ICR

No. 4
December 2020

TECHNICAL REPORTS OF THE
INSTITUTE OF CETACEAN RESEARCH
TEREP-ICR is published annually by the InsƟtute of Cetacean Research.
ISSN: 2433-6084
Copyright © 2020, The InsƟtute of Cetacean Research (ICR).

CitaƟonEnƟre issue:InsƟtute of Cetacean Research. 2020. Technical Reports of the InsƟtute of Cetacean Research (TEREP-ICR) No. 4. The InsƟtute of Cetacean Research, Tokyo, Japan, 79 pp.
Individual reports:‘Author Name’. 2020. ‘Report Ɵtle’. Technical Reports of the InsƟtute of Cetacean Research (TEREPICR) No. 4: ‘pp’-’pp’.

Editorial correspondence should be sent to:
Editor, TEREP-ICR
The InsƟtute of Cetacean Research,
4-5 Toyomi-cho, Chuo-ku, Tokyo 104-0055, Japan
Phone: +81(3) 3536 6521
Fax: +81(3) 3536 6522
E-mail: webmaster@icrwhale.org
TEREP-ICR is available online at www.icrwhale.org/TEREP-ICR.html

Cover photo: Stereoscopic microscope used for age determinaƟon based on counƟng of earplug
growth laminae (top); capillary sequencer ABI SeqStudio used for DNA sequencing analysis (middle);
ultra performance liquid chromatography-tandem mass spectrometer used for age determinaƟon
based on racemizaƟon analysis (boƩom).
Copyright © 2020, The InsƟtute of Cetacean Research (ICR).
TECHNICAL REPORTS OF THE
INSTITUTE OF CETACEAN RESEARCHTEREP-ICR
No. 4

The Ins�tute of Cetacean Research (ICR)
Tokyo, 2020

Technical Reports of the Ins�tute of Cetacean Research (2020)

pp. i

Foreword
It is a pleasure for me to introduce the fourth issue of the Technical Reports of the Ins�tute of Cetacean Research (TEREPICR-4). Following the change in Japanʼs whaling policy last year, the ICR has been designing and implemen�ng whale
research programs based on non-lethal methods. In the North Paciﬁc, ICR scien�sts have contributed to the design andimplementa�on of na�onal dedicated sigh�ng surveys and of the IWC-sponsored Paciﬁc Ocean Whale and Ecosystem
Research (POWER) surveys. In the Antarc�c, ICR scien�sts designed the new program called ʻJapanese Abundance andStock structure Surveys in the Antarc�cʼ (JASS-A), and the ﬁrst survey under this program was completed successfully
during the 2019/20 austral summer season. In addi�on, ICR scien�sts commenced the collec�on of biological samples
and data based on whales caught for commercial purposes in Japanʼs Exclusive Economic Zone since 1 July 2019.
Consistent with its stated objec�ves, TEREP-ICR describes and reports on the process, progress, and results of technical or scien�ﬁc research, or the state of technical or scien�ﬁc research programs conducted by the ICR, including thosecommenced recently.
ICR scien�sts were preparing for their par�cipa�on in several na�onal and interna�onal mee�ngs planned for 2020.
Due to the current Covid-19 pandemic aﬀec�ng the world, these mee�ngs were either cancelled or replaced by virtualmee�ngs (or e-mail discussions), and our scien�sts par�cipated in those virtual mee�ngs and discussions as much as
possible, given the circumstances.
Similar to previous TEREP-ICR issues, TEREP-ICR-3 was widely distributed to approximately 120 individual scien�sts
from Japan and 30 foreign countries. It was also distributed to approximately 190 research ins�tu�ons (including universi�es, research ins�tutes, public libraries, museums and aquariums), both in Japan and foreign countries. As a resultof dissemina�ng informa�on on ICRʼs research ac�vi�es through the TEREP-ICR, an increasing number of interna�onal
scien�sts are showing interest in conduc�ng research in collabora�on with the ICR. Based on this, I believe that TEREPICR is making contribu�on toward achieving its stated objec�ves. Furthermore, TEREP has been a good opportunity forour scien�sts to compile and summarize their research, conducted over the years, as a prior step before submi�ng theirworks for peer-review publica�ons.
Finally, I am pleased to announce that the reports published in TEREP are now indexed in the Japan Science and Technology Agencyʼs J-GLOBAL, which aims to link, expand and spark scien�ﬁc informa�on in Japan and the world.
I sincerely hope that this fourth issue of the TEREP-ICR will con�nue to contribute to an increased understanding of
the technical and research ac�vi�es conducted by the ICR among the na�onal and interna�onal scien�ﬁc communi�es.
Dr. Yoshihiro Fujise
Director General ICRTokyo, December 2020iTechnical Reports of the Ins�tute of Cetacean Research (2020)

pp. iii

Editorial
Welcome to the fourth issue of the Technical Reports of the Ins�tute of Cetacean Research (TEREP-ICR-4).
This issue contains eight technical reports and one commentary ar�cle. We con�nue with the series of reports summarizing the research ﬁndings on whales and the ecosystem in the Indo-Paciﬁc sector of the Antarc�c. This �me, Tamuraand colleagues focus on the ﬁndings of the ecological research on whales and their environment. Outlines of two important whale research programs, including results of the latest surveys, are included in this issue. Isoda and colleagues explain the program ʻJapanese Abundance and Stock structure Surveys in the Antarc�c (JASS-A)ʼ, while Matsuoka explains
the IWC-sponsored program ʻPaciﬁc Ocean Whale and Ecosystem Research (IWC-POWER)ʼ. The basic concept of gene�ctagging, and its applica�on for abundance es�mates in whales, are explained in the reports by Taguchi and Takahashi,
respec�vely. Works on earplug-based age determina�on and the es�ma�on of biological parameters have con�nued,
based on the samples and data collected during the former whale research programs under special scien�ﬁc permit. This�me, Bando and Maeda show the results of age determina�on and biological parameter es�mates of North Paciﬁc sei
whales. The ﬁrst Note by Goto and Oikawa describes Japanʼs system for monitoring of whale products sold in its domes�c market through DNA registra�on, and the second Note by Konishi and colleagues focuses on the preliminary resultsof satellite-monitored tracking of Antarc�c minke and ﬁn whales during the ﬁrst JASS-A survey.
In the commentary ar�cle, Yasunaga presents his views on the use of lethal and nonlethal techniques for the researchof large whales.
This fourth TEREP-ICR issue also includes sec�ons to outline the contribu�on of ICR scien�sts to interna�onal meetings in 2020, as well as their contribu�on in terms of peer-reviewed publica�ons up to December 2020.
I hope you will ﬁnd this fourth issue informa�ve and useful.
Dr. Luis A. Pastene
Editor TEREP-ICR
Tokyo, December 2020

iii

Technical Reports of the Ins�tute of Cetacean Research (2020)

p. v

Contents
Forewordi

Editorialiii

Contentsv

Technical Reports
Tamura, T., Matsuoka, K., Yasunaga, G. and Pastene, L.A. What do we know about whales and ecosystem in the Indo-Paciﬁc region of the Antarc�c? Part 2: summary of ecological studies

1

Isoda, T., Katsumata, T., Tamura, T., Matsuoka, K. and Pastene, L.A. An outline of the Japanese Abundance and Stock structure Surveys in the Antarc�c (JASS-A) including results of the ﬁrst survey
under this new research program
12
Matsuoka, K. An outline of the IWC-Paciﬁc Ocean Whale and Ecosystem Research (IWC-POWER)
including results of the 2019 survey
23
Taguchi, M. Gene�c tagging technique: basic concept and a case study by the Ins�tute of Cetacean
Research
35
Takahashi, M. Use of gene�c data for abundance es�mate purposes: a brief review of methods andcase studies by the Ins�tute of Cetacean Research
41
Bando, T. and Maeda, H. Progress in the research on earplug-based age determina�on and biologicalparameters of North Paciﬁc sei whales at the Ins�tute of Cetacean Research48

Technical Reports-Notes
Goto, M. and Oikawa, H. Japanʼs system for monitoring of whale products sold in its domes�c market
through DNA registra�on
54
Konishi, K., Isoda, T. and Katsumata, T. Satellite-monitored tracking of Antarc�c minke and ﬁn whales
under the ﬁrst JASS-A survey in the 2019/2020 austral summer season
57

Commentary
Yasunaga, G. Comments on the use of lethal and non-lethal techniques in the studies of large
whales59

Interna�onal mee�ngs

61

Peer-reviewed publica�ons

62v

Technical Reports of the Ins�tute of Cetacean Research (2020)

pp. 1–11

Technical Report (not peer reviewed)

What do we know about whales and ecosystem in the Indo-Paciﬁcregion of the Antarc�c? Part 2: summary of ecological studies
Tsutomu Tamura*, Koji Matsuoka, Genta Yasunaga 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 Ins�tute of Cetacean Research conducted whale research under special scien�ﬁc permit in the Antarc�cstar�ng from the austral summer season 1987/88. The research was conducted systema�cally under diﬀerent
research programs such as JARPA and JARPAII, and more recently, under NEWREP-A. These research programs
employed both lethal and non-lethal methods. NEWREP-A ceased a�er the 2018/19 austral summer season asa consequence of Japanʼs decision to withdraw from the Interna�onal Conven�on for the Regula�on of Whaling. Japanʼs whale research con�nues in the Antarc�c, using non-lethal methods only. This paper summarizes
the most relevant ecological outputs from Japanʼs whale research under special scien�ﬁc permit in the IndoPaciﬁc region of the Antarc�c.

INTRODUCTION
Japan conducted systema�c research on whales and theAntarc�c ecosystem for more than 30 years. The ﬁrst
research program was the Japanese Whale Research
Program under Special Permit in the Antarc�c (JARPA),
which was followed by JARPAII and subsequently by the
New Scien�ﬁc Whale Research Program in the Antarc�c
Ocean (NEWREP-A). The Ins�tute of Cetacean Research(ICR) was the ins�tu�on in charge of designing and implemen�ng those research programs. Tamura et al. (2017)
have provided details on the objec�ves, sampling andanaly�cal methodology of the three research programs.
Several interna�onal review workshops (e.g., IWC, 2015)
discussed and evaluated the large amount of data andresults from these research programs.
As a consequence of Japanʼs change in whaling policy,
the NEWREP-A ceased on 30 June 2019, the date of Japanʼs withdrawal from the Interna�onal Conven�on for
the Regula�on of Whaling (ICRW). From the 2019/20 austral summer season, Japan started whale research in the
Antarc�c using non-lethal methods. The new research
program is called the Japanese Abundance and Stockstructure Surveys in the Antarc�c program (JASS-A) (seeIsoda et al., this issue).
At this point, it was considered important to summarize the knowledge on whales and the Antarc�c ecosystem accumulated so far by Japanʼs whale research in
the Antarc�c. The objec�ve of this paper is to summarize

the most relevant ecological outputs from Japanʼs whale
research under special scien�ﬁc permit in the Indo-Paciﬁcregion of the Antarc�c.

SURVEYS, DATA AND SAMPLES
Surveys were conducted in the Indo-Paciﬁc region of the
Antarc�c, which correspond to the Interna�onal Whaling Commission (IWC) Management Areas III, IV, V andVI (Figure 1). Survey and sampling methodologies of
the Japanese whale research programs in the Antarc�cwere described in Pastene et al. (2014) and Tamura
et al. (2017). A list of data and samples collected by thesewhale research programs in the Antarc�c is available in
IWC (2015).

MAIN RESEARCH OUTPUTS ON WHALE ECOLOGY
Abundance and abundance trends of baleen whales
JARPA and JARPAII conducted vessel-based sigh�ng surveys under the Line Transect Method with the aim ofes�ma�ng abundance and abundance trends of large
whales in the Indo-Paciﬁc sector of the Antarc�c. Detailsof the methodology used are available in Tamura et al.
(2017) and Hakamada and Matsuoka (2017). The la�erauthors summarized the informa�on on abundance andabundance trend of Antarc�c minke whales, which is notrepeated here. Figure 2 shows pictures of the relevantbaleen whale species.
Baleen whale species other than the Antarc�c minke
whale were depleted by unregulated commercial whaling1Technical Reports of the Ins�tute of Cetacean Research (2020)

Figure 1.

Figure 2.

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

Baleen whale species in the Indo-Paciﬁc region of the Antarc�c: (a) Antarc�c minke whale; (b) humpback
whale; (c) ﬁn whale; (d) southern right whale; (e) Antarc�c blue whale.

in the past, and there is an interest to inves�gate the recovery of those species a�er protec�on. Fortunately, theabundance of large whale species such as humpback, ﬁn,southern right and blue whales has been increasing, albeit at diﬀerent rates over the past decades. An increasein the abundance of large baleen whales has implica�ons
for the Antarc�c ecosystem as a whole (Fujise and Pastene, 2018).
The scien�ﬁc evidences for the recovery of baleen
whale species are shown below.
Humpback whales
Abundance and abundance trend should be ideally es�mated on a biological stock basis. The IWC SC has iden�ﬁed seven breeding stocks of humpback whales in theSouthern Hemisphere, which are denominated with alphabe�c le�ers from ʻAʼ to ʻGʼ (IWC, 2011). The breeding
stocks occurring in the Indo-Paciﬁc sector of the Antarc�c
are Stock ʻCʼ (mainly in Area III), ʻDʼ (mainly in Area IV),
ʻEʼ (mainly in Area V), and ʻFʼ (mainly in Area VI) (Kanda
et al., 2014). There are some geographical overlaps be2

tween adjacent stocks in the Antarc�c.
Abundance and abundance trend es�mates basedon Japanese surveys have focused mainly on Areas IV(Breeding Stock D) and V (Breeding Stock E). In Area
IV the abundance was es�mated at 29,067 whales(CV=0.255) based on sigh�ng data collected in 2007/08;
in Area V the abundance was es�mated at 13,894 whales(CV=0.338) based on sigh�ng data collected in 2008/09(Hakamada and Matsuoka, 2014a).
Figure 3 shows the abundance trend of Breeding Stocks
D and E based on JARPA and JARPAII data. For comparison
purposes, the ﬁgures include data from the Interna�onal
Decade for Cetacean Research/Southern Ocean Whaleand Ecosystem Research (IDCR/SOWER) programs. The
ﬁgures show a clear increasing trend which is consistent
for JARPA, JARPAII and IDCR/SOWER survey data. Annual
rate of increase was es�mated at 13.6% (95% CI=8.4–
18.7%) and 14.5% (95% CI=7.6–21.5%) for Areas IV and V,
respec�vely. The rate of increase in Areas IV and V weresta�s�cally signiﬁcant (Hakamada and Matsuoka, 2014a).
The current abundance of Breeding Stock D in Area

What do we know about whales and ecosystem in the Indo-Pacific region of the Antarctic? Part 2

Figure 3.

Abundance es�mates for humpback whales in Areas IV and V (south of 60°S). Es�mates were based onsigh�ng data collected by JARPA and JARPAII between 1989/90 and 2008/09 primarily during January to
February. Es�mates from the IDCR-SOWER surveys (Branch, 2011) are shown for compara�ve purposes(ﬁlled circles). Ver�cal lines show 95% conﬁdence intervals (a�er Hakamada and Matsuoka, 2014a).

Figure 4.

Abundance trend of ﬁn whales in the Indo-Paciﬁc region of the Antarc�c, south of 60°S. Ver�cal lines show
95% conﬁdence intervals (a�er Matsuoka and Hakamada, 2014).

IV is close to that of its pre-exploita�on level (IWC HP:
h�ps://iwc.int/status).
Fin whales
There is limited informa�on on stock structure of ﬁnwhales in the Indo-Paciﬁc sector of the Antarc�c. For the
purpose of the abundance es�mates based on JARPA and
JARPAII surveys, the whole area was divided into westernarea (assuming an Indian Ocean stock in Area IIIE+IV)
and eastern area (assuming a Paciﬁc Ocean stock in Area
V+VIW). Figure 4 shows the abundance es�mates of ﬁnwhales in Areas IIIE+IV and V+VIW, plo�ed against thesurvey seasons. An increasing trend in abundance is evident for these two areas. For the western area the abundance south of 60°S was es�mated at 3,087 (CV=0.191)
in 1995/96, and 2,610 (CV=0.285) in 2007/08. For theeastern area the abundance south of 60°S was es�matedat 1,879 (CV=0.226) in 1996/97, and 14,981 (CV=0.298)
in 2008/09.
For the western area the increasing trend between
1995/96 and 2007/08 seasons was es�mated at 8.9%

(95% CI: −0.145%, 32.4%), while the trend in the eastern area between 1996/97 and 2008/09 was es�matedat 12.0% (95% CI: 2.6%, 21.5%). The es�mate for theeastern area was sta�s�cally signiﬁcant (Matsuoka andHakamada, 2014).
Southern right whales
During the JARPA and JARPAII surveys in Areas III–VI,
this species was mainly sighted in Area IV. In this Area,abundance es�mates (south of 60°S) ranged from 6(CV=0.761) in the 2003/04 season to 1,557 (CV=0.303) inthe 2007/08 season.
Figure 5 shows the abundance es�mates in Area IV
plo�ed against the survey seasons. The abundance trend
was es�mated at 5.9% (95% CI: −16.4%, 28.1%) between
1989/90 and 2007/08, which was not sta�s�cally signiﬁcant (Matsuoka and Hakamada, 2014).
Blue whales
There is limited informa�on on stock structure of blue
whales in the Antarc�c. Abundance of this species for the3Technical Reports of the Ins�tute of Cetacean Research (2020)

Figure 5.

Abundance es�mates of southern right whales
in Area IV (south of 60°S). Ver�cal lines show
95% conﬁdence intervals (a�er Matsuoka andHakamada, 2014).

Figure 6.

Abundance trend of blue whales in the IndoPaciﬁc region, south of 60°S. Ver�cal lines show
95% conﬁden�al intervals (a�er Matsuoka andHakamada, 2014).

Indo-Paciﬁc region of the Antarc�c (35°E–145°W), southof 60°S was 664 (CV=0.328) in 2005/06 and 2006/07
seasons. The abundance was es�mated at 1,223 whales(CV=0.345) in the 2007/08 and 2008/09 seasons.
Figure 6 shows the abundance es�mates for the whole
Indo-Paciﬁc region plo�ed against the survey seasons.
The abundance trend was es�mated at 8.2% (95% CI:
3.9%, 12.5%) between 1995/96 and 2008/09. This es�mate was not sta�s�cally signiﬁcant.
Future surveys and analyses
Dedicated sigh�ng surveys were planned for the seasons
2009/10–2013/14 under JARPAII. However, those surveys
could not be conducted due to external interferences by
an NGO (Nishiwaki et al., 2014). Subsequently, dedicatedsigh�ng surveys were conducted under the NEWREP-A
between 2015/16 and 2018/19 and data collected during these surveys are under analyses. From 2019/20
4

Figure 7.

Occupied area indices of Antarc�c minke and humpback whales in Area IV in three periods of JARPA and
JARPAII (Murase et al., 2014).

Japan started the JASS-A, which will be conducted fora tenta�ve period of eight years. The main objec�ve of
JASS-A is to es�mate the abundance of large whales in
the Indo-Paciﬁc region of the Antarc�c (Government ofJapan, 2019) (see Isoda et al., this issue). Data collectedduring NEWREP-A and JASS-A will be analyzed by ICR
scien�sts in conjunc�on with previous data in the samearea.
Geographical distribu�on of baleen whales
Substan�al increase in the abundance could have an
implica�on in the pa�ern of distribu�on of the species.
In the Indo sector of the Antarc�c (Area IV), humpback
whales have increased signiﬁcantly over the past decades, while the abundance of Antarc�c minke whales
has been rather stable. Under this scenario, we should
expect changes in the pa�ern of distribu�on of these two
species.

What do we know about whales and ecosystem in the Indo-Pacific region of the Antarctic? Part 2

Changes in the spa�al and temporal distribu�on patterns of Antarc�c minke and humpback whales in Area IV
were examined for the period of JARPA and JARPAII. The
spa�al distribu�on was es�mated using Generalized Addi�ve Models (GAM) (Murase et al., 2014). Presence orabsence of whales was used as a response variable while
seaﬂoor depth, distance from shelf break and longitude
were used as explanatory variables. Three �me periods
were deﬁned: ʻEarlyʼ (1989/90, 1991/92, 1993/94); ʻMiddleʼ (1995/96, 1997/98, 1999/00) and ʻLateʼ (2001/02,
2003/04, 2005/06).
Figure 7 shows the results of the GAM analyses. In theearly period, humpback whales were mainly distributedin the northern part of the survey area between longitudes 80°E and 100°E. In the middle period, humpback
whales expanded their habitat closer to the shelf break
between the longitudinal range of 80°E and 120°E. In
the late period, humpback whales were distributed in
the en�re survey area along the shelf break. While thespa�al distribu�on of humpback whales expanded during
the period of JARPA and JARPAII, no strong change wasobserved in the distribu�on of Antarc�c minke whales.
These results indicated a possible compe��on for habitat
between humpback and Antarc�c minke whales as abundance of humpback whales increased.
The pa�ern of spa�al distribu�on for other large
whales was examined by Matsuoka and Hakamada(2018). Based on JARPA and JARPAII data, there was acommon pa�ern for several whale species to concentrate
mainly in the sector between 80°E and 110°E, south of
60°S. This could be related to krill availability, as a highdensity of euphausiids was reported between 100°E and120°E in the 1999/00 JARPA survey (Murase et al., 2002).
Studies into the pa�ern of temporal (yearly) changes inrela�on to changes in environmental factors are underway.
Feeding ecology of Antarc�c minke whale
An overview of the procedures for examining stomach
content analyses by Japanese scien�sts was presented by
Tamura and Konishi (2014). The type and amount of prey
consumed by Antarc�c minke whales in the feeding season is fundamental for understanding the impact of theconsump�on on the ecosystem. The analyses of stomach
contents in Antarc�c minke whales have beneﬁted fromuseful sugges�ons and recommenda�ons from the IWC
SC.
Antarc�c minke whales feed mainly on Antarc�c krill(Euphausia superba) in oﬀshore waters, and on ice krill (E.
crystallorophias) (Figure 8) on the coastal shelf along the

Figure 8.

Ice krill (upper) and Antarc�c krill (lower).

Ross Sea and Prydz Bay (Tamura and Konishi, 2009; 2014).
The maximum weight of stomach contents of Antarc�c minke whales sampled by JARPA and JARPAII from1987/88 to 2010/11 were 142.4 kg (2.4% of body weight)
and 156.0 kg (3.4% of body weight) for immature males
and females, respec�vely. The ﬁgures were 387.0 kg(3.8% of body weight) and 326.9 kg (3.6% of body weight)
for mature males and females, respec�vely. The average
weight and rate of stomach contents per body weight(RSC) were 33.2±28.3 kg (RSC: 0.8%) and 37.6±29.1 kg(RSC: 0.9%) for immature males and females, respec�vely and 63.5±47.1 kg (RSC: 0.9%) and 71.8±54.3 kg(RSC: 0.9%) for mature males and females, respec�vely(Tamura and Konishi, 2014).
The daily prey consump�on by the whales in each
reproduc�ve status group was es�mated using energyrequirement and energy deposi�on methods. The daily
prey consump�ons of Antarc�c minke whales per capita
during the feeding season based on these two methods
were 95.1–127.0 and 182.6–250.3 kg for immature andmature males, 125.8–138.7 and 268.1–325.5 kg for immature and mature females, respec�vely. This is equivalent to 2.65–4.02% of their body weight. The total prey
consump�on of Antarc�c minke whale per capita during
the feeding season was 8.6–10.4 and 20.6–21.9 tons for
immature and mature males, and 11.3–12.5 and 32.2–
39.1 tons for immature and mature females, respec�vely(Tamura and Konishi, 2014).
Es�ma�on of the uncertain�es (e.g., allometric rela�onships, body weight of whales, energy values of preyspecies, assimila�on eﬃciency and length of feeding
period) in several components involved in es�ma�ng theamounts and types of prey consumed by North Paciﬁc baleen whales was based on sugges�ons from the IWC SC.
5

Technical Reports of the Ins�tute of Cetacean Research (2020)

Similar es�ma�ons of uncertain�es are being conducted
for Antarc�c minke whales.
In Areas III east and IV in the 2007/2008 season, theabundance of Antarc�c minke whales was es�mated at
9,406 and 14,739, respec�vely. In Area V and VI west in
the 2008/2009 season, the abundance was es�mated at
108,097 and 26,364, respec�vely (Hakamada and Matsuoka, 2014b). These abundance es�mates were used forinves�ga�ng the total consump�on of krill by Antarc�cminke whales.
In Areas III east and IV the seasonal (120 days) prey
consump�on of krill by Antarc�c minke whales wases�mated at 0.17–0.19 and 0.33–0.37 million tons,
respec�vely. In Areas V and VI west, the consump�on
was es�mated at 2.51–2.88 and 0.50–0.54 million tons,
respec�vely. The seasonal prey consump�on by Antarc�cminke whales in the total research area was es�matedat 3.51–3.98 million tons, amoun�ng to 7.6–8.6% of thekrill biomass es�mated by acous�c survey in the researcharea (Tamura and Konishi, 2014).
Energe�cs of Antarc�c minke whale
Whales generally accumulate energy as lipid in the blubber during the summer feeding period at high la�tudes,
and they spend the energy in migra�on and reproduc�onin low la�tude waters. The proxy of fat reserves in blubber have been used as an indica�on of body condi�on inwhale studies (e.g., Lockyer et al., 1985; 1986). The measurements of blubber thickness posi�vely correlate withlipid content in the whole body fat and have proven to bea dependable proxy for energy storage in whales (Lockyer
et al. 1985).
For the 18 years of JARPA period (1987/88–2004/05),
the annual trend in energy storage in the Antarc�c minke
whales was examined using blubber thickness (Figure 9)at two lateral measurement points, in addi�on to total

Figure 10.

6

fat weight in the whale body and girth measured at two
speciﬁed posi�ons.
Blubber thickness measurements were available for
4,689 whales; fat weight was available for 740 whales;
and half girth measurements were available for 4,681
animals. Regression analysis was carried out on blubber
thickness in 4,689 whales. The explanatory variables
were age, body weight, fetus length and total fat weight.
A large number of linear mixed-eﬀects sta�s�cal models
were inves�gated for each of the dependent variables,
and the Bayesian Informa�on Criterion (BIC) was used to
select the best model. All models examined had ʻyearʼ as
a possible explanatory variable. This regression analyses
clearly showed that blubber thickness had been decreasing in this period. The decrease per year in the JARPA
period was es�mated at approximately 0.02 cm for midlateral blubber thickness (Figure 10).
Similar analyses and results were found for fat weight,
which decreased 17 kg in the 18-year period (Figure 10).
Furthermore, the runs using ʻhalf girthʼ as another dependent variable also gave similar results. The total magnitude of the decline over these 18 years was 4% for girth

Figure 9.

Measurement of blubber thickness.

Temporal trends in body condi�on indicators of Antarc�c minke whales in the feeding season (a: blubber thickness; b: fatweight; c: half girth) for the Antarc�c minke whale in the feeding season (Konishi et al., 2008).

What do we know about whales and ecosystem in the Indo-Pacific region of the Antarctic? Part 2

measurements (Figure 10).
Similar analyses were conducted for stomach contents
weight (sieved contents of the forestomach). Regression
analyses using the weight of the stomach contents as thedependent variable, and explanatory variables such as
year, date, local �me, la�tude, sex and body length, wereconducted. A linear mixed-eﬀects analysis showed a 31%(95% CI=12.6–45.3%) decrease in the weight of stomach
contents over the 20 years since 1990/1991 (Konishi
et al., 2014).
The trend in stomach content was consistent withthat of the blubber thickness (Konishi and Walløe, 2015).
These results suggested that the food availability for Antarc�c minke whales may have declined in recent decades.
Monitoring of environmental pollutants
Pollutant analyses by Japanese scien�sts started under
the JARPA program for the objec�ve of ʻElucida�on of theeﬀect of environmental changes on cetaceansʼ and con�nued under the JARPAII program. The objec�ve involved
the elucida�on of the pa�ern of pollutant accumula�on
and the eﬀects of pollutants on cetaceans. Diﬀerent
kinds of pollutants were monitored, which included traceelements and organochlorine compounds (OCs). Traceelements included the toxic ones Hg, Cd, Pb and Ni (nonessen�al elements), and Cu, Zn, Fe and Mn (essen�al elements). Toxic elements induce adverse eﬀects in animals
and humans, and are more persistent than the essen�alelements. Essen�al elements are important for survival
and health of animals and humans.

Figure 11.

Details of the analy�cal procedures for pollutants were
given in Appendix 4 of Government of Japan (2005).
Regarding OCs, the levels of PCB and DDT in ﬁn andAntarc�c minke whales in the Antarc�c Ocean could be
the lowest among whale species in the world (Figure 11)(Yasunaga et al., 2006). The results during the JARPA/
JARPAII period suggested that levels of HCHs in the Antarc�c Ocean have varied from decreasing slightly to being
steady in the mid-1990s. HCB levels in large whales would
be aﬀected by trophic levels of its preys rather than byspa�al diﬀerences.
Regarding trace elements, the hepa�c Hg levels of Antarc�c minke whales of all age groups in Area IV decreased
signiﬁcantly over the research years, whereas that of
15–26 years old whales in Area V increased signiﬁcantly,
possibly indica�ng that food availability of Antarc�cminke whales in the 2000s may have diﬀered from thosein the 1980s and 1990s (Yasunaga et al., 2014).
Oceanography
Informa�on on oceanographic structure and its dynamics are important in interpre�ng changes in the Antarc�cecosystem. Changes in oceanographic condi�ons aﬀect
krill distribu�on and biomass and, in turn, the abundanceand distribu�on of whales. Changes in oceanographic
condi�ons might indicate an eﬀect of climate change. Details of the oceanographic survey can be found in Wada
(2019).
Watanabe et al. (2014) analyzed oceanographic observa�on data obtained by JARPA and JARPAII to clarify

PCB and DDT levels in the blubber of baleen whales (Yasunaga et al., 2006).
7

Technical Reports of the Ins�tute of Cetacean Research (2020)

physical oceanographic condi�ons in the JARPA/JARPAII
research area as a basis for understanding the habitat environment of whales. About 2,500 proﬁles were obtained
by XBT, XCTD and CTD observa�ons from 1990 to 2009.
Based on this data set, the averaged feature of the oceanographic structure in the research area for two decades
was described. In the area east of the Kerguelen Plateau,
the posi�on of the Southern Boundary (SB) changed on a
decadal �mescale. The SB is indicated by the 0°C contour
on 27.6σ 0. The southward shi� of the SB in the region wasobserved in the early 2000s, and the northward shi� wasobserved in the later 2000s (Figure 12).
Unlike the Antarc�c Peninsula region, the JARPA and
JARPAII temperature data showed no sta�s�cally signiﬁcant warming in the JARPA area for the two decades.
Inves�ga�on of the oceanographic condi�ons in the
Indo-Paciﬁc region of the Antarc�c is one of the secondary objec�ves of JASS-A. Data will be analyzed in conjunc�on with the large data sets produced by JARPA/JARPAII
and NEWREP-A in the Indo-Paciﬁc region of the Antarc�c.

Figure 12.

Temperature on the 27.6σ 0 isopycnal surface
for the Area IV (a) and Area V (b) (Watanabe
et al., 2014).

Figure 13.

8

Krill survey
Krill is a key prey species in the Antarc�c ecosystem, suppor�ng diﬀerent species of baleen whales, pinnipeds,
birds and ﬁsh. For this reason, the ICR has been conducting krill surveys alongside the whale sigh�ng surveys.
Sigh�ng of whales and krill surveys were carried out
concurrently in JARPAII and NEWREP-A. Details of the krill
survey can be found in Wada (2019).
Wada and Tamura (2014) presented results of krill biomass es�ma�ons based on the data from a quan�ta�veecho sounder (EK500; Simrad, Norway). Es�ma�ons weremade based on data obtained in Area IV in the 2007/08
JARPAII survey and in Area V in the 2008/09 JARPAII survey. The biomass es�mates obtained in Area IV was 12.5
million tons, and that in Area V was 24.0 million tons.
These es�ma�ons were useful to evaluate the impact of
the krill consump�on by three baleen whale species.
Regarding krill demography, informa�on on length frequency distribu�on and maturity stage of Antarc�c krill
are being obtained using samples from stomachs of Antarc�c minke whales taken in JARPA, JARPAII and NEWREPA (1989/90–2017/18). Figure 13 shows an example ofoutputs of such analyses.
Marine debris
In recent years marine debris has been recorded in the
sub-Antarc�c and Antarc�c islands. Marine debris causes
nega�ve eﬀects on whales through inges�on and by entanglement. JARPA, JARPAII and NEWREP-A conductedsystema�c monitoring of marine debris in whales andtheir environment in the Antarc�c. Details of the resultsof marine debris during JARPA/JARPAII can be found in
Isoda et al. (2018).
The density index (DI: number of marine debris per 100
nau�cal miles) of marine debris in the Antarc�c is lower
by two orders of magnitude in comparison to the North
Paciﬁc Ocean and its adjacent waters. Thus, the observa-

Length frequency distribu�on (le�) and maturity stage (right) of Antarc�c krill sampled from stomach
contents of Antarc�c minke whales in 2008/09 in the sector 130°E and 140°E.

What do we know about whales and ecosystem in the Indo-Pacific region of the Antarctic? Part 2

Figure 14.

Food web in the Antarc�c modeled in Mori and Bu�erworth (2006) and Moosa (2017).

�ons proved that the Antarc�c waters have a very low
density of marine debris on the sea surface. The level ofimpact of entanglements on Antarc�c minke whales was
low in comparison with other oceanic basins. Given the
low indices, the eﬀect of marine debris on whales in the
Antarc�c was expected to be limited at the present �me(Isoda et al., 2018).
Modelling work
An overview of the ICR work on ecosystem modelling in
the Antarc�c was presented by Hakamada and Tamura(2018). The ecological background for the modelling exercise was provided by Fujise and Pastene (2018).
Mori and Bu�erworth (2006) constructed a mul�species modelling in the Antarc�c Ocean using baleen
whales and seals as predators and krill as prey (Figure 14).
One of the main objec�ves of their modelling exercise
was to determine whether predator-prey interac�ons
alone can broadly explain observed popula�on trends
without the need for recourse to environmental change
hypotheses. Preliminary results indicated that this is likely
to be the case.
New ecosystem modelling studies are underway(Moosa, 2017).

CONCLUSIONSJapanʼs whale research programs under special permit
have provided important informa�on on the ecology of
Antarc�c minke whales and other baleen whale species
in the Indo-Paciﬁc region of the Antarc�c. The long �meseries of data produced by the research programs havebeen fundamental to the study and understanding ofabundance trends of several large whale species, thechanges in distribu�on of Antarc�c minke and humpback
whales, and the changes in nutri�onal condi�ons of the
Antarc�c minke whales. The research programs haveproduced new informa�on in helping to understand theoceanographic structure of the Indo-Paciﬁc region of the
Antarc�c.
Also, studies on the distribu�on, abundance and demography of the krill have commenced. Further, these

programs have provided informa�on on the level ofpollutant and marine debris in the Antarc�c, which fortunately are at low levels. Ecosystem models are being
developed to understand the predator-prey interac�on.

