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Vocalisations of Antarctic blue whales, Balaenoptera musculus intermedia, recorded during the 2001/2002 and 2002/2003 IWC/SOWER circumpolar cruises, Area V, Antarctica

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Abstract

Blue whale vocalisations recorded during the 2001/2002 and 2002/2003 International Whaling Commission-Southern Ocean Whale and Ecosystem Research (IWC/SOWER) cruises were analysed to determine the feasibility of using acoustic recordings for sub-species identification of the Antarctic blue whale (Balaenoptera musculus intermedia) and the pygmy blue whale (B.m. brevicauda). The research was conducted in IWC Area V, from latitude 60°S to the ice edge and between longitudes 130°E and 150°E on the Shonan Maru (2001/2002), and between 150°E and 170°W on the Shonan Maru No.2 (2002/2003). Data including 15 groups consisting of 42 animals, as well as opportunistic recordings of an unknown number of animals during evening sonobuoy stations were examined for this study. Vocalisations included long-duration 28Hz tonal sounds and relatively short-duration frequency-modulated sounds. The short-duration calls were similar to vocalisations recorded in the presence of blue whales in other locations worldwide. Not all recordings contained the longduration 28Hz call, considered to be a species-specific vocalisation of Antarctic blue whales. None of the sounds that have previously been attributed to pygmy blue whales were detected. The long-duration 28Hz tonal vocalisations included 3-unit calls, considered to be song phrases, as well as simple 28Hz sounds and 28Hz sounds followed by a downsweep. The centre and peak frequencies of the 28Hz tone for these three sound types were stable regardless of signal strength; however, for the 3-unit vocalisation, the presence and characteristics of their 2nd and 3rd units were variable. Examination of two distinct groups of simultaneously vocalising blue whales showed no evidence of temporally repeated patterns of vocalisations (song phrases). The results of this study suggest that the peak frequency of the 28Hz vocalisations may be used as a diagnostic feature to aid in discriminating between Antarctic blue whales and pygmy blue whales in the field; however, examination of vocalisations in relation to group size and behaviour are necessary to understand the circumstances in which the 28Hz vocalisations are produced.
INTRODUCTION
Two sub-species of blue whale are found in the Southern
Ocean, the Antarctic blue whale, Balaenoptera musculus
intermedia, and the pygmy blue whale, B.m. brevicauda.
There appears to be a general geographic segregation of the
sub-species in the austral mid-summer, with pygmy blue
whales occurring primarily north of 60°S and Antarctic blue
whales south of this latitude (Kato et al., 1995). Sub-species
discrimination in the field is problematic because it relies on
experienced observers noting relative body proportions and
details of the head shape. Population research related to the
conservation of large baleen whales requires accurate
species identification (IWC, 1995). Recent studies indicate
that monitoring of blue whale vocalisations may provide a
means of determining sub-species in the field (Ljungblad et
al., 1997; 1998; Stafford et al., 1999; 2001).
Sounds recorded in the presence of blue whales can be
divided into two categories: short-duration or long-duration
(Thompson and Cummings, 1996; Norris and Barlow,
2000). The short-duration vocalisations consist of individual
pulses and frequency-modulated (FM, typically downswept)
sounds of less than five seconds duration. These
vocalisations vary in frequency and duration and have been
recorded in the presence of blue whales in many locations
(Thompson and Cummings, 1996; Ljungblad et al., 1997;
Stafford et al., 2001). Short-duration sounds appear to be
common; however, they are underrepresented in the
literature.
Long-duration vocalisations are composed of one or more
units that are FM or amplitude-modulated (AM) sounds and
longer than five seconds (McDonald et al., Submitted). An
individual unit is defined as a continuous sound having
consistent characteristics; these vocalisation units are often
repeated in patterned sequences, or songs (Payne and
McVay, 1971; McDonald et al., Submitted). These song
units have been shown to vary geographically (Cummings
and Thompson, 1971; Edds, 1982; Thompson and Friedl,
1982; Alling et al., 1991; Thompson and Cummings, 1996;
Stafford et al., 1999; 2001). Preliminary examination of
sounds recorded in the presence of Antarctic blue whales
and pygmy blue whales in the Southern Hemisphere
indicate a similar geographic distribution of long-duration,
low-frequency song units (Clark and Fowler, 2001;
Ljungblad et al., 1997).
Recordings of pygmy blue whales off Madagascar show
repetitive sequences of 10-20s tonal sounds in the 25-45Hz
band (Ljungblad et al., 1998). Pygmy blue whale
vocalisations recorded off Australia consist of three separate
long tonal units in the 18-26Hz band (McCauley et al.,
2000). The long-duration sounds recorded in the presence of
Antarctic blue whales in the Antarctic consist of patterned
sequences of tonal sounds composed of three distinct units.
The first tone is centred at 28-29Hz with a duration of 8-12s.
A short 2s downsweep from 28-20Hz connects the first tonal
unit to the third, a slightly modulated tone (20-18Hz), that is
approximately 8-12s in duration (Ljungblad et al., 1998).
The three-unit vocalisation, or phrase, is usually repeated
J. CETACEAN RES. MANAGE. 7(1):13–20, 2005
13
Vocalisations of Antarctic blue whales, Balaenoptera musculus
intermedia, recorded during the 2001/2002 and 2002/2003
IWC/SOWER circumpolar cruises, Area V, Antarctica
SHANNON RANKIN
*
, DON LJUNGBLAD
+
, CHRIS CLARK
#
AND HIDEHIRO KATO
++
Contact e-mail: Shannon.Rankin@noaa.gov
ABSTRACT
Blue whale vocalisations recorded during the 2001/2002 and 2002/2003 International Whaling Commission-Southern Ocean Whale and
Ecosystem Research (IWC/SOWER) cruises were analysed to determine the feasibility of using acoustic recordings for sub-species
identification of the Antarctic blue whale (Balaenoptera musculus intermedia) and the pygmy blue whale (B.m. brevicauda). The research
was conducted in IWC Area V, from latitude 60°S to the ice edge and between longitudes 130°E and 150°E on the Shonan Maru
(2001/2002), and between 150°E and 170°W on the Shonan Maru No.2 (2002/2003). Data including 15 groups consisting of 42 animals,
as well as opportunistic recordings of an unknown number of animals during evening sonobuoy stations were examined for this study.
