Technical Reports of the Institute of Cetacean Research (2019)

pp. 1–8

Technical Report (not peer reviewed)

What do we know about whales and the ecosystem in the Indo-Paciﬁcregion of the Antarctic? Part 1:
Summary of results on assessment and management of Antarctic
minke whales
Tsutomu Tamura*, Koji Matsuoka and Luis A. Pastene
Institute of Cetacean Research, 4–5 Toyomi-cho, Chuo-ku, Tokyo 104–0055, Japan*Contact e-mail: tamura@cetacean.jp

ABSTRACTThe Institute of Cetacean Research has conducted whale research under special scientiﬁc permit in the Antarctic since the austral summer season of 1987/88. The research was conducted systematically under diﬀerent research programs such as JARPA and JARPAII, and more recently, under the NEWREP-A. These research
programs employed both lethal and non-lethal methods. NEWREP-A ceased after the 2018/19 austral summer
season as an eﬀect of Japanʼs decision to withdraw from the International Whaling Commission and start
commercial whaling within its Exclusive Economic Zone. This paper summarizes the most relevant outputs
from the Japanese research programs under special scientiﬁc permit in the Indo-Paciﬁc region of the Antarctic
related to the assessment and management of Antarctic minke whales.

INTRODUCTION
Japan conducted systematic research on whales and theAntarctic ecosystem for more than 30 years. The ﬁrst
research program was the Japanese Research Program
under Special Permit in the Antarctic (JARPA), which was
followed by JARPAII and subsequently by the New Scientiﬁc Whale Research Program in the Antarctic Ocean(NEWREP-A). The Institute of Cetacean Research (ICR)
has been the institution in charge of designing and implementing those research programs. Tamura et al. (2017)provided details on the objectives, sampling and analytical methodology of these three research programs.
Several international review workshops (e.g. IWC, 2015)
discussed and evaluated the large amount of data andresults from those research programs.
As an eﬀect of the change in the whaling policy of theGovernment of Japan, NEWREP-A ceased from 1 July
2019. From now on, Japan will conduct whale researchin the Antarctic based solely on non-lethal methods. At
this point, it was considered important to summarize theknowledge on whales and the Antarctic ecosystem accumulated so far by the Japanese whale research in the
Antarctic.
The objective of this paper was to summarize the most
relevant outputs from the Japanese research programs

under special scientiﬁc permit in the Indo-Paciﬁc regionof the Antarctic related to the assessment and management of Antarctic minke whales.

SURVEYS, DATA AND SAMPLES
Surveys were conducted in the Indo-Paciﬁc region of
the Antarctic, which correspond to the International
Whaling Commission (IWC)ʼs management Areas III, IV,
V and VI (Figure 1). Survey and sampling methodologiesof the Japanese whale research programs in the Antarctic were explained in Pastene et al. (2014) and Tamura
et al. (2017). A list of data and samples collected by theJapanese whale research programs in the Antarctic wasavailable from the IWC (2015).
The most relevant research outputs for assessmentand management (i.e. taxonomy/stock structure, abundance and biological parameters) of Antarctic minke
whales are presented in the next section.

MAIN RESEARCH OUTPUTS
Taxonomical aspects
Until relatively recently, only one species of minke whale
was thought to exist: Balaenoptera acutorostrata Lacèpéde, 1804. However, Rice (1998) reviewed morphological (e.g. Williamson, 1959; van Utrecht and van der
Spoel, 1962; Kasuya and Ichihara, 1965; Omura, 1975;1

Technical Reports of the Institute of Cetacean Research (2019)

Figure 1.

Research area of the JARPA, JARPAII and NEWREP-A in the Indo-Paciﬁc region of the Antarctic.

Figure 2.

External appearance of Antarctic minke (above) and dwarf minke (below) whales.