REFERENCES
Branch, T.A. 2011. Humpback whale abundance south of 60°S
from three complete circumpolar sets of surveys. J. Cetacean
Res. Manage. (special issue) 3: 53–69.
Fujise, Y. and Pastene, L.A. 2018. Cetaceans as indicators ofhistorical and current changes in the Antarc�c ecosystem.
Technical Reports of the Institute of Cetacean Research(TEREP-ICR) No. 2: 1–13.
Government of Japan. 2005. Plan for the Second Phase of theJapanese Whale Research Program under Special Permit in
the Antarc�c (JARPAII)—Monitoring of the Antarc�c ecosystem and development of new management objec�ves
for whale Resources. Paper SC/57/O1 presented to the
IWC Scien�ﬁc Commi�ee, June 2005 (unpublished). 99 pp.
[Available from the IWC Secretariat].
Government of Japan. 2019. Outline of a research program
to inves�gate the abundance, abundance trends and stock
structure of large whales in the Indo-Paciﬁc region of the
Antarc�c, including a survey plan for the 2019/20 australsummer season. Paper SC/68A/ASI08 presented to the
IWC Scien�ﬁc Commi�ee, May 2019 (unpublished). 16 pp.
[Available from the IWC Secretariat].
Hakamada, T. and Matsuoka, K. 2014a. Es�mates of abundance
and abundance trend of the humpback whale in Areas
IIIE-VIW, south of 60°S, based on JARPA and JARPAII sigh�ng
data (1989/90–2008/09). Paper SC/F14/J04 presented to theExpert Workshop to Review the Japanese JARPAII Special
Permit Research Programme, February 2014 (unpublished).
36 pp. [Available from the IWC Secretariat].
Hakamada, H. and Matsuoka, K. 2014b. Es�mates of abundance
and abundance trend of the Antarc�c minke whale in Areas
IIIE-VIW, south of 60°S, based on JARPA and JARPAII sigh�ng
data (1989/90–2008/09). Paper SC/F14/J03 presented to theExpert Workshop to Review the Japanese JARPAII Special
Permit Research Programme, February 2014 (unpublished).
41 pp. [Available from the IWC Secretariat].
Hakamada, T. and Matsuoka, K. 2017. Sigh�ng survey procedures for abundance es�mates of large whales in JARPA and9Technical Reports of the Ins�tute of Cetacean Research (2020)
JARPAII, and results for Antarc�c minke whales. Technical
Reports of the Institute of Cetacean Research (TEREP-ICR) No.
1: 28–36.
Hakamada, T. and Tamura, T. 2018. An overview of the Ins�tuteof Cetacean Research work on ecosystem modeling in the
Antarc�c and western North Paciﬁc. Technical Reports of theInstitute of Cetacean Research (TEREP-ICR) No. 2: 14–19.
Interna�onal Whaling Commission. 2011. Report of theworkshop on the Comprehensive Assessment of Southern
Hemisphere Humpback Whales. J. Cetacean Res. Manage.
(special issue) 3: 1–50.
Interna�onal Whaling Commission. 2015. Report of the Expert
Workshop to Review the Japanese JARPAII Special Permit
Research Programme. J. Cetacean Res. Manage. (Suppl.) 16:
369–409.
Isoda, T., Tamura, T. and Pastene, L.A. 2018. Using JARPA and
JARPAII pla�orms for inves�ga�ng the occurrence of marine
debris in the Indo-Paciﬁc region of the Antarc�c. Technical
Reports of the Institute of Cetacean Research (TEREP-ICR) No.
2: 64–73.
Kanda, N., Goto, M. and Pastene, L.A. 2014. Stock structureof humpback whales in the Antarc�c feeding grounds asrevealed by microsatellite DNA data. Paper SC/F14/J31
presented to the Expert Workshop to Review the Japanese
JARPAII Special Permit Research Programme, February 2014
(unpublished). 5 pp. [Available from the IWC Secretariat].
Konishi, K., Tamura, T., Zenitani, R., Bando, T., Kato, H. andWalløe, L. 2008. Decline in energy storage in the Antarc�cminke whale (Balaenoptera bonaerensis) in the Southern
Ocean. Polar Biol. 31: 1509–1520.
Konishi, K., Hakamada, T., Kiwada, H., Kitakado, T. and Walløe,
L. 2014. Decline in stomach contents in the Antarc�c minke
whale (Balaenoptera bonaerensis) in the Southern Ocean.
Polar Biol. 37: 205–215.
Konishi, K. and Walløe, L. 2015. Substan�al decline in energy storage and stomach fullness in Antarc�c minke whales(Balaenoptera bonaerensis) during the 1990's. J. Cetacean
Res. Manage. 15: 77–92.
Lockyer, C., McConnell, L.C. and Waters, T.D. 1985. Body condi�on in terms of anatomical and biochemical assessment ofbody fat in North Atlan�c ﬁn and sei whales. Can. J. Zool. 63:
2328–2338.
Lockyer, C. 1986. Body fat condi�on in Northeast Atlan�c ﬁnwhales, Balaenoptera physalus, and its rela�onship withreproduc�on and food resource. Can. J. Fish. Aquat. Sci. 43:
142–147.
Matsuoka, K. and Hakamada, T. 2014. Es�mates of abundance
and abundance trend of the blue, ﬁn and southern rightwhales in the Antarc�c Areas IIIE-VIW, south of 60°S, basedon JARPA and JARPAII sigh�ng data (1989/90–2008/09).
Paper SC/F14/J05 presented to the Expert Workshop to
Review the Japanese JARPAII Special Permit Research
Programme, February 2014 (unpublished). 36 pp. [Available

10from the IWC Secretariat].
Matsuoka, K. and Hakamada, T. 2018. Geographical distribu�on of whales in the Indo-Paciﬁc region of the Antarc�cbased on JARPA and JARPAII sigh�ng data collected in the
period 1987/88–2008/09. Technical Reports of the Instituteof Cetacean Research (TEREP-ICR) No. 2: 20–31.
Mori, A. and Bu�erworth, D.S. 2006. A ﬁrst step towards modelling the krill-predator dynamics of the Antarc�c ecosystem.
CCAMLR Science 13: 217–277.
Moosa, N. 2017. An updated model of the krill-predator dynamics of the Antarc�c ecosystem. MSc thesis, University of Cape
Town. 246 pp.
Murase, H., Matsuoka, K., Ichii, T. and Nishiwaki, S. 2002.
Rela�onship between the distribu�on of euphausiids andbaleen whales in the Antarc�c (35°E–145°W). Polar Biol. 25:
135–145.
Murase, H., Matsuoka, K., Hakamada, T. and Kitakado, T. 2014.
Preliminary analysis of changes in spa�al distribu�on of
Antarc�c minke and humpback whales in Area IV during
the period of JARPA and JARPAII from 1989 to 2006. Paper
SC/F14/J18 presented to the Expert Workshop to Review
the Japanese JARPAII Special Permit Research Programme,
February 2014 (unpublished). 17 pp. [Available from the IWC
Secretariat].
Nishiwaki, S., Ishikawa, H., Goto, M., Matsuoka, K. and Tamura,T. 2014. Review of general methodology and survey procedures under the JARPAII. Paper SC/F14/J02 presented to theExpert Workshop to Review the Japanese JARPAII Special
Permit Research Programme, February 2014 (unpublished).
76 pp. [Available from the IWC Secretariat].
Pastene, L.A., Fujise, Y. and Hatanaka, H. 2014. The Japanese
Whale Research Program under Special Permit in the
Antarc�c—Phase II (JARPAII): origin, objec�ves, research
progress made in the period 2005/06–2010/2011 and relevance for management and conserva�on of whales andthe ecosystem. Paper SC/F14/J01 presented to the Expert
Workshop to Review the Japanese JARPAII Special Permit
Research Programme, February 2014 (unpublished). 76 pp.
[Available from the IWC Secretariat].
Tamura, T. and Konishi, K. 2009. Feeding habits and prey consump�on of Antarc�c minke whale (Balaenoptera bonaerensis) in the Southern Ocean. J. Northw. Atl. Fish. Sci. 42: 13–25.
Tamura, T. and Konishi, K. 2014. Prey composi�on and consump�on rate by Antarc�c minke whales based on JARPA and
JARPAII stomach contents data. Paper SC/F14/J15 presented
to the Expert Workshop to Review the Japanese JARPAII
Special Permit Research Programme, February 2014 (unpublished). 20 pp. [Available from the IWC Secretariat].
Tamura, T., Matsuoka, K. and Pastene, L.A. 2017. An overview
of the research programs on large whales conducted by
the Ins�tute of Cetacean Research. Technical Reports of theInstitute of Cetacean Research (TEREP-ICR) No. 1: 1–14.
Wada, A. 2019. Dedicated whale sigh�ng vessels as a pla�orm

What do we know about whales and ecosystem in the Indo-Pacific region of the Antarctic? Part 2
for krill and oceanographic research in the Indo-Paciﬁc regionof the Antarc�c. Technical Reports of the Institute of Cetacean
Research (TEREP-ICR) No. 3: 21–31.
Wada, A. and Tamura, T. 2014. Es�ma�on of krill biomass basedon JARPAII acous�c surveys. Paper SC/F14/J19 presented to
the Expert Workshop to Review the Japanese JARPAII Special
Permit Research Programme, February 2014 (unpublished).
17 pp. [Available from the IWC Secretariat].
Watanabe, T., Okazaki, M. and Matsuoka, K. 2014. Results ofoceanographic analyses conducted under JARPA and JARPAII
and possible evidence of environmental changes. Paper
SC/F14/J20 presented to the Expert Workshop to Review
the Japanese JARPAII Special Permit Research Programme,
February 2014 (unpublished). 10 pp. [Available from the IWC

Secretariat].
Yasunaga, G., Fujise, Y., Zenitani, R., Honda, K. and Kato, H.
2006. Yearly trend of trace element accumula�on in liverof Antarc�c minke whale, Balaenoptera bonaerensis. Paper
SC/D06/J28 presented to the Intersessional Workshop to
Review Data and Results from Special Permit Research on
Minke Whales in the Antarc�c, December 2006 (unpublished). 23 pp. [Available from the IWC Secretariat].
Yasunaga, G., Bando, T. and Fujise, Y. 2014. Spa�al and temporalvaria�ons in organochlorine contaminants in the Antarc�cminke whale, Balaenoptera bonaerensis. Paper SC/F14/J29
presented to the Expert Workshop to Review the Japanese
JARPAII Special Permit Research Programme, February 2014
(unpublished). 13 pp. [Available from the IWC Secretariat].

11

Technical Reports of the Ins�tute of Cetacean Research (2020)

pp. 12–22

Technical Report (not peer reviewed)

An outline of the Japanese Abundance and Stock structure Surveys in
the Antarc�c (JASS-A) including results of the ﬁrst survey under thisnew research program
Tatsuya Isoda*, Taiki Katsumata, 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: isoda@cetacean.jp

ABSTRACTThis paper brieﬂy outlines the objec�ves, survey and analy�cal procedures and work schedule of a new research program on whales and the ecosystem in the Indo-Paciﬁc region of the Antarc�c (JASS-A=JapaneseAbundance and Stock structure Surveys in the Antarc�c) and summarizes the ﬁrst JASS-A survey conductedin the 2019/20 austral summer season. The main research objec�ves of JASS-A are i) the study of the abundance and abundance trends of large whale species, and ii) the study of the distribu�on, movement and stock
structure of large whale species. JASS-A also has ﬁve secondary research objec�ves related to oceanography,
marine debris and whale biology. JASS-A will be based on systema�c sigh�ng surveys u�lizing the Line Transect
Method, to be conducted alterna�vely in IWC Management Areas III, IV, V and VI by one or two specialized
vessels, during a tenta�ve period of eight austral summer seasons. Analyses related to main and secondary
objec�ves will be conducted based on new as well previous data collected by JARPA/JARPAII and NEWREP-A in
the same research area. The ﬁrst JASS-A survey was conducted in the western part of Area III (000°–015°E) for
25 days, from 13 January to 6 February 2020. Systema�c sigh�ng and oceanographic data were obtained. Alsobiopsy, photo-iden�ﬁca�on and satellite tracking experiments were conducted successfully.

INTRODUCTION
Long-term research surveys in the Antarc�c are scarce.
Circumpolar whale sigh�ng surveys were conductedduring the Interna�onal Decade for Cetacean Research/
Southern Ocean Whale and Ecosystem Research (IDCR/
SOWER) under the auspices of the Interna�onal Whaling Commission Scien�ﬁc Commi�ee (IWC SC) between 1978/79 and 2009/2010 (Matsuoka et al., 2003;
IWC, 2013). These surveys produced important sigh�ng
data to study the abundance and abundance trends oflarge whales in diﬀerent IWC Management Areas (Branch
and Bu�erworth, 2001; IWC, 2013).
The Japanese Whale Research Programs under Special
Permit in the Antarc�c (JARPA/JARPAII) and the New Scien�ﬁc Whale Research Program in the Antarc�c Ocean(NEWREP-A) involved comprehensive long-term systema�c surveys in the Indo-Paciﬁc region of the Antarc�c thatcollected biological informa�on from whales and datafrom its environment, using lethal (biological samplingof Antarc�c minke whale) and non-lethal (whale sigh�ng
surveys, oceanographic surveys, photo-iden�ﬁca�on,
biopsy sampling and satellite tagging of large whales) ap12

proaches. A comprehensive biological and environmental
data set, including abundance trends for several whale
species, was obtained from these surveys conducted between 1987/88 and 2018/19 (IWC, 2015).
Japan considered it important to con�nue whale andecosystem surveys in the Indo-Paciﬁc region of the Antarc�c through dedicated sigh�ng surveys and other nonlethal techniques to inves�gate primarily abundance,
abundance trends and stock structure of large whales.
For such an aim a new research program was designed
which is called JASS-A (Japanese Abundance and Stockstructure Surveys in the Antarc�c). The JASS-A program
was presented to the 2019 mee�ng of IWC SC (GOJ,
2019a), to the 2019 mee�ng of Commission for the Conserva�on of Antarc�c Marine Living Resources-Working
Group on Ecosystem Monitoring and Management(CCAMLR-EMM) (GOJ, 2019b), and to the mee�ng of theNorth Atlan�c Marine Mammal Commission Scien�ﬁcCommi�ee (NAMMCO SC) (GOJ, 2019c). JASS-A was posi�vely evaluated at these interna�onal forums.
The objec�ves of this paper are, i) to present a brief
outline of the objec�ves, methodology and researchschedule of JASS-A, and ii) to summarize the results of

An outline of JASS-A including results of the first survey under this new research program

the ﬁrst JASS-A survey in the Indo-Paciﬁc region of the
Antarc�c in the 2019/20 austral summer season.

OUTLINE OF JASS-A
Objec�ves of the JASS-A
The main research objec�ves (MO) of the JASS-A are:MO1: Study of the abundance and abundance trendsof large whale species in the Indo-Paciﬁc region of the
Antarctic
Abundance and abundance trends of whales in the
Antarc�c is essen�al for conserva�on and management
purposes. Many whale species were depleted in thepast. Some of them have shown signs of recovery in recent years, and it is important to monitor their recovery
process and how such recovery could aﬀect other whale
species in the ecosystem.
MO2: Study of the distribution, movement and stock
structure of large whale species in the Indo-Paciﬁc regionof the Antarctic
Stock structure informa�on is important to interpret
distribu�on and abundance data. Gene�c stocks aredemographically independent units and therefore eachstock will respond in a diﬀerent way to changes that haveoccurred in the ecosystem. Ideally abundance es�matesshould be based on the geographical and temporalboundaries of gene�c stocks.
JASS-A, in conjunc�on with the work already conducted under the previous research programs, will provide
informa�on to allow the determina�on of the status of
the stocks of large whales that are found in waters of
the Indo-Paciﬁc region of the Antarc�c in summer, andprovide the necessary scien�ﬁc background for future
policies on conserva�on and sustainable u�liza�on.
The secondary objec�ves (SO) of JASS-A are:SO1: Investigation of the oceanographic conditions in the
Indo-Paciﬁc region of the Antarctic
Oceanographic structure and dynamics provide important informa�on in interpre�ng changes in the Antarc�cecosystem. Changes in oceanographic condi�ons willaﬀect krill distribu�on and biomass and, in turn, theabundance and distribu�on of whales. Changes in oceanographic condi�ons can be related to climate changes.

SO2: Investigation of the spatial and temporal trends ofmarine debris on sea surface
Studies on marine debris in the Antarc�c are very scarcewith only a few records made in sub-Antarc�c, Antarc�cislands and the Antarc�c. It is important to con�nue withthis kind of survey to monitor the future trends in the occurrence of marine debris.
SO3: To conduct feasibility studies to evaluate the utilityof genetic data to estimate abundance
Systema�c sigh�ng surveys u�lizing the Line Transect
Method is the most used method to es�mate abundanceof whales. Basic line transect surveys however, are notalways appropriate, especially for rare species/popula�ons. Also in the case of the Antarc�c it is not possible
to u�lize line transect methodology for areas inside thepack-ice, e.g., polynyas, where whales are also distributed. Gene�c-based methods can assist in the abundancees�mates in such cases.
SO4: To continue with feasibility studies to evaluate
the utility of non-lethal techniques for whale biologicalresearch
During the NEWREP-A several studies were carried out toinves�gate the feasibility of novel non-lethal approaches
to address some of the main objec�ves of the NEWREP-A
on Antarc�c minke whales, e.g., progesterone analysis
in blubber to inves�gate reproduc�ve status (Inoueet al., 2019), and stable isotopes to inves�gate whalesʼ
prey items. There is the need to con�nue with the inves�ga�on on the u�lity of such techniques for large whales.
SO5: Feasibility study on the utility of Unmanned Aerial
Vehicle (UAV) for obtaining information relevant for abundance estimate of large whales
In par�cular this technique could be used to determine
the number of individuals in the schools, which is informa�on highly relevant for the abundance es�mates.
Research area, season, vessels and period
The research area of JASS-A will be comprised by IWC
Management Areas III, IV, V and VI, south of 60°S (Figure 1). This will allow for con�nuity and consistency withdata already collected by JARPA/JARPAII, NEWREP-A and
IDCR/SOWER surveys in those Management Areas.
The period of the annual surveys will be the australsummer season (January–February), which is the sameas in the previous JARPA/JARPAII, NEWREP-A and IDCR/
SOWER surveys.
Surveys will be carried out in principle by one or two spe13

Technical Reports of the Ins�tute of Cetacean Research (2020)

Figure 1.

Research area of JASS-A. The shaded area (000°–
015°E) indicates the surveyed area in the 2019/20
austral summer season.

cialized vessels, and the tenta�ve length for JASS-A will beeight years (2019/20–2026/27), a period required to coverhalf of each Area once (Table 1). The ﬁnal determina�on of
the period and number of vessels will depend on funding
availability and therefore some level of ﬂexibility is required.
Survey and analy�cal procedures
Abundance and abundance trends (MO1)
The approach will be systema�c vessel-based sigh�ng
surveys u�lizing the Line Transect Method. Surveys will be
designed and conducted following the protocols includedin the ʻRequirements and Guidelines for Conduc�ng Surveys and analyzing data within the Revised Management
Schemeʼ (IWC, 2012). Sigh�ng protocols will be the sameas those used in the former IDCR/SOWER surveys (Matsuoka et al., 2003). Two sigh�ng modes will be used forg(0) es�mates, Passing with abeam Closing mode (NSP)and Passing with Independent Observer mode (IO). The
OK model (Okamura and Kitakado, 2012) will be used toes�mate abundance taking g(0) es�mates into account.
As stated above, the new data will be analyzed in conjunc�on with previous data in those Areas (Table 1). Loglinear models will be used to es�mate abundance trends(see details in Hakamada et al., 2013).
Distribution, movement and stock structure (MO2)Distribu�on and movement of individual humpback, blue,
southern right and ﬁn whales will be inves�gated primarily by photo-id and gene�c tagging. Photo-id pictures in
conjunc�on with pictures obtained in previous programs
will be used for the matching exercise to inves�gate dis14

tribu�on and movement of those large whales. Studies
on stock structure in humpback, blue, southern right andﬁn whales will be based on gene�c analyses of previous
biopsy samples as well as addi�onal biopsy samples collected during the new research program. Reﬁnement of
the stock structure of the Antarc�c minke whales (e.g.,
Pastene and Goto, 2016) will con�nue under the newresearch program based on the large gene�c sample set
collected under JARPA/JARPAII and NEWREP-A.
Gene�c markers such as mtDNA control region sequencing and microsatellite DNA will be used. Standard
gene�c analyses based on hypothesis tes�ng, heterozygosity, diversity, and related measures will be used. Principal Component Analysis (PCA), Discriminant Analysis
of Principal Component (DAPC) as well the Bayesian approach STRUCTURE will also be used. The ﬁnal analy�cal
procedure will be determined depending on the amountand nature of the data collected for each species.
Oceanographic surveys (SO1)
Oceanographic surveys will be conducted at one sta�on
per day, at least, using eXpendable Conduc�vity, Temperature and Depth (XCTD). Oceanographic sta�ons will be
deployed at equal intervals along the track-lines deﬁned
for the sigh�ng surveys.
Spatial and temporal trend of marine debris (SO2)
Observa�on of marine debris on the sea surface will bemade by visual observa�on along the track-lines of sighting surveys, the same as was done in JARPA/JARPAII and
NEWREP-A. Details of the methodology can be found in
Isoda et al. (2018).
Feasibility studies to evaluate the utility of genetic data toestimate abundance (SO3)
Approaches based on mark-recapture of gene�cally iden�ﬁed individuals will be used. Individual iden�ﬁca�on will
be based on the genotype proﬁle of a set of microsatellite loci. Studies have already started for southern rightwhales based on biopsy sampling (see details in Pastene
et al., 2018). Other poten�al target species are the blueand humpback whales as a considerable number of biopsy samples are already available.
Antarc�c minke whales will also be a target species forpaternity analyses based on the substan�al amount ofgene�c and other biological data from JARPA/JARPAII and
NEWREP-A.

An outline of JASS-A including results of the first survey under this new research program
Table 1
An overview of previous surveys with abundance es�mates (AE) and tenta�ve plans for future surveys in Areas III, IV, V and VI under
the JASS-A (GOJ, 2019a).
No.

No.

1
2

1
2

3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18

3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18

19
20
21
22
23
24
25
26
27

1
2
3
4
5
6
7
8
9

28

1

29
30
31
32

1
2
3
4

33
34
35
36
37
38
39
40

1
2
3
4
5
6
7
8

Program

SeasonResearch area

Remarks

IIIW

IIIE

IVW

IVE

VW

VE

VIW

VIE

1987/88
1988/89

—
—

—
—

AE
—

—
—

—
—

—
AE

—
—

—
—

—
—

1989/90
1990/91
1991/92
1992/93
1993/94
1994/95
1995/96
1996/97
1997/98
1998/99
1999/2000
2000/01
2001/02
2002/03
2003/04
2004/05

—
—
—
—
—
—
—
—
—
—
—
—
—
—
—
—

—
—
—
—
—
—
AE
—
AE
—
AE
—
AE
—
AE
—

AE
—
AE
—
AE
—
AE
—
AE
—
AE
—
AE
—
AE
—

AE
—
AE
—
AE
—
AE
—
AE
—
AE
—
AE
—
AE
—

—
AE
—
AE
—
AE
—
AE
—
AE
—
AE
—
AE
—
AE

—
AE
—
AE
—
AE
—
AE
—
AE
—
AE
—
AE
—
AE

—
—
—
—
—
—
—
AE
—
AE
—
AE
—
AE
—
AE

—
—
—
—
—
—
—
—
—
—
—
—
—
—
—
—

—
—
—
—
—
—
—
—
—
—
—
—
—
—
—
—

JARPAII

2005/06
2006/07
2007/08
2008/09
2009/102010/112011/122012/132013/14

—
—
—
—
—
—
—
—
—

AE
—
AE
—
—
—
—
—
—

AE
—
AE
—
—
—
—
—
—

AE
—
AE
—
—
—
—
—
—

—
AE
—
AE
—
—
—
—
—

—
AE
—
AE
—
—
—
—
—

—
AE
—
AE
—
—
—
—
—

—
—
—
—
—
—
—
—
—

—
—
—
—
Canceled
Canceled
Canceled
Canceled
Canceled

JASS

2014/15

—

—

AE

—

—

—

—

—

—

NEWREP-A

2015/162016/172017/18
2018/19

—
—
—
—

—
—
—
AE

—
—
—
—

AE
—
—
—

—
AE
—
—

—
—
AE
—

—
—
AE
—

—
—
—
—

—
—
—
—

JASS-A

2019/20
2020/212021/222022/232023/242024/252025/262026/27

Plan
—
—
—
—
—
—
—

—
—
—
—
—
—
—
Plan

—
—
Plan
—
—
—
—
—

—
—
—
Plan
—
—
—
—

—
—
—
—
Plan
—
—
—

—
—
—
—
—
Plan
—
—

—
—
—
—
—
—
Plan
—

—
Plan
—
—
—
—
—
—

JARPA

Feasibility studies on the utility of non-lethal techniques
to obtain biological and feeding ecology information fromlarge whales (SO4)

Progesterone analyses of blubber (SO4-A)
Inoue et al. (2019) studied the feasibility of analyses ofprogesterone in blubber as an indicator of the reproduc15

Technical Reports of the Ins�tute of Cetacean Research (2020)
Table 2
Tenta�ve schedule for the work on main and secondary objec�ves of JASS-A.
SeasonMO1 Abundance and trends
Blue whale
Fin whale
Antarc�c minke whale
Humpback whale
Southern right whale
Sperm whale
Southern bo�lenose whale
MO2 Stock structureBlue whale
Fin whale
Antarc�c minke whale
Humpback whale

2019/20

2020/21○○2021/22○○2022/23

○
○

○
○
○

2024/25○○2025/262026/27

○
○
○

○
○
○
○
○
○
○
○
○

○
○

Secondary Objec�ves
Oceanography
Marine debris
Abundance gene�cs
Feasibility non-lethal (progesterone)
Feasibility non-lethal (stable isotope)
UAV experiments

2023/24

○5

○1
○3
○7
○9

○2
○4
○6
○8
○ 10
○

1

Preliminary results oceanography; 2Complete the work, oceanography; 3Preliminary results marine debris; 4Complete the work,
marine debris; 5Complete the work, southern right whale; 6Complete the work, humpback and Antarc�c minke whale; 7Preliminary
results for humpback whale; 8Complete the work, humpback whale and Antarc�c minke whale; 9Preliminary results for Antarc�cminke, ﬁn and southern right whales; 10Complete the work, Antarc�c minke, ﬁn and southern right whales

�ve status of Antarc�c minke whales. The study was possible because informa�on on the reproduc�ve status ofeach sample used was available (from lethal sampling)
for comparison purposes. Eﬀorts will be made to collectbiopsy samples on an opportunis�c basis so that the progesterone study for this species can con�nue. Anothercandidate species for this study is the humpback whales
for which a substan�al number of biopsy samples are
already available.
Stable isotopes analyses (SO4-B)
Prey items of large baleen whales will be inves�gatedbased on previous samples collected lethally, and biopsy samples (old and new samples) and stable isotope
analyses (see details of the method in Mitani et al., 2006).
Priority species for this study will the Antarc�c minke, ﬁn
and southern right whales.
Feasibility study on the utility of Unmanned Aerial Vehicle(UAV) for obtaining information relevant for abundance
estimate of large whales (SO5)
Several types of UAV will be tested to inves�gate the u�l16

ity of this technique for obtaining informa�on relevant
for abundance es�mates of large whales including thenumber of whales in a school.
As noted above, MO and SO data will be analyzed inconjunc�on with the large data sets produced by JARPA/
JARPAII and NEWREP-A in the Indo-Paciﬁc region of the
Antarc�c.
Research schedule
Table 2 shows the research schedule of JASS-A with proposed dates for progressing and comple�ng the workof MO and SO. Once again some degree of ﬂexibility isrequired on the schedule proposed in this table.
Organiza�on of JASS-A and opportuni�es for researchcollabora�on
Scien�sts from the Ins�tute of Cetacean Research (ICR)
will play the leading role in order to pursue the researchac�vi�es and achieve the research objec�ves of JASS-A,
in collabora�on with scien�sts from other domes�c research organiza�ons such as the Na�onal Research Ins�-An outline of JASS-A including results of the first survey under this new research programtute of Far Seas Fisheries, and the Tokyo University of Marine Science and Technology. A domes�c steering grouphas been formed to coordinate the research ac�vi�es.
Qualiﬁed external scien�sts will be welcome to par�cipate in the ﬁeld and analy�cal works of JASS-A. Qualiﬁed
external scien�sts can submit ﬁeld or analy�cal research
proposals for considera�on of the domes�c steeringgroup. To facilitate the process, the steering group will
prepare guidelines for the submission process.

SUMMARY OF THE FIRST JASS-A SURVEY
Research area
The research area covered by the survey was the westernpart of Area IIIW (000°–015°E), south of 60°S (Figure 1).

Figure 2.

Research area (000°–015°E) and track-line of the
JASS-A survey in 2019/20. The survey commenced
at 68°40ʹS 000°00ʹ and ended at 63°06ʹS 15°00ʹE.

Figure 3.

The area was divided into northern and southern strata.
The boundary between southern and northern strata wasdeﬁned by a line 45 n.miles from the ice edge (Figure 2).
In the northern and southern strata, the survey tracklines consisted of a zigzag course changing direc�on at
5°00ʹ and 2°30ʹ longitudinal degree intervals in a 10 degrees longitudinal band respec�vely. A randomized startpoint for survey tracks was used.
Research vessel and sigh�ng mode
The dedicated sigh�ng vessel (SV) Yushin-Maru No. 2(747GT, 69.6 m) was engaged in the survey. Three experienced Japanese researchers on line transect surveys,
biopsy sampling, photo-id, satellite tagging experiments,and oceanographic survey, were aboard. The vessel was
equipped with a top barrel pla�orm (TOP, 19.5 m), Independent Observer Pla�orm (IOP, 13.5 m) and an upper
bridge pla�orm (UBP, 11.5 m). For NSP mode, there were
two primary observers on the TOP and there was open
communica�on between the UBP and the TOP. For IO
mode, there were two primary observers on the TOP and
one primary observer on the IOP. The observers on the
TOP or IOP communicated to the UBP independently. Two
primary observers (captain and helmsman) and researchers were at the upper bridge, regardless of the researchmode (Figure 3). These observers conducted searchingfor cetaceans by using angle board and binoculars withre�cles (7x), which include the distance es�mate scales.
Research eﬀort in the research area
The dedicated sigh�ng vessel was engaged in the survey
for 25 days. The survey commenced on 13 January at
posi�on 68°40ʹS 000°00ʹ and ended on 6 February at posi�on 63°06ʹS 15°00ʹE covering the predetermined transects. The total searching distance was 1,447.9 n. miles(2,681.5 km), including 650.3 n. miles covered in NSP
mode and 797.6 n. miles in IO mode. The survey coverageSigh�ng ac�vity at the TOP (le�); conﬁrma�on and tracking of whales including iden�ﬁca�on of duplicatesat the UBP (middle); Researcher recording a sigh�ng and inpu�ng data at the UBP (right).
17Technical Reports of the Ins�tute of Cetacean Research (2020)
Table 3
Number of sigh�ngs in the research area, by stratum and species.
Western part of Area IIIW: 000°–015°E
SpeciesBlue whale
Fin whale
Like ﬁnAntarc�c minke whale
Like minke
Humpback whale
Like humpback
Baleen whale
Sperm whale
Killer whale
Southern bo�lenose whale
Ziphiidae
Uniden�ﬁed whale

Figure 4.

Southern stratum

Northern stratum

Prim.

Prim.

Second.

Second.

Prim.

Total

Second.

Sch.

Ind.

Sch.

Ind.

Sch.

Ind.

Sch.

Ind.

Sch.

Ind.

Sch.

Ind.

Sch.

Ind.

11
7
0
51
5
13
0
2
0
1
4
1
1

11
16
0
106
6
22
0
3
0
1
10
2
1

2
1
1
14
1
12
0
1
0
0
0
0
0

2
6
3
15
1
16
0
1
0
0
0
0
0

6
63
2
48
4
62
2
5
5
0
3
13
3

7
113
3
73
4
123
2
6
5
0
12
22
3

0
1
0
6
0
3
0
1
0
0
0
0
0

0
1
0
9
0
7
0
1
0
0
0
0
0

17
70
2
99
9
75
2
7
5
1
7
14
4

18
129
3
179
10
145
2
9
5
1
22
24
4

2
2
1
20
1
15
0
2
0
0
0
0
0

2
7
3
24
1
23
0
2
0
0
0
0
0

19
72
3
119
10
90
2
9
5
1
7
14
4

20
136
6
203
11
168
2
11
5
1
22
24
4

Posi�on of primary sigh�ngs of Antarc�c minke, humpback, ﬁn, and blue whales with informa�on of the surveyed transects.

was 71% in the northern stratum and 83% in the southern
stratum. The total experimental �me for photo-id, biopsy
sampling, satellite tagging and distance and angle experiment was 26 hours and 49 minutes.
Whale sigh�ng in the research area
Four baleen whale species, blue (19 schools/20 individuals), ﬁn (72/136), Antarc�c minke (119/203), humpback(90/168) whales and at least three toothed whale species, sperm (5/5), southern bo�lenose (7/22) and killer(1/1) whales, were sighted (Table 3).
Antarctic minke whales
This species was the most frequently sighted. Density
18

Sub-total

index (DI: schools sighted/100 n.miles searching distance)
based on primary sigh�ngs was 0.04 in the northern stratum and 0.14 in the southern stratum. In this survey, thesigh�ngs of Antarc�c minke whales were abundant near
the ice edge. In addi�on, many sigh�ngs occurred in thewestern side of the northern stratum (Figure 4).
Humpback whales
This species was the second most frequently sighted. It
was widely distributed in the research area (Figure 4). The
DI based on primary sigh�ngs was 0.06 in the northern
stratum and 0.03 in the southern stratum. The distribu�on was concentrated between 61°S and 62°S. The
pa�ern of distribu�on and density of this species in thisAn outline of JASS-A including results of the first survey under this new research program

survey seems to be diﬀerent from those in past surveys
in the Area III.
Fin whales
This species was the third most frequently sighted. It
was found mainly north of 64°S (Figure 4). The numberof sigh�ngs were larger than in previous surveys in thesame area.
Blue whales
This species was found mainly in the southern stratumof the research area (Figure 4). In this survey, 13 schools(13 individuals) were distributed in the southern stratum,
similar to the situa�on in previous surveys. A total of 6
schools (7 individuals) was distributed in the northern
stratum.
Duplicate sigh�ngs
Duplicates were recorded for a total of 75 sigh�ngs (involving several whale species) during the IO mode survey,
and these data will be used to es�mate whale abundance
while taking es�mated g(0) into considera�on.

Figure 5.

Photo-id experiments on humpback whales from the bow deck of the vessel (le�); details of the ventral
ﬂuke pigmenta�on (right).

Figure 6.

Biopsy sampling of blue whale (le�) and skin/blubber sample of blue whale obtained by biopsy sampling(right).

Figure 7.