Vocalisations included long-duration 28Hz tonal sounds and relatively short-duration frequency-modulated sounds. The short-duration calls
were similar to vocalisations recorded in the presence of blue whales in other locations worldwide. Not all recordings contained the long-
duration 28Hz call, considered to be a species-specific vocalisation of Antarctic blue whales. None of the sounds that have previously been
attributed to pygmy blue whales were detected. The long-duration 28Hz tonal vocalisations included 3-unit calls, considered to be song
phrases, as well as simple 28Hz sounds and 28Hz sounds followed by a downsweep. The centre and peak frequencies of the 28Hz tone for
these three sound types were stable regardless of signal strength; however, for the 3-unit vocalisation, the presence and characteristics of
their 2
nd
and 3
rd
units were variable. Examination of two distinct groups of simultaneously vocalising blue whales showed no evidence of
temporally repeated patterns of vocalisations (song phrases). The results of this study suggest that the peak frequency of the 28Hz
vocalisations may be used as a diagnostic feature to aid in discriminating between Antarctic blue whales and pygmy blue whales in the field;
however, examination of vocalisations in relation to group size and behaviour are necessary to understand the circumstances in which the
28Hz vocalisations are produced.
KEYWORDS: BLUE WHALE; COMMUNICATION; VOCALISATION; ANTARCTIC; MANAGEMENT PROCEDURE; SURVEY-
ACOUSTIC; POPULATION ASSESSMENT; ACOUSTICS; DISTRIBUTION; SOWER
*
Southwest Fisheries Science Center, NOAA, US National Marine Fisheries Service, 8604 La Jolla Shores Drive, La Jolla, CA 92038, USA.
+
Ljungblad Associates, PO Box 6, Elk Mountain, WY 82324, USA.
#
Bioacoustics Research Program, Cornell Laboratory of Ornithology, Cornell University, 159 Sapsucker Woods Rd, Ithaca, NY 14850, USA.
++
Large Cetacean Section, National Research Institute of Far Seas Fisheries, 5-7-1 Orido, Shimizu, Japan.
every 70-80s, at intervals between 40-50s (Ljungblad et al.,
1998). There is a high degree of variability in the presence
and intensity of the three individual units, and therefore we
use the terms ‘3-unit vocalisation’ to describe vocalisations
with all three units intact, ‘28Hz downsweep’ to describe
vocalisations with the first two units intact, and ‘28Hz tone’
to describe vocalisations where only the first unit is intact.
In addition to studying the distinct variation in
vocalisations of the two sub-species of blue whale for
accurate species identification in the field, knowledge of the
behavioural contexts of these sounds is needed for long-
term vocal and population studies. The blue whale
component of the International Whaling Commission’s
SOWER (Southern Ocean Whale and Ecosystem Research)
programme obtains videos, photographs, biopsies and
acoustic recordings of blue whales in the field. This study
examines recordings and behavioural information obtained
in the presence of blue whales during two seasons of
SOWER cruises to provide a preliminary examination of the
variability associated with the 3-unit vocalisation, and its
effect on blue whale population studies in the Southern
Ocean.
METHODS
Data collected by the authors during the 2001/2002 SOWER
cruise from the vessel Shonan Maru and the 2002/2003
SOWER cruise from the vessel Shonan Maru No.2 were
used for this study. The research area surveyed was in IWC
Area V
1
between 130°E-150°E (2001/2002) and 150°E-
170°W (2002/2003), and extending from 60°S to the ice
edge (Fig. 1). Line-transect visual observations of cetaceans
were conducted between 06:00 and 18:00 local time,
weather permitting, using a visual observation team
consisting of three tiers of experienced observers
(Anonymous, 2002b). Summaries of the methods are given
in Ensor et al. (2002 and 2003). Briefly, the visual
observation team was responsible for sighting and positively
identifying whales, estimating group sizes and obtaining
biopsy samples, video tapes, and photo-identification
photographs. An acoustician was responsible for obtaining
recordings in the vicinity of blue whales and collecting
opportunistic evening recordings.
The primary acoustic recording method used expendable
DIFAR (Direction Finding and Ranging) AN/SSQ 53B
sonobuoys. These were deployed in close proximity to
sighted blue whales and monitored for as long as time
permitted for a minimum of 30 minutes. Opportunistic
recordings were also made while drifting in the evenings.
The sonobuoy radio signal was received via the ship
antenna, which was coupled to an ICOM IC-R100 single
channel receiver. This output was connected to a Sony DAT
TCD-D7 recorder (flat frequency response from 5Hz to
24kHz) or a Sony mini-disk MZ-R700 recorder (frequency
response 20Hz-20kHz ± 3dB). Recordings were later
digitised to a Sony PCG-FX120 computer (sample rate
48kHz) using the software program ISHMAEL (Mellinger,
2001) and analysis was performed using Spectra-Plus
software. All vocalisations attributed to blue whales with a
strong signal-to-noise ratio (SNR) from the 2001/2002
season were measured (44.1kHz sample rate, 32768 point
FFT size, 90% overlap, Hamming window). Only long-
duration calls with a strong SNR from the 2002/2003 season
were measured (5512Hz sample rate, 8192 point FFT size,
90% overlap, Hanning window).
All high-quality vocalisations attributed to blue whales
were categorised according to their frequency and duration
characteristics. The short-duration FM calls included
amplitude-modulated downsweeps, high-frequency
downsweeps, low-frequency downsweeps, high-frequency
upsweeps, low-frequency upsweeps and complex calls. The
long-duration sounds included the 3-unit vocalisations,
28Hz downsweeps and 28Hz tonal vocalisations.
Measurements were made of lowest frequency, highest
frequency, centre frequency (for tonal sounds), start
frequency, end frequency, frequency shift, peak frequency,
and duration for all vocalisations and vocalisation units.
Measurement of the time between the deployment of the
sonobuoy and detection of the first 3-unit vocalisation was
made to examine feasibility of using these vocalisations for
in situ species identification.
Temporal patterns of vocalisations were examined for a
series of recordings (10.5 hours total) associated with blue
whale sightings on 23 January 2003. All times are given as
local times at sea. Bearings to each vocalisation were
obtained using DIFAR signal processing. This was
performed using an automatic MATLAB function within
Ishmael that executes a series of commands for de-
multiplexing the DIFAR signal (software developed by
Greenridge Sciences, Inc.), and determines the bearing to a
sound source (software designed by M. McDonald).