Best, 1985) and genetic (e.g. Wada et al., 1991; Arnason
et al., 1993; Pastene et al., 1994) data collected fromextant minke whale populations and re-speciﬁed two
species, the Antarctic minke whale B. bonaerensis Burmeister, 1867, which is restricted to the Southern Hemisphere, and the common minke whale B. acutorostrata
Lacèpéde, 1804, which is distributed globally.
In the Southern Hemisphere the common minke whale
is known as ʻdiminutiveʼ or ʻdwarfʼ minke whale (Best,
1985; Arnold et al., 1987). Currently, the dwarf minke
whale is considered an un-named sub-species of B.
acutorostrata. A large amount of biological and genetic
information was obtained from 16 dwarf minke whalescaught by earlier JARPA surveys in the Indo-Paciﬁc regionof the Antarctic. Earlier genetic analyses based on thosesamples and mitochondrial DNA (mtDNA) showed thatdwarf and North Paciﬁc minke whales were more similar to each other than they were to the Antarctic minke
whale (Wada et al., 1991; Pastene et al., 1994).
2

Kato and Fujise (2000) reported on the morphology,
growth and life history of dwarf minke whales based on
16 animals sampled by JARPA in the Indo-Paciﬁc regionof the Antarctic. Results of their analyses conﬁrmed several of the morphological and morphometric characters
reported previously, and the striking diﬀerences with
the Antarctic minke whale. The mean body length at
physical maturity was estimated at 7.0 m and 6.6 m forfemales and males, respectively. The age and length atsexual maturity in females was 7–10 years and 6.0–6.5 m,
respectively. Furthermore, the authors stated that theconception of the dwarf minke whale is highly concentrated in mid-winter. Some additional morphologicaland morphometric analyses of minke whales worldwide,
including dwarf minke whales, were conducted by Nakamura et al. (2014) and Kato et al. (2015).
On the basis of the morphological diﬀerences documented by Best (1985) and Arnold et al. (1987) andgenetic diﬀerences (Wada et al., 1991) the IWC Scien-

What do we know about whales and the ecosystem in the Indo-Pacific region of the Antarctic? Part 1

tiﬁc Committee (IWC SC) recognized the existence of two
southern minke whales and agreed that the two minke
whales in the Southern Hemisphere should deﬁnitively
be considered separately for management purposes(IWC, 1991). In 1993, after examining the genetic information provided by Pastene et al. (1994), the IWC SC
recommended the inclusion of the dwarf minke whalein the Schedule, so that catch limits for Antarctic minke
whales recognise the distinction between the two southern minke whales (IWC, 1994).
In 2000, the IWC SC again recognised the two species but deferred a decision on other nominal taxa until
the completion of a worldwide review of genetic andnon-genetics information for minke whales (IWC, 2001).
Subsequent worldwide genetic analyses of minke whales
provided further evidence for the separation of the two
species, B. bonaerensis and B. acutorostrata, and at least
three sub-species of the common minke whale as recognized by Rice (1998) using mtDNA sequences (Pastene
et al., 2010) and microsatellite DNA (Glover et al., 2013).
While additional genetic and morphological/morphometric studies are required to further elucidate the taxonomic status within B. acutorostrata, including that of thedwarf minke whale, there is agreement on the taxonomic
status of B. bonaerensis. Figure 2 shows the external
morphological appearance of Antarctic minke and dwarf
minke whales.
Stock structure of the Antarctic minke whale
Detailed information on stock structure of the Antarctic
minke whale was provided by Taguchi et al. (2017). The
main ﬁndings are summarized here.
The primary data sources for studies of stock structure in this species were from the JARPA and JARPAII
programmes, both of which had stock identity as one of
their objectives. Studies based upon these samples have
proved to be more useful than those from the commercial period, given the wider geographical coverage andthe more random sampling design. In contrast, the commercial whaling samples were taken mainly from areas ofhigh density near the ice-edge (Pastene, 2006). Initially,
the JARPA genetic studies on stock structure were basedon mtDNA Restriction Fragment Length Polymorphism(RFLP), and considerable genetic heterogeneity in Areas
IV and V was found (Pastene et al., 1993; 1996).
Consideration of a suite of information from mtDNA
and microsatellite DNA (Pastene et al., 2006), morphometrics for 10 external measurements (Fujise, 1995;
Hakamada, 2006) and mean length at physical maturity(Bando et al., 2006) led to the conclusion that Antarctic