Satellite tagging and biopsy sampling of ﬁn (le�) and Antarc�c minke (right) whales.
19Technical Reports of the Ins�tute of Cetacean Research (2020)

Sigh�ng survey in the transit area
The sigh�ng survey was conducted between 10°S and theresearch area, excluding the areas of foreign countriesʼ
EEZs. Only NSP mode was used. The searching eﬀort was
2,539.9 n.miles and total sigh�ngs included blue (3/4), ﬁn(9/14), Antarc�c minke (5/8), humpback (12/23), sperm(9/10), southern bo�lenose (2/8) and killer whales TypeA (1/17) whales.
Other research ac�vi�es
Sighting distance and angle experiment
The sigh�ng distance and angle experiment was conducted in order to evaluate the accuracy of sigh�ng
distance and angle provided by primary observers. The
results of this experiment will be used for the calcula�onof abundance es�mates. A training for this experiment
was carried out on 15 January. The actual experiment was
conducted on 30 January and 144 trials were completed
successfully.

These data will be registered into the ICR photo-id catalogue (e.g., Matsuoka and Pastene, 2014).
Biopsy sampling for large whales (Figure 6)
A total of 29 biopsy samples were collected, including
10 blue, 11 ﬁn, and 8 Antarc�c minke whales, using theLarsen system (Larsen, 1998). Satellite tags were a�achedon eight biopsied Antarc�c minke whales and seven of
the 11 biopsied ﬁn whales. Biopsy samples were stored at−20°C. These samples will be used in gene�c analyses andstudies to evaluate the u�lity of non-lethal techniques forwhale biological research.

Photo-id (Figure 5)
A total of 20 blue, 14 humpback and 5 killer whales (TypeA) were successfully photo-iden�ﬁed during the survey.

Figure 9.
Figure 8.

Oceanographic survey by XCTD.

Figure 10.
20

Oceanographic observa�on sta�ons (XCTD cas�ng
points).

Image of two blue whales taken using a small UAV in the research area.

An outline of JASS-A including results of the first survey under this new research program

Satellite tagging (Figure 7)
The satellite-monitored tags (SPOT6, Wildlife Computers,
Redmond, Washington, USA) were deployed with the Air
Rocket Transmi�er System (ARTS) (LK-ARTS, Skutvik, Norway). The details of the deployment system and protocols
is described in Konishi et al. (2020). The tags were successfully deployed on 10 ﬁn and 8 Antarc�c minke whales(see Konishi et al., this issue).
Oceanographic survey
Oceanographic survey was conducted by XCTD (Figure 8)at 75 sta�ons on the survey track-lines, including transit
survey (Figure 9). Oceanographic data will be analyzed to
study the oceanographic structure of the research area
and the rela�onship with whale distribu�on (Watanabe
et al., 2014). In addi�on, an Argo ﬂoat, under the Argo
oceanographic program (JAMSTEC, 2020), was successfully deployed on 4 January, during the transit to the
Antarc�c.
Marine debris observation
A total of three marine debris objects, comprising twoplas�c bo�les and a ﬁshing buoy, were observed during
the survey in the research area. These data will be registered into the ICR database and reported in the future(e.g., Isoda et al., 2018).
Feasibility study on the utility of UAV
A preliminary experiment was conducted for collec�ng
aerial images of whale using a small UAV (Phantom 4
Pro; DJI, Shenzhen, China). A total of two blue (Figure 10)
and three humpback whales were photographed using
the UAV. These data will be registered into the photo-id
catalogue of ICR.
Data and samples collected in this survey were validated and stored at the ICR, and will be analyzed in conjunc�on with data and samples obtained during past research
programs in the Antarc�c (JARPA, JARPAII, NEWREP-A),
in the context of the primary and secondary objec�vesof the JASS-A. Data will be available to the na�onal andinterna�onal scien�ﬁc community through establisheddata access protocols of ICR.

ACKNOWLEDGEMENTS
We thank the captain, crew members and researchers
who par�cipated in the 2019/20 JASS-A survey for theireﬀort in the collec�on of data and samples. Thanks also
to Yoshihiro Fujise (ICR), project leader of the cruise,for valuable sugges�ons and the staﬀ of ICR for logis�c

support. Kyodo Senpaku Co. Ltd. is acknowledged forlogis�c arrangements and support. Thanks also to Kenji
Konishi (ICR) for planning and assis�ng the satellite tagging experiments. Finally, we thank the Fisheries Agencyof Japan for the funding support for the JASS-A program.

REFERENCES
Branch, T.A. and Bu�erworth, D.S. 2001. Es�mates of abundance south of 60°S for cetacean species sighted frequently
on the 1978/79 to 1997/98 IWC/IDCR-SOWER sigh�ng surveys. J. Cetacean Res. Manage. 3 (3): 251–270.
Government of Japan. 2019a. Outline of a research program
to inves�gate the abundance, abundance trends and stock
structure of large whales in the Indo-Paciﬁc region of the
Antarc�c, including a survey plan for the 2019/20 australsummer season. Paper SC/68a/ASI8 presented to the IWC
Scien�ﬁc Commi�ee, May 2019 (unpublished). 16 pp.
[Available from the IWC Secretariat].
Government of Japan. 2019b. Outline of a research program
to inves�gate the abundance, abundance trends and stock
structure of large whales in the Indo-Paciﬁc region of the
Antarc�c, including a survey plan for the 2019/20 australsummer season. Paper WG-EMM-2019/68 presented to
CCAMLR mee�ng of Working Group on Ecosystem Monitoringand Management, June–July 2019 (unpublished). 16 pp.
Government of Japan. 2019c. Outline of a research program
to inves�gate the abundance, abundance trends and stock
structure of large whales in the Indo-Paciﬁc region of the
Antarc�c, including a survey plan for the 2019/20 australsummer season (revised version of document SC/68a/ASI08
presented to the IWC SC 2019 mee�ng). Paper SC/26/NPR-JP
presented to the 26th mee�ng of the NAMMCO Scien�ﬁcCommi�ee, October-November 2019 (unpublished). 16 pp.
Hakamada, T., Matsuoka, K., Nishiwaki, S. and Kitakado, T. 2013.
Abundance es�mates and trends for Antarc�c minke whales(Balaenoptera bonaerensis) in Antarc�c Areas IV and V basedon JARPA sigh�ng data. J. Cetacean Res. Manage. 13 (2):123–151.
Interna�onal Whaling Commission. 2012. Requirements andguidelines for conduc�ng surveys and analyzing data within
the Revised Management Scheme. J. Cetacean Res. Manage.
(Suppl.) 13: 507–517.
Interna�onal Whaling Commission. 2013. Report of the
Scien�ﬁc Commi�ee. J. Cetacean Res. Manage. (Suppl.) 14:
1–102.
Interna�onal Whaling Commission. 2015. Report of the Expert
Workshop to Review the Japanese JARPAII Special Permit
Research Programme. J. Cetacean Res. Manage. (Suppl.) 16:
369–409.
Inoue, S., Yasunaga, G. and Pastene, L.A. 2019. Comparison ofprogesterone concentra�ons in blubber and plasma amongfemale Antarc�c minke whales of known reproduc�ve status.
Fish Sci 85: 971–977.
21

Technical Reports of the Ins�tute of Cetacean Research (2020)
Isoda, T., Tamura, T. and Pastene, L.A. 2018. Using JARPA and
JARPAII pla�orms for inves�ga�ng the occurrence of marine
debris in the Indo-Paciﬁc region of the Antarc�c. Technical
Reports of the Institute of Cetacean Research (TEREP-ICR) No.
2: 64–73.
Japan Agency for Marine-Earth Science and Technology. 2020.
Japan Argo-part of the integrated global observa�on strategy.
h�p://www.jamstec.go.jp/J-ARGO/index_j.html. Accessed 1
May 2020.
Konishi, K., Isoda, T., Bando, T., Minamikawa, S. and Kleivane, L.
2020. Antarc�c minke whales ﬁnd ice gaps along the ice edgein foraging grounds of the Indo-Paciﬁc sector (60°E and 140°E)of the Southern Ocean. Polar Biol. 43 (4): 343–357.
Larsen, F. 1998. Development of a biopsy system primarily foruse on large cetaceans. Paper SC/50/O15 presented to the
IWC Scien�ﬁc Commi�ee, May 1998 (unpublished). 8 pp.
[Available from the IWC Secretariat].
Matsuoka, K., Ensor, P., Hakamada, T., Shimada, H., Nishiwaki,S., Kasamatsu, F. and Kato, H. 2003. Overview of minke
whale sigh�ngs surveys conducted on IWC/IDCR and SOWER
Antarc�c cruises from 1978/79 to 2000/01. J. Cetacean Res.
Manage. 5 (2): 173–201.
Matsuoka, K. and Pastene, L.A. 2014. Summary of photoiden�ﬁca�on informa�on of blue, southern right and humpback whales collected by JARPA/JARPAII. Paper SC/F14/J34
presented to the IWC Scien�ﬁc Commi�ee, February 2014

22

(unpublished). 11 pp. [Available from the IWC Secretariat].
Mitani, Y., Bando, T., Takai, N. and Sakamoto, W. 2006. Pa�erns
of stable carbon and nitrogen isotopes in the baleen of common minke whale Balaenoptera acutorostrata from the western North Paciﬁc. Fish Sci. 72: 69–76.
Okamura, H. and Kitakado, T. 2012. Abundance es�matesof Antarc�c minke whales using the OK method. Paper
SC/64/IA2 presented to IWC Scien�ﬁc Commi�ee, June 2012
(unpublished). 24 pp. [Available from the IWC Secretariat].
Pastene, L.A. and Goto, M. 2016. Gene�c characteriza�on andpopula�on gene�c structure of the Antarc�c minke whale
Balaenoptera bonaerensis in the Indo-Paciﬁc region of theSouthern Ocean. Fish Sci. 82: 873–886.
Pastene, L.A., Goto, M., Acuna, P., Taguchi, M., Hakamada, T.
and Matsuoka, K. 2018. Popula�on iden�ty, site-ﬁdelity,
movement ranges and preliminary es�mates of abundance ofsouthern right whales in the Antarc�c Indian sector inferredfrom gene�c markers. Document WG-EMM-18/18 presented
to the CCAMLR Working Group on Ecosystem Monitoring andManagement, July 2018. 13 pp.
Watanabe, T., Okazaki, M. and Matsuoka, K. 2014. Results ofoceanographic analyses conducted under JARPA and JARPAII
and possible evidence of environmental changes. Paper
SC/F14/J20 presented to the JARPAII review Workshop,
February 2014 (unpublished). 10 pp. [Available from the IWC
Secretariat].

Technical Reports of the Ins�tute of Cetacean Research (2020)

pp. 23–34

Technical Report (not peer reviewed)

An outline of the IWC-Paciﬁc Ocean Whale and Ecosystem Research(IWC-POWER) including results of the 2019 survey
Koji Matsuoka*
Institute of Cetacean Research, 4–5 Toyomi-cho, Chuo-ku, Tokyo 104–0055, Japan*Contact e-mail: matsuoka@cetacean.jp

ABSTRACTThis paper outlines the objec�ves, research area and general methodology of the Interna�onal Whaling Commission-Paciﬁc Ocean Whale and Ecosystem Research Program (IWC-POWER). It also provides the main resultsof the 2019 survey conducted under this research program. The IWC-POWER is designed and implemented by
the IWC Scien�ﬁc Commi�ee (SC) in special partnership with the Government of Japan. The long-term objec�ve of the IWC-POWER is to ʻprovide informa�on to allow determina�on of the status of popula�ons (and thus
stock structure is inherently important) of large whales that are found in the North Paciﬁc waters and provide
the necessary scien�ﬁc background for appropriate conserva�on and management ac�onsʼ. To fulﬁll this objec�ve, the IWC-POWER originally iden�ﬁed short and medium-term ac�vi�es and priori�es. The IWC-POWER
is close to successfully comple�ng its ﬁrst-phase (2010–2021) related to the short-term priori�es, and the IWC
SC will start organizing the second phase (from 2022 onward). It has also updated the medium-term priori�es
for this second phase. The 2019 survey was conducted successfully between 3 July and 25 September 2019
in the Gulf of Alaska, within the U.S. Exclusive Economic Zone by the Japanese R/V Yushin Maru No. 2. Blue(17 schools/19 individuals), ﬁn (266/458), sei (9/20), common minke (5/5), humpback (173/402), gray (6/15),
sperm (20/20), Bairdʼs beaked (2/37), Cuvierʼs beaked (3/5) and killer (53/264) whales were observed during
the survey. Photo-iden�ﬁca�on data were collected from 16 blue, 51 ﬁn, 30 humpback, 6 gray and 19 killer
whales. A total of 75 biopsy samples were collected from 12 blue, 45 ﬁn, 4 sei, 12 humpback and 2 gray whales.
A total of 229 sonobuoys were deployed, for a total of over 820 monitoring hours. The main species detectedby the acous�c survey included ﬁn, sperm, killer, blue, humpback, North Paciﬁc right and sei whales. A total of
42 objects of marine debris were also observed.

INTRODUCTION
The Interna�onal Whaling Commission-Paciﬁc Ocean
Whale and Ecosystem Research (IWC-POWER) programis an interna�onal research eﬀort in the North Paciﬁc coordinated by the IWC and designed by the IWC Scien�ﬁcCommi�ee (SC) in special partnership with the Government of Japan. Scien�sts from the Ins�tute of Cetacean
Research (ICR) par�cipate regularly in the IWC-POWER
program, both in designing and implemen�ng the surveys. The IWC-POWER in the North Paciﬁc follows the series of IWC Interna�onal Decade for Cetacean Research/
Southern Ocean Whale and Ecosystem Research (IDCR/
SOWER) surveys that had been conducted in the Antarc�c since 1978.
The long-term objec�ve of the IWC-POWER is to ʻprovide informa�on to allow determina�on of the status of
popula�ons (and thus stock structure is inherently impor-tant) of large whales that are found in the North Paciﬁcwaters and provide the necessary scien�ﬁc backgroundfor appropriate conserva�on and management ac�onsʼ.
The ﬁrst survey of this program was conducted in 2010and subsequent surveys have taken place on an annual
basis.
The IWC SC is close to comple�ng the ﬁrst phase of the
IWC-POWER, which is related to the short-term priori�es, and now is preparing for the second phase related
to medium term priori�es, based on the results of theﬁrst phase.
The objec�ves of this document are: i) to present anoutline of the IWC-POWER program including its objec�ves, research area, and general methodology; and ii) topresent the results of the 2019 IWC-POWER survey basedon Matsuoka et al. (2020).

23

Technical Reports of the Ins�tute of Cetacean Research (2020)

OUTLINE OF THE IWC-POWER
Objec�ves
The IWC-POWER has the following long-term objec�ve:
ʻProvide informa�on to allow determina�on of the status of popula�ons (and thus stock structure is inherently
important) of large whales that are found in North Paciﬁcwaters and provide the necessary scien�ﬁc backgroundfor appropriate conserva�on and management ac�ons.
The programme will primarily contribute informa�on onabundance and trends in abundance of popula�ons oflarge whales and try to iden�fy the causes of any trends
should these occur. The programme will learn from boththe successes and weaknesses of past na�onal and interna�onal programmes and cruises, including the IDCR/
SOWER programmeʼ. (IWC, 2011).
To a�ain this long-term objec�ve, the IWC-POWER hasu�lized short and medium-term ac�vi�es and priori�es.
The short-term priori�es involved the deﬁni�on and implementa�on of surveys in areas of the Central and Eastern North Paciﬁc (east of 170°E) (see sec�on on researcharea below). This region had not been surveyed since the�me of commercial whaling in the 1970ʼs (see Annex D
of IWC, 2012). A�er reviewing current knowledge of theregion, a list of medium-term priori�es by species forthe program was agreed to by the IWC SC (see Table 1 of
IWC, 2013). This list was updated recently in view of the
prepara�on of the second phase of IWC-POWER (see sec�on ʻWHAT NEXT?ʼ).
Research area for the short-term priori�es
The research area was comprised between north of 20°N
and south of the Bering Sea, and between 170°E and135°W, including the Exclusive Economic Zone (EEZ) of
the United States (U.S. EEZ) and Canada (Canadian EEZ).

Figure 1.

24

The research areas covered by IWC-POWER in
2010–2019 (EEZs are shown by blue do�ed line).
The yellow indicates the most recently surveyed
area.

Figure 1 shows a map of the survey areas covered since
2010.
Survey season and research vessels
To collect consistent seasonal sigh�ng data, the annual
surveys were conducted in the middle of summer, during July to September (e.g., Matsuoka et al., 2011). The
following research vessels were engaged in the annual
survey:Kaiko Maru (860.25 GT) in 2010;
Yushin Maru No. 3 (742 GT) from 2011–2016; andYushin Maru No. 2 (747 GT) from 2017–2019.
These vessels were prepared with specialized equipment and personnel for visual survey of cetaceans.
Sigh�ng survey procedure
Survey time
Research eﬀort began 60 minutes a�er sunrise and ended
60 minutes before sunset, with a maximum of 12-hour research day (maximum 06:00–19:00, including 30 minutes
for meal �me for lunch and supper, when surveying in IO
mode; see below).
Number of primary observers
Sigh�ng eﬀort was conducted by the boatswain and topmen from the TOP barrel and by two primary observers(the helmsman and captain) and at least four secondaryobservers (oﬃcer-on-watch, three researchers, and thechief engineer) from the upper bridge. Sigh�ng ac�vi�esaboard the ship were classiﬁed into two principal types:
On-eﬀort and Oﬀ-eﬀort. On-eﬀort ac�vi�es were �mes
when full search eﬀort was executed and condi�ons (suchas weather and sea state) were within acceptable parameters to conduct research. Oﬀ-eﬀort ac�vi�es were all ac�vi�es that were not On-eﬀort when no primary observers were in the TOP barrel (e.g., during dri�ing, Top Down
(TD) or transi�ng along the trackline due to bad weather
condi�ons). All sigh�ngs recorded during On-eﬀort wereclassiﬁed as Primary sigh�ngs. All other sigh�ngs wereconsidered as Secondary sigh�ngs.
Survey modes
Passing with abeam closing mode (NSP): Two topmen
were on eﬀort from the TOP barrel at all �mes. There wasopen communica�on between the upper bridge and thebarrel. The observers on the upper bridge communicatedwith the topmen only to clarify sigh�ng informa�on. The
upper bridge observers did not distract the topmen fromtheir normal search procedure unless they were directed
to do so by the cruise leader (IWC, 2020a).

An outline of the IWC-Pacific Ocean Whale and Ecosystem Research (IWC-POWER) including results of the 2019 survey

Independent Observer Mode (IO): This is also in effect Passing Mode. Two topmen were observing fromthe TOP barrel and two from the IO barrel at all �mes.
Communica�ons were essen�ally one-way, with topmen
from the TOP and IO pla�orms repor�ng informa�on to
the upper bridge in isola�on from each other to ensurethat no sigh�ng informa�on was exchanged between
the TOP and IO barrel observers. The observers on theupper bridge would communicate with the topmen only
to clarify sigh�ng informa�on and would not direct thetopmen to disrupt their normal search procedure unlessdirected to do so by the cruise leader. Immediately a�er
a sigh�ng was detected from the barrel, the topman
relayed informa�on to observers on the upper bridge.
A�er the sigh�ng informa�on was relayed to the upper
bridge observers, the topman responsible for the sigh�ng
con�nued his normal searching pa�ern. Observers on theupper bridge located the sigh�ng made by the topman
and decided whether it would be possible to conﬁrm species and conduct a school size count before the sighted
whale(s) passed abeam of the vessel. The topmen gave
no further informa�on to the upper bridge unless thewhale school resurfaced within their normal searchingarea. A designated researcher on the upper bridge recorded the species and es�mated number of whales inthe school when the sigh�ng passed abeam of the vessel.
This was done in consulta�on with other upper bridge
observers/researchers (Anon, 2019a).

of the shape of the blow, surfacing and other behaviouralpa�erns were suﬃcient to iden�fy whales. This judgement was made only by the cruise leader or other designated researcher. Iden�ﬁca�on of species was recordedas ʻprobableʼ where mul�ple cues were insuﬃcient to beabsolutely conﬁdent of iden�ﬁca�on (recorded as ʻlikeʼ).
This usually occurred when blows and surfacing pa�erns
could be conﬁrmed, but the whaleʼs body could not beclearly seen. Details of recording procedures during sightings can be found in Anon (2019a).

Approach to the whale schools
During both NSP or IO modes, and at the abeam �me,the vessel altered the course to approach the whale(s),and the speed was increased to 15 knots to hasten theclosure. Vessel speed was decreased when the school
was close, usually within 0.2 to 0.4 n.miles from the ini�alsigh�ng posi�on. A�er the school was approached, thespecies, number of animals in the school, es�mated bodylength(s), number of calves present, and behaviour were
determined and recorded. Following this, other ac�vi�eswould normally be conducted (�me allowing and at thediscre�on of the cruise leader), such as photography fornatural marking (Photo-ID) studies and biopsy sampling.
All sigh�ngs detected during these ac�vi�es were classiﬁed as secondary sigh�ngs (Anon, 2019a).

Es�mated Distance and angle experiments
This experiment has been conducted every year since2010. The experiment is designed to calibrate and iden�fy any biases in individual observersʼ es�ma�on of angle
and distance. The experiment should be conducted during weather and sea condi�ons representa�ve of the condi�ons encountered during the survey. In 2015, a major
improvement was made increasing the number of target
buoys to improve accuracy of the observerʼs es�mates(IWC, 2020b).

Species identiﬁcation
Species iden�ﬁca�on was based on agreed guidelines(IWC, 2020a). Posi�ve iden�ﬁca�on of species was basedon mul�ple cues and usually required clear observa�onof the whaleʼs body. Occasionally, repeated observa�ons

School size determination
Schools where the number of animals, or an accuratees�mated range of the number of animals was determined, were classiﬁed as conﬁrmed schools. Data fromthe conﬁrmed schools can be used to determine a meanschool size. Normally, schools believed to be conﬁrmed
for school size were approached to within 1 n.mile forlarge whales and to within 0.3 n.miles for common minke
whales. Allowing for context-speciﬁc diﬀerences (i.e.,environmental condi�ons and animal behaviour), every
eﬀort was made to be consistent with regard to the
maximum �me spent on iden�ﬁca�on of species andconﬁrma�on of numbers. Normally, if the sigh�ng wasthought to be common minke whales, no more than 20
minutes (a�er closure has been completed) was spenton conﬁrma�on. This reduces the poten�al for confusion
with other whale sigh�ngs in the vicinity (IWC, 2020a).

Photo-ID data collec�onThis experiment has been conducted since 2010. As
appropriate and decided by the cruise leader, research�me is allocated for photo-iden�ﬁca�on and/or video recording of large whales. The priority species were North
Paciﬁc right (highest priority), gray, blue and sei whales.
Medium-priority species included ﬁn, Brydeʼs, common
minke, sperm and killer whales. Generally, large whales
were approached within 15–20 m. Adults, juveniles, andfemales accompanied by calves were approached for25Technical Reports of the Ins�tute of Cetacean Research (2020)

photo-iden�ﬁca�on. Photo-ID experiments involved aminimum of one photographer and a maximum of three
on the bow, with addi�onal photographers in the TOP
barrel and IO barrel or upper bridge (Anon, 2019a).
Biopsy samplingThis experiment has been conducted since 2010 to analyze stock structure and movement for each species. The
priority species for biopsy sampling were the same as forphoto-iden�ﬁca�on (see above). The Larsen samplingsystem was used to collect samples within approximately
5 to 30 m of the bow of the vessel (Larsen, 1998).
Acous�c monitoring
Passive acous�cs monitoring using sonobuoys wereintroduced to detect North Paciﬁc right whalesʼ calls in
the Bering Sea and a part of the Gulf of Alaska, and was
conducted successfully from 2017 to 2019. A sonobuoy isa free-ﬂoa�ng, expendable, short-term passive acous�clistening device that transmits signals in real �me via VHF
radio waves to a receiver on a vessel. Sonobuoys were
deployed approximately every 2–2.5 hours, or approximately every 20–25 n.miles, from 06:00 to 18:00 (ship�me) to obtain an evenly-sampled cross-survey census
of marine mammal vocaliza�ons. Sonobuoys were monitored in real-�me by the acous�cian, and presence ofspecies-speciﬁc call types was noted (Crance et al., 2017).

Marine debris observa�onThis observa�on has been conducted since 2010. Dataon ﬂoa�ng marine debris were observed and recorded.
Observa�on of marine debris was limited to the ﬁrst 15
minutes of each hour, as �me permi�ed (so as not tointerfere with marine mammal observa�ons). In addi�on,marine debris were photographed if items were par�cularly large and/or could poten�ally lead to large whale
entanglements. For all recorded marine debris items, observers recorded angle, distance and �me of ini�al sighting, IWC code and a descrip�on. Photographs of items
were archived and are available to interested scien�sts.
Data recording and format
Research data collected during the survey (weather, effort, sigh�ng and distance experiment data) were enteredby researchers using the ʻonboard data collec�ng systemʼ
developed by ICR (ICR, 2013). An English language version
is now available.

RESULTS OF THE 2019 IWC-POWER SURVEY
In this sec�on, the main results of the 2019 IWC POWER
survey are summarized, based on Matsuoka et al. (2020).
I�nerary
The survey was conducted between 3 July and 25 September, 2019 in the Gulf of Alaska, within the U.S. Exclusive Economic Zone. The i�nerary is shown in Table 1.

Table 1The 2019 survey i�nerary.

26

Date

Event

5-Jul
6-Jul
6-Jul
11-Jul11-Jul
14-Jul16-Jul17-Jul18-Jul12-Aug
14-Aug
8-Sep
12-Sep14-Sep16-Sep16-Sep26-Sep28-SepPre-cruise mee�ng at Shiogama, Japan
Vessel departed Shiogama
Started transit survey to Dutch Harbor (D.H.)
Vessel enter U.S. EEZ (14:53, 48°08.3ʹN, 177°53.3ʹE)
Passed the dateline at 48-57.0N (23:03)
Vessel arrived at D.H., Alaska, USA
Pre-cruise mee�ng at D.H.
Vessel le� D.H. and started transit survey to the start point of Western stratum
Vessel started surveying the Western stratum
Vessel ﬁnished surveying the Western stratum and moved to the star�ng point of the Eastern stratum
Vessel started the survey of Eastern stratum
Vessel completed the Eastern stratum and started transit survey back to Kodiak (via NPRW Cri�cal Habitat)
Vessel completed transit survey
Vessel arrived Kodiak, Alaska, USA
Post-cruise mee�ng
Vessel le� Kodiak and started the transit survey to Shiogama
Vessel completed the transit survey
Vessel arrived Shiogama, JapanAn outline of the IWC-Pacific Ocean Whale and Ecosystem Research (IWC-POWER) including results of the 2019 survey

IWC SC for this survey:
Koji Matsuoka (Japan)–Cruise Leader/Chief Scien�st,sigh�ng, photo-ID;
Jessica Crance (USA)–acous�cs, photo-ID;
James Gilpatrik (USA)–sigh�ng, photo-ID data management, marine debris;
Isamu Yoshimura (Japan)–sigh�ng data, marine debris
and biopsy sample managements.

Figure 2.

Research area, transit and survey track lines withstart and end points for the 2019 IWC-POWER
cruise. D.H.: Dutch Harbor.

Figure 3.

Photography of the R/V Yushin Maru No. 2.

Table 2
Speciﬁca�ons of the R/V Yushin Maru No. 2.
Call sign
Length overall [m]Molded breadth [m]Gross tonnage (GT)Barrel height [m]IO barrel height [m]Upper bridge height [m]Bow height [m]Engine power [PS/kW]

JPPV
69.61
11.5
747
19.5
13.5
11.5
6.5
5303/3900

Research area
The research area was the northern Gulf of Alaska, between 170°W and 135°W, and comprised the U.S. EEZ(Figure 2).
Research vessel and scien�ﬁc personnel
The R/V Yushin Maru No. 2 was used for this survey. The
vessel is shown in Figure 3 and its speciﬁca�ons in Table
2.
Four interna�onal researchers were nominated by the

Searching eﬀort
A total of 1,030.7 n.miles (NSP: 534.0 n.miles, IO: 496.7 n.
miles) and 1,088.1 n.miles (NSP: 557.4 n.miles, IO:
530.7 n.miles) were surveyed in the western and eastern
strata, respec�vely. A total of 420.0 n.miles was surveyed
during transit surveys (e.g., transit from Dutch Harbor tothe most western point and transit between the end andstart points around Kodiak Island). The eﬀort (�me anddistance) spent on sigh�ng and several experiments areshown in Table 3.
Sigh�ng summary
In the research area, blue (17 schools/19 individuals), ﬁn(266/458), sei (9/20), common minke (5/5), humpback(173/402), gray (6/15), sperm (20/20), Bairdʼs beaked(2/37), Cuvierʼs beaked (3/5) and killer (53/264) whales
were sighted. Several dolphin species were also sighted(Table 4).
Gray whale (Eschrichtius robustus)
A total of 6 schools (15 individuals) were sighted in theWestern stratum, south of Kodiak Island (5 to 10 n.miles
oﬀ Dangerous Cape, south of Ugak Island, water depth
between 60 m and 80 m) (Figure 4a, b). Several graywhales were feeding (mud trails were seen). No motherand calf pairs were seen. Sea temperatures ranged from
12.9°C to 15.7°C.
Blue whale (Balaenoptera musculus)
Blue whales were widely distributed in the oﬀshore partof the Western and Eastern strata near EEZ border (Figure
5a, b). A total of 17 schools (19 individuals, including two
mother and calf pairs) were sighted (Table 4). All blue
whales were sighted in deep water of depths over 1,000 m.
Sea surface temperatures of the sigh�ng posi�ons were
between 10.9°C and 15.6°C.
Fin whale (Balaenoptera physalus)
Fin whales were the most frequently encounteredbaleen whale species in the research area, and they
were widely distributed throughout both the Westernand Eastern strata. Several high-density ﬁn whale areas
were observed, around Kodiak Island as well as the eastern part of the Eastern stratum (Figure 6). A total of 266
27Technical Reports of the Ins�tute of Cetacean Research (2020)
Table 3
Summary of the searching eﬀort (�me and distance) and experimental �me (hours) by each survey with the area code conductedduring 2019 survey. Dutch Harbor (D.H.), High Sea (H.S.), Research Area (R.A.).

Area

Area
Code

Shiogama to D.H. 1
H.S.
Shiogama to D.H. 11
D.H. to R.A

Leg
No.

Start

End

Start

Date

Date

End

Time

Time

1 6 Jul.

11 Jul.

—

14:53

6:00

1 11 Jul.

13 Jul.

U.S. EEZ

—

18:00

78

841 18-Jul.

18-Jul.

U.S. EEZ—

12:00

Western stratum 76

14:53
7:20

101 18-Jul.

26-Jul.

U.S. EEZ 114 12:00

13:13

Detour Shumagin 80
844 26-Jul.
IslandU.S. EEZ — 13:13

8:13

Western stratum 76

8-Aug.

116 27-Jul.

U.S. EEZ 126 8:13

27-Jul.

80

846 8-Aug.

8-Aug.

U.S. EEZ—

10:22

NPRW Cri�cal
Habitat80

847 8-Aug.

9-Aug.

U.S. EEZ—

10:38

Detour Kodiak
Island80

846 9-Aug.

10 Aug.

U.S. EEZ—

8:59

Western stratum 76

10:22
10:38

128 10 Aug. 12 Aug.

U.S. EEZ 134 8:59
W. to E. Stratum
Eastern stratum

849 12 Aug. 14 Aug.

U.S. EEZ—

77

201 14 Aug. 8-Sep.

U.S. EEZ 224 7:42
R.A to Kodiak
NPRW Cri�cal
Habitat
R.A to Kodiak

854 8-Sep.

10-Sep.

—

13:22

79

855 10-Sep. 12-Sep.

U.S. EEZ—

79

854 12-Sep. 13-Sep.
—

12

2 18-Sep. 22-Sep.

U.S. EEZ—

Kodiak to
Shiogama

22 23-Sep. 26-Sep.

H.S.

—

Total

28

Dist.
(n.m.)

TimeDist.
(n.m.)

TimeTime

0:00:00

0

26:35:14

322.32

26:35:14

322.32

0:00:00

0:00:00

0:00:00

0

0:00:00

0

0:00:00

0

0:00:00

0:00:00

0:00:00

0

0:00:00

0

0:00:00

0

0:00:00

2:43:00

20:02:43

228.61

19:35:18

224.71

39:38:01453.32

10:08:11

0:00:00

1:19:2215.3

0:00:00

0

1:19:2215.3

0:00:00

0:00:00

21:16:38

243.97

17:25:52

199.16

38:42:30

443.13

14:14:47

0:00:00

1:01:1611.64

0:00:00

0

1:01:1611.64

0:00:00

0:00:00

9:24:46106.280:00:00

0

9:24:46106.280:36:39

0:00:00

7:02:3280.2

0:00:00

0

7:02:3280.2

0:33:50

0:00:00

5:23:1161.45

6:22:33

72.78

11:45:44

134.23

2:23:12

0:00:00

8:57:11100.980:00:00

0

8:57:11100.98

1:45:04

0:00:00

49:00:30

557.35

46:33:50

530.73

95:34:20 1,088.08

25:33:44

7:06:35

0:00:00

0

0:00:00

0

0:00:00

0

0:00:00

0:00:00

9:21:39105.56

0:00:00

0

9:21:39105.56

0:00:00

0:00:00

0:00:00

0

0:00:00

0

0:00:00

0

0:00:00

0:00:00

0:00:00

0

0:00:00

0

0:00:00

0

0:00:00

0:00:00

0:00:00

0

9:23:32

115.03

9:23:32

18:00

Kodiak to
Shiogama

7:55

Time

18:00

U.S. EEZ

18:00

Dist.
(n.m.)

10:01

79

13:22

Time

7:42

U.S. EEZ

10:01

NSP+IO

13:31

81

13:31

IO

Es�mated angle
and distancetraining/
experiment

8:52

Detour Kodiak
Island8:52

NSP

Photo-ID,
Biopsy16:25
115.03

0:00:00

0:00:00

6:00

12:00

6-Jul.

26-Sep. 132:49:48 1,511.30 125:56:19 1,464.70 258:46:07 2,976.10

55:15:27

9:49:35

6:00

12:00

An outline of the IWC-Pacific Ocean Whale and Ecosystem Research (IWC-POWER) including results of the 2019 survey
Table 4
Number of sigh�ngs for all species observed in the research area (Original trackline and transit tracklines) by eﬀort mode. NSP:
Normal Passing with abeam closing Mode; IO: Independent Observer Mode (IO), OE: Top down (TD) and dri�ing (DR). Numbers of
Individuals includes the number of calves.
SpeciesBlue whale
Fin whale
Like ﬁn
Sei whale
Like seiCommon minke whale
Humpback whale
Like humpback
Gray whale
Sperm whale
Bairdʼs beaked whale
Cuvierʼs beaked whale
Ziphiidae
Killer whale
Paciﬁc white-sided dolphin
Habour porpoise
Dalli type Dallʼs porpoise
Unid. type Dallʼs porpoise
Unid. small cetacean

NSP

IO

OE

Total

Sch.

Ind.

Calf

Sch.

Ind.

Calf

Sch.

Ind.

Calf

Sch.

Ind.