Bearings of individual vocalisations allowed the detection
of distinct groups of vocalising whales, so that patterns of
vocalisations could be examined within and between
groups. It was not possible to use DIFAR to distinguish
individuals within groups due to the close association and
variable movement patterns of animals. The paucity of
recording tapes available during the 2001/2002 season
necessitated recording at the lowest possible sampling rate
to maximise the recording time (with a sample rate of
32kHz, the frequency response of the Sony TCD-D7 was 20-
14,500Hz ± 1dB). This eliminated the multiplexed DIFAR
signal and so bearings could not be obtained for these data.
RESULTS
Recordings were made in the vicinity of 12 blue whale
groups (31 animals total) during the 2001/2002 season and
in the vicinity of three blue whale groups (11 animals total)
for the 2002/2003 season (Table 1). Blue whale sounds were
detected during 14 of these 15 groups.
Blue whale encounters
2001/2002
Between 6 and 8 January 2002, a total of nine sightings of
blue whales were observed within an area bounded by
64°18’S and 64°29’S and 136°29’E and 137°24’E, near the
northern margin of belts of the pack ice. Seven of the groups
(totalling 14 animals) were determined to be Antarctic blue
whales; photo and/or video and biopsy attempts were made
for these groups (Table 1). The eighth was a group of three
animals observed at night that was not approached and was
classified as undetermined blue whales. A distant group of
two animals determined to be ‘like’ blue whales were
sighted outside of a larger congregation of blue whales, but
these animals were not approached. During all encounters,
sounds attributed to Antarctic blue whales were recorded,
although for most encounters these sounds were not
detected within the first hour of recording (Table 1).
From 21-31 January 2002, three groups of Antarctic blue
whales (totalling 12 animals) and three groups of
unidentified blue whales (six animals total) were sighted.
14 RANKIN
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VOCALISATIONS OF ANTARCTIC BLUE WHALES
1
For a description of IWC Areas, see Donovan (1991).
Photos and/or video and acoustic recordings were obtained
for all but the undetermined groups of whales, and biopsy
attempts were made for the groups identified on 29 and 31
January (Table 1). Very few vocalisations were recorded
during these encounters.
2002/2003
On 23 January 2003, two groups of Antarctic blue whales
(totalling 8 animals) were sighted in the outer margin of the
pack ice, in the vicinity of 67°07’S and 166°54’E (Table 1).
All animals appeared to be feeding on krill patches.
Photography, biopsy attempts and acoustic recordings were
undertaken during this sighting. Acoustic recordings began
at 17:10 and continued throughout the night in the location
of the scattered blue whale sightings. An additional sighting
of three animals was detected in the middle of the night (24
January) and confirmed the continued presence of blue
whales in the area. A detailed examination of DIFAR
processing of the acoustic behaviour of these groups is
described below.
Characteristics of blue whale vocalisations
A total of 85 hours of recordings were made from the
Shonan Maru during the 2001/2002 cruise, with over 33
hours of recordings in areas of blue whale sightings. All
recordings were monitored for the presence of sounds that
could be attributed to blue whales, and over 42 hours of
recordings contained blue whale vocalisations. A total of
193 short FM vocalisations and 261 long-duration tonal
vocalisations (including 3-unit, 28Hz downsweep and 28Hz
tonal vocalisations) were measured from the recordings for
this survey.
J. CETACEAN RES. MANAGE. 7(1):13–20, 2005
15
Fig. 1. Locations of acoustic detections of blue whales within the study area. The open squares represent recordings from blue
whale sightings; the closed circles represent opportunistic evening sonobuoy stations with blue whale acoustic detections.
A total of 38.7 hours of recordings were made from the
Shonan Maru No.2 during the 2002/2003 survey, including
11 hours in the vicinity of blue whale sightings. Sounds that
could be attributed to blue whales were detected in nearly 26
hours of recordings, however only recordings associated
with sightings were examined. A total of 92 long-duration
tonal vocalisations with high SNRs were measured from this
survey; short FM vocalisations were recorded, but not
measured.
The most common short-duration FM vocalisation was
the simple high-frequency downsweep from 76.3-40.0Hz,
with a mean signal duration of 2.7 seconds (n=132, Table 2,
Fig. 2(a)). The amplitude-modulation found in the pulsed
downsweep appeared to be caused by propagation, and the
basic frequency and duration characteristics closely
resemble those of the high-frequency downsweep (Fig.
2(b)). The low-frequency downsweep (n=4), low-frequency
upsweep (n=7) and high-frequency upsweep sounds (n=4)
were relatively uncommon compared to the high-frequency
downsweep vocalisations (Table 2).
Although the complex sounds were variable in nature,
several similar types were frequently observed. The most
common complex vocalisations were variations on the high-
frequency downsweep, with one or more inflection points
(Fig. 2(c)). Other less common complex vocalisations are
short, high-frequency downsweeps, variable FM sounds and
‘concave’ sounds (Fig. 3).
The long-duration calls were divided into three categories
as described in the Introduction: the 3-unit vocalisation; the
28Hz downsweep; and the 28Hz tone (Fig. 4). The 3-unit
vocalisation consisted of a tone at 27.7Hz lasting an average
of 8.3 seconds, occasionally followed by a brief downsweep
of variable duration, to a typically FM moan from 19.5-
19.1Hz with an average duration of 6.9 seconds (Table 3).
The 28Hz downsweep consisted of a moan at 27.7Hz of a
variable duration followed by a downsweep to
approximately 19.1Hz. Measurements of the entire sample
of vocalisations (‘All’) were compared with a sub-sample of
high-quality vocalisations (‘Best’); the centre and peak
frequency of the 28Hz tone varied little, regardless of the
vocalisation type or signal quality. For vocal animals, at
least 30 minutes elapsed between sonobuoy deployment and
initial detection of vocalisations associated with the
Antarctic blue whales (defined as the acoustic identification
time) (Table 1).
DIFAR analyses of vocalisations
DIFAR analyses of recordings from 23 January 2003
allowed differentiation between the calls from several
groups of blue whales (sightings 11 and 12; Fig. 5).