minke whales in the feeding grounds between Areas
IIIE and VIW do not comprise a single stock (IWC, 2008).
Rather, the results are consistent with the occurrenceof at least two genetic stocks in these feeding grounds,
which are probably related to the two proposed breedingareas in the eastern Indian Ocean and western South Paciﬁc (Kasamatsu et al., 1995). The following names havebeen proposed for these stocks: Eastern Indian Ocean
Stock (I-Stock) and Western South Paciﬁc Ocean Stock(P-Stock) (Pastene, 2006). The analyses of the JARPA dataalso suggested an area of transition in the region around150°–165°E, across which there is an as yet undetermined level and range of mixing (IWC, 2008).
Given additional data from JARPAII, the analysesof stock structure were reﬁned in two ways: the ﬁrst
involved additional laboratory work for additional
genetic markers, and the second involved modiﬁed
analytical measures. All the Antarctic minke whales
taken by JARPAII between 2005/06 and 2010/11 weresequenced for a 340 bp-segment of the control region of
the mtDNA (instead of the mtDNA RFLP used in JARPA),
and genotyped using 12 microsatellite loci (instead of
the six used in JARPA). Results of the heterogeneity test
for both markers showed statistically signiﬁcant genetic
diﬀerences between whales in the two sectors, western(35°–130°E) and eastern (165°E–145°W), conﬁrming that
diﬀerent stocks inhabit the Indian and Paciﬁc sectors of
the Antarctic (the I- and P-stocks). Microsatellite DNA
analyses showed more dispersal amongst males thanfemales and some degree of annual variation (Pastene
and Goto, 2016).
Schweder et al. (2011) developed an integrated approach for estimating longitudinal segregation of two
stocks using various sources of data: morphometric,
microsatellite and mtDNA data. This approach revealed
that the soft boundary (or transition area) suggested
previously could vary by year and sex. A joint likelihood
function was deﬁned to estimate mixing proportions
and apply statistical tests without assuming any baseline
populations. The approach was originally applied to the
JARPA data (Schweder et al., 2011) and subsequently to
JARPA and JARPAII data (Kitakado et al., 2014). The resultsof this approach conﬁrmed the occurrence of at least two
stocks (I and P) in the Indo Paciﬁc sector of the Antarctic.
Furthermore, the results indicated that the spatial distribution of the two stocks had soft boundaries in Area
IVE (100°–130°E) and VW (130°–165°E), which changes
by year. Results also suggested possible sex diﬀerencesin the pattern of distribution of the two stocks (Kitakado
et al., 2014).
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Technical Reports of the Institute of Cetacean Research (2019)

Figure 3.

Hypothesis on stock structure of the Antarctic minke whale. The upper ﬁgure shows the encounter ratesof Antarctic minke whales in 10° squares of latitude and longitude in waters 0°–30°S during October(Kasamatsu et al., 1995). The high sighting densities in the eastern Indian Ocean and western South Paciﬁc
could correspond to breeding grounds of this species. At least two stocks (I- and P-stocks) occur in the research area of JARPA and JARPAII, which mix through a transition area. The transition area and the mixing
rate appears to change by year and sex (Pastene and Goto, 2016).

Consequently, the structure of Antarctic minke whale
in Areas IIIE to VIW appears to be more complex thanoriginally thought. In summary, the IWC SC agreed(IWC, 2008) that there are (at least) two stocks with awide mixing area that may change by year and sex. The
extent of the I-stock to the west of 35°E and that of Pstock to the east of 145°W cannot be investigated due toa lack of appropriate samples.
The identiﬁcation of spatial and temporal boundaries
of stocks has been important for the interpretation of
the biological parameters estimated of this species in the
Antarctic (i.e. Punt et al., 2014; Bando, 2017), and should
also be important for the interpretation of abundance
and abundance trends in the near future.
Figure 3 shows a schematic representation of the stock
structure hypothesis derived from diﬀerent analyses
based on JARPA and JARPAII data.
Abundance of Antarctic minke whales
Detailed information on abundance estimates of Antarc4

tic minke whales was provided by Hakamada and Matsuoka (2017). The main ﬁndings are summarized here.
Hakamada et al. (2013) estimated abundance andabundance trends of Antarctic minke whales in the Indian sector (Area IV) and Paciﬁc sector (Area V) based onJapanese dedicated sighting surveys (JARPA and JARPAII),
under the assumption of g(0)=1. Abundance estimates
for the Indian sector range from 16,562 (CV=0.542) in
1997/98 to 44,945 (CV=0.338) in 1999/00, while thosefor the Paciﬁc sector range from 74,144 (CV=0.329) in
2004/05 to 151,828 (CV=0.322) in 2002/03. Estimates of
the annual rates of increase in abundance are 1.8% with
a 95% CI of [−2.5%, 6.0%] for the Indian sector and 1.9%
with a 95% CI of [−3.0%, 6.9%] for the Paciﬁc sector.
Adjustments to allow for the g(0) being less than 1
were made by the application of a regression model, developed from the results of the Okamura–Kitakado (OK)method estimate of minke whale abundance from the
IDCR/SOWER surveys, which provides estimates of g(0)from the statistics of the minke whale school size distri-What do we know about whales and the ecosystem in the Indo-Pacific region of the Antarctic? Part 1