Calf

5
145
6
3
0
3
116
1
6
8
2
1
2
36
0
1
50
3
1

6
260
11
4
0
3
234
1
15
8
37
1
4
156
0
1
277
13
3

1
0
0
0
0
0
0
0
0
0
2
0
0
9
0
0
1
0
0

9
112
13
6
1
2
49
6
0
12
0
2
3
17
0
0
34
1
0

10
186
17
16
2
2
132
14
0
12
0
4
3
108
0
0
196
4
0

1
0
0
0
0
0
0
0
0
0
0
0
0
5
0
0
5
0
0

3
9
1
0
0
0
8
0
0
0
0
0
0
0
1
0
2
0
0

3
12
2
0
0
0
36
0
0
0
0
0
0
0
4
0
32
0
0

0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
4
0
0

17
266
20
9
1
5
173
7
6
20
2
3
5
53
1
1
86
4
1

19
458
30
20
2
5
402
15
15
20
37
5
7
264
4
1
505
17
3

2
0
0
0
0
0
0
0
0
0
2
0
0
14
0
0
10
0
0

schools (458 individuals) of ﬁn whales were observedin the research area. No mother and calf pairs were observed. A total of 20 schools and 30 individuals of “Likeﬁn” were recorded; these looked like ﬁn whale blows but
could not be conﬁrmed (Table 4). Most ﬁn whales weresighted in deep water of depths over 1,000 m (Figure 6).
Sea temperatures ranged from 9.9°C to 16.7°C.
Sei whale (Balaenoptera borealis)
A total of 9 schools (20 individuals) of sei whales wereobserved (Table 4). Observed mean school size was 2.22.
Although sei whales were the third most frequently
encountered baleen whale species through the surveys,
almost all sei whale sigh�ngs were in the southern partof the Western stratum. There was a notable lack of sei
whales in the Eastern stratum (Figure 7). This is similar
to results seen during a previous POWER survey in 2012,
where no sei whales were sighted in the U.S. EEZ (Eastern
Stratum of current survey). Sea surface temperatures
ranged from 11.2°C to 13.0°C, which was lower than therange for ﬁn whales.
Common minke whale (Balaenoptera acutorostrata)
Common minke whales were the rarest baleen whale
species in the research area, and were only sighted very
close to shore (Figure 8). A total of 5 schools (5 indi-viduals, all solitary schools) were observed (Table 4). Sea
temperature at sigh�ng loca�ons ranged from 10.8°C
to 16.9°C. Because of the diﬃculty in seeing their blow
and small body, observa�ons of this species were morediﬃcult than in Antarc�c waters. Common minke whaleblows are very small and are diﬃcult to spot in rough sea
surface condi�ons (e.g., wind speed over 12 knots). During this survey, sea states averaged 4–5 on the Beaufort
scale, which is assumed to be too rough for sigh�ng common minke whales.
Humpback whale (Megaptera novaeangliae)
Humpback whales were the second most frequently
encountered baleen whale species in the research area,
primarily in the Western stratum. High-density areasof this species were observed in waters north-east andsouth-east of Kodiak Island, and along the most eastern
trackline along the con�nental shelf (near 135°W). There
was a lack of sigh�ngs between 150°W and 138°W in the
Eastern stratum (Figure 9). A total of 173 schools (402individuals) were observed in the research area. There
were no mother and calf pairs. In some cases, red colored
fecal ma�er was observed indica�ng that the whales
were feeding on krill. Sea temperatures ranged from
10.0°C to 15.7°C.
29Technical Reports of the Ins�tute of Cetacean Research (2020)

Figure 4a.

The searching eﬀort (thin line) and sigh�ng posi�ons (white circles) of gray whales during the 2019
IWC-POWER cruise.

Figure 5a.

The searching eﬀort (thin line) and sigh�ng posi�ons (blue circles) of blue whales during the 2019
IWC-POWER cruise.

Figure 4b.

Three gray whales observed during the 2019
IWC-POWER survey.

Figure 5b.

Two blue whales observed during the 2019 IWC
POWER survey.

Sperm whale (Physeter macrocephalus)
Sperm whales were widely distributed throughout theWestern and Eastern strata where the water depth wasover 1,000 m (Figure 10). A total of 20 schools (20 individuals) were sighted (Table 4). All schools observed were
solitary individuals (probably large male). Sperm whales
were recorded in waters with SST ranging from 10.9°C to
16.7°C.
Bairdʼs beaked whale (Berardius bairdii)
A total of 2 schools (37 individuals) were sighted(Table 4). Both Bairdʼs beaked whale schools were sighted
along the most eastern trackline in the Eastern stratum
along the con�nental slope oﬀ Prince of Wales Island(water depths over 1,000 m). Sea temperatures ranged
from 13.9°C to 14.4°C.
Killer whale (Orcinus Orca)
Killer whales were mainly sighted in the Western stratum (Figure 11). A total of 50 schools (248 individuals,
including 13 calves) were sighted (Table 4). They weresighted in waters with SST ranging from 10.9°C to 16.2°C.
30

Iden�ﬁca�on of duplicated sigh�ngs
A total of 154 resigh�ngs (duplicates) was made during
IO Mode. Of these, for ﬁn whales, there were 60 ʻDeﬁnite
duplicates,ʼ 1 ʻprobable duplicateʼ and 13 ʻNot duplicates.ʼ
For sei whales, there were 12 ʻDeﬁnite duplicatesʼ and 4
ʻNot duplicates.ʼ For humpback whales, there were 34
ʻDeﬁnite duplicatesʼ and 8 ʻNot duplicates.ʼ
Photo-ID photographs
Photo-iden�ﬁed species included: blue (15 schools/16
individuals), ﬁn (64/51), humpback (25/30), gray (2/6)
and killer (8/19) whales (Table 5). Bairdʼs beaked whales(20 individuals) and Dallʼs (24) porpoises were also photographed. Images collected during the cruise were uploaded to the IWC/ICR master photographic database in
Adobe Lightroom (LR) (Anon, 2019b). Preliminary coding
was completed for all cetacean images (7,762), including
the alloca�on of species name, sigh�ng number, school
size and biopsy eﬀort.

An outline of the IWC-Pacific Ocean Whale and Ecosystem Research (IWC-POWER) including results of the 2019 survey

Figure 6.

The searching eﬀort (thin line) and sigh�ng posi�ons (red circles) of ﬁn whales during the 2019
IWC-POWER cruise.

Figure 9.

The searching eﬀort (thin line) and sigh�ng posi�ons (black circles) of humpback whales during the2019 IWC-POWER cruise.

Figure 7.

The searching eﬀort (thin line) and sigh�ng posi�ons (yellow circles) of sei whales during the 2019
IWC-POWER cruise.

Figure 10.

The searching eﬀort (thin line) and sigh�ng posi�ons (red triangles) of sperm whales during the2019 IWC-POWER cruise.

Figure 8.

The searching eﬀort (thin line) and sigh�ng posi�ons (pink circles) of common minke whales during
the 2019 IWC-POWER cruise.

Figure 11.

The searching eﬀort (thin line) and sigh�ng posi�ons (black cross) of killer whales during the 2019
IWC-POWER cruise.

Biopsy sampling
Biopsy samples were collected from 75 individual whales:
12 blue, 45 ﬁn, 4 sei, 12 humpback, 2 gray whales(Table 6). Every biopsy encounter was documented photographically. All biopsy samples were catalogued andstored in cryo-vials frozen at a temperature of −30°C
on the vessel. These samples will be used for moleculargene�cs analyses. Figure 12 shows an event of biopsy
sampling.

31

Technical Reports of the Ins�tute of Cetacean Research (2020)
Table 5
Summary of the number of Photo-IDʼd individuals by each species.
Blue

Fin

Humpback

Gray

Killer

Total

Western Stratum
Eastern Stratum

9
7

16
35

20
10

6
0

5
14

56
66

Sub-total (US-EEZ)

16

51

30

6

19

122

Sub-total (High Sea)

0

0

0

0

0

0

Total

16

51

30

6

19

122

Table 6
Summary of the number of biopsy samples collected by each species.
Blue*

Fin

Sei

Humpback

Gray

Total

Western Stratum
Eastern Stratum

6
6

12
33

4
0

10
2

2
0

34
41

Sub-total (U.S. EEZ)

12

45

4

12

2

75

Sub-total (High Sea)

0

0

0

0

0

0

Total

12

45

4

12

2

75

Blue* Including one mother and calf pair.

corded ʻon eﬀortʼ (i.e., during the ﬁrst 15 minutes of eachhour) and 1 item was recorded during ʻoﬀ eﬀort.ʼ

WHAT NEXT?
The following tasks are iden�ﬁed for the immediate
future of IWC-POWER.

Figure 12. Biopsy sampling for sei whale during the IWC-POWER
survey.

Acous�c monitoring
A total of 229 sonobuoys were deployed for a total ofover 820 monitoring hours. Species detected included ﬁn(119 buoys, 56.1% detec�on), sperm (112, 52.8%), killer(76, 35.8%), blue (54, 25.5%), humpback (47, 22.2%),
North Paciﬁc right (10, 4.7%), and sei (4, 1.9%) whales.
Other species/events detected included Bairdʼs beaked
whales (3, 1.4%), Paciﬁc white-sided dolphins (3, 1.4%),
earthquakes (33, 15.6%), and unknown calls/signals (12,5.7%).
Marine debris observa�on
A total of 42 marine debris objects were observed including 9 single ﬁshing ﬂoats. A total of 41 items were re32

Comple�on of the short-term priori�es
To complete the short-term priori�es, the IWC SC reiterated the importance of comple�ng the Bering Sea survey
areas. However, given the diﬃcul�es previously experienced, it was important to consider a backup plan forthe 2020 cruise. If the Russian area cannot be coveredin 2020, then every eﬀort should be made to cover thisin 2021, given its importance to mee�ng the objec�vesof the IWC-POWER program (IWC, 2020c). Also, several
analy�cal tasks based on data collected in the ﬁrst phase
under the short-term priori�es were recommended.
Design of the next phase
The IWC SC agreed on revised priori�es for the mediumterm (Table 7). To develop a detailed plan for the second
phase of IWC-POWER, it is important to complete theanalyses of the data collected during the ﬁrst phase(short-term).
The IWC SC will hold a pre-mee�ng just before the2021 Scien�ﬁc Commi�ee Mee�ng in order to develop
a detailed proposal for a workshop to design the next

An outline of the IWC-Pacific Ocean Whale and Ecosystem Research (IWC-POWER) including results of the 2019 survey
Table 7
Sugges�ons for updated medium-term priori�es based upon results from Phase 1 for IWC-POWER (*refers to likelihood of obtaining an abundance es�mate at least in some areas **refers to likelihood of obtaining biopsy and/or photo-ID data from encounteredschools) (IWC, 2020c).
Ini�al priority/feasibilityRa�onale/comments

Blue whale (High)
Medium direct*, highopportunis�c**•Deple�on level (i.e. highly depleted based on catch history)•Ini�al abundance es�mates from IWC-POWER (s�ll being ﬁnalised) suggest it remains heavily depleted.
•Results of analyses of exis�ng samples (27 IWC-POWER samples available) in conjunc�on with other samples (e.g. samples collectedunder Japanese na�onal programmes) important in addressing popula�on structure in context of idea/scope of occasional focussed
cruises, especially samples from the west (see (4) below).
•Poten�al for some blue whale focussed cruises in speciﬁc areas (including direc�onal acous�cs) should be considered (e.g. Gulf of Alaska)
as well as con�nuing opportunis�c studies.
•Con�nued collabora�on with exis�ng photo-id work e.g. US and Japanese na�onal programmes is important (42 individuals available
from IWC-POWER)–and Japan west exis�ng samples.
•Consider telemetry studies

Fin whale (High)High direct*, highopportunis�c**•Deple�on level (i.e. high based upon catch history)•Ini�al abundance es�mates from IWC-POWER (s�ll being ﬁnalised) suggest some recovery•Results of gene�c analyses important to contribute to future survey strategy and future Comprehensive Assessment (e.g. is there
evidence of more than one stock from the exis�ng 124 biopsy samples that cover waters from 170°E to 135°W?).
•Work in Russian Federa�on waters provided appropriate permits can be obtained is important•Co-ordina�on with na�onal programmes in Japan, Korea and USA needed including exis�ng samples

Right whale (High)
Medium direct*, highopportunis�c**•Deple�on level: (i.e. highly depleted based on catch history)•S�ll cri�cally low numbers in east (from US studies and IWC-POWER)•Feasibility of collec�ng biopsy and photo-ID data high if targeted and using acous�cs•Feasibility of obtaining abundance in east from line-transect low given such small numbers; may be much higher in west e.g. Sea of
Okhotsk and southeast of Kamchatka Peninsula where popula�on is at least 10X larger or more
•Although new area, considera�on should be given to a targeted survey in Sea of Okhotsk-high feasibility and priority to obtain good abundance, photo-id and biopsy data provided appropriate permits can be obtained from the Russian Federa�on.

Sei whale (Medium)
High direct*, highopportunis�c**•Deple�on level: (i.e. high based on catch history)•Ini�al abundance es�mates from IWC-POWER (s�ll being ﬁnalised) and Japan suggest some recovery•IWC-POWER has provided valuable informa�on for the ongoing Comprehensive Assessment (the 2020 backup cruise will provide biopsy
samples from a poorly covered area)•Results of that CA will help focus future IWC-POWER medium-term strategy and priority for this species-e.g. (a) possible focussed biopsy
sampling in postulated coastal stock areas and (b) frequency and scope dedicated abundance surveys in ʻpelagicʼ area to examine trends

Humpback whale (Medium)
High direct*, highopportunis�c**•Good informa�on already available from SPLASH and na�onal programmes suggests overall high abundance (gene�c and photo-ID markrecapture) hence medium priority•Con�nue to contribute to exis�ng gene�c and photo-ID databases.
•Ongoing Comprehensive Assessment will assess status and poten�al deple�on of [sub-] popula�ons.
•Abundance es�mates from IWC-POWER (s�ll being ﬁnalised) can provide interes�ng ʻsnapshotʼ es�mates to compare with mark-recapture es�mates by popula�on/feeding aggrega�on•The results of the CA will assist in developing medium-term strategy and priority for this species within IWC-POWER

Sperm whale (Medium)•Deple�on level: (unknown but possibly high given catch history)
Medium direct* and medium •Lack of good informa�on on popula�on structure and status at present although good distribu�onal data from IWC-POWER
opportunis�c**•Obtaining abundance es�mates from visual surveys can be problema�c due to long dive �mes and other issues•Combined acous�c (towed array)/visual surveys have been successful for sperm whales however feasibility in the context of IWC-POWER
depends on availability of equipment and prac�cality in light of other priori�es•Possibility of using towed acous�c arrays in some years should be considered
Gray whale (Medium)Low direct*, highopportunis�c**•There are ASW hunts but that primary data sources to evaluate those are from other visual, gene�c and photo-ID programmes (e.g. US,
Mexico, Sakhalin Island)–hence medium priority•Main IWC-POWER contribu�on is in obtaining biopsy/photo-ID in areas outside those programmes for comparison and informa�on on
popula�on structure•Sharing of data with other programmes should con�nue

Brydeʼs whale (Medium)
High direct*, highopportunis�c**•Suggest low priority for ﬁrst six or so years of next phase of POWER because:•Recently completed IR shows good popula�on status and apparently low level of threats•Removing from target species allows a great reduc�on in size of priority research area to north of 40°N
•If required, a targeted survey or surveys could be designed towards end of 10-year period (e.g. from 2027)

Common minke whale (Low) •Deple�on level (probably low east/central based upon catch history) and in west dealt with by na�onal programmes
Suggest only opportunis�c•However, if Okhotsk Sea is able to be covered for high priority species (e.g. right whales) then would provide valuable informa�on incl.
biopsy samples•If permission granted by Russian Federa�on then consider modifying present ʻacceptableʼ condi�ons as at the present high range they
are unsuitable for es�ma�ng abundance for this species

33

Technical Reports of the Ins�tute of Cetacean Research (2020)

phase of the IWC-POWER program. The emphasis will be
on par�cipa�on from all range states and the availabilityof analyses/data. The workshop should include considera�on of more methodologically focused cruises in designated years (e.g., use of towed acous�c arrays, telemetry
work, use of SeaGlider, etc.).
The ICR scien�sts will par�cipate ac�vely in the discussions regarding the second phase of IWC-POWER. This
research program has contributed and will con�nue tocontribute important informa�on relevant for the conserva�on and management of large whales in the North
Paciﬁc.

ACKNOWLEDGEMENTS
I thank all interna�onal researchers for their hard work
during the IWC-POWER cruises. Thanks also to the crewsof the respec�ve vessels that par�cipated in the annual
surveys for their eﬀort and assistance. I also thank scien�sts of the IWC-POWER Steering Group for planning anddesigning the cruises and for their technical support. My
gra�tude to the IWC and the Governments of Japan andthe USA for ﬁnancial support. Acknowledgement to Luis
A. Pastene (ICR) for his assistance in the prepara�on ofthis document.

REFERENCES
Anon. 2019a. 2019 IWC/POWER Cruise, Informa�on for
Researchers. 55 pp. [Available from the IWC Secretariat].
Anon. 2019b. IWCʼs IDCR/SOWER and POWER cruise
Photographic Database Manual. 21 pp. [Available from the
IWC Secretariat].
Crance, J.L., Berchok, C.L. and Kea�ng, J.L. 2017. Gunshot callproduc�on by the North Paciﬁc right whale, Eubalaena japonica, in the southeastern Bering Sea. Endang. Species Res.
34: 251–267.
Ins�tute of Cetacean Research. 2013. The Outline of the ICR

34

Accurate Informa�on System (ICRAS). 13 pp. [Available fromthe IWC Secretariat].
Interna�onal Whaling Commission. 2011. Report of theIntersessional Mee�ng on the North Paciﬁc Survey
Programme. J. Cetacean Res. Manage. (Suppl.) 12: 413–420.
Interna�onal Whaling Commission. 2012. Report of theWorkshop on Planning for an IWC Co-ordinated North Paciﬁc
Research Cruise Programme. J. Cetacean Res. Manage.
(Suppl.) 13: 369–391.
Interna�onal Whaling Commission. 2013. Report of theTechnical Advisory Group (TAG) Mee�ng on the Short andMedium Term Objec�ves and Plans for the IWC-POWER
Cruises. J. Cetacean Res. Manage. (Suppl.) 14: 341–355.
Interna�onal Whaling Commission. 2020a. Report of the
Planning Mee�ng for the 2019 IWC-POWER Cruise. J.
Cetacean Res. Manage. (Suppl.) 21: 333–345.
Interna�onal Whaling Commission. 2020b. Report of the 2018
Mee�ng of the IWC-POWER Technical Advisory Group (TAG).
J. Cetacean Res. Manage. (Suppl.) 21: 311–332.
Interna�onal Whaling Commission. 2020c. Report of theMee�ng of the IWC-POWER Technical Advisory Group (TAG),
January 2020, Tokyo, Japan. Paper SC/68B/Rep01 presented
to the IWC Scien�ﬁc Commi�ee, May 2020 (unpublished). 23
pp. [Available from the IWC Secretariat].
Larsen, F. 1998. Development of a biopsy system primarily foruse on large cetaceans. Paper SC/50/O15 presented to the
IWC Scien�ﬁc Commi�ee, May 1998 (unpublished). 7 pp.
[Available from the IWC Secretariat].
Matsuoka, K., Hakala, S., Kim, H.W., Aki, M. and Shinyasiki,Y. 2011. 2010 IWC/Japan Joint Cetacean Sigh�ng Survey
Cruise in the North Paciﬁc. Paper SC/63/O5 presented to the
IWC Scien�ﬁc Commi�ee, June 2011 (unpublished). 40 pp.
[Available from the IWC Secretariat].
Matsuoka, K., Crance, J., Gilpatrick, J., Yoshimura, I. and Ohkoshi,
C. 2020. Cruise report of the 2019 IWC-Paciﬁc Ocean Whaleand Ecosystem Research (IWC-POWER). Paper SC/68B/ASI/20
presented to the IWC Scien�ﬁc Commi�ee, May 2019 (unpublished). 57 pp. [Available from the IWC Secretariat].

Technical Reports of the Ins�tute of Cetacean Research (2020)

pp. 35–40

Technical Report (not peer reviewed)

Gene�c tagging technique: basic concept and a case study by theIns�tute of Cetacean Research
Mioko Taguchi*
Institute of Cetacean Research, 4–5 Toyomi-cho, Chuo-ku, Tokyo 104–0055, Japan*Contact e-mail: taguchi@cetacean.jp

ABSTRACTThe studies on biological conserva�on require contemporary demographic es�mates, e.g., migra�on rate,eﬀec�ve popula�on size and stock structure. Although these es�mates can be obtained through standard
popula�on gene�cs analyses, such studies could fail to infer them because of recent evolu�onary and biogeographical events. Alterna�ve approaches to obtain such parameters include individual iden�ﬁca�on andkinship inferences using gene�c data, also called ʻgene�c taggingʼ, which has become a common technique inthe ﬁelds of ecology and conserva�on during the last several decades. This paper presents the basic concept ofgene�c individual iden�ﬁca�on and kinship inference. It subsequently outlines the parentage analysis for thewestern North Paciﬁc common minke whale conducted by the Ins�tute of Cetacean Research as a case study.

INTRODUCTION
The studies on biological conserva�on in animal species
require informa�on on contemporary popula�on parameters, e.g., eﬀec�ve popula�on size, migra�on rate andpopula�on structure. The es�mates of these parameters
can be obtained through tradi�onal popula�on gene�cs
analyses. However, these analyses o�en fail to infer themin cases where the popula�ons have diverged recently,or if changes in gene ﬂow and eﬀec�ve popula�on size
have occurred. Therefore, such es�mates have been
mainly obtained by individual-based tagging researchusing conven�onal tagging or photo-iden�ﬁca�on (e.g.,
Calambokidis et al., 2001; Johnson et al., 2009; Mizroch
and Rice, 2013; Urian et al., 2015).
Individual animals can also be iden�ﬁed using gene�cdata, generally called ʻgene�c taggingʼ (Palsbøll, 1999).
This has become a common technique to obtain the contemporary es�mates in the ecological and conserva�onﬁelds during the last several decades (e.g., humpback
whales, Megaptera novaeangliae, Palsbøll et al., 1997;
North Paciﬁc right whales, Eubalaena japonica, Wadeet al., 2011). In addi�on, in recent years, the extended gene�c tagging incorpora�ng informa�on on kinship amongindividuals is being used to inves�gate contemporary
popula�on dynamics (e.g., Kanda et al., 2014; Bravington
et al., 2016; Ohashi et al., in-press). The methods forabundance es�mates based on two types of gene�c tags
(individual iden�ﬁca�on and kinship) are summarized by

Takahashi (this issue).
This paper presents the basic concept of gene�cindividual iden�ﬁca�on and kinship inference, with abrief summary of a case study by the Ins�tute of Cetacean Research (ICR), which is the parentage analysis for
the North Paciﬁc common minke whale (Balaenoptera
acutorostrata).

BASIC CONCEPT OF GENETIC TAGGING TECHNIQUES
Individual iden�ﬁca�on
Sexual reproduc�on guarantees that, in the main, eachindividual has a unique genotype. There are some excep�ons, however, such as iden�cal twins. This characteris�callows each individual to be tagged and, as a consequence, it can be determined if two gene�c samples
come from the same individual. Figure 1 shows an example of individual iden�ﬁca�on using microsatellite
genotype data. In this example, the four samples are derived from three whales. Samples A and B have the same
genotypes and therefore the two samples are assumed
from the same whale, while samples C and D have diﬀerent genotypes sugges�ng they are from diﬀerent whales.
Probability of identity
The probability of iden�ty (I) is the probability that two
unrelated individuals in the popula�on share the same
genotype. This es�mate requires informa�on on allele
frequencies in the popula�on, because common alleles
35

Technical Reports of the Ins�tute of Cetacean Research (2020)

Figure 1.

Example data involving four microsatellite loci genotyped for each of four whales. Genotypes are represented by 6-digit codes, and each allele is coded by 3 digits. The results of individual iden�ﬁca�on are
shown in the column ʻMatching DNA proﬁleʼ.

Figure 2.

Example of sex determina�on in common minke whales through mul�plex ﬂuorescent PCR using the SRY
locus (black) located on the Y chromosome and a microsatellite locus (blue) as the internal control: (a)failure to amplify PCR fragments; (b) male; (c) female.

are much more likely to be shared than rare ones. Thiscan be es�mated for a popula�on with random ma�ng
according to the formula derived by Paetkau and Strobeck(1994):

Iˆ =



pi4 +

i
k  i

∏


(2 pi p j ) 2 

j >i




where, pi and pj are the frequencies of the ith and jthalleles at the kth locus in the popula�on. In prac�ce, it
is desirable to aim for a probability of shared genotypeiden�ty between any two individuals (I) of <0.05.

36

Molecular sex determination
Sex informa�on is useful not only to assist in individual
iden�ﬁca�on but also for a biological interpreta�on of
the results of analyses.
The PCR-based technique has been proposed for sex
determina�on using the presence or absence of genes on
the sex chromosomes. The tes�s-determining SRY genes
are male-speciﬁc in mammals and can be detected by
PCR ampliﬁca�on using speciﬁc primers. The ICR gene�cteam developed the primer set that ampliﬁes the SRY
gene to determine the sex of cetaceans, using a microsatellite locus as the internal control (Abe et al., 2001).

Genetic tagging technique: Basic concept and a case study by the Institute of Cetacean Research

Figure 3.

Example of pedigree containing ten non-inbred individuals. The ﬁgure shows the kinship categories andthe values for three IBD coeﬃcients (κ 0, κ 1 and κ 2). The three IBD coeﬃcients can be also summarized by
the kinship coeﬃcients (Φ) which is the probability that a random allele from one individual is IBD to arandom allele from the other. The Φ is also equivalent to the inbreeding coeﬃcient of their oﬀspring.

Figure 2 shows an example of gene�c sex determina�onusing this method. In case of male, a PCR fragment of SRY
gene is observed between 150 and 155bps (Figure 2b),while the fragment is absent in case of female or PCR
failure. We can dis�nguish female (Figure 2c) from PCR
failure (Figure 2a) by the presence of the PCR fragment of
one microsatellite locus as an internal control.
Kinship inference
Individuals that are biologically related share genes thatare iden�cal-by-descent (IBD), i.e., iden�cal copies of agene segrega�ng from a common ancestor within thedeﬁned pedigree. IBD is a fundamental concept that
underlies kinship inference (Thompson, 2013). Here, thebasic concept of the IBD coeﬃcient is explained, which
provides an introduc�on to the ideas of IBD. It is followedby the simple principle of parentage analysis which is aunique applica�on of kinship inferences.
IBD coeﬃcients
Two non-inbred diploid individuals can share none,
one, or two alleles IBD. In case of the pedigree shown in
Figure 3, full sibling individuals 4 and 5 share one allele
IBD (A3 which is inherited from their father) while unrelated individuals 3 and 6 share no allele IBD. The monozygotic twins 9 and 10 share two alleles IBD (A3 and A6, which

Table 1Pa�erns of allele sharing IBD for a full sibling pair (individuals
4 and 5 in Figure 3).
Possiblegenotypes

A1A3

A1A4

A2A3

A2A4

A1A3
A1A4
A2A3
A2A4

2
1
1
0

1
2
0
1

1
0
2
1

0
1
1
2

are inherited from father and mother, respec�vely). The
probabili�es of each of the events are summarized by
the IBD coeﬃcients, i.e., κ 0, κ 1 and κ 2 (Cockerham, 1971),
which are shown in Figure 3.
Table 1 shows an example on how to determine the
IBD coeﬃcients for a category of full siblings. Consider
that the genotypes of the parents at a locus are A1A2 andA3A4 (individuals 1 and 2 in Figure 3). Each oﬀspring canhave one of the four following genotypes: A1A3, A1A4,
A2A3 or A2A4 (individuals 4 and 5 in Figure 3). Out of thesixteen ways to pair two oﬀsprings, the dyad can share 2
alleles that are IBD in four ways, 1 allele in eight ways and0 alleles in four ways. Thus, the IBD coeﬃcients, κ 0, κ 1 andκ 2, for full siblings are 0.25 (4/16), 0.50 (8/16) and 0.25
(4/16), respec�vely. The IBD coeﬃcients for other kinshipcategories can be calculated in the same manner.
37

Technical Reports of the Ins�tute of Cetacean Research (2020)

Figure 4.

Principle of parentage analysis using the exclusion approach. The numbers under the peaks are the size of
PCR fragment in base pairs which can be interpreted as alleles.

Parentage analyses
Parentage analysis is a unique applica�on in which
the most likely parents of a target oﬀspring are sought
amongst eligible candidates. The basic concepts of theparentage analysis are grouped into six categories according to Jones et al. (2010): (1) Exclusion, (2) Categorical
Alloca�on, (3) Frac�onal Alloca�on, (4) Full Probability
Parentage Analysis, (5) Parental Reconstruc�on, (6) Sibship Reconstruc�on. In this sec�on, categories (1) (theconceptually simplest approach) and (2) (the most commonly used approach), are described.
Exclusion approachThis approach uses rules of Mendelian inheritance for
diploid organisms wherein an oﬀspring inherits oneof two alleles at each locus from each of its parents. A
simple exercise for the exclusion approach using a single
microsatellite locus is shown in Figure 4.
In this exercise, the mother shows the genotype of
200/204, and the genotypes of her three oﬀspring 1, 2
and 3 were 196/200, 200/208 and 204/210, respec�vely.
Here, a male with genotype of 196/208 can be excluded
as a father candidate for oﬀspring 3 since no allele fromthe male is observed in oﬀspring 3. On the other hand,
the male is s�ll a father candidate for oﬀspring 1 because
the alleles 196 and 200 can be found from the male andmother, respec�vely. The same is true for oﬀspring 2.
This approach is powerful when there are few candidate parents and mul�ple highly polymorphic markers
are available. In prac�ce, most of the studies based on
the exclusion approach actually require at least two mismatching loci between the candidate and the oﬀspring to38

account for typing errors or muta�ons.
Categorical Alloca�on approach
As in the case of exclusion method, this approach requires at least one focal oﬀspring and a set of candidate
parents. The Categorical Alloca�on approach was developed to resolve situa�ons in which complete exclusion
may not be feasible (Meagher and Thompson, 1986). The
main beneﬁt of this approach is to choose the single most
likely parent from a group of non-excluded puta�ve parents. The logic stems from the observa�on that diﬀerent
genotypes of parent may diﬀer in their probability of having produced the genotype of the focal oﬀspring (Meagher and Thompson, 1986). This approach also has theadvantage of handling genotyping errors or muta�ons.
Currently, most of the Categorical Alloca�on approaches use a likelihood approach (e.g., Marshall et al., 1998).
However, a Bayesian approach (e.g., Nielsen et al., 2001)
can also be used (see also Takahashi, this issue).
Genotyping errors and their eﬀects
Microsatellite DNA marker is one of the most common
gene�c markers not only for individual iden�ﬁca�onand kinship analyses, but also for the standard analyses
on popula�on gene�cs. However, microsatellite DNA is
known as the error-prone marker in its genotyping, whichis recognized to have a serious impact on gene�c individual iden�ﬁca�on as well as kinship inferences (reviewedin Hoﬀman and Amos, 2005).
The genotyping of a microsatellite allele depends onthe microsatellite proﬁle, and requires strict rules to bedeﬁned in advance. Typical microsatellite proﬁles are

Genetic tagging technique: Basic concept and a case study by the Institute of Cetacean Researchcharacterized by a succession of peaks with growing intensity due to stu�er band. Researchers o�en suﬀer fromseveral types of ambiguous microsatellite proﬁles in genotyping, e.g., null alleles, overlapping of stu�er bands,
split peak, large allele dropout or short allele dominanceand false alleles. Some factors are known to be involvedin the ambiguous proﬁles, e.g., varia�on in DNA sequences, low quality or quan�ty of template DNA, biochemicalar�fact and human error (Hoﬀman and Amos, 2005).
Figure 5 illustrates two types of genotyping error thatare likely to induce false paternity and consequently bias
the biological conclusions. Consider in the example of
Figure 5 that the male is the real father of the oﬀspring.
The real genotype for male is 196/208 without error: theoﬀspring inherited allele 200 from his mother and allele
208 from his father. In the cases with genotyping errors,

Figure 5.

Simple example of parentage analysis by the
Exclusion approach. Arrows show the genotyping
errors due to false allele (red) and large allele dropout (blue).

Figure 6.

the male would incorrectly be excluded as the father ofthe oﬀspring because allele 208 is mistyped as allele 202.
In order to avoid this type of incorrect paternity exclusion, it is be�er to allow one or several gene�c mismatches in the parentage analysis depending on the probabilityof iden�ty and the calculated error rate.

A CASE STUDY ON PARENTAGE BY THE INSTITUTEOF CETACEAN RESEARCH
Parent-oﬀspring inference in North Paciﬁc common
minke whales
The parent-oﬀspring pair analysis was carried out based ona type of Categorical Alloca�on approach, using a total of
4,707 common minke whales in the western North Paciﬁc(see Tiedemann et al. (2017) for the details of the analy�cal procedures). Each sample was genotyped at 16 microsatellite loci. The mitochondrial DNA control region haplotypes as well as biological informa�on, e.g., sampling dateand posi�on, sex, sexual maturity and body length, wereused, if available, to assist the interpreta�on of the results.
The parent-oﬀspring analyses inferred a total of 40
and 13 parent-oﬀspring pairs for the O and J stock (Gotoet al., 2017), respec�vely . The forty parent-oﬀspring pairsof O stocks were widely distributed through the Paciﬁcside of Japan (Figure 6). A total of 17 pairs of O stock animals were found between coastal and oﬀshore waters. In
most of the pairs, the oﬀspring were found near the coast
while their parents were in oﬀshore waters. This is consistent with the pa�ern of migra�on with sexual segrega�ondescribed previously for western North Paciﬁc common
minke whales (Hatanaka and Miyashita, 1997).
The thirteen parent-oﬀspring pairs of J stocks were
distributed within and between the Sea of Japan and the
Paciﬁc side of Japan (Figure 6). Four of them were foundbetween the Sea of Japan side and the Paciﬁc side. The

Distribu�on of parent-oﬀspring pairs of O (le�) and J (right) stocks in western North Paciﬁc common minke
whales. Blue square: parent; orange circle: mature oﬀspring; red cross: immature or unknown maturity
stage oﬀspring.
39

Technical Reports of the Ins�tute of Cetacean Research (2020)

results implied that the J stock individuals occurring in
the Sea of Japan and the Paciﬁc side of Japan were derived from the same stock.
Future work
The development of the new gene�c marker which is lesserror-prone, that is, Single Nucleo�de Polymorphisms(SNPs) marker is on-going at the ICR. The ICR gene�cteam will perform analyses using SNPs genotypes, notonly for western North Paciﬁc common minke whales,
but also for other whale species such as Brydeʼs, ﬁn, rightand blue whales. The results of these analyses will aid in
obtaining the contemporary demographic es�mates ofthese whale species using gene�c data.