Determination of the bearing angles to the sound source
using DIFAR analysis was performed on 1,069 vocalisations
(208 long-duration, 861 short-duration) during the 11 hours
of recordings. Close association of several animals within a
group prevented identification of the vocalising animal in
most cases; sound source for all sightings is for the group
and not an individual animal (where group size >1). The
similar DIFAR bearing angles of the blue whales (160°) and
the ship (167°) at 17:45 indicate that animals biopsied at this
time were vocalising (Fig. 5(a)). The discontinuity in the
ship’s course at 18:15 (Fig. 5(b)) occurred as the ship
returned to course and speed after biopsy sampling;
Fig. 2. Spectrogram of the most common short-duration FM
vocalisations (5kHz sample rate, 2048 point FFT): (a) high-
frequency downsweep; (b) amplitude-modulated downsweep; and
(c) a complex variation of the high-frequency downsweep.
16 RANKIN
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VOCALISATIONS OF ANTARCTIC BLUE WHALES
Fig. 3. Spectrogram of uncommon short-duration FM vocalisations
(5kHz sample rate, 2048 point FFT): (a) high frequency upsweep;
(b) short high frequency downsweep; (c) variable high frequency
downsweep; and (d) concave vocalisation.
excessive noise during this manoeuvre temporarily
precluded DIFAR processing. At 18:15 the ship’s true
course of 5° closely matched that determined by the
sonobuoy. At 18:45 the ship’s course crossed 0° in front of
the sonobuoy.
Movement of the ship and the subsequent loss of the
sonobuoy signal led to a gap in the recordings from 19:00
until 23:00. Recordings continued while the vessel was
drifting for the remainder of the night, and DIFAR bearing
angles suggest that a large congregation of vocalising blue
whales separated into two distinct groups at about 00:30
(Fig. 6(a)). At 01:20 on 24 January a group of three blue
whales (Fig. 6(b), 350°) were seen feeding next to the ship.
DIFAR angles show that this group may have produced
occasional short-duration FM vocalisations, but was not a
part of either consistently vocalising group (40° and 150°).
Between 00:39 and 02:49 the sounds from the two
simultaneously vocalising blue whale groups (denoted A
and B) had a high SNR and well-defined DIFAR bearing
angles. Both short-duration FM and long-duration tonal
vocalisations from the two groups were plotted over time to
identify patterns in temporal variation (Fig. 7). However,
there was no apparent temporal pattern for the short-
duration or long-duration 28Hz vocalisations. An expanded
view of the common high-frequency downsweeps (HFDN)
does not suggest countercalling between groups Aand B.
DISCUSSION
This study shows that the 3-unit vocalisation is a
geographically distinct call associated with Antarctic blue
whales in the Southern Ocean south of 60°S. The 3-unit
vocalisations recorded during 2001/2002 are consistent with
previous results for sounds attributed to Antarctic blue
whales in the Antarctic (Anonymous, 2002a; Clark and
Fowler, 2001; Ljungblad et al., 1998). The dataset presented
here represents the largest analysis to date of these calls.
None of the characteristic sounds attributed to pygmy blue
whales in Madagascar (Ljungblad et al., 1998) or Chile
(Cummings and Thompson, 1971) were detected in the 247
hours of recordings. During the two cruises, all whales
visually identified at the sub-species level were considered
to be Antarctic blue whales. Genetic analysis of biopsies
obtained on these cruises is underway and will hopefully
confirm that these sounds were indeed produced by
Antarctic blue whales. The association of specific calls
exclusively to Antarctic blue whales provides a step towards
in situ acoustic sub-species identification. As noted earlier,
real-time identification currently relies on visual inspection
by experienced observers. However, whales do not vocalise
continuously, which limits the value of the technique. In
addition, should time be limited, identifications in real-time
may not be feasible due to the processing time required for
a single identification. Nonetheless, the technique is a
valuable tool, particularly when used in conjunction with
genetic analysis and visual identification methods.
To use vocalisation for species identification one must be
able to positively detect the call. There appears to be
variability in the presence and characteristics of the 2
nd
(inter-tone downsweep) and 3
rd
(19Hz tone) unit of this
vocalisation (Table 3). It is clear that blue whales produce
both 28Hz tonal and 28Hz downsweep vocalisations, in
addition to the 3-unit calls previously examined. Even with
a high SNR, an overlap of the multi-path signals of long-
J. CETACEAN RES. MANAGE. 7(1):13–20, 2005
17
Fig. 4. Spectrogram of three long-duration blue whale vocalisations
associated with different Antarctic blue whales (48kHz sample rate,
decimation ratio 4:1, 32768 point FFT): (a) 3-unit vocalisation
including 28Hz tone followed by an inter-tone interval and a 19Hz
tone; (b) 28Hz tone plus downsweep; and (c) a 28Hz tone. All were
sufficiently intense to suggest detection of the entire signal.
Fig. 5. DIFAR bearings for blue whales during biopsy attempts for sighting number 11 on 23 January 2003. Two distinct groups could
be identified, one at 260° and the other at 160°. All magnetic bearing angles were converted to true angles for comparison.
duration calls can make it difficult to determine the
characteristics of individual sounds. With increasingly faint
vocalisations, it may be difficult to distinguish the 3-unit
vocalisations from the 28Hz tonal and 28Hz downsweeps.
The primary consistent feature is the tone centred at 27.7Hz;
peak frequency varies little among these three vocalisations
regardless of the signal intensity. If all three long-duration
28Hz vocalisations can be positively, and exclusively, linked
to Antarctic blue whales, then this research suggests that it
may be possible to attribute any long (6-12s) 28Hz tonal
vocalisation south of 60°S to Antarctic blue whales.
Research to date suggests this to be the case. Sub-species
identification based on detection of the 28Hz tonal
vocalisation is feasible for most groups of Antarctic blue
whales, assuming a minimum one hour recording time.
Future efforts should include deployment of sonobuoy
arrays to localise calling animals so that comparisons with
visual detection and genetic sampling of individual calling
animals can be conducted.
Previous studies suggested that the long-duration, low-
frequency sounds produced by blue whales are songs
(Anonymous, 2002a), or patterned series of repeated
vocalisations. The 2.5 hour sample of two vocalising groups
within the larger congregation of scattered blue whales,
recorded on 23 January 2003, does not suggest that the
vocalisations were repeated in patterned series within a
given group, or between the two groups (Fig. 7). The
comparisons of vocalisations with the various blue whale
18 RANKIN
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VOCALISATIONS OF ANTARCTIC BLUE WHALES
Fig. 6. DIFAR bearings for blue whales during evening recording in location of sighting numbers 11 and 12 for 23 January 2003 and
sighting number 1 on 24 January 2003. All magnetic bearing angles were converted to true angles for comparison. The discontinuities
in the ships’ bearing angle are associated with repositioning.