bution in a stratum. With this adjustment, abundance
estimates increased by an average of 32,333 (106%) for
the Indian sector and 89,245 (86%) for the Paciﬁc sector,
while the estimates of annual rates of increase and their
95% CIs changed slightly to 2.6% [−1.5%, 6.9%] for the Indian sector and 1.6% [−3.4%, 6.7%] for the Paciﬁc sector.
See Figure 4 for the abundance trends in the two sectors.
In 2012 the IWC SC agreed to a new best abundance
estimate for Antarctic minke whales in Antarctic open waters south of 60°S, based on IDCR/SOWER sighting data.
The estimates were 720,000 based on the sighting datacollected during the CPII (1985/86–1990/91) with 95% CI(512,000, 1,012,000), and 515,000 based on the sighting
data collected during the CPIII (1992/93–2003/04) with
95% CI (361,000, 733,000). No signiﬁcant statistical diﬀerences were found between the CPII and CPIII estimates(IWC, 2013).
The estimates in the Indian sector (Area IV) were55,237 (CV: 0.17) in CPII and 59,677 (CV: 0.34) in CPIII. In
the Paciﬁc sector (Area V) were 300,214 (CV: 0.13) in CPII
and 183,915 (CV: 0.11) in CPIII.
By considering the 95% CIs of the estimates it can besuggested that the stocks of Antarctic minke whales are

Figure 4.

broadly stable with at most a slight decline, therefore theconclusion on trends is similar to that derived from JARPA
and JARPAII data in Areas IV and V.
Abundance of Antarctic minke whales should be interpreted in the near future in the context of additional
information on stock structure of the species in the IndoPaciﬁc region of the Antarctic.
Biological parameters of Antarctic minke whale
Detailed information on biological parameter estimatesof Antarctic minkes whale was provided by Bando (2017).
The main ﬁndings are summarized here.
Estimates of biological parameters of Antarctic minke
whales based on samples obtained during the JARPA, aresummarized in Table 1 (IWC, 2008; Bando, 2017).
The grouping of samples for the estimates of biologicalparameters considered the information on stock structure summarized above.
Age dependent natural mortality was estimated fromthe Statistical Catch-at-Age (SCAA) model. The pattern
was similar for the I and P stocks with natural mortality
being higher in young and old animals. It was calculated
as 0.048 (for age=15) to 0.107 (for age=35) for the I-

The best-case estimates of annual abundance of Antarctic minke whale in the Indian sector (Area IV) andPaciﬁc sector (Area V) together with their 95% CIs. The IDCR/SOWER estimates for a common northernboundary for CPII and CPIII are shown by the open triangles. Conﬁdence intervals include allowance for
additional variance. The dashed curves indicate the 95% CIs for the exponential model (after Hakamada
et al., 2013).

Table 1
Summary of biological parameters of Antarctic minke whales estimated according to one stock structure hypothesis(after IWC, 2008).

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Technical Reports of the Institute of Cetacean Research (2019)

Figure 5.

Time-trajectories of total (1+) population size (upper panels), age-speciﬁc natural mortality (center panels), and total (1+) population size relative to carrying capacity (lower panels) for three ways to model natural mortality (the Siler model, autoregressive and piecewise linear) of two stocks of Antarctic minke whale(from Punt et al., 2014). Biological data obtained from both JARPA and JARPAII were used in the SCAA.

stock and 0.046 (for age=15) to 0.103 (for age=35) forP-stock (Figure 5) (Punt et al., 2014). Time-trajectories
derived from SCAA showed that total population size of
the Antarctic minke whale increased until 1970ʼs andthen declined until 2000ʼs for both stocks (Figure 5) (Puntet al., 2014).

CONCLUSIONSThe Japanese whale research programs under special
permit have provided important information on stock
structure, biological parameters and abundance of Antarctic minke whales in the Indo-Paciﬁc region of the Antarctic. Such information is key for the assessment of thespecies and the application of management procedures
such as the IWCʼs Revised Management Procedure (RMP)
in the future. Continuation of the systematic monitoringof the stocks of Antarctic minke whales is recommended.

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