REFERENCES
Abe, H., Goto, M., Pastene, L.A., Dewa, K. and Naito, E. 2001.
Prac�cal use of mul�plex ﬂuorescent PCR for cetacean sex
iden�ﬁca�on. Mar. Mam. Sci. 17(3): 657–664.
Bravington, M.V., Skaug, H.J. and Anderson, E.C. 2016. Close-kin
mark-recapture. Statist. Sci. 31(2): 259–274.
Calambokidis, J., Steiger, G.H., Straley, J.M., Herman, L.M.,
Cerchio, S., Salden, D.R., Jorge, U.R., Jacobsen, J.K., Ziegesar,
O.V., Balcomb, K.C., Gabriele, C.M., Dahlheim, M.E., Uchida,S., Ellis, G., Mlyamura, Y., de guevara Paloma Ladrón,P., Yamaguchi, M., Sato, F., Mizroch, S.A., Schlender, L.,
Rasmussen, K., Barlow, J. and Ii, T.J.Q. 2001. Movements
and popula�on structure of humpback whales in the North
Paciﬁc. Mar. Mam. Sci. 17: 769–794.
Cockerham, C.C. 1971. Higher order probability func�ons ofiden�ty of alleles by descent. Genetics 69(2): 235–246.
Goto, M., Taguchi, M. and Pastene, L.A. 2017. Distribu�onand movement of ʻOʼ and ʻJʼ stock common minke whales
in waters around Japan based on gene�c assignment methods. Technical Reports of the Institute of Cetacean Research(TEREP-ICR) No.1: 37–43.
Hatanaka, H. and Miyashita, T. 1997. On the feeding migra�on of the Okhotsk Sea-West Paciﬁc stock of minke whales,es�mates based on length composi�on data. Rep. int. Whal.
Commn 47: 557–564.
Hoﬀman, J.I. and Amos, W. 2005. Microsatellite genotyping
errors: detec�on approaches, common sources and consequences for paternal exclusion. Mol. Ecol. 14: 599–612.
Johnson, M., Aguilar, de Soto N. and Madsen, P.T. 2009. Studying
the behaviour and sensory ecology of marine mammals using
acous�c recording tags: a review. Mar. Ecol. Prog. Ser. 395:
55–73.
Jones, A.G., Small, C.M., Paczolt, K.A. and Ra�erman, N.L. 2010.
A prac�cal guide to methods of parentage analysis. Mol. Ecol.
Resour. 10: 6–30.
Kanda, N., Goto, M. and Pastene, L.A. 2014. Paternity analysis
on Antarc�c minke whales using JARPA and JARPAII samples.

40

Paper SC/F14/J7 presented to the IWC/SC Review Workshopof the Japanese Whale Research Program under Special
Permit in the Antarc�c-Phase II (JARPAII), February 2014
(unpublished). 4 pp. [Available from the IWC Secretariat].
Marshall, T.C., Slate, J., Kruuk, L.E.B. and Pemberton, J.M. 1998.
Sta�s�cal conﬁdence for likelihood-based paternity inferencein natural popula�ons. Mol. Ecol. 7: 639–655.
Meagher, T.R. and Thompson, E. 1986. The rela�onship between single parent and parent pair gene�c likelihoods ingenealogy reconstruc�on. Theor. Popul. Biol. 29(1): 87–106.
Mizroch, S.A. and Rice, D.W. 2013. Ocean nomads: Distribu�onand movements of sperm whales in the North Paciﬁc shownby whaling data and Discovery marks. Mar. Mam. Sci. 29:
E136–E165.
Nielsen, R., Ma�la, D.K., Clapham, P.J. and Palsbøll, P.J. 2001.
Sta�s�cal approaches to paternity analysis in natural popula�ons and applica�ons to the north Atlan�c humpback whale.
Genetics 157: 1673–1682.
Ohashi, Y., Goto, M., Taguchi, M., Pastene, L.A. and Kitakado,
T. Evalua�on of a paternity method based on microsatellite
DNA genotypes for es�ma�ng the abundance of the Antarc�cminke whales, Balaenoptera bonaerensis, in the Indo-Paciﬁcregion of the Antarc�c. Cetacean Population Studies (inpress).
Palsbøll, P.J. 1999. Gene�c tagging: contemporary molecular
ecology. Biol. J. Linn. Soc. Lond. 68: 3–22.
Palsbøll, P.J., Allen, J., Berube, M., Clapham, P.J., Feddersen,
T.P., Hammond, P.S., Hudson, R.R., Jorgensen, H., Katona, S.,
Larsen, A.H., Larsen, F., Lien, J., Ma�la, D.K., Sigurjonsson, J.,
Sears, R., Smith, T., Sponer, R., Stevick, P. and Oien, N. 1997.
Gene�c tagging of humpback whales. Nature 388(6644):
767–769.
Paetkau, D. and Strobeck, C. 1994. Microsatellite analysis ofgene�c varia�on in black bear popula�ons. Mol. Ecol. 3:
489–495.
Thompson, E.A. 2013. Iden�ty by Descent: Varia�on in Meiosis,
Across Genomes, and in Popula�ons. Genetics 194(2):301–326.
Tiedemann, R., Tiedemann, M.R., Goto, M., Taguchi, M. andPastene, L.A. 2017. Finding parent-oﬀspring pairs among
western North Paciﬁc common minke whales. Paper SC/67a/
SDDNA1 presented to the IWC Scien�ﬁc Commi�ee,
May 2017 (unpublished). 18 pp. [Available from the IWC
Secretariat].
Urian, K., Gorgone, A., Read, A., Balmer, B., Wells, R.S., Berggren,P., Durban, J., Eguchi, T., Rayment, W. and Hammond, P.S.
2015. Recommenda�ons for photo-iden�ﬁca�on methods
used in capture-recapture models with cetaceans. Mar.
Mam. Sci. 31: 298–321.
Wade, P.R., Kennedy, A., LeDuc, R., Barlow, J., Carre�a, J.,
Shelden, K., Perryman, W., Pitman, R., Robertson, K., Rone, B.,
Salinas, J.C., Zerbini, A., Brownell, R.L. and Clapham, P.J. 2011.
The world's smallest whale popula�on? Biol. Lett. 7: 83–85.

Technical Reports of the Ins�tute of Cetacean Research (2020)

pp. 41–47

Technical Report (not peer reviewed)

Use of gene�c data for abundance es�mate purposes: a brief review
of methods and case studies by the Ins�tute of Cetacean Research
Megumi Takahashi*
Institute of Cetacean Research, 4–5 Toyomi-cho, Chuo-ku, Tokyo 104–0055, Japan*Contact e-mail: takahashi@cetacean.jp

ABSTRACT

Sigh�ng surveys by the line transect method is the most used approach to es�mate abundance in whales.
Sigh�ng surveys have some limita�ons associated with bad weather and sea condi�ons, and with the diﬃculty
to cover all areas of distribu�on of the stocks. Given these limita�ons, alterna�ve methods are being inves�gated for es�ma�ng the abundance of whales. This paper presents an overview of the current gene�c methods for abundance es�ma�on in whales. These methods are based on the mark-recapture (MR) approach andcan be divided into two groups: i) those based on the history of individual iden�ﬁca�on (genotyping basedon biopsy samples), and ii) those based on kinship informa�on including paternity analysis and close-kin MR(based on biopsy samples and/or catches). This paper also presents informa�on on case studies conducted by
the Ins�tute of Cetacean Research.

INTRODUCTION
Abundance es�ma�on is one of the most importantsources of informa�on required for the conserva�onand management of whales. The analysis of sigh�ng
data collected by the line transect (LT) method (Buckland
et al., 2001) is the most common approach for es�mating the abundance of whales. However, the use of thismethod has some limita�ons. For example, bad weather
and sea condi�ons can prevent the surveys, while someareas are inaccessible to the surveys (e.g., polynyas in the
Antarc�c or territorial waters of other countries).
The mark-recapture (MR) method (Hammond, 1986;
Pollock et al., 1990) based on Discovery-type tags isanother method used for abundance es�ma�on. However, this method depends on whaling opera�ons for therecovery of the tags, and this has been limited in recent
years. MR methods can be used for individual whales
iden�ﬁed by natural marks (photo-iden�ﬁca�on), however not all species have dis�nc�ve marks for individual
iden�ﬁca�on, and the method would not be prac�cal forabundant popula�ons.
Given these limita�ons, alterna�ve methods for abundance es�ma�on of whales are being explored. In recent
years, several methods for abundance es�ma�on basedon gene�c individual iden�ﬁca�on and kinship inference
have been reported. This paper presents an overview of
those methods and outlines some case studies conduct-

ed by the Ins�tute of Cetacean Research (ICR).

BASIC CONCEPT OF MARK-RECAPTURE
The MR method is widely used in ecology, par�cularly
to es�mate popula�on size and some biological parameters such as survival rates. The MR method relies on
sampling and re-sampling individual animals. A capture
history simply describes whether or not an animal was
captured in a series of sampling occasions during a periodof data collec�on. It is usually represented by a series of
1 (captured) and 0 (not captured). The key idea of the MR
method is to determine the ra�o of marked individuals
to unmarked individuals in the target popula�on, i.e.,es�mate p (capture probability) from the propor�on ofmarked animals that are recaptured. Figure 1 shows thesitua�on of marks and mark-recaptures in two samplingoccasions (two-sample es�mator).
In Figure 1, M is the number of individuals marked in a
popula�on in the ﬁrst year; c is the total number captured
in the second year (with and without a mark); R is thenumber of captured in the second year that are marked(recaptured); and N is the size of the popula�on at the�me of marking and release.
The simplest MR method is the ʻPetersen methodʼ also
known as ʻLincoln methodʼ (Seber, 1982):
R
M
=
.
c
N
41

Technical Reports of the Ins�tute of Cetacean Research (2020)

The propor�on of marked animals in the second �me
R/c is an es�mate of the probability of capturing individual (p̂), so that:
M
.
Nˆ =
pˆ

Chapmanʼs modiﬁca�on of Petersenʼs two-sample es�mator (Chapman, 1951) is more appropriate in situa�onsof small sample sizes, and is expressed as follows:N=(M +1)(c +1)
− 1.
R +1

Here,
M=number of individuals captured in the ﬁrst year;
c=number of individuals captured in the second year;
R=number of individuals captured in both years.
Mul�ple sampling occasions
In the case of the two-sample es�mator in Figure 1, theassump�on of ʻclosed popula�onʼ is adopted (births,
immigra�on, emigra�on and death are not considered).
For mul�ple sampling occasions involving a long periodof �me, the assump�on of ʻopen popula�onʼ is more appropriate, and immigra�on, emigra�on, birth and deathshould be considered (Figure 2).
In the case of mul�ple sampling occasions, the JollySeber (JS) model (Jolly, 1965; Seber, 1965), an open
popula�on model, is the most appropriate. This model
is very ﬂexible and, in addi�on, can provide es�mates ofsurvival, recruitment and popula�on growth rates.
There are a number of formula�ons based on the JS fores�ma�ng abundance and related parameters, such as
the ʻPOPAN,ʼ Link-Barker and Pradel recruitment, and the
Burnham JS and Pradel-λ formula�ons. In par�cular, thePOPAN formula�on (Schwarz and Arnason, 1996) can provide abundance and net births by using the full likelihood
approach. Net birth refers to all animals that enter a�ersampling occasion i and survive un�l sampling occasion i+1.

ABUNDANCE ESTIMATES BASED ON GENETIC
DATA
The main advantages of using gene�c tags for individual
iden�ﬁca�on and abundance es�mates based on MR are
that: i) they are permanent and ii) they exist in all individuals. It is important that the gene�c approach for individual iden�ﬁca�on and kinship inferences are reliable.
The gene�c basis for individual iden�ﬁca�on and kinshipinferences is shown in Taguchi (this issue).

42

Figure 1.

Diagram showing the basic idea of the MR method
based on individual iden�ﬁca�on of humpback
whales by photographic matching.

Methods based on the history of individual iden�ﬁca�on
The most common approach for gene�c individual iden�ﬁca�on is based on biopsy sampling and genotyping,based on a number of nuclear marker loci. One of themost used markers is microsatellite DNA. If two biopsy
samples, taken in diﬀerent occasions/loca�ons present
the same genotype, then it is assumed that those biopsies were taken from the same individual (see Taguchi,
this issue).
Some studies are presented here as examples ofabundance es�mates in whale popula�ons based on
biopsies, gene�c individual iden�ﬁca�on and the MR
methods. Palsbøll et al. (1997) conducted an analysis ofsix microsatellite loci for 3,060 skin samples collectedfrom humpback whales throughout the North Atlan�c.
They detected 2,368 unique genotypes and, because
the samples with iden�cal genotypes were of the same
sex, they considered that the 3,060 samples represented
2,368 individual whales. There was a total of 692 ʻrecaptures.ʼ The abundance and its conﬁdence intervals (CI)were es�mated based on the data collected in 1992 and1993 and the two-sample es�mators. The abundance
was es�mated at 4,894 (95% CI: 3,374–7,123) for males
and 2,804 (95% CI: 1,776–4,463) for females.
Based on 354 biopsy samples collected from the
New Zealand subantarc�c southern right whales during the austral winter from 1995 to 1998, Carroll et al.
(2011) were able to amplify 302 samples with 9–13 microsatellite loci. They iden�ﬁed 235 unique individuals
during these four winter surveys. They es�mated thesuperpopula�on abundance using the POPAN JS model,

Use of genetic data for abundance estimate purposes

Figure 2.

Diagram showing the situa�on of mul�ple sampling under the assump�on of open popula�on. A popula�on changes in
number and density due to births, deaths, immigra�on and emigra�on. Note that, φi represent the probability of (apparent)
survival un�l the next sampling occasion; pi is the probability of being captured and re-captured; Bi is the net number of
new individuals joining the study area (modiﬁed from h�p://www.phidot.org/so�ware/mark/docs/book/pdf/chap12.pdf ).

which provided an es�mate of 910 non-calf whales (95%
CI: 641–1,354) in 1998.
Methods based on kinship informa�on
An addi�onal advantage of DNA tags is that they contain
informa�on about kinship among individuals. Hence, kinship rela�onship can be used for abundance es�mates
based on the classic MR using ʻrecapture of selfʼ to ʻrecapture of closely-related kinʼ. Several studies used DNA
proﬁles to detect instances of paternity or other kinshiprela�onships in whale popula�ons (see Taguchi, thisissue). For this approach, samples obtained from biopsy
sampling as well as from catches can be used.
Paternity analysis
For the aim of paternity inference, Marshall et al. (1998)
derived likelihood ra�os with codominant markers taking account of typing error, and deﬁned a sta�s�c Δ for
resolving paternity. The likelihood ra�o can be wri�en as:
P ( D | H1 )L ( H1 , H 2 | D ) =,P( D | H 2 )In this formula�on, P(D|Hi) is the probability of obtaining data D, i.e., genotypes of oﬀspring, mother andpoten�al father at a par�cular locus, under hypothesis Hi(i=1, 2) . The hypothesis of H1is that the poten�al father
is the true father, and this is tested against hypothesis H2
wherein the poten�al father is an unrelated individual
selected at random from the popula�on.
Nielsen et al. (2001) presented a new method for paternity analysis in natural popula�ons based on genotypic
data that can take the sampling frac�on of puta�ve parents into account. They developed an approach for es�ma�ng parentage probabili�es for paternity assignment,
which is a Bayesian alterna�ve to the method developedby Marshall et al. (1998). They used the term ʻparentage
probabilityʼ to describe the posterior probability that apar�cular individual/puta�ve father might be the actual
father of a known oﬀspring. They applied the method togenotypic data (six microsatellite DNA loci) collected from
North Atlan�c humpback whales.
The method of Nielsen et al. (2001) is brieﬂy describedhere following the nota�ons of the authors. They notedthat the objec�ve was to es�mate the posterior probability that a par�cular individual might be the father ofa known oﬀspring. Let Ij (i) indicate the event that the jthpoten�al father is the father of the ith oﬀspring. The ithmaternal genotype is Mi, the associated genotype of theoﬀspring is Oi, the genotype of jth poten�al father is Fj,
and A is the matrix of allelic frequencies for all loci. If nof N males on the breeding ground were sampled, theposterior probability of paternity could be calculated as:Pr( I j (i ) | M i , F, A, N )=Pr(Oi | M i , Fj ).
n
 j=1 Pr(Oi | M i , Fj ) + ( N − n)Pr(Oi | M i , A)

In this formula�on Pr(Oi|Mi, Fj) is the shorthand nota�on for Pr(Oi|Mi, Fj, Ij(i)). ∑nj=1 Pr(Oi|Mi, Fj) refers to thesum of the probability that ith fetus has jth poten�alfather as itʼs father when ith maternal genotype and jth
poten�al father are given. Assuming Mendelian segrega�on and independence among loci, the probability of anobserved oﬀspring genotype, given the maternal genotype and the genotype of a par�cular poten�al father
Pr(Oi|Mi, Fj), could be calculated using standard methods(e.g., Thompson, 1975; 1976). Likewise, Pr(Oi|Mi, A)
can be calculated assuming Hardy-Weinberg equilibrium and independence among loci (linkage equilibrium).
(N−n)Pr(Oi|Mi, A) refers to the probability of the ith
43Technical Reports of the Ins�tute of Cetacean Research (2020)

fetusʼs genotype given the ith motherʼs genotype. In es�ma�ng the posterior probability that a poten�al father isthe father of an oﬀspring, assump�ons need to be maderegarding the prior probability of a poten�al father being
the father. In the absence of other informa�on, it is assumed that the prior probability that a par�cular maleis the father is 1/N, where N is the number of poten�ally
breeding males in the breeding area (Nielsen et al., 2001).
Here, assuming independence among oﬀspring, thelikelihood func�on for N can be calculated as:
L( N ) ∝∏ Pr(O | M , F, A, N )
i

i

i

=

1
 ( N − n)Pr(Oi | M i , A) +
N
N

∏ 
i



n
j=1


Pr(Oi | M i , F j )  ,


N≥ n.

The maximum-likelihood es�mate of N and the conﬁdence interval is provided, for example, using parametricbootstrapping. Note that the es�mate based on this approach such as parent-oﬀspring genotypes is an es�mateof the current (i.e., at the �me of sampling) male popula�on size. To es�mate the total abundance of the target
popula�on/species, addi�onal informa�on is required.
For example, the propor�ons of males and females, andthe sexual maturity of whales from the sample, under theassump�on that the sample was representa�ve of thetrue popula�on.
Based on the method above, the maximum-likelihoodes�mate of N was 6,540 breeding male humpback whales(95% CI: 3,700–17,000) using parametric bootstrapping(Nielsen et al., 2001).
CKMR method
CKMR is a recent extension of the ordinary MR methods
used to es�mate animal abundance and other popula�onparameters. The CKMR approach was ﬁrst described bySkaug (2001) working on microsatellite data (10 loci) of
North Atlan�c common minke whales. He derived an es�mator for the total popula�on size based on the numberof parent-oﬀspring pairs (POPs).
Subsequently Bravington et al. (2016a) extended theclassic MR framework (Lincoln-Petersen type es�mator
mainly) by incorpora�ng close-kin informa�on. Whereas
ordinary MR methods only consider the subsequentiden�ﬁca�on of the same animal as a recapture, CKMR
expands this by also viewing the gene�c iden�ﬁca�on ofa rela�ve (e.g., parent-oﬀspring, full-sibling, half-sibling)
as a recapture.
Below is the simplest version of the CKMR by Bravington et al. (2016a). This is a two-sample es�mator for the44

adult popula�on size, which is the close-kin version of the
Lincoln-Petersen popula�on size es�mator:
 = 2 nJ n A .
(1)
N
A
H
Consider a sample of nJ juveniles and nA adults fromthe same popula�on. Each sampled juvenile can be considered a ʻmarkʼ of its two adult parents. The genotypeof each of the nJ juvenile samples is compared to each of
the nA adult samples, to check if a ʻmarkʼ is recaptured.
The probability that the adult happens to be one of thejuvenileʼs two parents is 2/NA, where NA is adult popula�on size. The expected number of parent-oﬀspring pairs
across all nA·nJ comparisons is then 2nA nJ/NA. Equa�ng
this to the observed number of parent-oﬀspring pairs
gives the CKMR version of the Lincoln-Petersen es�mator. Hence, if the en�re set of nJ ×nA comparisons yields H
which is the number of parent-oﬀspring pairs, then adultabundance can be es�mated as eq. (1).
In a more realis�c se�ng, some types of kinship probabili�es are likely to be present, and therefore, an explicitsta�s�cal MR model is required. That is, the prior probability that a pair is a POP is set by a popula�on dynamics
model, which accounts for pair-speciﬁc data, and which
includes demographic parameters such as adult abundance, individual survival probability, age of maturity, percapita birth rate and probably age-speciﬁc fecundi�es.
For example, probability that individual i with covariates zi (date of capture for example) is the mother ofindividual j with covariates zi is represented as:

P (i is j ′s mother| zi , z j ) i′s reproductive output in j ′s year of birth = E 
Total female reproductive output
in j ′s year of birth| zi , z jwhere E() is the expected value and the absolute abundance (of females) now enters implicitly through thedenominator.
Not only parent-oﬀspring (PO) but also other kinshiprela�onships such as half-sibling can be considered in the
CKMR. A pairwise comparison between individuals i andj gives rise to a kinship category Kij taking values from a
discrete set k={PO, HS, . . .} (see also Taguchi (this issue)).
These probabili�es are the building blocks of the pseudolikelihood shown below.
Demographic parameters can be es�mated from thelog-likelihood summed over all pairwise comparisons.
The joint distribu�on of {Kij; 1 ≤ i < j ≤ n} is too complicated to permit the construc�on of a full likelihood. Instead,a pseudo-likelihood approach (Skaug, 2001; Bravington
et al., 2016a) which involves only the marginal probabili-

Use of genetic data for abundance estimate purposes�es of the Kij in eq. (2) was used.
lP (θ ) = ∑1 ≤ i < j ≤ n log P ( K ij = kij | zi , z j ; θ ) .

(2)

Let θ denote the vector of all parameters that govern
the basic quan��es, for example, total popula�on size
(males and females): N, individual survival probability: φ,per capita average birth rate: β and capture probability at
a given sampling occasion: p).
The CK greatly expands the scope of MR because avariety of samples can be used, including biopsy samples
and samples from catches. However, age informa�on isessen�al for precise CKMR es�mates because it iden�ﬁes the �me of ʻmarkingʼ, which is the year the sampled
animal was born.
Bravington et al. (2016b) used the approach above forthe case of the southern blueﬁn tuna, using microsatellite
data at 20–25 loci.

CASE STUDIES BY THE ICR
Based on the histories of individual iden�ﬁca�on
Pastene et al. (2018) examined a total of 157 biopsy samples of southern right whales collected by IDCR/SOWER
and JARPA/JARPAII surveys in austral summer between
1987/88 to 2013/14 in the longitudinal sector between
80°E and 135°E (Figure 3). These southern right whales
are associated with the South West Australia popula�on.
Biopsy samples were examined with 14 microsatellite
DNA loci. A�er removing four duplicates, the sample
sizes for the individual iden�ﬁca�on became 153. Eight
incidences of individual matching (ʻmark-recaptureʼ)
were detected (four males and four females). Individual
matching by mul�locus genotypes was supported by mitochondrial DNA analyses (same haplotype), sex determina�on (same sex), and in two cases where pictures wereavailable, by photo-iden�ﬁca�on. Es�mate of abundance

Figure 3.

Geographical distribu�on of southern right whales
in Area IV examined by Pastene et al. (2018). Red:
females; Blue: males.

was based on the POPAN model implemented in RMark(Laake, 2013). This preliminary analysis considered theassump�on of constant apparent survival, constantprobability of capture and constant probability of entry
through the years.
The results of the preliminary analyses showed similar annual abundance es�mates to those obtained bysigh�ng data in the same area and a similar period. For
example, the es�mate of abundance by the gene�cʻmark-recaptureʼ method was 1,619 (95% CI: 868–3,151)
individuals for 2015/16, similar to the most recent(2007/08) sigh�ng survey abundance es�mate of 1,557(95% CI: 871–2,783) individuals using the LT method in
the same area.
Future analyses
The analysis conducted on southern right whales assumed constant apparent survival rate, constant capture
probability, and constant probability of entry throughthe years. Some addi�onal inves�ga�ons are required toevaluate the eﬀect of changing these assump�ons on theabundance es�mates. State-space models and hierarchical Bayesian approach have already been proposed as aconvenient and ﬂexible framework for specifying stochas�c models for the dynamics of wild animal popula�ons(Gimenez et al., 2009; Kéry and Schaub, 2011; Hendrix
et al., 2012), and these models could be used for southern right whales in the future. Such models consider individual diﬀerences of the survival rate. Furthermore, thedetec�on/capture probability, as a random eﬀect can beevaluated by the state-space model.
Gene�c databases for Antarc�c blue and humpback
whales are available at the ICR, and the abundance es�ma�on approach for southern right whales can also be
used for these other species.
One issue with mark-recapture analyses, regardlessof which method of individual iden�ﬁca�on is used, isthat the study area must provide adequate opportuni�es for ʻcapturingʼ all individuals within a popula�on.
However, this is hardly feasible because many popula�ons are widely and unevenly distributed. Therefore, forobtaining more accurate abundance es�mates, applying
analy�cal techniques that take account of unsampledloca�ons are also suggested for future analyses (e.g., Frasier et al., 2020). Also, more informa�on on the biologicalfeatures of the species, such as migra�on and habitat use
pa�erns, are required.
Paternity analyses
Kanda et al. (2014) conducted a study on paternity in45

Technical Reports of the Ins�tute of Cetacean Research (2020)

Antarc�c minke whales based on samples collected by
JARPA and JARPAII surveys. Speciﬁcally, genotypic data ofa maximum of 12 microsatellite DNA loci were preparedfrom 137 fetuses collected from females captured during
the 2003/04 JARPA survey, and their poten�al fathers
were sought among 1,779 males collected from 2001/02
JARPA to 2010/11 JARPAII surveys.
One mother-fetus-father trio was detected using
CERVUS (Marshall et al., 1998), a computer program forassignment of parents to their oﬀspring using gene�cmarkers. This single match was used to tenta�vely es�mate the mature male number in the Eastern Indian
Ocean Stock (I stock) of Antarc�c minke whales using thePetersen mark-recapture method modiﬁed by Chapman.
Only mature males (n=677) (sexual maturity criteria ofKato, 1987) were used in the es�mates.
The abundance es�mate of the mature male component of the stock was 46,782 (CV=0.572) animals.
Assuming the mature/immature ra�o and the sex ra�owere 1 : 1, respec�vely, the total stock size was es�matedas 187,128 animals. This es�mate was higher than the
JARPA/JARPAII (51,474) and the IDCR (151,174) sigh�ng
surveys es�mates in similar areas. Kanda et al. (2014)
noted that their analysis es�mated total stock size, while
the JARPA/JARPAII and IDCR analyses es�mated the number of individuals distributed in the research area basedon sigh�ng surveys, which would explain the diﬀerences.
Ohashi et al. (in-press) further examined the popula�on size of mature males of Antarc�c minke whales
based on paternity analysis. Their analyses were basedon biological and gene�c (microsatellite DNA at 12 loci)data collected by JARPAII in the Indo-Paciﬁc region of the
Antarc�c. A total of 2,126 Antarc�c minke whales taken in
the austral seasons 2006/07, 2008/09, 2009/10, 2010/11
and 2011/12 were used in the analyses. Two assump�ons
on stock structure were considered, a single stock in the
research area and two stocks in the research area (the I
stock and the Western South Paciﬁc Stock (P stock)).
For abundance es�ma�on, the approach of Nielsen
et al. (2001) outlined above was used. For the hypothesisof a single stock, the abundance of mature males wases�mated as 68,874 (90% CI: 42,625–122,779). Using the
propor�on of male and immature whales of 0.436 and
0.254, respec�vely, the es�mate of total popula�on was
211,600 (90% CI: 130,953–377,210). This es�mate waslower than that from the IDCR/SOWER sigh�ng surveys(413,202) and of the same order of JARPA/JARPAII es�mates in Areas IV and V (263,711).

46

Future analyses
Ohashi et al. (in-press) reported that there are some technical issues in their analyses on Antarc�c minke whales
that should be improved in the future. Firstly, there is aneed to op�mize the use of the available gene�c data andsolve issues, for example, on genotyping errors and missing allele informa�on for some loci. The second issue isrelated to the assump�ons of the stock structure hypothesis. The current hypothesis on stock structure of Antarc�c minke whales suggests a transi�on sector shared by
the two stocks rather than a hard longitudinal boundary
between them as assumed in the paternity analysis. Thistransi�on sector should be considered in future paternity
analyses.
In addi�on, Ohashi et al. (in-press) noted that future
analyses should consider not only the rela�onship between mother/fetus and father, but also some other kinship rela�onships (e.g., half-sibling) following the work of
Bravington et al. (2016a) outlined above. Therefore, thenext challenge for the ICR research on this topic is the applica�on of the CKMR method for abundance es�ma�on,which op�mizes the use of the available gene�c data.
Comparison of abundance between MR and LT methods
As noted by Kanda et al. (2014) and Pastene et al. (2018),es�mates of abundance, using the ordinary MR approachbased on gene�c data, refer to the total popula�on/stock
size. The es�mates based on sigh�ng data under the LT
approach, refer to the abundance of animals in a speciﬁcarea and a speciﬁc period of a survey. Then, the diﬀerence in es�mates will depend on the sigh�ng survey coverage and on the movement of individuals into and out of
the sigh�ng survey area (Hammond, 2010).

ACKNOWLEDGEMENTS
I thank T. Hakamada (ICR) for technical assistance on themethodology outlined in this paper, and L.A. Pastene(ICR) for his assistance in preparing this report.

REFERENCES
Bravington, M.V., Skaug, H.J. and Anderson, E.C. 2016a.
Close-kin mark-recapture. Stat. Sci. 31: 259–274.
Bravington, M.V., Grewe, P.M. and Davies, C.R. 2016b. Absoluteabundance of southern blueﬁn tuna es�mated by close-kin
mark-recapture. Nat. Commun. 7: 1–8.
Buckland, S.T., Anderson, D.R., Burnham, K.P., Laake, J.L.,
Borchers, D.L. and Thomas, L. 2001. Introduction to DistanceSampling: Estimating Abundance of Biological Populations.
Oxford University Press, Oxford. 432 pp.
Chapman, D.G. 1951. Some proper�es of the hypergeometricUse of genetic data for abundance estimate purposes
distribu�on with applica�ons to zoological sample censuses.
University of California Publications in Statistics 1: 131–160.
Carroll, E.L., Patenaude, N.J., Childerhouse, S.J., Kraus, S.D.,
Fewster, R.M. and Baker, C.S. 2011. Abundance of the New
Zealand subantarc�c southern right whale popula�on es�mated from photo-iden�ﬁca�on and genotype mark-recapture. Mar. Biol. 158: 2565–2575.
Frasier, T., Petersen, S., Postma, L., Johnson, L., Heide-Jørgensen,
M.P. and Ferguson, S. 2020. Abundance es�ma�on from gene�c mark-recapture data when not all sites are sampled: An
example with the bowhead whale. Glob. Ecol. Conserv. 22:
e00903.
Gimenez, O., Bonner, S.J., King, R., Parker, R.A., Brooks, S.P.,
Jamieson, L.E., Grosbois, V., Morgan, B.J.T. and Thomas,
L. 2009. WinBUGS for popula�on ecologists: Bayesianmodeling using Markov Chain Monte Carlo methods. pp.
883–915. In: D.L. Thomson, E.G. Cooch and M.J. Conroy (eds.)
Modeling Demographic Processes in Marked Populations.
Environmental and ecological sta�s�cs 3. Springer, New York.
1132 pp.
Hammond, P.S. 1986. Es�ma�ng the size of naturally marked
whale popula�ons using capture-recapture techniques. Rep.
int. Whal. Commn (special issue) 8: 252–282.
Hammond, P.S. 2010. Es�ma�ng the abundance of marine
mammals. pp. 42–67. In: I.L. Boyd, W.D. Bowen and S. Iverson(eds). Marine Mammal Ecology and Conservation: a handbook of techniques. Oxford University Press, New York. 431
pp.
Hendrix, A.N., Straley, J., Gabriele, C.M. and Gende, S.M. 2012.
Bayesian es�ma�on of humpback whale (Megaptera novaeangliae) popula�on abundance and movement pa�erns insoutheastern Alaska. Can. J. Fish. Aquat. Sci. 69: 1783–1797.
Jolly, G.M. 1965. Explicit es�mates from capture-recapture
data with both death and immigra�on-stochas�c model.
Biometrika 52: 225–247.
Kanda, N., Goto, M. and Pastene, L.A. 2014. Paternity analysis
on Antarc�c minke whales using JARPA and JARPAII samples.
Paper SC/F14/J07 presented to the IWC SC JARPAII Review
Workshop, February 2014 (unpublished). 4 pp. [Available
from the IWC Secretariat].
Kato, H. 1987. Density dependent changes in growth parameters of the southern minke whale. Sci. Rep. Whales Res. Inst.
38: 47–73.
Kéry, M. and Schaub, M. 2011. Bayesian Population Analysis
Using WinBUGS: A Hierarchical Perspective. Academic Press,

Waltham, Massachuse�s. 554 pp.
Laake, J.L. 2013. RMark: An R Interface for analysis of capturerecapture data with MARK. AFSC Processed Rep. 2013-01, 25
p. Alaska Fish. Sci. Cent., NOAA, Natl. Mar. Fish. Serv., 7600
Sand Point Way NE, Sea�le WA 98115.
Marshall, T.C., Slate, J., Kruuk, L.E.B. and Pemberton, J.M. 1998.
Sta�s�cal conﬁdence for likelihood-based paternity inferencein natural popula�ons. Mol. Ecol. 7: 639–655.
Nielsen, R., Ma�la, D.K., Clapham, P.J. and Palsbøll, P.J. 2001.
Sta�s�cal approaches to paternity analysis in natural popula�ons and applica�ons to the north Atlan�c humpback whale.
Genetics 157: 1673–1682.
Ohashi, Y., Goto, M., Taguchi, M., Pastene, L.A. and Kitakado,
T. Evalua�on of a paternity method based on microsatellite
DNA genotypes for es�ma�ng the abundance of Antarc�cminke whales (Balaenoptera bonaerensis) in the Indo-Paciﬁcregion of the Antarc�c. Cetacean Population Studies (inpress).
Palsbøl, P.J., Allen, J., Bérubé, M., Clapham, P.J., Feddersen,
T.P., Hammond, P.S., Hudson, R.R., Jørgensen, H., Katona, S.,
Larsen, A.H., Larsen, F., Lien, J., Ma�la, D.K., Sigurjónsson, J.,
Sears, R., Smith, T., Sponer, R., Stevick, P. and Øien, N. 1997.
Gene�c tagging of humpback whales. Nature 388: 767–769.
Pastene, L.A., Hakamada, T., Acuña, P., Taguchi, M., Goto, M.,
Matsuoka, K. and Nishiwaki, S. 2018. Site-ﬁdelity and movement ranges of southern right whales in Antarc�c Area IV inferred from gene�c tagging. Paper SC/67B/SH/06 presented
to the IWC Scien�ﬁc Commi�ee, May 2018 (unpublished). 10
pp. [Available from the IWC Secretariat].
Pollock, K.H., Nichols, J.D., Brownie, C. and Hines, J.E. 1990.
Sta�s�cal inference for capture-recapture experiments.
Wildl. Monogr. 6: 288–295.
Schwarz, C.J. and Arnason, A.N. 1996. A general methodology
for the analysis of capture-recapture experiments in open
popula�ons. Biometrics 52: 860–873.
Seber, G.A.F. 1965. A note on the mul�ple-recapture census.
Biometrika 52: 249–259.
Seber, G.A.F. 1982. The estimation of animal abundance andrelated parameters. 2nd Edition. Griﬃn, London. 654 pp.
Skaug, H.J. 2001. Allele-sharing methods for es�ma�on of popula�on size. Biometrics 57: 750–756.
Thompson, E.A. 1975. The es�ma�on of pairwise rela�onships.
Ann. Hum. Genet. 39: 173–188.
Thompson, E.A. 1976. Inference of genealogical structure. Soc.
Sci. Inform. 15: 477–526.