Fig. 7. Temporal variation of vocalisation types for the two blue whale groups from 00:30 until 3:00, 24 January 2003. An expanded
view of HF DN vocalisations is provided for clarity. HF=high frequency; LF=low frequency; DN=downsweep; UP=upsweep.
sightings (Table 3) suggest that there is considerable
variation in the vocal behaviour of different groups. The
sightings in the 2001/2002 study differed in group size,
behaviour and habitat. Unfortunately, the locations of
vocalising animals in relation to the sonobuoy could not be
determined due to problems in recording the DIFAR signal.
This severely limited our ability to examine temporal
patterns, or the presence of song during the 2001/2002
cruise. Further research combining visual and acoustic
studies (with functional DIFAR) on different blue whale
groups is necessary to understand the circumstances in
which the temporal patterns considered to be ‘song’ are
produced.
Successful use of bottom-mounted hydrophones to
monitor whale song in the Pacific (McDonald et al., 1995;
Stafford et al., 1999) and Atlantic Oceans (Clark, 1995) led
to deployment of similar hydrophones in the Southern
Ocean to monitor the blue whale population year-round
(Sirovic et al., 2004). Minimum abundance can be estimated
through noting the ranges of individual singing whales;
however, our results suggest that only a small proportion of
the blue whale population may be singing. The ability to
relate geographically distinct vocalisations (song units) to
an index of abundance relies heavily on their behavioural
contexts. These concerns are essentially the same as those
for using acoustics as a method for species identification.
The short FM vocalisations recorded here are similar to
sounds associated with blue whales in other regions. With
the exception of the high-frequency downsweep, most short
FM vocalisations are uncommon. Groups of vocalising
whales were noted to produce both short-duration FM and
long-duration 28Hz vocalisations. During extended biopsy
attempts during the 2001/2002 survey there was an apparent
overall increase in vocalisations. The inability to confirm
the vocalising group using DIFAR software limits this to a
simple speculation. This should be examined for other close
approaches, as this may influence the ability of acoustics to
determine species identification.
Clearly there is great variation in the vocal behaviour of
different blue whale groups; however, we cannot yet explain
these differences. The structure of the 3-unit vocalisation
appears to be highly variable, but 27.7Hz peak frequency is
stable even over great distances. The 3-unit vocalisations,
and the other 28Hz vocalisations do not always occur in
patterned series or ‘songs’, and some whale groups are not
vocal. More information must be gathered on the variations
in vocalisations by age, sex, season, time of day, group
composition and behaviour. These data can only be obtained
by integrated in situ studies of blue whales.
The IWC has stated that there is a need for a dedicated
blue whale study in the Southern Ocean to combine visual
and acoustics surveys with biopsy, photo-identification and
J. CETACEAN RES. MANAGE. 7(1):13–20, 2005
19
satellite tagging of individuals in order to determine the
winter breeding grounds. Blue whales have been known to
frequent the ice edge between 150°E and 165°E (Kato et al.,
1995). The relatively high populations of blue whales in this
area during the 2001/2002 and 2002/2003 confirm that this
is an ideal location for deployment of a series of bottom-
mounted hydrophones for recording of vocalisations,
coinciding with future shipboard populations surveys.
ACKNOWLEDGMENTS
This work could not have been accomplished without the
hard work and dedication of the captain and crews of the
Shonan Maru and the Shonan Maru No.2. George Murphy
and the US Navy provided surplus sonobuoys for use in this
project. Financial support was provided by the International
Whaling Commission, The Office of Naval Research and
the US Navy. Special thanks to Jay Barlow, Robert
Brownell, Kathy Kane, Steve Rankin and the Southwest
Fisheries Science Center for their continued support and
Paul Ensor, Jay Barlow, Erin Oleson, Kate Stafford and
Mark McDonald for their comments on earlier versions of
this manuscript, and Richard Cosgrove for assistance with
the graphics. This work could not have been accomplished
without the dedication and guidance of Paul Ensor with
whom it was an honour to work.
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20 RANKIN
et al.:
VOCALISATIONS OF ANTARCTIC BLUE WHALES
... Z-calls, produced year-round by ABWs, are highly stereotypic, low-frequency vocalizations, typically composed of three units within the frequency range of 18-28 Hz (Ljungblad et al., 1998;Rankin et al., 2005). This study used the daily acoustic presence of ABWs based on previous automated detections of Z-call vocalizations as described in detail in Thomisch et al. (2016). ...
... For the majority of marine mammal species, the vocal repertoire has been described (Rankin et al., 2005) and many feature unique (with regard to species) and comprehensive (with regard to seasonal and contextual coverage) vocalizations. ...
... Nevertheless, for some species, not all calls are equally suitable to be included in SDMs. Some call types may occur in rare behavioural contexts (Klinck et al., 2010) only, or may have strong similarities to the calls produced by other species, such as the FM call in baleen whale species (Rankin et al., 2005;Thomisch, 2017). ...
Article
Full-text available
Aim: Species distribution models (SDMs) are essential tools in ecology and conservation. However, the scarcity of visual sightings of marine mammals in remote polar areas hinders the effective application of SDMs there. Passive acoustic monitoring (PAM) data provide year-round information and overcome foul weather limitations faced by visual surveys. However, the use of PAM data in SDMs has been sparse so far. Here, we use PAM-based SDMs to investigate the spatiotemporal distribution of the critically endangered Antarctic blue whale in the Weddell Sea. Location: The Weddell Sea. Methods: We used presence-only dynamic SDMs employing visual sightings and PAM detections in independent models. We compared the two independent models with a third combined model that integrated both visual and PAM data, aiming at leveraging the advantages of each data type: the extensive spatial extent of visual data and the broader temporal/environmental range of PAM data. Results: Visual and PAM data prove complementary, as indicated by a low spatial overlap between daily predictions and the low predictability of each model at detections of other data types. Combined data models reproduced suitable habitats as given by both independent models. Visual data models indicate areas close to the sea ice edge (SIE) and with low-to-moderate sea ice concentrations (SIC) as suitable, while PAM data models identified suitable habitats at a broader range of distances to SIE and relatively higher SIC. Main Conclusions: The results demonstrate the potential of PAM data to predict year-round marine mammal habitat suitability at large spatial scales. We provide reasons for discrepancies between SDMs based on either data type and give methodological recommendations on using PAM data in SDMs. Combining visual and PAM data in future SDMs is promising for studying vocalized animals, particularly when using recent advances in integrated distribution modelling methods.
