,
J.
(
2007
). “
Individual recognition: It is good to be different
,”
Trends Ecol. Evol.
22
(
10
),
529
537
.
50.
Van Dommelen
,
W. A.
(
1990
). “
Acoustic parameters in human speaker recognition
,”
Lang. Speech
33
(
3
),
259
272
.
51.
Watkins
,
W. A.
, and
Schevill
,
W. E.
(
1977
). “
Sperm whale codas
,”
J. Acoust. Soc. Am.
62
,
1485
1490
.
52.
Weilgart
,
L.
, and
Whitehead
,
H.
(
1988
). “
Distinctive vocalizations from mature male sperm whales (Physeter macrocephalus)
,”
Can. J. Zool.
66
,
1931
1937
.
53.
Weilgart
,
L.
, and
Whitehead
,
H.
(
1993
). “
Coda communication by sperm whales (Physeter macrocephalus) off the Galapagos Islands
,”
Can. J. Zool.
71
,
744
752
.
54.
Weilgart
,
L.
, and
Whitehead
,
H.
(
1997
). “
Group-specific dialects and geographical variation in coda repertoire in South Pacific sperm whales
,”
Behav. Ecol. Sociobiol.
40
,
277
285
.
55.
Weilgart
,
L.
,
Whitehead
,
H.
, and
Payne
,
K.
(
1996
). “
A colossal convergence
,”
Am. Sci.
84
(
3
),
278
287
.
56.
Whitehead
,
H.
(
2003
).
Sperm Whales: Social Evolution in the Ocean
(
The University of Chicago
,
Chicago, IL
),
431
pp.
57.
Whitehead
,
H.
,
Antunes
,
R.
,
Gero
,
S.
,
Wong
,
S. N. P.
,
Engelhaupt
,
D.
, and
Rendell
,
L.
(
2012
). “
Multilevel societies of female sperm whales (Physeter macrocephalus) in the Atlantic and Pacific: Why are they so different?
,”
Int. J. Primatol.
33
,
1142
1164
.
58.
Whitehead
,
H.
, and
Weilgart
,
L.
(
1991
). “
Patterns of visually observable behavior and vocalizations in groups of female sperm whales
,”
Behaviour
118
,
275
296
.
59.
Wisniewska
,
D. M.
,
Johnson
,
M.
,
Beedholm
,
K.
,
Wahlberg
,
M.
, and
Madsen
,
P. T.
(
2012
). “
Acoustic gaze adjustments during active target selection in echolocating porpoises
,”
J. Exp. Biol.
215
,
4358
4373
.
60.
Zimmer
,
W. M. X.
,
Johnson
,
M. P.
,
D'Amico
,
A.
, and
Tyack
,
P. L.
(
2003
). “
Combining data from a multisensory tag and a passive sonar to determine the diving behaviour of a sperm whale (Physeter macrocephalus)
,”
IEEE J. Ocean. Eng.
28
(
1
),
13
28
.
61.
Zimmer
,
W. M. X.
,
Tyack
,
P. L.
,
Johnson
,
M. P.
, and
Madsen
,
P. T.
(
2005
). “
Three-dimensional beam pattern of regular sperm whale clicks confirms bent-horn hypothesis
,”
J. Acoust. Soc. Am.
117
(
3
),
1473
1485
.
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