47

Technical Reports of the Ins�tute of Cetacean Research (2020)

pp. 48–53

Technical Report (not peer reviewed)

Progress in the research on earplug-based age determina�on andbiological parameters of North Paciﬁc sei whales at the Ins�tute of
Cetacean Research
Takeharu Bando1, * and Hikari Maeda2
1

Institute of Cetacean Research, 4–5 Toyomi-cho, Chuo-ku, Tokyo 104–0055, Japan
Japan Fisheries Research and Education Agency, 2–12–4 Fukuura, Kanazawa-ku, Yokohama-shi, Kanagawa
236–8648, Japan

2*Contact e-mail: bando@cetacean.jp

ABSTRACTThis study summarizes the progress in the research on earplug-based age determina�on and es�ma�on of
biological parameters of the western North Paciﬁc sei whales sampled by the JARPNII/NEWREP-NP surveys
between 2002 and 2018. A total of 1,622 whales (744 males and 878 females) were sampled in the western
North Paciﬁc north of 35°N and between the Paciﬁc coast of Japan and 170°E. Earplugs were read for whales
collected up to 2016 (n=1,354). Age informa�on was obtained for 887 whales (65.5%). Earplug readability was
higher for mature (69.7%) than immature (52.1%) whales, and no diﬀerence between sexes was observed.
The growth curves for males and females were Lt =14.14 (1−e−0.174(t+6.650)) and Lt =15.17 (1−e−0.150(t+7.407)),
respec�vely. The age at sexual maturity for males and females were 6.7 (SE=0.29) and 6.9 (SE=0.27) years,
respec�vely. The body length at sexual maturity was es�mated as 12.72 m (SE=0.12) for males and 13.31 m(SE=0.12) for females. The annual ovula�on rate was es�mated as 0.744/year. Substan�al biological informa�on was obtained for the North Paciﬁc sei whale during the 17 year of JARPNII and NEWREP-NP. The analysesof biological data will contribute to the management of this whale species in the North Paciﬁc.

INTRODUCTION
Sei whale (Balaenoptera borealis) is widely distributed
throughout the world. As with other baleen whale species, they undertake seasonal migra�on spending thesummer feeding season in subpolar waters and returning
to low la�tude waters to calve in winter (Horwood, 2017).
In the North Paciﬁc, sei whales (Figure 1) distributewidely in the oﬀshore area associated with the produc�ve polar and subarc�c fronts during the summer feeding
season (Murase et al., 2014).
The second phase of the Japanese Whale Research
Program under Special Permit in the western North
Paciﬁc (JARPNII) started in 2000 (GOJ, 2000; 2002), andwas completed in 2016. The sei whale was added as
one of the target species for sampling in 2002. The New
Scien�ﬁc Whale Research Program in the western North
Paciﬁc (NEWREP-NP) started in 2017 (GOJ, 2017) and also
had the sei whale as one of the target species. During theannual surveys, biological data and samples such as bodylength measurements and reproduc�ve organs, wereobtained systema�cally. A total of 1,622 sei whales were48

sampled during a 17 year survey period. The NEWREP-NP
was completed in 2018.
Age data is one of the most important informa�on forstock assessment and management of large whales. Agees�ma�on based on coun�ng of growth layers accumulated in the earplugs is considered the most reliable tool
for age determina�on in baleen whales (Lockyer, 1984a;

Figure 1.

A sei whale feeding on plankton near the sea surface
in the western North Paciﬁc.

Age determina�on and biological parameters of North Paciﬁc sei whales at the Ins�tute of Cetacean Research

Maeda et al., 2016). During JARPNII and NEWREP-NP, earplugs were collected from each whale sampled by experienced biologists in order to increase the readability. Someage-related biological parameters are known to change in
response to changes in abundance, food availability orcompe��on among whale species (Gambell, 1973; Katoand Sakuramoto, 1991; Lockyer, 1984b). Therefore, monitoring of biological parameters is indispensable for theassessment and management of whale stocks.
This study summarized the progress in the researchon earplug-based age determina�on and es�ma�on of
the biological parameters of the western North Paciﬁc sei
whales sampled by the JARPNII and NEWREP-NP surveys
between 2002 and 2018.

est 1 cm, from the �p of the snout to the notch of the
ﬂukes using a stainless measure. Sexual maturity in males
was determined by examining the histological samples
from the tes�s. Males with seminiferous tubules over
100 µm diameter, sperma�d or open lumen in the tubules
were determined to be sexually mature (Masaki, 1976;
Lockyer, 1984b). Maturity of females was preliminarily
determined in the ﬁeld by the presence or absence of at

MATERIALS AND METHODS
Whale sampling
A total of 1,622 (744 males and 878 females) sei whales
were sampled by JARPNII and NEWREP-NP during
2002–2018. The geographical distribu�on of the sampledwhales is shown in Figure 2. Table 1 shows the number ofsampled whales by year and sexual maturity status.
Figure 2.

Biological data
Body length was measured in a straight line to the near-

Research area of JARPNII and NEWREP-NP andsigh�ng posi�ons of sei whales sampled during the
2002–2018 surveys.

Table 1
Number of sei whales sampled by JARPNII and NEWREP-NP in the western North Paciﬁc in the period 2002–2018, by year, sex andsexual maturity status.
Year

Male

Female

Total

Immature

MatureUnknown

Total

Immature

Mature

Total

2002
2003
2004
2005
2006
2007
2008
2009
2010
2011
2012
2013
2014
2015
2016
2017
2018

3
4
11
6
18
21
6
10
12
8
12
15
7
9
10
13
12

12
19
36
45
30
33
38
36
31
46
32
29
31
20
28
50
50

0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1

15
23
47
51
48
54
44
46
43
54
44
44
38
29
38
63
63

4
4
14
14
14
16
17
9
10
13
11
6
11
15
16
15
17

20
23
39
35
38
30
39
45
47
28
45
50
41
46
36
56
54

24
27
53
49
52
46
56
54
57
41
56
56
52
61
52
71
71

39
50
100
100
100
100
100
100
100
95
100
100
90
90
90
134
134

Total

177

566

1

744

206

672

878

1622
49

Technical Reports of the Ins�tute of Cetacean Research (2020)

Tm = g − 0.5 +var(tm ) =hBisected earplug surface of sei whale with light anddark laminae of growth layer groups (GLGs). The
scale shows a 1 mm interval.

least one corpus luteum or corpus albicans in both sidesof the ovaries, and conﬁrmed later in the laboratory bycoun�ng number of corpora.
Earplug sampling and age determina�on
Earplugs were collected from all sampled animals, following the method developed for baleen whale (Omura,
1963; Maeda et al., 2016). The le� and right earplugs
were collected and immediately ﬁxed in 10% formalinun�l age determina�on. In the laboratory, the surface
along the central axis of the earplug was cut using a sharp
blade, then it was placed on a wet stone to expose theneonatal line and growth layers (Figure 3). Growth layers
were counted under water using stereoscopic microscope. A growth layer group (GLG) was deﬁned as one
pair of light and dark laminae in the core and considered
as one year of age.
Es�ma�on of biological parameters
Growth curve
To es�mate growth curve, the von Bertalanﬀy growthmodel was ﬁ�ed to the body length and age as:

Lt = L∞ (1 − e− K (t −t0 ) )where
Lt is the body length at age t,
L ∞ is the asympto�c length,
K is the growth rate coeﬃcient, andt0 is the theore�cal �me at zero length.
Age at sexual maturity
Age at sexual maturity (tm) was es�mated by the following equa�on (Cooke, 1984).

50

 Ia 

a=ga  N 
M a Ia N ( N − 1)a=g

Figure 3.

h

2
a

a

where
Ma is the number of mature animals in age a,
Ia is the number of immature animals in age a,
Na is the total number of animals in age a,
g is the age of the youngest mature animal in the sample,
andh is the age of the oldest immature animal in the sample.
Body length at sexual maturity
Body length at sexual maturity (Lm) was es�mated by
the following equa�on (Cooke, 1984; Kato, 1992). Whencalcula�ng, body length data was rounded to the nearest
0.1 m.
Lm = j − 0.05 + 0.1
var(lm ) = qkk Ib b= jb

  N 

M b Ib N ( N − 1)
b= j

2
b

b

where
Mb is the number of mature animals in body length b,
Ib is the number of immature animals in body length b,
Nb is the total number of animals in body length b,j is the body length of the smallest mature animal in thesample, andk is the body length of the largest immature animal in thesample.
Annual ovulation rate
The annual ovula�on rate was es�mated by applying
linear regression analysis between age and the total number of corpora (corpora lutea and albican�a). The regression line was ﬁ�ed to ages 14 years and older because
all females were sexually mature at the age of 14 years
in this study.

RESULTS AND DISCUSSIONSAge readability
Earplugs were read for whales collected up to 2016
(n=1,354). Age informa�on was obtained for 887 whales(65.5%). The readability of earplugs varied dependingon the maturity status. Readability of sexually immatureindividuals was 55.9% for males and 48.9% for females(Table 2). Readability of earplugs in sexually mature animal was higher, 72.5% and 67.4% for males and females,

Age determina�on and biological parameters of North Paciﬁc sei whales at the Ins�tute of Cetacean Research
Table 2
Earplug-age readability of sei whales sampled during 2002–2016 surveys, by sex and sexual maturity status.
Maturity
statusMale

FemaleNumber ofwhalesReadable
earplugs

Immature
Mature

152
466

85
338

55.9
72.5

174
562

85
379

48.9
67.4

Total

618

423

68.4

736

464

63.0

Figure 4.

Readability Number of(%)
whalesReadable
earplugs

Total
Readability Number of(%)
whalesReadable
earplugs

Readability
(%)

326
1028

170
717

52.1
69.7

1354

887

65.5

Rela�onship between body length and age in sei whales. The solid lines show the von Bertalanﬀy growth
curves.

respec�vely (Table 2). Readability of all samples was
65.5%, which was higher than for western North Paciﬁc
common minke whales (44.1%) (Maeda et al., 2016), andcomparable to Brydeʼs whales (65.2%) (Bando, 2018),
which were based on samples collected by JARPN and
JARPNII.
Growth curve
For both sexes, the growth rate was high at younger ages
and stabilized a�er 15 years (Figure 4). The following vonBertalanﬀy growth curves were es�mated:

Lt = 14.14 (1 − e −0.174(t+6.650) )Male :
Female : Lt = 15.17 (1 − e −0.150(t+7.407) )Masaki (1976) es�mated growth curves of sei whales
based on samples collected from the commercial whaling
in the North Paciﬁc in the 1970s. However, the es�matedcurve was incomplete due to the lack of young individuals. During JARPNII and NEWREP-NP, samples from a wide
range of ages, including young individuals, were collected, and for the ﬁrst �me a reasonable growth curve
could be obtained for western North Paciﬁc sei whales.
Age at sexual maturity
Sexually mature males ﬁrst appeared at the age of 4 years,

and from 13 years old onwards all animals were sexually mature (Figure 5). Tm for males was es�mated as 6.7
years (SE=0.29). For females, sexually mature whales ﬁrst
appeared at the age of 3 years, and from 14 years old onwards all animals were sexually mature (Figure 5). Tm wases�mated as 6.9 years (SE=0.27). Masaki (1976) reported
the Tm of sei whales from the commercial whaling in the
North Paciﬁc in the 1970s as 2.5 and 6.5 years for males
and females, respec�vely. However, these values were
underes�mated as the ages of younger animals are o�enes�mated as less than their actual ages. This is because
the so�er and smaller earplug of young animals, together
with a lack of neonatal layers, in some cases, make it difﬁcult to recognize growth layers (Masaki, 1976).
Body length at sexual maturity
The rela�onship between sexual maturity and bodylength is shown in Figure 6. Sexually mature male ﬁrst
appeared at body length of 12.2 m, and from 13.6 m onwards all animals were sexually mature. Lm was es�matedas 12.72 m (SE=0.12). For females, sexually mature
whales ﬁrst appeared at the body length of 12.6 m, andfrom 14.0 m all animals were sexually mature (Figure 6).
Lm was es�mated as 13.31 m (SE=0.12). Masaki (1976)es�mated Lm of male sei whales as 12.9 m based on the51

Technical Reports of the Ins�tute of Cetacean Research (2020)

Figure 5.

Sexual maturity status by age and sex in sei whales. Numbers in the bars show the numbers of samples
examined.

Figure 6.

Sexual maturity status by body length and sex for sei whales. Numbers in the bars show the numbers ofsamples examined.

commercial whaling samples from the North Paciﬁc,which was slightly higher than the value es�mated in thisstudy. However, Masaki (1976) determined the sexual
maturity of males by tes�s weight rather than histologicalobserva�on, which might have inﬂuenced the results. For
females, a similar value was es�mated from commercial
whaling samples (13.4 m) (Masaki, 1976).
Annual ovula�on rate
The corpus luteum and albicans ﬁrst appeared at the ageof 3 years and the number of corpora increased linearlya�er the age of 14 years (Figure 7). Annual ovula�on rate
was es�mated as 0.744. The es�mated value means thatthe majority of sei whales give birth (or ovula�on occurs)
three �mes in four years. This value was higher than
0.604 which was es�mated from the 1970s commercial
whaling samples (Masaki, 1976).

CONCLUDING REMARKS
Substan�al biological informa�on was obtained for the
North Paciﬁc sei whale during the 17 years of JARPNII and
NEWREP-NP. In par�cular, more age data from young in52

Figure 7.

Ji�er plot showing the rela�onship between age
and number of corpora in sei whales. The linear
regression line was ﬁ�ed to ages 14 years and over.

dividuals were obtained, resul�ng in improved es�matesof biological parameters than those obtained in the period of commercial whaling. The analyses of biological data

Age determina�on and biological parameters of North Paciﬁc sei whales at the Ins�tute of Cetacean Research

will contribute to the management of this whale species
in the North Paciﬁc.

ACKNOWLEDGEMENTS
Our sincere thanks to all researchers and crews who collected data and samples during the JARPNII and NEWREPNP surveys. We thank the members of the Ins�tute of
Cetacean Research (ICR) and Japan Fisheries Researchand Educa�on Agency for useful sugges�ons as well as assistance in the laboratory work. H. Kato (Tokyo Universityof Marine Science and Technology and ICR) and Yoshihiro
Fujise (ICR) encouraged us to conduct this study and provided valuable ideas for the analyses.

REFERENCES
Bando, T. 2018. Earplug-based age determina�on and es�ma�on of biological parameters in North Paciﬁc Brydeʼs whales
based on samples collected by JARPNII. Technical Reports of
the Institute of Cetacean Research (TEREP-ICR) No. 2: 32–37.
Cooke, J.C. 1984. The es�ma�on of mean ages at sexual maturity from age sample. Paper SC/36/O22 presented to
IWC Scien�ﬁc Commi�ee, May 1984 (unpublished). 8 pp.
[Available from the IWC Secretariat].
Gambell, R. 1973. Some eﬀects of exploita�on on reproduc�onin whales. J. Reproduct. Fert. (Suppl.) 19: 533–553.
Government of Japan. 2000. Research plan for cetacean studies
in the western North Paciﬁc under special permit (JARPNII)(Feasibility study plan for 2000 and 2001). Paper SC/52/O1
presented to the IWC Scien�ﬁc Commi�ee, June 2000 (unpublished). 68 pp. [Available from the IWC Secretariat].
Government of Japan. 2002. Research plan for cetacean studies
in the western North Paciﬁc under special permit (JARPNII).
Paper SC/54/O2 presented to the IWC Scien�ﬁc Commi�ee,
April 2002 (unpublished). 115 pp. [Available from the IWC
Secretariat].

Government of Japan. 2017. Revised research plan for New
Scien�ﬁc whale Research Program in the western North
Paciﬁc NEWREP-NP). Paper SC/67a/SCSP10 presented to the
IWC Scien�ﬁc Commi�ee, May 2017 (unpublished). 147 pp.
[Available from the IWC Secretariat].
Horwood, J. 2017. Sei whale Balaenoptera borealis. pp. 845–
847. In: B. Würsig, J.G.M. Thewissen and K. Kovacs (eds.)
Encyclopedia of Marine Mammals, Third Edition. Academic
Press, San Diego. 1190 pp.
Kato, H. 1992. Body length, reproduc�on and stock separa�onof minke whales oﬀ northern Japan. Rep. int. Whal. Commn
42: 443–453.
Kato, H. and Sakuramoto, K. 1991. Age at sexual maturity ofsouthern minke whales: a review and some addi�onal analyses. Rep. int. Whal. Commn 41: 331–337.
Lockyer, C. 1984a. Age determina�on by means of the ear plugin baleen whales. Rep. int. Whal. Commn 34: 692–696.
Lockyer, C. 1984b. Review of baleen whale (Mys�ce�) reproduc�on and implica�ons for management. Rep. int. Whal.
Commn (special issue) 6: 27–50.
Maeda, H., Bando, T., Kishiro, T., Kitakado, T. and Kato, H. 2016.
Basic informa�on of earplug as age character of common
minke whales in western North Paciﬁc. Paper SC/F16/JR53
presented to the JARPNII special permit expert panel review
workshop, February 2016 (unpublished). 10 pp. [Available
from the IWC Secretariat].
Masaki, Y. 1976. Biological studies on the North Paciﬁc sei
whale. Bull. Far Seas Fish. Res. Lab. 14: 1–114.
Murase, H., Hakamada, T., Matsuoka, K., Nishiwaki, S., Inagake,D., Okazaki, M., Toji, N. and Kitakado, T. 2014. Distribu�on ofsei whales (Balaenoptera borealis) in the subarc�c-subtropical transi�on area of the western North Paciﬁc in rela�on to
oceanic fronts. Deep Sea Res. II 107: 22–28.
Omura, H. 1963. An improved method for collec�on of ear plugs
from baleen whales. Norsk Hvalfangst-Tidende 10: 279–283.

53

Technical Reports of the Ins�tute of Cetacean Research (2020)

pp. 54–56

Technical Report-Note (not peer reviewed)

Japanʼs system for monitoring of whale products sold inits domes�c market through DNA registra�on
Mutsuo Goto* and Hiroyuki Oikawa
Institute of Cetacean Research, 4–5 Toyomi-cho, Chuo-ku, Tokyo 104–0055, Japan*Contact e-mail: goto@cetacean.jp
As in the case of Norway and Iceland, Japan has beenusing a DNA-based system for monitoring (tracking
origins of) large whale products sold in the retail market(Figure 1). Such a task is entrusted to the Ins�tute of
Cetacean Research (ICR) by the Government of Japan(Fisheries Agency of Japan) as an outsourced program
every year.
The system is composed of two main components, i) a
DNA register including gene�c proﬁles of all large whales
legally taken (including bycatches) and imported into
Japan, and ii) systema�c surveys of large whale products
sold in the retail market. The objec�ve of this system is toprevent illegal take of large whales and illegal import oflarge whale products by comparing DNA proﬁles of large
whale products obtained in the market with those ﬁled in
the DNA register.
The Japanese DNA register (JDR) is established based ontechnical speciﬁca�ons similar to those of the Norwegian
DNA register (Glover et al., 2012). It contains DNA proﬁlesof large whales sampled from diﬀerent sources:i)

Scien�ﬁc whaling conducted under special permit
in the Antarc�c (JARPA/JARPAII and NEWREP-A).
This category includes Antarc�c minke whales
collected between 1987/88 and 2018/19, and ﬁnwhales between 2005/06 and 2011/12.
ii) Scien�ﬁc whaling conducted under special permit in the western North Paciﬁc (JARPN/JARPNII

Figure 1.

54

Flow chart of the DNA monitoring system of large
whale products sold in the Japanese retail market.

and NEWREP-NP). This category includes common minke whales collected between 1994 and2019; Brydeʼs whales between 2000 and 2019;sei whales between 2002 and 2019; and sperm
whales between 2000 and 2013.
iii) Commercial whaling within Japanʼs Exclusive
Economic Zone (EEZ). This category includes common minke, Brydeʼs and sei whales collected since
2019.
iv) Bycatches in set nets. This category includes mainly common minke whales sold/used domes�cally(register system established since 1 July 2001).
v) Imports. This category includes North Atlan�c common minke whales from Norway and North Atlan�cﬁn whales from Iceland since 2008.
By the end of 2019, DNA proﬁles of a total of 19,000
individual whales were registered in the JDR.
In the cases of scien�ﬁc whaling and current commercial whaling, samples for gene�c analysis have been collected by researchers from each whale. Samples include
skin samples (two or three pieces of 5×5×5 mm kept
frozen or in 99% ethanol). In addi�on, a large amountof informa�on is collected using established protocols
from every whale sampled including species, catch dateand loca�on (longitude, la�tude), body length, sex andmaturity status.
The Japanese regula�on on bycatches of large whales(established from 1 July 2001) requires that all animals
should be registered with a DNA proﬁle before any
products derived from a bycaught animal are sold in
the market. Details of the regula�on and procedurecan be found in the following web page: h�p://www.
icrwhale.org/pdf/higekujira.pdf. Skin or muscle samples(5×5×5 cm) are taken by the ﬁsherman who intends tosell the bycaught whale and sent to the laboratory at ICR(as frozen samples). Together with the gene�c samples,the ﬁshermen should provide the following informa�onin accordance with an established protocol: species, dateand loca�on of the bycatch, type of set net, body length

Japanʼs system for monitoring of whale products sold in its domestic market through DNA registration
Table 1
The number of whale products by sampling sites in the 2017
survey.
City/Town

Meat

Blubber

Ventralgrooves

Total

SapporoSendai
Niigata
Kanazawa
NotoAnamizu
ToyamaHimi
Nagoya
Kochi
Kobe
Osaka
Taiji
Nachikatsuura
Hiroshima
Shimonoseki
Fukuoka
Nagasaki

23
27
3
3
5
1
4
3
13
14
10
12
5
2
11
18
16
14

2
2
10
3
5
1
0
5
1
11
8
5
6
5
0
6
5
6

7
6
3
1
1
0
0
1
5
9
3
7
2
2
4
10
14
10

32
35
16
7
11
2
4
9
19
34
21
24
13
9
15
34
35
30

Total

184

81

85

350

Figure 2.

Sampling sites in the 2017 DNA market survey.

and sex of the bycaught whale to be used (sold).
The following gene�c markers are used in the JDR:i)

An approximately 500 bp fragment of the 5ʹ-endof the mitochondrial DNA (mtDNA) control region,
which is used for the purpose of iden�ﬁca�on of
the origin of the whale products (species iden�ﬁca�on), through phylogene�c analyses constructed by ʻtestʼ and ʻtypeʼ sequences;ii) Microsatellite DNA (genotyping with 13–17 lociin each species), which is used for the purpose ofindividual iden�ﬁca�on; andiii) Y chromosome DNA, which is used for the purposeof sex determina�on.

For further details, see Pastene and Goto (2006).
Regarding sampling in the market, one or two technicians, who are familiar with whale products and market
opera�ons, carry out the sampling of whale products
at retail shops each year. On an annual basis, around
350 samples are collected between September andDecember throughout Japan, including around 18 ci�esor towns. Table 1 shows the ci�es/towns surveyed andthe whale products sampled in 2017. Figure 2 shows
the geographical distribu�on of the sampling site in the

Figure 3.

An example of species iden�ﬁca�on by phylogene�c analysis of ʻtestʼ and ʻtypeʼ sequences of thewhale products iden�ﬁed as Brydeʼs whale in the2017 market survey. SH: Southern Hemisphere; NA:
North Atlan�c; and NP: North Paciﬁc. 17SS+3 digits
indicates the market samples collected in the 2017
survey.

2017 survey. The sampling a�empts to cover the whole
Japanese archipelago, but the method of sampling is nota random procedure. The following informa�on is collected for each whale product sampled: kind of whale
55

Technical Reports of the Ins�tute of Cetacean Research (2020)
Table 2
Results of species and individual iden�ﬁca�on of the 2017 market survey. The numbers of individual
whales are shown in parenthesis. NG: Failed DNA extrac�on. NP: North Paciﬁc and NA: North Atlan�c.
Species iden�ﬁca�on

N

Antarc�c minke whale
Common minke whale: NP

110
87

Common minke whale: NA
Fin whale: NA
Sei whale
Brydeʼs whale
Bairdʼs beaked whale
Striped dolphin
NG
Total

9
40
84
12
6
1
1
350

Individual iden�ﬁca�on
Not conducted*
Special permit: Oﬀshore
Special permit: Coastal
BycatchImport from Norway
Import from Iceland
Special permit: Oﬀshore
Special permit: Oﬀshore
Not conducted**
Not conducted**
—
Total

N
—
1 (1)
28 (13)58 (37)9 (7)40 (35)84 (66)12 (11)———232 (170)*Individual iden�ﬁca�on of Antarc�c minke whales was not conducted because of the low probabilityof illegal products thereof being introduced into the Japanese market.
**Small cetaceans are not subject to the JDR.

product, sampling locality, date, weight and price. Oncesampled, the products are sent to the laboratory at ICR
for gene�c analysis.
The laboratory work for the whale product samples
were conducted with designated gene�c markers for the
JDR described above.
Standard phylogene�c analyses of ʻtestʼ (whale products sampled in the market) and ʻtypeʼ (whale species in
the JDR) mtDNA sequences are conducted to determine
the species of the sampled whale products. Figure 3
shows an example of species iden�ﬁca�on by phylogene�c analysis in the case of Brydeʼs whales sampled in
the 2017 survey.
For individual iden�ﬁca�on, matching of microsatellite genotypes of ʻtestʼ and ʻtypeʼ samples was conductedusing the computer program ʻCervus 3.0 so�ware packageʼ (Kalinowski et al., 2007).
The results of the Japanese market survey in 2017
are shown in Table 2. All 350 market samples, but one,were successfully iden�ﬁed for the species and the origin(area) by the mtDNA analysis. One whale product (fried
whale blubber or ʻKoroʼ in Japanese) failed in the DNA
extrac�on. The microsatellite analysis revealed that the
232 collected samples included 170 individual whales
origina�ng from the Northern Hemisphere. The gene�c
proﬁles of the ʻtestʼ and ʻtypeʼ samples matched in all
the cases (Table 2). Individual iden�ﬁca�on for Antarc�cminke whales was not conducted because of the low

56

probability of illegal products thereof being introduced
into the Japanese market.
Results of the comparisons between ʻtestʼ and ʻtypeʼ
samples are summarized in the annual report to the Fisheries Agency of the Government of Japan.
Considering results of the work obtained so far, it canbe concluded that the DNA-based system is an eﬀec�vetool for monitoring (tracking origins of) large whale products in the Japanese retail market.

ACKNOWLEDGEMENTS
We thank Luis A. Pastene (ICR) for his assistance in preparing this document.

REFERENCES
Glover, K.A., Haug, T., Oien, N., Walløe, L., Lindblom, L.,
Seliussen, B.B. and Skaug, H.J. 2012. The Norwegian minke
whale DNA register: a data base monitoring commercial
harvest and trade of whale products. Fish Fish. 13: 313–332.
Kalinowski, S.T., Taper, M.L. and Marshall, T.C. 2007. Revising
how the computer program CERVUS accommodates genotyping error increases success in paternity assignment. Mol. Ecol.
16: 1099–1106.
Pastene, L.A. and Goto, M. 2006. Report of the Scien�ﬁcCommi�ee. Annex N. Report of the working group on stockdeﬁni�on. Appendix 3. Status of the Japanese DNA register
for large whales. J. Cetacean Res. Manage. (Suppl.) 8:
255–258.

Technical Reports of the Ins�tute of Cetacean Research (2020)

pp. 57–58

Technical Report-Note (not peer reviewed)

Satellite-monitored tracking of Antarc�c minke and ﬁn whales underthe ﬁrst JASS-A survey in the 2019/2020 austral summer season
Kenji Konishi*, Tatsuya Isoda and Taiki Katsumata
Institute of Cetacean Research, 4–5 Toyomi-cho, Chuo-ku, Tokyo 104–0055, Japan*Contact e-mail: konishi@cetacean.jp
skin biopsy samples for gene�c analyses were obtained
from the same satellite-tagged whales. The details of thepla�orm and equipment for satellite-monitored tagging
were described in Konishi and Isoda (2019) and Konishi
et al. (2020).
During the ﬁrst JASS-A survey, the tag deployments
were conducted from the end of January to early February 2020 in the western part of the Interna�onal Whaling
Commission (IWC) Management Area III, in the longitudinal sector between 000°E and 015°E. The tags weresuccessfully deployed on eight Antarc�c minke and ten
ﬁn whales (Table 1). Tracking informa�on was received
from ﬁve Antarc�c minke (Figure 1) and nine ﬁn (Figure 2) whales. In the case of Antarc�c minke whale PTT
ID 181817, which was tagged on 2 February 2020, track-

The Japanese Abundance and Stock structure Surveys in
the Antarc�c (JASS-A) commenced in the 2019/20 australsummer season. The main research objec�ves of JASS-A
are 1) the study of the abundance and abundance trendsof large whale species, and 2) the study of the distribu�on, movement and stock structure of large whale species (GOJ, 2019; Isoda et al., this issue). One of the techniques proposed for the research on main objec�ve 2)
was satellite tagging.
Implantable satellite-monitored tags (SPOT6-type;Wildlife Computers, Redmond, Washington, USA) wereused for tracking Antarc�c minke and ﬁn whales during
the ﬁrst JASS-A survey in the 2019/20 austral summer
season on board of the research vessel Yushin Maru
No.2 (YS2) (see Isoda et al., this issue). When possible,

Table 1
Summary of the satellite-monitored tagging for Antarc�c minke and ﬁn whales in the 2019/20 JASS-A survey.
No.

Date ofdeployment

Species

Schoolsize

La�tude(°S)

Longitude(°E)

Es�mated bodylength (m)*PTT ID

Biopsy samples

1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18

2020/1/21
2020/1/24
2020/1/24
2020/1/26
2020/1/26
2020/1/27
2020/1/30
2020/2/1
2020/2/1
2020/2/1
2020/2/1
2020/2/1
2020/2/2
2020/2/2
2020/2/2
2020/2/3
2020/2/4
2020/2/6

Fin
Fin
Fin
Fin
Fin
Fin
Ant. Minke
Ant. Minke
Ant. Minke
Fin
Fin
Fin
Ant. Minke
Ant. Minke
Ant. Minke
Ant. Minke
Ant. Minke
Fin

1
2
1
4
3
5
2
1
1
2
6
6
2
1
1
1
1
3

67.3
61.6
61.2
61.5
61.9
64.4
68.8
68.7
68.5
68.4
68.4
68.4
68.5
68.6
68.5
68.4
65.8
63.0

11.0
7.6
7.3
5.9
5.7
4.2
12.8
13.7
14.1
14.3
14.5
14.5
14.6
15.0
14.5
13.5
13.4
15.1

23.1
20.1
19.7
19.4
20.5
19.5
7.6
7.5
7.9
19.6
20.1
20.3
7.6
7.0
6.2
6.4
5.7
18.8

66629
66641
66628
181812
181818181816
181813
181811
181821
181819181824
181820181817
181815
18181066627
181822
181814

Y
N
Y
Y
Y
N
Y
Y
Y
N
Y
Y
Y
Y
Y
Y
Y
Y

*Body lengths of whales were es�mated by the researcher on board.
Tagging experiments in No. 11 and 12 were from the same school.
57

Technical Reports of the Ins�tute of Cetacean Research (2020)

ing was s�ll ongoing as of mid-August 2020. This whale
showed a long longitudinal movement before star�ng
migra�on to the north (Figure 1).
These tracking data will be analyzed in conjunc�onwith previous data obtained under NEWREP-A to inves�gate distribu�on, movement and stock structures ofthese whales under main objec�ve 2) of JASS-A.

ACKNOWLEDGEMENTS

Figure 1.

Tracking lines of ﬁve Antarc�c minke whales tagged
during the ﬁrst JASS-A survey, as of 13th August2020. The lines are plo�ed using all ARGOS Loca�onClasses.

The authors thank the crew and researchers of the ﬁrst
JASS-A survey for their work on whale tagging and biopsy
sampling. The authors also thank the Fisheries Agency ofJapan for the ﬁnancial support of JASS-A and to the Ins�tute of Cetacean Research for logis�cal support. Thanks
also to Luis A. Pastene for his assistance in preparing thisreport.

REFERENCES

Figure 2.

58

Tracking lines of nine ﬁn whales tagged during theﬁrst JASS-A survey. The lines are plo�ed using all
ARGOS Loca�on Classes.

Government of Japan. 2019. Outline of a research program
to inves�gate the abundance, abundance trends and stock
structure of large whales in the Indo-Paciﬁc region of the
Antarc�c, including a survey plan for the 2019/20 australsummer season. Paper SC/68a/ASI8 presented to the IWC
Scien�ﬁc Commi�ee, May 2019 (unpublished). 16 pp. [Paper
available from the IWC Secretariat].
Konishi, K. and Isoda, T. 2019. The pla�orm and equipment forsatellite-monitored tagging experiments in NEWREP-A and
NEWREP-NP. Technical Reports of the Institute of Cetacean
Research (TEREP-ICR) No. 3: 52–54.
Konishi, K., Isoda, T., Bando, T., Minamikawa, S. and Kleivane,
L. 2020. Antarc�c minke whales ﬁnd ice gaps along the iceedge in foraging grounds of the Indo-Paciﬁc sector (60°E and140°E) of the Southern Ocean. Polar Biol. 43 (4): 343–357.

Technical Reports of the Ins�tute of Cetacean Research (2020)

pp. 59–60

Commentary
The views expressed here are those of the author and do not necessarily reﬂect the views of the Institute of Cetacean
Research

Comments on the use of lethal and non-lethal techniques inthe studies of large whales
Genta Yasunaga
Institute of Cetacean Research, 4–5 Toyomi-cho, Chuo-ku, Tokyo 104–0055, Japan
Japan designed and implemented whale research programs under special scien�ﬁc permit in both the Antarc�c and western North Paciﬁc. These research programs
ceased on 30 June 2019 when Japan withdrew from theInterna�onal Conven�on for the Regula�on of Whaling(ICRW). The research programs involved both lethal andnon-lethal techniques. One of the issues frequently discussed among scien�sts was of the need to take whales
for scien�ﬁc purposes. I thought it would be appropriate
to express my comments on this topic now that the research programs have ceased.
In my view, the following aspects are relevant when
making a decision on lethal take of whales for scien�ﬁc purposes: i) whether the samples and data to becollected are required for conserva�on and managementof the whale stock (s); ii) the availability of non-lethal
techniques for ge�ng the same kind of samples anddata; iii) the eﬀect of the lethal take on the whale stock(s) involved.
Regarding i) above, it is recognized that considerable
biological informa�on is required for the appropriate
conserva�on and management of whale resources. For
example, samples and data are required to inves�gate
relevant parameters such as growth, reproduc�on, recruitment, natural mortality, abundance, distribu�on,
feeding ecology, environmental factors, interac�ons withother species, and the inter-annual variability of thesefactors (Ohsumi et al., 2007). Some of this informa�on
can be obtained only through the lethal sampling ofwhales, for example earplugs for age es�ma�on andsexual organs for inves�ga�on of maturity and pregnancy.
These basic biological data are important for es�mating several parameters such as age and length at sexual
maturity, natural mortality, recruitment and so forth,which are important parameters in popula�on dynamics
models. Lethal sampling is also required to inves�gate
stomach contents (qualita�ve and quan�ta�ve) which,in turn, allows us to inves�gate the feeding impact of thespecies (stock) in the ecosystem.