... PSD slope-The spectral energy of the LFC2 was also estimated as the maximum gradient (i.e., steepness of the rise) of the PSD slope of the audio file between 15 and 19 Hz (Equation (8), Figure 1, Table 1). The descending steepness of the PSD slope at the higher frequency limit of the LFC2 could unfortunately not be used due to the frequent presence of Antarctic blue whale z-calls at the same frequency [35]. The spectral energies of the HFC8 and HFC9 were also estimated as the maximum ascending minus the minimum descending steepness of the PSD slope of the audio file at the respective lower and higher frequency limits of HFC8 and HFC9 (Equation (8)). ...
... To identify individual F20P in the PDS, audio files were processed in 2s-sliding windows (i.e., with 1.5 s overlap). The resulting 2s-signal was bandpass-filtered (IIR filter order 20) between 15 and 26 Hz (i.e., upper frequency limit at 26 Hz to reduce energy input of blue whale Z-calls [35]) and the kurtosis of the signal was computed (Kurt). The bandpass-filtered signal was also processed with a Teager-Kaiser energy (TKE) operator algorithm to enhance low SNR impulse signals [41]. ...
... To identify individual F20P in the PDS, audio files were processed in 2s-sliding wi dows (i.e., with 1.5 s overlap). The resulting 2s-signal was bandpass-filtered (IIR filter o der 20) between 15 and 26 Hz (i.e., upper frequency limit at 26 Hz to reduce energy inp of blue whale Z-calls [35]) and the kurtosis of the signal was computed (Kurt). The ban pass-filtered signal was also processed with a Teager-Kaiser energy (TKE) operator alg rithm to enhance low SNR impulse signals [41]. ...
Article
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The growing availability of long-term and large-scale passive acoustic recordings open the possibility of monitoring the vocal activity of elusive oceanic species, such as fin whales (Balaenoptera physalus), in order to acquire knowledge on their distribution, behavior, population structure and abundance. Fin whales produce low-frequency and high-intensity pulses, both as single vocalizations and as song sequences (only males) which can be detected over large distances. Numerous distant fin whales producing these pulses generate a so-called chorus, by spectrally and temporally overlapping single vocalizations. Both fin whale pulses and fin whale chorus provide a distinct source of information on fin whales present at different distances to the recording location. The manual review of vast amounts of passive acoustic data for the presence of single vocalizations and chorus by human experts is, however, time-consuming, often suffers from low reproducibility and in its entirety, it is practically impossible. In this publication, we present and compare robust algorithms for the automatic detection of fin whale choruses and pulses which yield good performance results (i.e., false positive rates < 3% and true positive rates > 76%) when applied to real-world passive acoustic datasets characterized by vast amounts of data, with only a small proportion of the data containing the target sounds, and diverse soundscapes from the Southern Ocean.
... Blue and fin whales produce low-frequency and high intensity calls that may be detectable up to distances of 200-1700 km (Clark et al. 1995;Širović et al. 2007;Samaran et al. 2010;Thomisch et al. 2019;Shabangu et al. 2020a). Antarctic blue whales produce two different call types, namely a stereotyped Z-call (Rankin et al. 2005) and a D-call (Oleson et al. 2007a). Z-calls have three components that are frequency modulated and range from 18 to 26 s in duration (Rankin et al. 2005). ...
... Antarctic blue whales produce two different call types, namely a stereotyped Z-call (Rankin et al. 2005) and a D-call (Oleson et al. 2007a). Z-calls have three components that are frequency modulated and range from 18 to 26 s in duration (Rankin et al. 2005). The first component occurs at frequency ~ 27 Hz and is 8-12 s long, the second component's frequency downsweeps from ~ 27 Hz to 20 Hz and is 2 s in duration, and the third component is somewhat frequency modulated from 20 Hz to ~ 18 Hz and is 8-12 s in duration (Rankin et al. 2005). ...
... Z-calls have three components that are frequency modulated and range from 18 to 26 s in duration (Rankin et al. 2005). The first component occurs at frequency ~ 27 Hz and is 8-12 s long, the second component's frequency downsweeps from ~ 27 Hz to 20 Hz and is 2 s in duration, and the third component is somewhat frequency modulated from 20 Hz to ~ 18 Hz and is 8-12 s in duration (Rankin et al. 2005). These Z-calls can occur Fig. 1 Location of autonomous acoustic recorders (AARs) off the west coast of South Africa. ...
Article
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Antarctic blue and fin whales were once abundant in the southeastern Atlantic Ocean, yet their occurrence and ecology in this region is still poorly understood. Seasonal acoustic occurrence and behaviour of Antarctic blue and fin whales off the South African west coast were determined using bio-acoustic data collected through two autonomous acoustic recorders between December 2015 and January 2017. Blue whale Z-calls were detected year-round with a peak in July, while fin whale 20 Hz pulses were detected seasonally with a peak in June by a recorder deployed at 1118 m water depth. Blue and fin whale calls were detected seasonally with a similar peak in May by a recorder deployed at 4481 m water depth. The blue whale 27 Hz chorus, and blue and fin whale 18–28 Hz chorus followed a similar trend as the seasonal acoustic occurrence of individual Z-calls and 20 Hz pulses. A maximum detection range of 800 km estimated by acoustic propagation modelling suggests that detected calls originate from whales within the South African west coast waters. Random forest models classified month of the year, wind speed, log-transformed chlorophyll-a, and sea surface temperature anomaly as the most important predictors of blue and fin whale acoustic occurrence and behaviour. Our study highlights the South African west coast as an important year-round habitat and seasonal breeding or overwintering habitat of these whales. Additionally, the year-round acoustic occurrence in this region supports the notion that blue whale migration patterns are more dynamic than previously perceived.
... Unlike songs, there is no obvious geographic variation in the D-call time-frequency shape. They have been recorded in association with blue whales in every ocean in which blue whale songs have been recorded (Barlow et al., 2021;Buchan et al., 2021;Ljungblad et al., 1997;McDonald et al., 2001McDonald et al., , 2006Mellinger & Clark, 2003;Rankin et al., 2005;Samaran et al., 2010;Schall et al., 2019). ...
Article
Monitoring the presence of blue whale (Balaenoptera musculus ssp.) stereotyped calls has been a widely used method to assess the different populations' distribution worldwide. All blue whale populations also produce nonstereotyped vocalizations, or D‐calls. Here, we monitored the presence of D‐calls in long‐term records from a large hydrophone array located in the open southern Indian Ocean, using an automated detection method and manual validation of the detections. D‐calls were detected at all sites of the array, which extends from 24°S to 56°S, but the majority of them were detected at the two southernmost sites. We observed a latitudinal shift in their seasonal occurrence, with more D‐calls in the north during austral autumn and winter and more in the south during austral spring. The geographical occurrence of D‐calls compared to that of songs indicates that blue whale acoustic behavior switches from a song‐intensive and sparse‐D‐call emission in the north to song‐moderate and more intensive D‐call emissions in the south. These findings support the hypothesis that both call types are used for different purposes, as D‐calls are mainly detected around foraging grounds and songs in wintering grounds. Monitoring both call types might therefore be a relevant acoustic indicator of blue whale behavior.
... It is likely pygmy blue whales were commonly detected along the 110 • E meridian due to their intense, low frequency calls which propagate well through the ocean. Pygmy blue whale downsweeps could not be discriminated from Antarctic blue whale D-calls (Rankin et al., 2005). Pygmy blue whale downsweeps are a suspected signal used whilst travelling, as when migrating around southwest Australia the downsweeps have been recorded simultaneously with visual confirmation to be travelling and not feeding (Recalde-Salas et al., 2014). ...
Article
The first extensive physical and biological observations of the Indian Ocean were made from 1959 to 1965, during a ship-based International Indian Ocean Expedition (IIOE-1). Decades later in 2019, the 110°E meridian was revisited during the second International Indian Ocean Expedition (IIOE-2). The aim of this study, as part of a large number of related studies, was to examine the occurrence of cetaceans and seabirds along the 110°E meridian from temperate to tropical waters (39.5–11.5°S). Cetaceans and seabirds were actively scanned for across a four-week period spanning austral autumn to winter. Acoustic recordings of vocalising cetaceans were made using directional and omnidirectional sonobuoys (n = 87 deployments). In total, seven cetacean sightings (six baleen whale, one toothed whale), 186 seabird sightings and 242 cetacean acoustic detections were recorded. The baleen whale species detected acoustically were assigned to Antarctic blue whales (Balaenoptera musculus intermedia; Z-calls), pygmy blue whales (B. m. brevicauda; south east Indian Ocean 3-component calls, D-calls), fin whales (B. physalus; 20 Hz-pulses), southern right whales (Eubalaena australis; upcalls), Antarctic minke whales (B. bonaerensis; bio-duck calls) and an unknown signal; a “spot call”. The toothed whale species detected acoustically were Kogia sperm whales (Kogia sp.) and delphinid whilstes. Pygmy blue whales were detected across Subantarctic to Tropical Surface Waters, and were the most commonly detected cetacean. There was some delineation in other cetaceans: the spot call was detected in Subantarctic and Subtropical Surface Water (south of 23°S); fin whales in Subtropical Surface Water (between 23° and 30.5°S); and Antarctic minke whales in Tropical Surface Water (between 14° and 23°S). Data were not collected on cetaceans during IIOE-1, so data here represent baseline occurrence along 110°E for future studies. A total of 22 seabird species were sighted, including, petrels, storm petrels, albatrosses, tropicbirds, terns, shearwaters, boobies, frigatebirds, gannets, gulls, skuas and prions. Soft-plumaged petrels (Pterodroma mollis) were observed across all water masses and were the most commonly sighted seabird. There was some delineation of seabird species; albatrosses were sighted south of the Subtropical Front (south of 32°S); flesh-footed shearwaters (Ardenna carneipes) in Subantarctic and Subtropical Surface Waters (south of 27°S); and tropicbirds in Tropical Surface Water (north of 20°S). The occurrence of highly mobile species is particularly important to investigate as the waters in the eastern Indian Ocean have been warming faster than in the Pacific and Atlantic Oceans.
... Worldwide, blue whales B. musculus are known to produce frequency-modulated downswept calls that are often referred to as D-calls (e.g. Berchok et al., 2006;Gavrilov et al., 2011;Mccauley et al., 2000;McDonald et al., 2001;Miller et al., 2014;Rankin et al., 2005;Recalde-Salas et al., 2014;Schall et al., 2020). While blue whale song has the apparent advantage of carrying information about population (McDonald et al., 2006), it is believed that only males produce song, and that these calls are associated with the breeding season. ...
Article
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An automated algorithm for passive acoustic detection of blue whale D-calls was developed based on established deep learning methods for image recognition via the DenseNet architecture. The detector was trained on annotated acoustic recordings from the Antarctic, and performance of the detector was assessed by calculating precision and recall using a separate independent dataset also from the Antarctic. Detections from both the human analyst and automated detector were then inspected by an independent judge to identify any calls missed by either approach and to adjudicate whether the apparent false-positive detections from the automated approach were actually true positives. A final performance assessment was conducted using double-observer methods (via a closed-population Huggins mark–recapture model) to assess the probability of detection of calls by both the human analyst and automated detector, based on the assumption of false-positive-free adjudicated detections. According to our double-observer analysis, the automated detector showed superior performance with higher recall and fewer false positives than the original human analyst, and with performance similar to existing top automated detectors. To understand the performance of both detectors we inspected the time-series and signal-to-noise ratio (SNR) of detections for the test dataset, and found that most of the advantages from the automated detector occurred at low and medium SNR.