Regarding ii) above, scien�sts should make eﬀorts
to develop new non-lethal techniques to obtain the required informa�on for conserva�on and management. In
this sense, scien�sts of the Ins�tute of Cetacean Research(ICR) have commenced several research projects to inves�gate the sexual maturity status of females throughprogesterone levels in blubber (Inoue et al., 2019). They
have also started researching the feasibility of the DNA
methyla�on proﬁling in skin to obtain age informa�onof whales (Tanabe et al., 2020; Goto et al., 2020). Skinand blubber can be obtained through biopsy sampling.
The Interna�onal Whaling Commission Scien�ﬁc Commi�ee (IWC SC) recommended that a ﬁeld and analy�cal
protocol should be developed to assist the evalua�on of
the u�lity of novel non-lethal techniques (IWC, 2016).
In response to this recommenda�on, scien�sts from the
ICR developed a protocol consis�ng of several ques�onsthat should be responded to in order to evaluate novel
non-lethal techniques (Mogoe, 2018). The protocol consists of four ques�ons: Ques�on 1 is whether �ssues orother kinds of samples can be obtained by a non-lethal
technique. Ques�on 2 is whether a suﬃcient number ofsamples for sta�s�cal analyses can be obtained by the
non-lethal technique. Ques�on 3 is whether the samples
obtained by the non-lethal technique can produce scien�ﬁc informa�on comparable to that produced by alethal sampling technique. Ques�on 4 is whether thecost of obtaining the samples is reasonable for producing
relevant scien�ﬁc informa�on. Ques�on 1 and Ques�on
2 are technical in nature. Ques�on 3 is analy�cal while
Ques�on 4 is of a logis�cal nature. If the answers to
all these ques�ons are aﬃrma�ve, then the non-lethal
technique can be considered appropriate and feasible(Mogoe, 2018).
Regarding iii) above, only abundant species and stocks
can be considered for lethal sampling. Furthermore, popula�on dynamics models should be applied to evaluate
the eﬀect on the stocks of the number of whales taken
lethally. For example, in the case of the former Japanese59Technical Reports of the Ins�tute of Cetacean Research (2020)

whale research under special scien�ﬁc permit in the Antarc�c, the eﬀect of the take of 333 whales on the stocksof Antarc�c minke whales was examined based on Hi�erFi�er model, and results suggested no nega�ve eﬀect on
the stocks (GOJ, 2018).
In summary, the issue of lethal versus non-lethal techniques to study whales is not a ʻblack and-whiteʼ ma�er.
In making a decision on the need for lethal sampling, theu�lity of the target data and samples for conserva�onand management, the availability/feasibility of non-lethal
techniques to obtain the same samples, as well as thestatus of the stock (s) should be considered.

REFERENCES
Goto, M., Kitakado, T. and Pastene, L.A. 2020. A preliminary
study of epigene�c es�ma�on of age of the Antarc�c minke
whale Balaenoptera bonaerensis. Cetacean Population
Studies 2: 5–14.
Government of Japan. 2018. Research Plan for New
Scien�ﬁc Whale Research Program in the Antarc�c Ocean(NEWREP-A). Available at: h�ps://www.jfa.maﬀ.go.jp/j/whale/
pdf/151127newrep-a.pdf.

60Inoue, S., Yasunaga, G. and Pastene, L.A. 2019. Comparison ofprogesterone concentra�ons in blubber and plasma amongfemale Antarc�c minke whales of known reproduc�ve status.
Fish Sci 85: 971–977.
Interna�onal Whaling Commission. 2016. Report of the Expert
Panel of the ﬁnal review on the western North PaciﬁcJapanese Special Permit Programme (JARPNII). Paper SC/66b/
Rep06 presented to the IWC Scien�ﬁc Commi�ee, June 2016
(unpublished). 96 pp. [Available from the IWC Secretariat].
Mogoe, T. 2018. Field and analy�cal protocol for the evalua�on of novel non-lethal techniques in the Japanese whale
research programs. Technical Reports of the Institute of
Cetacean Research (TEREP-ICR) No. 2: 50–55.
Ohsumi, S., Goto, M. and Otani, S. 2007. Necessity of combining
lethal and non-lethal methods for whale popula�on researchand their applica�on in JARPA. Paper SC/59/O2 presented to
the IWC Scien�ﬁc Commi�ee, May 2007 (unpublished). 7 pp.
[Available from the IWC Secretariat].
Tanabe, A., Shimizu, R., Osawa, Y., Suzuki, M., Ito, S., Goto, M.,
Pastene, L.A., Fujise, Y. and Sahara, H. 2020. Age es�ma�on
by DNA methyla�on in the Antarc�c minke whale. Fish Sci
86: 35–41.

Technical Reports of the Ins�tute of Cetacean Research (2020)

pp. 61

International meetingsPar�cipa�on of scien�sts from the Ins�tute of Cetacean Research inInterna�onal Mee�ngs in 2020
Annual mee�ng of the Interna�onal Whaling
Commission Scien�ﬁc Commi�ee (IWC SC)
The Interna�onal Whaling Commission (IWC) is an interna�onal body set up by the terms of the Interna�onal Conven�on for the Regula�on of Whaling (ICRW), which was signed
in Washington, D.C., United States, on 2 December 1946, toʻprovide for the proper conserva�on of whale stocks andthus make possible the orderly development of the whaling industryʼ. One of the important subsidiary bodies of the
IWC is the Scien�ﬁc Commi�ee (SC), which meets annually.
Originally, the 2020 mee�ng of the IWC SC was planned
for Cambridge, United Kingdom, between 12 and 24 May.
The in-person mee�ng was cancelled due to the Covid-19
pandemic. Instead, the mee�ng was carried out througha series of virtual sessions in the same period. A total ofnine scien�sts from the ICR par�cipated in the virtualmee�ngs (Fujise, Kato, Pastene, Tamura, Matsuoka, Hakamada, Taguchi, Isoda and Katsumata). They presented
a total of 11 documents: six documents at the Standing
Working Group on Abundance Es�mates, Stock Status
and Interna�onal Cruises (ASI), three as Intersessional
Steering Group (ISG) reports, one at the ad hoc Working
Group on Photo-ID (PH), and one at the Sub-Commi�ee
on Conserva�on Management Plans (CMP).
The report of the IWC SC mee�ng can be found on thewebsite of the IWC (h�ps://iwc.int/home).
In 2020, ICR scien�sts also par�cipated in the Mee�ngof the IWC-POWER Technical Advisory Group (TAG) andthe 2020 IWC-POWER Planning Mee�ng, held at the Fisheries Agency of Japan (FAJ)ʼs Crew House, Tokyo, Japan
from 18 to 21 January. On 20 April, a virtual mee�ng washeld to discuss the eﬀect of the Covid-19 pandemic on
the logis�cal aspects of the 2020 IWC-POWER survey.
Kato, Pastene, Matsuoka, Hakamada, Taguchi and Takahashi from the ICR par�cipated in these mee�ngs.
The report of these mee�ngs can be found on the website of the IWC (h�ps://iwc.int/home).

Annual mee�ng of the Conven�on on the Conserva�on of Antarc�c Marine Living Resources–
Working Group on Ecosystem Monitoring andManagement (CCAMLR-EMM)
The Conven�on on the Conserva�on of Antarc�c Marine
Living Resources (CCAMLR) is part of the Antarc�c Treaty

System. The Conven�on was opened for signature on 1 August 1980 and entered into force on 7 April 1982, thereby
establishing the Commission for the Conserva�on of Antarc�c Marine Living Resources. Its goal is to preserve marine life and environmental integrity in and near Antarc�ca.
It was established in large part in response to concerns thatan increase in krill catches in the Southern Ocean could
have a serious impact on popula�ons of other marine life,
which are dependent upon krill for food. The CCAMLR has
a Scien�ﬁc Commi�ee and several Working Groups, including the Working Group on Ecosystem Monitoring andManagement (EMM), which meet annually.
The 2020 CCAMLR EMM mee�ng was planned originally for Kochi, India, between 6 and 17 July. The in-personmee�ng was cancelled due to the Covid-19 pandemic.
Instead, an e-mail group held virtual discussions between
6 and 13 July, on three main topics: Risk Assessment Approach, Krill Management Approach, and Research andMonitoring Plan. One scien�st from the ICR par�cipatedin the mee�ng (Pastene).
The report of the e-mail group discussion can be foundon the website of the CCAMLR (h�ps://www.ccamlr.org/).

Annual mee�ng of the North Paciﬁc Marine
Science Organiza�on (PICES)The North Paciﬁc Marine Science Organiza�on (PICES) is anintergovernmental science organiza�on established in 1992
to promote and coordinate marine scien�ﬁc research in the
North Paciﬁc Ocean and its adjacent seas, and to provide amechanism for informa�on and data exchange among scien�sts of member countries. The present members are Canada, Japan, Peopleʼs Republic of China, Republic of Korea,
the Russian Federa�on, and the United States of America.
The 2020 mee�ng of the PICES was originally plannedfor Qingdao, China, between 22 October and 1 November.
The in-person mee�ng was cancelled due to the Covid-19
pandemic. Instead, a business mee�ng of the Sec�on on
Marine Bird and Mammals (S-MBM) was carried out virtually on 16–17 October. One scien�st from the ICR par�cipated in the mee�ng (Tamura). He presented the documentʻEs�ma�on of prey consump�on by marine mammals in the
PICES regions—Update of Hunt et al. (2000)—ʼ. The businessmee�ng discussed research projects for the next ﬁve years.
The report of the mee�ng can be found on the website
of PICES (h�ps://mee�ngs.pices.int/).
61

Technical Reports of the Ins�tute of Cetacean Research (2020)

pp. 62–79

Peer-reviewed publications

List of peer-reviewed publica�ons based on the Ins�tute of
Cetacean Research (ICR)ʼs surveys up to 2020
This sec�on presents a list of peer-reviewed publica�ons
based on data collected by surveys conducted under
special scien�ﬁc permit (JARPA/JARPAII/NEWREP-A and
JARPN/JARPNII/NEWREP-NP), including both lethal andnon-lethal techniques. Peer-reviewed publica�ons basedon these surveys are focused mainly on topics related toassessment and management of large whales. However
samples and data collected by the surveys have also been
useful to carry out studies of a more academic-oriented
nature. Publica�ons based on such studies are also listed
here.
This sec�on also includes a list of peer-reviewed publica�ons resul�ng from other surveys and research ac�vi�es, diﬀerent from special scien�ﬁc permit surveys.
Publica�ons having as a ﬁrst author a non-ICR scien�st commonly followed a data request or collabora�onresearch agreement with ICR. In a few cases, externalscien�sts used published data from ICR surveys in theiranalyses and publica�ons, without a formal agreementwith ICR. These cases are indicated by an asterisk (*).

JARPA/JARPAII/NEWREP-A surveys
1989 (2)Kato, H., Hiroyama, H., Fujise, Y. and Ono, K. 1989.
Preliminary report of the 1987/88 Japanese feasibility study of the special permit proposal for Southern
Hemisphere minke whales. Rep. int. Whal. Commn 39:
235–248.
Nakamura, T., Ohnishi, S. and Matsumiya, Y. 1989. A
Bayesian cohort model for catch-at-age data obtained
from research takes of whales. Rep. int. Whal. Commn
39: 375–382.
1990 (8)Bu�erworth, D.S. and Punt, A.E. 1990. Some preliminary
examina�ons of the poten�al informa�on content
of age-structure data from Antarc�c minke whale research catches. Rep. int. Whal. Commn 40: 301–315.
Ichii, T. 1990. Distribu�on of Antarc�c krill concentra�ons
exploited by Japanese krill trawlers and minke whales.
Proc. NIPR Symp. Polar Biol. 3: 36–56.
Itoh, S., Takenaga, F. and Tsuyuki, H. 1990. Studies on lipidsof the Antarc�c minke whale. I. The fa�y acid composi�ons of the minke whale blubber oils caught on 1987/88
62

season. Yukagaku 39 (7): 486–490 (in Japanese).
Kasamatsu, F., Kishino, H. and Hiroyama, H. 1990. Es�ma�on of the number of minke whale (Balaenoptera
acutorostrata) schools and individuals based on the
1987/88 Japanese feasibility study data. Rep. int. Whal.
Commn 40: 239–247.
Kato, H., Fujise, Y., Yoshida, H., Nakagawa, S., Ishida, M.
and Tanifuji, S. 1990. Cruise report and preliminary
analysis of the 1988/89 Japanese feasibility study ofthe special permit proposal for southern hemisphereminke whales. Rep. int. Whal. Commn 40: 289–300.
Kato, H., Kishino, H. and Fujise, Y. 1990. Some analyses on
age composi�on and segrega�on of southern minke
whales using samples obtained by the Japanese feasibility study in 1987/88. Rep. int. Whal. Commn 40:
249–256.
Nagasaki, F. 1990. The Case for Scien�ﬁc Whaling. Nature
334: 189–190.
Tanaka, S. 1990. Es�ma�on of natural mortality coeﬃcient of whales from the es�mates of abundance andage composi�on data obtained from research catches.
Rep. int. Whal. Commn 40: 531–536.
1991 (9)Bergh, M.O., Bu�erworth, D.S. and Punt, A.E. 1991. Further examina�on of the poten�al informa�on content
of age-structure data from Antarc�c minke whale
research catches. Rep. int. Whal. Commn 41: 349–361.
Ichii, T. and Kato, H. 1991. Food and daily food consump�on of southern minke whales in the Antarc�c. Polar
Biol 11 (7): 479–487.
Kasamatsu, F., Kishino, H. and Taga, Y. 1991. Es�ma�onof southern minke whale abundance and school size
composi�on based on the 1988/89 Japanese feasibility
study data. Rep. int. Whal. Commn 41: 293–301.
Kato, H., Fujise, Y. and Kishino, H. 1991. Age structure andsegrega�on of southern minke whales by the data obtained during Japanese research take in 1988/89. Rep.
int. Whal. Commn 41: 287–292.
Kato, H. and Miyashita, T. 1991. Migra�on strategy ofsouthern minke whales in rela�on to reproduc�ve
cycles es�mated from foetal lengths. Rep. int. Whal.
Commn 41: 363–369.
Kato, H., Zenitani, R. and Nakamura, T. 1991. Inter-reader
calibra�on in age readings of earplugs from southern

List of peer-reviewed publications based on the Institute of Cetacean Research (ICR)ʼs surveys up to 2020

minke whale, with some notes of age readability. Rep.
int. Whal. Commn 41: 339–343.
Kishino, H., Kato, H., Kasamatsu, F. and Fujise, Y. 1991.
Detec�on of heterogeneity and es�ma�on of popula�on characteris�cs from the ﬁeld survey data: 1987/88
Japanese feasibility study of the Southern Hemisphereminke whales. Ann. Inst. Statist. Math. 43 (3): 435–453.
Nakamura, T. 1991. A new look at a Bayesian cohortmodel for �me-series data obtained from researchtakes of whales. Rep. int. Whal. Commn 41: 345–348.
Wada, S., Kobayashi, T. and Numachi, K. 1991. Gene�cvariability and diﬀeren�a�on of mitochondrial DNA in
minke whales. Rep. int. Whal. Commn (special issue)
13: 203–215.
1992 (2)Nakamura, T. 1992. Simula�on trials of a Bayesian cohortmodel for �me-series data obtained from researchtakes of whales. Rep. int. Whal. Commn 42: 421–427.
Tanaka, S., Kasamatsu, F. and Fujise, Y. 1992. Likely precision of es�mates of natural mortality rates fromJapanese research data for Southern Hemisphereminke whales. Rep. int. Whal. Commn 42: 413–420.
1993 (7)Fujise, Y., Ishikawa, H., Saino, S., Nagano, M., Ishii, K.,
Kawaguchi, S., Tanifuji, S., Kawashima, S. and Miyakoshi H. 1993. Cruise report of the 1991/92 Japanese research in Area IV under the special permit for Southern
Hemisphere minke whales. Rep. int. Whal. Commn 43:
357–371.
Hasunuma, R., Ogawa, T., Fujise, Y. and Kawanishi,
Y. 1993. Analysis of selenium metabolites in urine
samples of minke whale (Balaenoptera acutorostrata)
using ion exchange chromatography. Comp. Biochem.
Physiol. 104C (1): 87–89.
Itoh, S., Takenaga, F. and Tsuyuki, H. 1993. Studies onlipids of the Antarc�c minke whale. II. The fa�y acidcomposi�ons of the blubber oils of minke whale anddwarf minke whale caught on 1988/89 and 1989/90
seasons. Yukagaku 42 (12): 1007–1011 (in Japanese).
Iwata, H., Tanabe, S., Sakai, N. and Tatsukawa, R. 1993.
Distribu�on of persistent organochlorines in the oceanic air and surface seawater and the role of ocean on
their global transport and fate. Environ. Sci. Technol.
27: 1080–1098.
Kasamatsu, F., Yamamoto, Y., Zenitani, R., Ishikawa, H.,
Ishibashi, T., Sato, H., Takashima, K. and Tanifuji, S.
1993. Report of the 1990/91 southern minke whale
research cruise under scien�ﬁc permit in Area V. Rep.

int. Whal. Commn 43: 505–522.
Nakamura, T. 1993. Two-stage Bayesian cohort modelfor �me-series data to reduce bias in the es�mate ofmean natural mortality rate. Rep. int. Whal. Commn
43: 343–348.
Pastene, L.A., Kobayashi, T., Fujise, Y. and Numachi, K.
1993. Mitochondrial DNA diﬀeren�a�on in Antarc�cminke whales. Rep. int. Whal. Commn 43: 349–355.
1994 (3)Kimoto, H., Endo, Y. and Fujimoto, K. 1994. Inﬂuence ofinteresteriﬁca�on on the oxida�ve stability of marine
oil triacylglycerols. JAOCS 71 (5): 469–473.
Pastene, L.A., Fujise, Y. and Numachi, K. 1994. Diﬀeren�a�on of mitochondrial DNA between ordinary anddwarf forms of southern minke whale. Rep. int. Whal.
Commn 44: 277–281.
Yoshioka, M., Okumura, T., Aida, K. and Fujise, Y. 1994. A
proposed technique for quan�fying muscle progesterone content in the minke whales (Balaenoptera acutorostrata). Can. J. Zool. 72: 368–370.
1995 (3)
Fukui, Y., Mogoe, T., Terawaki, Y., Ishikawa, H., Fujise,
Y. and Ohsumi, S. 1995. Rela�onship between physiological status and serum cons�tuent values in minke
whales (Balaenoptera acutorostrata). Journal of Reproduction and Development 41 (3): 203–208.
Ishikawa, H. and Amasaki, H. 1995. Development andphysiological degrada�on of tooth buds and development of rudiment of baleen plate in Southern minke
whale, Balaenoptera acutorostrata. J. Vet. Med. Sci. 57(4): 665–670.
Kasamatsu, F., Nishiwaki, S. and Ishikawa, H. 1995. Breeding areas and southbound migra�ons of southernminke whales Balaenoptera acutorostrata. Mar. Ecol.
Prog. Ser. 119: 1–10.
1996 (7)Bakke, I., Johansen, S., Bakke, O. and El-Gewely, M.R.
1996. Lack of popula�on subdivision among the minke
whales (Balaenoptera acutorostrata) from Icelandic
and Norwegian waters based on mitochondrial DNA
sequences. Marine Biology 125: 1–9.
Bu�erworth, D.S. and Geromont, H.F. 1996. On the provision of advice on the eﬀect on stock(s) of scien�ﬁcpermit catches, with par�cular reference to proposed
research catches of minke whales from Antarc�c Area
IV. Rep. int. Whal. Commn 46: 653–655.
Bu�erworth, D.S., Punt, A.E., Geromont, H.F., Kato, H. and63

Technical Reports of the Ins�tute of Cetacean Research (2020)

Miyashita, T. 1996. An ADAPT approach to the analysis
of catch-at-age informa�on for Southern Hemisphereminke whales. Rep. int. Whal. Commn 46: 349–359.
Fukui, Y., Mogoe, T., Jung, Y.G., Terawaki, Y., Miyamoto, A.,
Ishikawa, H., Fujise, Y. and Ohsumi, S. 1996. Rela�onships among morphological status, steroid hormones,
and post-thawing viability of frozen spermatozoa ofmale minke whales (Balaenoptera acutorostrata).
Marine Mammal Science 12 (1): 28–37.
Iga, K., Fukui, Y., Miyamoto, A., Ishikawa, H. and Ohsumi,
S. 1996. Endocrinological observa�ons of female minke
whales (Balaenoptera acutorostrata). Marine Mammal
Science 12 (2): 296–301.
Matsuoka, K., Fujise, Y. and Pastene, L.A. 1996. A sigh�ng
of a large school of the pygmy right whale, Caperea
marginata in the southeast Indian Ocean. Marine
Mammal Science 12 (4): 594–597.
Pastene, L.A., Goto, M., Itoh, S. and Numachi, K. 1996.
Spa�al and temporal pa�erns of mitochondrial DNA
varia�on in minke whale from Antarc�c Areas IV and V.
Rep. int. Whal. Commn 46: 305–314.
1997 (3)Aono, S., Tanabe, S., Fujise, Y., Kato, H. and Tatsukawa,R. 1997. Persistent organochlorines in minke whale(Balaenoptera acutorostrata) and their prey species
from the Antarc�c and the North Paciﬁc. Environmental Pollution 98: 81–89.
Fukui, Y., Mogoe, T., Ishikawa, H. and Ohsumi, S. 1997a.
Factors aﬀec�ng in vitro matura�on of minke whale(Balaenoptera acutorostrata) follicular oocytes. Biology of Reproduction 56: 523–528.
Fukui, Y., Mogoe, T., Ishikawa, H. and Ohsumi, S. 1997b.
In vitro fer�liza�on of in vitro matured minke whale(Balaenoptera acutorostrata) follicular oocytes. Marine Mammal Science 13 (3): 395–404.
1998 (7)
Ichii, T., Shinohara, N., Fujise, Y., Nishiwaki, S. and Matsuoka, K. 1998. Interannual changes in body fat condi�onindex of minke whales in the Antarc�c. Mar. Ecol. Prog.
Ser. 175: 1–12.
Ito, S., Takenaga, F. and Tsuyuki, H. 1998. Lipids in Antarc�c minke whales III. Site and sexual diﬀerences ofmuscle lipids. J. Jpn. Oil Chem. Soc. 47 (2): 191–194 (inJapanese).
Kida, M.Y. 1998. Morphology of the tracheobronchial
tree and the route of the pulmonary artery in the fetal
minke whale (Balaenoptera acutorostrata). Okajimas
Folia Anat. Jpn.75 (5): 251–258.
64

Mogoe, T., Fukui, Y., Ishikawa, H. and Ohsumi, S. 1998a.
Morphological observa�ons of frozen-thawed spermatozoa of Southern minke whales (Balaenoptera acutorostrata). Journal of Reproduction and Development 44(1): 95–100.
Mogoe, T., Fukui, Y., Ishikawa, H. and Ohsumi, S. 1998b.
Eﬀects of diluent composi�on and temperature on
mo�lity and viability a�er liquid storage and cryopreserva�on of minke whale (Balaenoptera acutorostrata) spermatozoa. Marine Mammal Science 14(4): 854–860.
Nagasawa, K., Yamada, T.K. and Ishikawa, H. 1998. Measurements of the skeleton of a minke whale (Balaenoptera acutorostrata) from the Antarc�c Ocean. Bulletinof the Yamagata Prefectural Museum 20: 1–19.
Terabayashi, T. and Kawanishi, Y. 1998. Naturally occurring ganglioside lactones in minke whale brain. Carbohydrate Research 307 (3): 281–290.
1999 (4)Bannister, J.L., Pastene, L.A. and Burnell, S.R. 1999.
First record of movement of a southern right whale(Eubalaena australis) between warm water breeding
grounds and the Antarc�c Ocean, South of 60°S. Marine Mammal Science 15 (4): 1337–1342.
Bu�erworth, D.S., Punt, A.E. Geromont, H.F., Kato, H. andFujise, Y. 1999. Inferences on the dynamics of Southern
Hemisphere minke whales from ADAPT analyses ofcatch-at-age informa�on. J. Cetacean Res. Manage. 1(1): 11–32.
Ishikawa, H., Amasaki, H., Dohguchi, H., Furuya, A. andSuzuki, K. 1999. Immunohistological distribu�ons ofﬁbronec�n, tenascin, type I, III and IV collagens, andlaminin during tooth development and degenera�on infetuses of minke whale, Balaenoptera acutorostrata. J.
Vet. Med. Sci. 61 (3): 227–232.
Thomson, R.B., Bu�erworth, D.S. and Kato, H. 1999. Has
the age at transi�on of southern hemisphere minke
whales declined over recent decades? Marine Mammal Science 15 (3): 661–682.
2000 (5)Asada, M., Horii, M., Mogoe, T., Fukui, Y., Ishikawa, H.
and Ohsumi, S. 2000. In vitro matura�on and ultrastructural observa�on of cryopreserved minke whale(Balaenoptera acutorostrata) follicular oocytes. Biology of Reproduction 62 (2): 253–259.
Kasamatsu, F., Matsuoka, K. and Hakamada, T. 2000. Interspeciﬁc rela�onships in density among the whale
community in the Antarc�c. Polar Biol 23 (7): 466–473.

List of peer-reviewed publications based on the Institute of Cetacean Research (ICR)ʼs surveys up to 2020

Mogoe, T., Suzuki, T., Asada, M., Fukui, Y., Ishikawa, H. andOhsumi, S. 2000. Func�onal reduc�on of the Southernminke whale (Balaenoptera acutorostrata) tes�s during the feeding season. Marine Mammal Science 16(3): 559–569.
Rosenbaum, H.C., Brownell, R.L., Brown, M.W., Schaeﬀ, C.,
Portway, V., White, B.N., Malik, S., Pastene, L.A., Pateneude, N.J., Baker, C.S., Goto, M., Best, P.B., Clapham,
P.J., Hamilton, P., Moore, M., Payne, R., Rowntree, V.,
Tynan, C.T., Bannister, J.L. and DeSalle, R. 2000. Worldwide gene�c diﬀeren�a�on of Eubalaena: ques�oning
the number of right whale species. Molecular Ecology
9: 1793–1802.
Wei, H. and Fukui, Y. 2000. Fer�lizability of ovine, bovine
or minke whale (Balaenoptera acutorostrata) spermatozoa intracytoplasmically injected into bovine oocytes. Zygote 8 (3): 267–274.
2001 (3)Asada, M., Tetsuka, M., Ishikawa, H., Ohsumi, S. andFukui, Y. 2001. Improvement on in vitro matura�on,
fer�liza�on and development of minke whale (Balaenoptera acutorostrata) oocytes. Theriogenology 56 (4):521–533.
Asada, M., Wei, H., Nagayama, R., Tetsuka, M., Ishikawa,
H., Ohsumi, S. and Fukui, Y. 2001. An a�empt at intracytoplasmic sperm injec�on of frozen-thawed minke
whale (Balaenoptera bonaerensis) oocytes. Zygote 9(3): 299–307.
Suzuki, T., Mogoe, T., Asada, M., Miyamoto, A., Tetsuka,
M., Ishikawa, H., Ohsumi, S. and Fukui, Y. 2001. Plasmaand pituitary concentra�ons of gonadotropins (FSHand LH) in minke whales (Balaenoptera acutorostrata)
during the feeding season. Theriogenology 55 (5):1127–1141.
2002 (6)Abe, H. and Goto, M. 2002. The applica�on of microsatellite DNA for determining popula�on structure of minke
whale. pp. 109–113. In: Nishimura, A. (Ed). Technical
Reports of the Hokkaido National Fisheries ResearchInstitute No.5. Hokkaido Na�onal Fisheries ResearchIns�tute, Hokkaido.
Kunito, T., Watanabe, I., Yasunaga, G., Fujise, Y. and Tanabe, S. 2002. Using trace elements in skin to discriminate the popula�ons of minke whale in southern hemisphere. Marine Environmental Research 53: 175–197.
Murase, H., Matsuoka, K., Ichii, T. and Nishiwaki, S. 2002.
Rela�onship between the distribu�on of euphausiids
and baleen whales in the Antarc�c (35°E–145°W).

Polar Biol 25 (2): 135–145.
Pastene, L.A., Goto, M. and Kanda, N. 2002. The u�lity of
DNA analysis for the management and conserva�on oflarge whales. Fish Sci 68 (Suppl. I): 286–289.
Suzuki, M., Ishikawa, H., Otani, S., Tobayama, T., Katsumata, E., Ueda, K., Uchida, S., Yoshioka, M. and Aida,
K. 2002. The characteris�cs of adrenal glands and its
hormones in cetacean. Fish Sci 68 (Suppl. I): 272–275.
Urashima, T., Sato, H., Munakata, J., Nakamura, T., Arai, I.,
Saito, T., Tetsuka, M., Fukui, Y., Ishikawa, H., Lyndersen,
C. and Kovacs, K.M. 2002. Chemical characteriza�on of
the oligosaccharides in beluga (Delphinapterus leucas)
and minke whale (Balaenoptera acutorostrata) milk.
Comparative Biochemistry and Physiology Part B 132:
611–624.
2003 (5)Hayashi, K., Nishida, S., Yoshida, H., Goto, M., Pastene,
L.A. and Koike, H. 2003. Sequence varia�on of the DQB
allele in the cetacean MHC. Mammal Study 28 (2):89–96.
Matsuoka, K., Watanabe, T., Ichii, T., Shimada, H. andNishiwaki, S. 2003. Large whale distribu�ons (south of
60°S, 35°E–130°E) in rela�on to the Southern Boundary of the Antarc�c Circumpolar Current. pp. 26–30.
In: Antarctic Biology in a Global Context. Proceedingsof the VIIIth SCAR Interna�onal Biology Symposium, 27
August-1 September 2001. Vriji Universiteit, Amsterdam, The Netherlands, Backhuys Publisheres, Leiden.
Nishida, S., Hayashi, K., Pastene, L.A., Goto, M., Kanda, N.
and Koike, H. 2003. Polymorphic analysis of cetacean
MHC - A case study on the minke whales -. Mammalian
Science (Suppl.) 3: 75–78 (in Japanese).
Nishida, S., Pastene, L.A., Goto, M. and Kanda, N. 2003.
SRY gene structure and phylogeny in the cetacean species. Mammal Study 28 (1): 57–66.
Ohishi, K., Zenitani, R., Bando, T., Goto, Y., Uchida, K.,
Maruyama, T., Yamamoto, S., Miyazaki, N. and Fujise, Y.
2003. Pathological and serological evidence of Brucella-infec�on in baleen whales (Mys�ce�) in the western
North Paciﬁc. Comp. Immun. Microbiol. Infect. Dis. 26(2): 125–136.
2004 (8)Amemiya, K., Iwanami, Y., Kobayashi, T., Terao, T., Fukui,Y., Ishikawa, H., Ohsumi, S., Hirabayashi, M. and Hochi,
S. 2004. Acquirement of oocyte-ac�va�ng factor in
Antarc�c minke whale (Balaenoptera bonaerensis)
spermatogenic cells, assessed by meiosis resump�onof microinseminated mouse oocytes. J. Mamm. Ova
65Technical Reports of the Ins�tute of Cetacean Research (2020)

Res. 21: 149–156.
Branch, T.A., Matsuoka, K. and Miyashita, T. 2004. Evidence for increase in Antarc�c blue whales based on
Bayesian modeling. Marine Mammal Science 20 (4):726–754.
Fujihira, T., Kinoshita, M., Sasaki, M., Ohnishi, M., Ishikawa, H., Ohsumi, S. and Fukui, Y. 2004. Compara�vestudies on lipid analysis and ultrastructure in porcine
and southern minke whale (Balaenoptera bonaerensis)
oocytes. Journal of Reproduction and Development 50(5): 525–532.
Fukui, Y., Togawa, M., Abe, N., Takano, Y., Asada, M.,
Okada, A., Iida, K., Ishikawa, H. and Ohsumi, S. 2004.
Valida�on of the sperm quality analyzer and the hypoosmo�c swelling test for frozen-thawed ram and minke
whale (Balaenoptera bonarensis) spermatozoa. Journal of Reproduction and Development 50 (1): 147–154.
Ikumi, S., Sawai, K., Takeuchi, Y., Iwayama, H., Ishikawa,
H., Ohsumi, S. and Fukui, Y. 2004. Interspecies soma�ccell nuclear transfer for in vitro produc�on of Antarc�cminke Whale (Balaenoptera bonaerensis) Embryos.
Cloning and Stem Cells 6 (3): 284–293.
Iwayama, H., Hochi, S., Kato, M., Hirabayashi, M., Kuwayama, M., Ishikawa, H., Ohsumi, S. and Fukui, Y. 2004.
Eﬀects of cryodevice type and donorʼs sexual maturity
on vitriﬁca�on of minke whale (Balaenoptera bonaerensis) oocytes at germinal vesicle-stage. Zygote 12 (4):333–338.
Muranishi, Y., Sasaki, M., Hayashi, K., Abe, N., Fujihira, T.,
Ishikawa, H., Ohsumi, S., Miyamoto, A. and Fukui, Y.
2004. Rela�onship between the appearance of preantral follicles in the fetal ovary of Antarc�c minke whales(Balaenoptera bonaerensis) and hormone concentra�ons in the fetal heart, umbilical cord and maternal
blood. Zygote 12 (2): 125–132.
Tetsuka, M., Asada, M., Mogoe, T., Fukui, Y., Ishikawa, H.
and Ohsumi, S. 2004. The pa�ern of ovarian development in the prepubertal Antarc�c minke whale (Balaenoptera bonaerensis). Journal of Reproduction andDevelopment 50 (4): 381–389.
2005 (6)Ishikawa, H. and Shigemune, H. 2005. Improvements
in more humane killing methods of Antarc�c minke
whales, Balaenoptera bonaerensis, in the Japanese
Whale Research Program under Special Permit in the
Antarc�c Sea (JARPA). Jpn. J. Zoo Wildl. Med. 10 (1):
27–34.
Iwayama, H., Ishikawa, H., Ohsumi, S. and Fukui, Y. 2005.
A�empt at in vitro matura�on of minke whale (Balae66

noptera bonaerensis) oocytes using a portable CO2
incubator. Journal of Reproduction and Development
51 (1): 69–75.
Matsuoka, K., Pitman, R. and Marques, F. 2005. A note ona pigmy right whale (Caperea marginata) sigh�ng inthe southwestern Paciﬁc Ocean. J. Cetacean Res. Manage. 7(1): 71–73.
Matsuoka, K., Hakamada, T., Kiwada, H., Murase, H. andNishiwaki, S. 2005. Abundance increases of large baleen whales in the Antarc�c based on the sigh�ng survey during Japanese Whale Research Program (JARPA).
Global Environmental Research 9 (2): 105–115.
Nikaido, M., Sasaki, T., Makino, H., Goto, M., Kanda, N.,
Pastene, L.A. and Okada, N. 2005. Phylogene�c reconstruc�on of baleen whales and detec�on of their pastextensive radia�on event by the SINE inser�on analysis. Fossils 77: 22–28 (in Japanese).
Sasaki, T., Nikaido, M., Hamilton, H., Goto, M., Kato, H.,
Kanda, N., Pastene, L.A., Cao, Y., Fordyce, R.E., Hasegawa, M. and Okada, N. 2005. Mitochondrial phylogene�cs and evolu�on of mys�cete whales. Systematic
Biology 54 (1): 77–90.
2006 (9)Fujihira, T., Kobayashi, M., Hochi, S., Hirabayashi, M.,
Ishikawa, H., Ohsumi, S. and Fukui, Y. 2006. Developmental capacity of Antarc�c minke whale (Balaenoptera bonaerensis) vitriﬁed oocytes following in vitro
matura�on, and parthenogene�c ac�va�on or intracytoplasmic sperm injec�on. Zygote 14 (2): 89–95.
Honda, K., Aoki, M. and Fujise, Y. 2006. Echochemical approach using mercury accumula�on of Antarc�c minke
whale, Balaenoptera bonaerensis, as tracer of historical change of Antarc�c marine ecosystem during 1980–
1999. Bull. Environ. Contam. Toxicol. 76: 140–147.
Iwanami, K., Mita, H., Yamamoto, Y., Fujise, Y., Yamada,T. and Suzuki, T. 2006. cDNA-derived amino acid sequences of myoglobins from nine species of whales
and dolphins. Comparative Biochemistry and Physiology, Part B 145: 249–256.
Kobayashi, T., Amemiya, K., Takeuchi, K., Tsujioka, T.,
Tominaga, K., Hirabayashi, M., Ishikawa, H., Fukui,
Y. and Hochi, S. 2006. Contribu�on of spermatozoal
centrosomes to the microtubule-organizing centre in
Antarc�c minke whale (Balaenoptera bonaerensis).
Zygote 14 (1): 45–51.
Konishi, K. 2006. Characteris�cs of blubber distribu�on
and body condi�on indicators for Antarc�c minke
whale (Balaenoptera bonaerensis). Mammal Study 31(1): 15–22.