Article
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Passive acoustic monitoring (PAM) is commonly used to obtain year-round continuous data on marine soundscapes harboring valuable information on species distributions or ecosystem dynamics. This continuously increasing amount of data requires highly efficient automated analysis techniques in order to exploit the full potential of the available data. Here, we propose a benchmark, which consists of a public dataset, a well-defined task and evaluation procedure to develop and test automated analysis techniques. This benchmark focuses on the special case of detecting animal vocalizations in a real-world dataset from the marine realm. We believe that such a benchmark is necessary to monitor the progress in the development of new detection algorithms in the field of marine bioacoustics. We ultimately use the proposed benchmark to test three detection approaches, namely ANIMAL-SPOT, Koogu and a simple custom sequential convolutional neural network (CNN), and report performances. We report the performance of the three detection approaches in a blocked cross-validation fashion with 11 site-year blocks for a multi-species detection scenario in a large marine passive acoustic dataset. Performance was measured with three simple metrics (i.e., true classification rate, noise misclassification rate and call misclassification rate) and one combined fitness metric, which allocates more weight to the minimization of false positives created by noise. Overall, ANIMAL-SPOT performed the best with an average fitness metric of 0.6, followed by the custom CNN with an average fitness metric of 0.57 and finally Koogu with an average fitness metric of 0.42. The presented benchmark is an important step to advance in the automatic processing of the continuously growing amount of PAM data that are collected throughout the world's oceans. To ultimately achieve usability of developed algorithms, the focus of future work should be laid on the reduction of the false positives created by noise.
Article
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Classification of the acoustic repertoires of animals into sound types is a useful tool for taxonomic studies, behavioral studies, and for documenting the occurrence of animals. Classification of acoustic repertoires enables the identification of species, age, gender, and individual identity, correlations between sound types and behavior, the identification of changes in vocal behavior over time or in response to anthropogenic noise, comparisons between the repertoires of populations living in different geographic regions and environments, and the development of software tools for automated signal processing. Techniques for classification have evolved over time as technical capabilities have expanded. Initially, researchers applied qualitative methods, such as listening and visually discerning sounds in spectrograms. Advances in computer technology and the development of software for the automatic detection and classification of sounds have allowed bioacousticians to quickly find sounds in recordings, thus significantly reducing analysis time and enabling the analysis of larger datasets. In this chapter, we present software algorithms for automated signal detection (based on energy, Teager–Kaiser energy, spectral entropy, matched filtering, and spectrogram cross-correlation) as well as for signal classification (e.g., parametric clustering, principal component analysis, discriminant function analysis, classification trees, artificial neural networks, random forests, Gaussian mixture models, support vector machines, dynamic time-warping, and hidden Markov models). Methods for evaluating the performance of automated tools are presented (i.e., receiver operating characteristics and precision-recall) and challenges with classifying animal sounds are discussed.
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Calling blue and fin whales have been tracked using relative travel times and amplitudes from both direct and multipath arrivals to a seafloor array of seismometers. Calls of three fin whales swimming in the same general direction, but several kilometers apart, are believed to represent communication between the whales because of signature differences in call character, an alternating call pattern, and coordination of call and respiration times. Whale call tracks, call patterns, call character, and swimming speeds were examined during periods with and without the presence of noise. Noise sources included airguns, when the whales were subject to sound levels of up to 143 dB P-P (peak-to-peak) re: 1 pPa over the 10 to 60-Hz band, and transits of merchant ships, when the whales received continuous levels up to 106 dB rms re: I / • Pa over the 10 to 60-Hz band (115 dB P-P). Whale responses associated with these noises remain arguable. ¸ 1995 Acoustical Society of America.
Article
Blue whales were seen often in the Indian Ocean. They occur over a wide latitudinal range even in mid-summer when they are normally thought to be feeding in the Antarctic. The blue whales seen in lower latitudes are pygmy blue whales, Balaenoptera musculus brevicauda. In mid-summer they are segregated from the true blue whales B. m. musculus (being found in more northerly waters) although the sub-species cannot be distinguished at sea due to difficulties in identification. The present analysis also suggests pygmy blue whales are found in the W region of the South Pacific and the E region of the South Atlantic. -from Authors
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Reviews the origins of the current stock boundaries used by the International Whaling Commission, intended to provide the background to the IWC decision to fund work on the use of biochemical techniques to examine stock identity as a priority in its Comprehensive Assessment programme. -from Author
Article
Recordings of low?frequency sounds were made in the presence of blue whales (Balaenoptera musculus) in 1998 and 1999 in waters off northern Peru during a marine mammal research cruise. Recordings were made on DAT from sonobuoys deployed near vocalizing whales. Photographic identifications and biopsy samples were collected from several animals that were recorded. Over 20 signals were extracted and analyzed from DAT segments that contained the best signal quality. All signals from the 1998 encounter consisted of rapid, frequency modulated (FM) downsweeps (begin frequency x?=94, S.D.=13.6 Hz; end frequency x?=59, S.D.=7.8 Hz; frequency range x?=41.7, S.D.=14.9) of relatively short duration (x?=1.2, S.D.=0.3 s). Approximately half of the signals exhibited a characteristic s?shape pattern when examined spectrographically. Long duration sweeps and pulses, typical of blue whales sounds from the eastern North and Central Pacific, were not detected. The vocalizations recorded in this study are compared to recordings of short duration blue whale sounds made elsewhere, including off California, Baja California, and southwestern Australia. In general, short duration blue whale sounds from these other areas are more variable and/or have different frequency characteristics. The function of short duration, FM blue whale sounds is unknown.
Article
Through January-April 2000 research was carried out off the Rottnest trench to search for blue or pygmy blue whales. A consortium of researchers carried out aerial surveys, boat based studies and acoustical measures. Historical records led us to believe that a Western Australian population of pygmy blue whales (Balaenopteridae musculus brevicauda, sub species of the true blue whale, B. m. musculus) existed, while a preliminary boat survey in 1994 suggested that some of these animals aggregated in the Rottnest trench west of Perth. This was confirmed in the early 2000 observations, in 30 days boat based searching 17 pygmy blue whales were sighted. Five thousand acoustic records were made, almost all of which had blue/pygmy blue whale calling in, some having up to six animals calling at once. Although of a slightly different format, recorded call components were of a similar character to those described from other populations. Also common were impuslive 'clicking' calls which were shorter than the 12-23 s blue whale call components and of low to very low frequency (< 1 Hz to 20 Hz). The literature suggests these are produced by fin whales but none were sighted. The low frequency (< 100 Hz) sea noise spectra from a series of 90 s recordings made every 10 minutes for 33.5 days was dominated was dominated by blue whale calling.