List of peer-reviewed publications based on the Institute of Cetacean Research (ICR)ʼs surveys up to 2020

Mori, M. and Bu�erworth, D.S. 2006. A ﬁrst step towards
modelling the krill–predator dynamics of the Antarc�cecosystem. CCAMLR Science 13: 217–277.
Nikaido, M., Hamilton, H., Makino, H., Sasaki, T., Takahashi, K., Goto, M., Kanda, N., Pastene, L.A. and Okada,N. 2006. Baleen whale phylogeny and a past extensive
radia�on event revealed by SINE inser�on analysis.
Mol. Biol. Evol. 23 (5): 866–873.
Rock, J., Pastene, L.A., Kaufman, G., Forestell, P., Matsuoka, K. and Allen, J. 2006. A note on East Australia
Group V Stock humpback whale movement between
feeding and breeding areas based on photo-iden�ﬁca�on. J. Cetacean Res. Manage. 8 (3): 301–305.
Yabuki, T., Suga, T., Hanawa, K., Matsuoka, K., Kiwada, H.
and Watanabe, T. 2006. Possible source of the Antarc�c bo�om water in the Prydz Bay Region. Journal of
Oceanography 62 (5): 649–655.
2007 (7)Asada, M., Tetsuka, M., Ishikawa, H., Ohsumi, S. andFukui, Y. 2007. Ultrastructural changes during matura�on and cryopreserva�on of follicular oocytes of Antarc�c minke whales (Balaenoptera bonaerensis). Jpn. J.
Zoo. Wildl. Med. 12 (1): 51–66.
Branch, T.A., Staﬀord, K.M., Palacios, D.M., Allison, C.,
Bannister, J.L., Burton, C.L.K., Cabrera, E., Carlson, C.A.,
Galle� Vernazzani, B., Gill, P.C., Hucke-Gaete, R., Jenner, K.C.S., Jenner, Mn. M., Matsuoka, K., Mikhalev,
Y.A., Miyashita, T., Morrice, M.G., Nishiwaki, S., Sturrock, V.J., Tormosov, D., Anderson, R.C., Baker, A.N.,
Best, P.B., Borsa, P., Brownell Jr, R.L., Childerhouse,
S., Findlay, K.P., Gerrode�e, T., Ilangakoon, A.D., Joergensen, M., Kahn, B., Ljungblad, D.K., Maughan, B.,
Mccauley, R.D., Mckay, S., Norris, T.F., Oman Whaleand Dolphin Research Group, Rankin, S., Samaran, F.,
Thiele, D., Van Waerebeek, K. and Warneke, R.M. 2007.
Past and present distribu�on, densi�es and movements of blue whales in the Southern Hemisphere andadjacent waters. Mammal Rev. 37 (2): 116–175.
LeDuc, R.G., Dizon, A.E., Goto, M., Pastene, L.A., Kato,
H., Nishiwaki, S. and Brownell, R.L. 2007. Pa�erns ofgene�c varia�on in southern hemisphere blue whales,
and the use of assignment test to detect mixing on
the feeding grounds. J. Cetacean Res. Manage. 9 (1):73–80.
Nagai, H., Mogoe, T., Ishikawa, H., Hochi, S., Ohsumi, S.
and Fukui, Y. 2007. Follicle size-dependent changes
in follicular ﬂuid components and oocyte diameter in
Antarc�c minke whales (Balaenoptera bonaerensis).
Journal of Reproduction and Development 53 (6):1265–1272.
Nishida, S., Goto, M., Pastene, L.A., Kanda, N. and Koike,
H. 2007. Phylogene�c rela�onships among cetaceansrevealed by Y-chromosome sequences．Zoological Science 24 (7): 723–732．
Onbe, K. Nishida, S., Sone, E., Kanda, N., Goto, M., Pastene, L.A., Tanabe, S. and Koike, H. 2007. Sequence varia�on in the Tbx4 gene in marine mammals．Zoological
Science 24 (5): 449–464.
Pastene, L.A., Goto, M., Kanda, N., Zerbini, A.N., Kerem,D., Watanabe, K., Bessho, Y., Hasegawa, M., Nielsen,
R., Larsen, F. and Palsbøll, P.J. 2007. Radia�on andspecia�on of pelagic organisms during periods ofglobal warming: the case of the common minke whale,
Balaenoptera acutorostrata. Molecular Ecology 16:1481–1495.
2008 (4)Ishikawa, H. and Shigemune, H. 2008. Compara�ve experiment of whaling grenades in the Japanese Whale
Research Program under Special Permit (JARPA and
JARPN). Jpn. J. Zoo Wildl. Med. 13 (1): 21–28.
Konishi, K., Tamura, T., Zenitani, R., Bando, T., Kato, H. andWalløe, L. 2008. Decline in energy storage in the Antarc�c minke whale (Balaenoptera bonaerensis) in theSouthern Ocean. Polar Biol 31 (2): 1509–1520.
Ohishi, K., Fujise, Y. and Maruyama, T. 2008. Brucella spp.
in the western North Paciﬁc and Antarc�c cetaceans: areview. J. Cetacean Res. Manage. 10 (1):67–72.
Yunoki, K., Ishikawa, H., Fukui, Y. and Ohnishi, M. 2008.
Chemical proper�es of epidermal lipids, especiallysphingolipids, of the Antarc�c minke whale. Lipids 43(2): 151–159.
2009 (2)Ono, N., Yamaguchi, T., Ishikawa, H., Arakawa, M.,
Takahashi, N. Saikawa, T. and Shimada, T. 2009. Morphological varie�es of the Purkinje ﬁber network inmammalian hearts, as revealed by light and electron
microscopy. Arch Histol Cytol 72 (3): 139–149.
Tamura, T. and Konishi, K. 2009. Feeding habits and prey
consump�on of Antarc�c minke whale (Balaenoptera
bonaerensis) in the Southern Ocean. J. Northw. Atl.
Fish. Sci. 42: 13–25.
2010 (15)
Fujise, Y., Nishiwaki, S. and Ishikawa, S. 2010. Overview of
the Japanese Whale Research Program under Special
Permit in the Antarc�c (JARPA). Bull. Jpn. Soc. Fish.
Oceanogr. 74: 43–44 (in Japanese).
67Technical Reports of the Ins�tute of Cetacean Research (2020)

Fujise, Y., Hatanaka, H. and Ohsumi, S. 2010. Changes
in the Antarc�c marine ecosystem as revealed by the
JARPA research: What has happened to the Antarc�cminke whale stocks? Ottar (Tromsø Museum) 280:
29–34 (in Norwegian).
Glover, K.A., Kanda, N., Haug, T., Pastene, L.A., Øien, N.,
Goto, M., Seliussen, B.B. and Skaug, H.J. Migra�on of
Antarc�c minke whales to the Arc�c. PLoS ONE 5 (12):e15197.
Ishikawa, H. 2010. Animal Welfare - Introduc�on -. Jpn. J.
Zoo. Wildl. Med. 15 (1): 1–3 (in Japanese).
Ishikawa, H. 2010. Lethal Research for Wildlife. Jpn. J.
Zoo. Wildl. Med. 15 (1): 9–14 (in Japanese).
Kokubun, N., Yasuma, H., Murase, H. and Naganobu, M.
2010. Distribu�on of seabirds in the Ross Sea, Antarc�ca. Bull. Jpn. Soc. Fish. Oceanogr. 74: 42–43 (inJapanese).
Matsuoka, K., Kiwada, K., Murase, H. and Nishiwaki, S.
2010. Whale species composi�on and their distribu�on pa�erns in the Ross Sea, Antarc�ca. Bull. Jpn. Soc.
Fish. Oceanogr. 74: 44–45 (in Japanese).
Murase, H., Matsuoka, K., Hakamada, T., Nishiwaki, S.,
Naganobu, M. and Kitakado, T. 2010. Es�ma�on of
spa�al distribu�on pa�ern of cetacean using environmental covariates in the Ross Sea, Antarc�ca. Bull. Jpn.
Soc. Fish. Oceanogr. 74: 47–48 (in Japanese).
Naganobu, M. and Murase, H. 2010. Ecological linkage
between oceanographic environment and distribu�onof top predators and their preys in the Ross Sea region.
Bull. Jpn. Soc. Fish. Oceanogr. 74: 48–49 (in Japanese).
Naganobu, M., Murase, H., Nishiwaki, S., Yasuma, H.,
Matsukura, R., Takao, Y., Taki, K., Hayashi, T., Watanabe,Y., Yabuki, T., Yoda, Y., Noiri, Y., Kuga, M., Yoshikawa, K.,
Kokubun, N., Iwami, T., K., Itoh, K., Goto, M., Isoda, T.,
Matsuoka, K., Tamura, T. and Fujise, Y. 2010. Structureof marine ecosystem of the Ross Sea, Antarc�ca—overview and synthesis of the results of the Japanese mul�disciplinary study by Kaiyo-Maru and JARPA—. Bull.
Jpn. Soc. Fish. Oceanogr. 74: 1–12.
Naganobu, M., Nishiwaki, S., Nagai, N. and Tohyama, D.
2010. Mul�disciplinary ecological survey by the R/V
Kaiyo Maru and JARPA in the Ross Sea, Antarc�ca. Bull.
Jpn. Soc. Fish. Oceanogr. 74: 34–35 (in Japanese).
Pastene, L. A., Acevedo, J., Goto, M., Zerbini, A.N., Acuna,
P. and Aguayo-Lobo, A. 2010. Popula�on structure andpossible migratory links of common minke whales,
Balaenoptera acutorostrata, in the Southern Hemisphere. Conserv Genet 11 (4): 1553–1558．
Tamura, T., Konishi, K., Nishiwaki, S., Taki, K., Hayashi, T.
and Naganobu, M. 2010. Feeding ecology of the Ant68

arc�c minke whales in the Ross Sea, Antarc�ca. Bull.
Jpn. Soc. Fish. Oceanogr. 74: 46–47 (in Japanese).
Yabuki, T., Noiri,Y., Murase, H., Kiwada, H., Matsuoka, K.
and Naganobu, M. 2010. Physical and chemical studyof hydrographic feature based on the R/V Kaiyo Maru
cruise and JARPA in the Ross Sea, Antarc�ca. Bull. Jpn.
Soc. Fish. Oceanogr. 74: 35–36 (in Japanese).
Zenitani, R. and Kato, H. 2010. The growth of baleen
plates in Antarc�c minke whales, with special reference to the V-shape notch appearing at the outer edgeof the plates. Nippon Suisan Gakkaishi 76 (5): 870–876(in Japanese with English abstract).
2011 (1)Matsuoka, K., Hakamada, T., Kiwada, H., Murase, H. andNishiwaki, S. 2011. Abundance es�mates and trends
for humpback whales (Megaptera novaeangliae) in
Antarc�c Areas IV and V based on JARPA sigh�ng data.
J. Cetacean Res. Manage. (special issue) 3: 75–94.
2012 (1)*Sremba, A.L., Hancock-Hanser, B., Branch, T.A., LeDuc,
R.L. and Baker, C.S. 2012. Circumpolar diversity andgeographic diﬀeren�a�on of mtDNA in the cri�callyendangered Antarc�c blue whale (Balaenoptera musculus intermedia). PLoS ONE 7 (3): e32579.
2013 (6)Glover, K.A., Kanda, N., Haug, T., Pastene, L.A., Øien, N.,
Seliussen, B.B., Sørvik, A.G.E. and Skaug, H.J. 2013.
Hybrids between common and Antarc�c minke whales
are fer�le and can back-cross. BMC Genetics 14: 25.
Hakamada, T., Matsuoka, K., Nishiwaki, S. and Kitakado,
T. 2013. Abundance es�mates and trends for Antarc�cminke whales (Balaenoptera bonaerensis) in Antarc�c
Areas IV and V for the period 1989/90–2004/05. J. Cetacean Res. Manage. 13 (2): 123–151.
Kitakado, T., Lockyer, C. and Punt, A.E. 2013. A sta�s�cal
model for quan�fying age-reading errors and its applica�on to the Antarc�c minke whales. J. Cetacean Res.
Manage. 13 (3): 181–190.
Murase, H., Kitakado, T., Hakamada, T., Matsuoka, K.,
Nishiwaki, S. and Naganobu, M. 2013. Spa�al distribu�on of Antarc�c minke whales (Balaenoptera bonaerensis) in rela�on to spa�al distribu�ons of krill in theRoss Sea, Antarc�ca. Fish. Oceanogr. 22 (3): 154–173.
Sasaki, M., Amano, Y., Hayakawa, D., Tsubota, T., Ishikawa,
H., Mogoe, T., Ohsumi, S., Tetsuka, M., Miyamoto, A.,
Fukui, Y., Budipitojo, T. and Kitamura, N. 2013. Structure and steroidogenesis of the placenta in the Antarc-

List of peer-reviewed publications based on the Institute of Cetacean Research (ICR)ʼs surveys up to 2020

�c minke whale (Balaenoptera bonaerensis). Journal of
Reproduction and Development 59 (2): 159–167.
Shibata, Y., Matsuishi, T., Murase, H., Matsuoka, K.,
Hakamada, T., Kitakado, T. and Matsuda, H. 2013. Effects of stra�ﬁca�on and misspeciﬁca�on of covariates
on species distribu�on models for abundance es�ma�on from virtual line transect survey data. Fish Sci 79(4): 559–568.
2014 (3)Konishi, K., Hakamada, T., Kiwada, H., Kitakado, T. andWalløe, L. 2014. Decrease in stomach contents in the
Antarc�c minke whale (Balaenoptera bonaerensis) in
the Southern Ocean. Polar Biol 37 (2): 205–215.
Punt, A.E. 2014. A summary history of the applica�onof sta�s�cal catch-at-age analysis (SCAA) to Antarc�cminke whales. J. Cetacean Res. Manage. 14: 81–92.
Punt, A.E., Hakamada, T., Bando, T. and Kitakado, T. 2014.
Assessment of Antarc�c minke whales using sta�s�cal
catch-at-age analysis (SCAA). J. Cetacean Res. Manage.
14: 93–116.
2015 (2)Konishi, K. and Walløe, L. 2015. Substan�al decline inenergy storage and stomach fullness in Antarc�c minke
whales (Balaenoptera bonaerensis) during the 1990ʼs.
J. Cetacean Res. Manage. 15: 77–92.
Yasunaga, G., Fujise, Y., Zenitani, R., Tanabe, S. and Kato,
H. 2015. Spa�al and temporal varia�on of PCBs and organochlorine pes�cides in the Antarc�c minke whales,
Balaenoptera bonaerensis, in the period 1987–2005.
Chemosphere 126: 11–17.
2016 (2)Pastene, L.A. and Goto, M. 2016. Gene�c characteriza�on and popula�on gene�c structure of the Antarc�cminke whale Balaenoptera bonaerensis in the IndoPaciﬁc region of the Southern Ocean. Fish Sci 82 (6):873–886.
Wada, N., Yamanaka, S., Shibato, J., Rakwal, R., Hirako, S.,
Iizuka, Y., Kim, H., Matsumoto, A., Kimura, A., Takenoya,F., Yasunaga, G. and Shioda, S. 2016. Behavioral andomics analyses study on poten�al involvement of
dipep�de balenine through supplementa�on in dietof senescence-accelerated mouse prone 8. Genomics
Data 10: 38–50.
2017 (3)Isoda, T. and Tamura, T. 2017. Report of a White Adélie
Penguin Pygoscelis adeliae in the Antarc�c Ocean. J.

Yamashina Inst. Ornithol. 48 (2): 74–77.
*Leduc, R.G., Archer, F.I., Lang, A.R., Mar�en, K.K., Hancock-Hanser, B., Torres-Florez, J.P., Hucke-Gaete, R.,
Rosenbaum, H.C., VanWaerebeek, Bownell, R.L. andTaylor, B.L. 2017. Gene�c varia�on in blue whales inthe eastern paciﬁc: implica�ons for taxonomy and useof common wintering grounds. Molecular Ecology 26:
740–751.
Yasunaga, G., Pastene, L.A., Bando, T., Hakamada, T. andFujise, Y. 2017. Age es�ma�on of Antarc�c minke
whales Balaenoptera bonaerensis based on aspar�cacid racemiza�on technique. Fish Sci 86 (6): 947–954.
2019 (1)Inoue, S., Yasunaga, G. and Pastene, L.A. 2019. Comparison of progesterone concentra�ons in blubber
and plasma among female Antarc�c minke whales ofknown reproduc�ve status. Fish Sci 85 (6): 971–977.
2020 (6)Cunen, C., Walløe, L. and Hjort, N.L. 2020. Focused model
selec�on for linear mixed models with an applica�on
to whale ecology. Ann. Appl. Stat. 14 (2): 872–904.
Goto, M., Kitakado, T. and Pastene, L.A. 2020. A preliminary study of epigene�c es�ma�on of age of the
Antarc�c minke whale Balaenoptera bonaerensis. Cetacean Population Studies 2: 5–14.
Konishi, K., Isoda, T., Bando, T., Minamikawa, S. and Kleivane, L. 2020. Antarc�c minke whales ﬁnd ice gaps along
the ice edge in foraging grounds of the Indo-Paciﬁc sector (60°E and 140°E) of the Southern Ocean. Polar Biol
43 (4): 343–357.
Matsuoka, K. and Hakamada, T. 2020. Density distribu�on of several major whale species in the Indo-Paciﬁcregion of Antarc�c using JARPA and JARPAII sigh�ng
data obtained through 1987/88–2008/09 seasons. Cetacean Population Studies 2: 15–38.
Murase, H., Palka, D., Punt, A.E., Pastene, L.A., Kitakado,T., Matsuoka, K., Hakamada, T., Okamura, H., Bando,T., Tamura, T., Konishi, K., Yasunaga, G., Isoda, T. andKato, H. 2020. Review of the assessment of two stocksof Antarc�c minke whales (eastern Indian Ocean andwestern South Paciﬁc). J. Cetacean Res. Manage. 21:
95–122.
Tanabe, A., Shimizu, R., Osawa, Y., Suzuki, M., Ito, S.,
Goto, M., Pastene, L.A., Fujise, Y. and Sahara, H. 2020.
Age es�ma�on by DNA methyla�on in Antarc�c minke
whales. Fish Sci 86 (1): 35–41.

69

Technical Reports of the Ins�tute of Cetacean Research (2020)

JARPN/JARPNII/NEWREP-NP surveys
1996 (3)Bu�erworth, D.S., Geromont H.F. and Wada S. 1996.
Further analysis of allele frequency data to provide
es�mates of the extent of mixing between the various
North Paciﬁc minke whale stocks, and their implica�ons for the status of the “O”stock. Rep. int. Whal.
Commn 46: 443–451.
Kuramochi, T., Machida, M., Araki, J., Uchida, A., Kishiro,
T. and Nagasawa, K. 1996. Minke whales (Balaenoptera acutorostrata) are one of the major ﬁnal hosts ofAnisakis simplex (Nematoda: Anisakidae) in the Northwestern North Paciﬁc Ocean. Rep. int. Whal. Commn
46: 415–419.
Miyashita, T. and Fujise, Y. 1996. Abundance es�mate of
the western North Paciﬁc minke whale in sub-area 9
with notes on the results of dedicated surveys. Rep. int.
Whal. Commn 47: 543–551.
1997 (6)Aono, S., Tanabe, S., Fujise, Y., Kato, H. and Tatsukawa,R. 1997. Persistent organochlorines in minke whale(Balaenoptera acutorostrata) and their prey species
from the Antarc�c and the North Paciﬁc. Environmental Pollution 98: 81–89.
Araki, J., Kuramochi, T., Machida, M., Nagasawa, K. andUchida, A. 1997. A note on the parasite fauna of thewestern North Paciﬁc minke whale (Balaenoptera acutorostrata). Rep. int. Whal. Commn 47: 565–567.
Goto, M. and Pastene, L.A. 1997. Popula�on structureof western North Paciﬁc minke whale based on an
RFLP analysis of mtDNA control region. Rep. int. Whal.
Commn 47: 531–537.
Hatanaka, H. and Miyashita, T. 1997. On the feeding migra�on of the Okhotsk Sea-West Paciﬁc stock of minke
whales, es�mates based on length composi�on data.
Rep. int. Whal. Commn 47: 557–564.
Hatanaka, H. 1997. A short review of hypotheses onminke whale stock structure in the Northwesternpart of the North Paciﬁc. Rep. int. Whal. Commn 47:
553–555.
Pastene, L.A., Goto, M., Itoh, S., Wada, S. and Kato, H.
1997. Intra- and inter-oceanic pa�erns of mitochondrial DNA varia�on in the Brydeʼs whale, Balaenoptera
edeni. Rep. int. Whal. Commn 47: 569–574.
1998 (4)
Lindstrøm, U., Fujise, Y., Haug, T. and Tamura, T. 1998.
Feeding habits of western North Paciﬁc minke whales,
70

Balaenoptera acutorostrata, as observed in JulySeptember 1996. Rep. int. Whal. Commn 48: 463–469.
Pastene, L.A., Goto, M. and Kishino, H. 1998. An es�mateof mixing propor�on of ʻJʼ and ʻOʼ stocks minke whales
in sub-area 11 based on mitochondrial DNA haplotype
data. Rep. int. Whal. Commn 48: 471–474.
Tamura, T., Fujise, Y. and Shimazaki, K. 1998. Diet of minke
whales Balaenoptera acutorostrata in the Northwestern part of the North Paciﬁc in summer 1994 and 1995.
Fish Sci 64 (1): 71–76.
Uchida, A., Kawakami, Y., Yuzu, S., Kishikawa, S., Kuramochi, T., Araki, J., Machida, M. and Nagasawa, K.
1998. Prevalence of parasites and histopathology ofparasi�sa�on in minke whales (Balaenoptera acutorostrata) from the western North Paciﬁc Ocean andSouthern Sea of Okhotsk. Rep. int. Whal. Commn 48:
475–479.
1999 (2)Le, T.H.L., Takahashi, S., Saeki, K., Nakatani, N., Tanabe,S., Miyazaki, N. and Fujise, Y. 1999. High percentage ofbutyl�n residues in total �n in the livers of cetaceansfrom Japanese coastal waters. Environmental Science& Technology 33 (11): 1781–1786.
Mar�nez, I. and Pastene, L.A. 1999. RAPD-typing ofcentral and eastern North Atlan�c and western North
Paciﬁc minke whales, Balaenoptera acutorostrata. ICES
Journal of Marine Science 56 (5): 640–651.
2000 (2)Punt, A.E., Bu�erworth, D.S. and Wada, S. 2000. On theuse of allele frequency data within a Bayesian framework: evalua�on of rela�ve probabili�es of alterna�ve
stock structure hypotheses for North Paciﬁc minke
whales. J. Cetacean Res. Manage. 2 (2): 151–158.
Uchida, A. and Araki, J. 2000. The ectoparasites and endoparasites in the minke whale, Balaenoptera acutorostrata from the western North Paciﬁc Ocean. J. Jpn.
Vet. Med. Assoc. 53 (2): 85–88 (in Japanese).
2001 (1)Okamura, H., Matsuoka, K., Hakamada, T., Okazaki, M.
and Miyashita, T. 2001. Spa�al and temporal structureof the western North Paciﬁc minke whale distribu�oninferred from JARPN sigh�ngs data. J. Cetacean Res.
Manage. 3 (2): 193–200.
2002 (5)Abe, H. and Goto, M. 2002. The applica�on of microsatellite DNA for determining popula�on structure of minke

List of peer-reviewed publications based on the Institute of Cetacean Research (ICR)ʼs surveys up to 2020

whale. pp. 109–113. In: Nishimura, A. (Ed). Technical
Reports of the Hokkaido National Fisheries ResearchInstitute No.5. Hokkaido Na�onal Fisheries ResearchIns�tute, Hokkaido.
Cui, G., Punt, A.E., Pastene, L. A. and Goto, M. 2002.
Bayes and Empirical Bayes approaches to addressing
stock structure ques�ons using mtDNA data, with anillustra�ve applica�on to North Paciﬁc minke whales. J.
Cetacean Res. Manage. 4 (2): 123–134.
Ohsumi, S. and Tamura, T. 2002. Dietary studies on baleen whales in the North Paciﬁc. Fish Sci 68 (Suppl. I):
260–263.
Pastene, L.A., Goto, M. and Kanda, N. 2002. The u�lity of
DNA analysis for the management and conserva�on oflarge whales. Fish Sci 68 (Suppl. I): 286–289.
Tamura T. and Fujise Y. 2002. Geographical and seasonalchanges of the prey species of minke whale in the
Northwestern Paciﬁc. ICES Journal of Marine Science59 (3): 516–528.
2003 (4)Hayashi, K., Nishida, S., Yoshida, H., Goto, M., Pastene,
L.A. and Koike, H. 2003. Sequence varia�on of the DQB
allele in the cetacean MHC. Mammal Study 28 (2):89–96.
Nishida, S., Pastene, L.A., Goto, M. and Kanda, N. 2003.
SRY gene structure and phylogeny in the cetacean species. Mammal Study 28 (1): 57–66.
Nishida, S., Hayashi, K., Pastene, L.A., Goto, M., Kanda, N.
and Koike, H. 2003. Polymorphic analysis of cetacean
MHC - A case study on the minke whales -. Mammalian
Science (Suppl.) 3: 75–78 (in Japanese).
Ohishi, K., Zenitani, R., Bando, T., Goto, Y., Uchida, K.,
Maruyama, T., Yamamoto, S., Miyazaki, N. and Fujise, Y. 2003. Pathological and serological evidence of
Brucella-infec�on in baleen whales (Mys�ce�) in thewestern North Paciﬁc. Comp. Immun. Microbiol. Infect.
Dis 26 (2): 125–136.
2004 (5)Kim, E.Y., Iwata, H., Fujise, Y. and Tanabe, S. 2004. Searching for novel CYP members using cDNA library from aminke whale liver. Marine Environmental Research 58:
495–498.
Ohishi. K., Takishita, K., Kawato, M., Zenitani, R., Bando,T., Fujise, Y., Goto, Y., Yamamoto, S. and Maruyama,
T. 2004. Molecular evidence of new variant Brucellain North Paciﬁc common minke whales. Microbes andInfection 6: 1199–1204.
Ohishi, K., Takishita, K., Kawato, M., Maruyama, T.,

Zenitani, R., Bando, T., Fujise, Y., Goto, Y. and Yamamoto, S. 2004. Molecular characteriza�on of Brucellafrom North Paciﬁc common minke whale. Oceans'04:
499–504.
Watanabe, H., Mogoe, T., Asada, M., Hayashi, K., Fujise,Y., Ishikawa, H., Ohsumi, S., Miyamoto, A. and Fukui,
Y. 2004. Rela�onship between serum sex hormone
concentra�ons and histology of seminiferous tubulesof captured baleen whales in the western North Paciﬁcduring the feeding season. Journal of Reproduction andDevelopment 50 (4): 419–427.
Zharikov, K., Fujise, Y., Tamura, T., Kiwada, H., Bando, T.,
Konishi, K. and Isoda, T. 2004. Distribu�on and feedingof large cetaceans based on the data of JARPN II - 2003
May, 17-July, 9. Marine Mammals of the Holarctic
2004: 207–210.
2005 (5)Birukawa, N., Ando, H., Goto, M., Kanda, N., Pastene, L.A.,
Nakatsuji, H., Hata, H. and Urano, A. 2005. Plasma andurine levels of electrolytes, urea and steroid hormones
involved in osmoregula�on of cetaceans. Zoological
Science 22 (11): 1245–1257.
Nikaido, M., Sasaki, T., Makino, H., Goto, M., Kanda, N.,
Pastene, L.A. and Okada, N. 2005. Phylogene�c reconstruc�on of baleen whales and detec�on of their pastextensive radia�on event by the SINE inser�on analysis. Fossils 77: 22–28 (in Japanese).
Niimi, S., Watanabe, M.X., Kim, E.Y., Iwata, H., Yasunaga,G., Fujise, Y. and Tanabe, S. 2005. Molecular cloning
and mRNA expression of cytochrome P4501A1 and
1A2 in the liver of common minke whales (Balaenoptera acutorostrata). Marine Pollution Bulletin 51 (8):784–793.
Ohishi, K., Takishita, K., Kawato, M., Zenitani, R., Bando,T., Fujise, Y., Goto, Y., Yamamoto, S. and Maruyama,
T. 2005. Chimeric structure of omp2 of Brucella from
North Paciﬁc common minke whales (Balaenoptera
acutorostrata). Microbiol. Immunol. 49 (8): 789–793.
Sasaki, T., Nikaido, M., Hamilton, H., Goto, M., Kato, H.,
Kanda, N., Pastene, L.A., Cao, Y., Fordyce, R.E., Hasegawa, M. and Okada, N. 2005. Mitochondrial phylogene�cs and evolu�on of mys�cete whales. Systematic
Biology 54 (1): 77–90.
2006 (5)
Iwanami, K., Mita, H., Yamamoto, Y., Fujise, Y., Yamada,T. and Suzuki, T. 2006. cDNA-derived amino acid sequences of myoglobins from nine species of whales
and dolphins. Comparative Biochemistry and Physiol71

Technical Reports of the Ins�tute of Cetacean Research (2020)

ogy, Part B 145: 249–256.
Kanda, N., Goto, M. and Pastene, L.A. 2006. Gene�ccharacteris�cs of western North Paciﬁc sei whales,
Balaenoptera borealis, as revealed by microsatellites.
Marine Biotechnology 8: 86–93.
Mitani, Y., Bando., T., Takai., N. and Sakamoto, W. 2006.
Pa�erns of stable carbon and nitrogen isotopes in thebaleens of common minke whale Balaenoptera acutorostrata from the western North Paciﬁc. Fish Sci 72(1): 69–76.
Nikaido, M., Hamilton, H., Makino, H., Sasaki, T., Takahashi, K., Goto, M., Kanda, N., Pastene, L.A. and Okada,N. 2006. Baleen whale phylogeny and a past extensive
radia�on event revealed by SINE inser�on analysis.
Molecular Biology and Evolution 23 (5): 866–873.
Ohishi, K., Maruyama, T., Ninomiya, A., Kida, H., Zenitani,
R., Bando, T., Fujise, Y., Nakamatsu, K., Miyazaki, N. and
Boltunov, A.N. 2006. Serologic inves�ga�on of inﬂuenza a virus infec�on in cetaceans from the western
North Paciﬁc and the Southern Oceans. Marine Mammal Science 22 (1): 214–221.
2007 (11)
Fukui, Y., Iwayama, H., Matsuoka, T., Nagai, H., Koma,
N., Mogoe, T., Ishikawa, H., Fujise, Y., Hirabayashi, M.,
Hochi, S., Kato, H. and Ohsumi, S. 2007. A�empt at
intracytoplasmic sperm injec�on of in vitro matured
oocytes in common minke whales (Balaenoptera acutorostrata) captured during the Kushiro coast survey.
Journal of Reproduction and Development. 53 (4):945–952.
Kanda, N., Goto, M., Kato, H., McPhee, M.V. and Pastene,
L.A. 2007. Popula�on gene�c structure of Brydeʼswhales (Balaenoptera brydei) at the inter-oceanic andtrans-equatorial levels. Conservation Genetics 8 (4):853–864.
Konishi, K. and Tamura, T. 2007. Occurrence of the minimal armhook squids Berryteuthis anonychus (Cephalopoda: Gona�dae) in the stomachs of common minke
whales Balaenoptera acutorostrata in the western
North Paciﬁc. Fish Sci 73 (5): 1208–1210.
Murase, H., Tamura, T., Kiwada, H., Fujise, Y., Watanabe,
H., Ohizumi, H., Yonezaki, S., Okamura, H. and Kawahara, S. 2007. Prey selec�on of common minke (Balaenoptera acutorostrata) and Brydeʼs (Balaenoptera
edeni) whales in the western North Paciﬁc in 2000 and
2001. Fish. Oceanogr. 16 (2): 186–201.
Niimi, S., Kim, E.Y., Iwata, H., Watanabe, M.X., Yasunaga,G., Fujise, Y. and Tanabe, S. 2007. Iden�ﬁca�on andhepa�c expression proﬁles of cytochrome P450 1–4
72

isozymes in common minke whales (Balaenoptera acutorostrata). Comparative Biochemistry and Physiology,
Part B 147: 667–681.
Nishida, S., Goto, M., Pastene, L.A., Kanda, N. and Koike,
H. 2007. Phylogene�c rela�onships among cetaceansrevealed by Y-chromosome sequences. Zoological
Science 24 (7): 723–732.
Onbe, K. Nishida, S., Sone, E., Kanda, N., Goto, M., Pastene, L.A., Tanabe, S. and Koike, H. 2007. Sequencevaria�on in the Tbx4 Gene in marine mammals. Zoological Science 24 (5): 449–464.
Pastene, L.A., Goto, M., Kanda, N., Zerbini, A.N., Kerem,D., Watanabe, K., Bessho, Y., Hasegawa, M., Nielsen,
R., Larsen, F. and Palsbøll, P.J. 2007. Radia�on andspecia�on of pelagic organisms during periods ofglobal warming: the case of the common minke whale,
Balaenoptera acutorostrata. Molecular Ecology 16:1481–1495.
Urashima, T., Kobayashi, M., Asakuma, S., Uemura, Y.,
Arai, I., Fukuda, K., Saito, T., Mogoe, T., Ishikawa, H. andFukui, Y. 2007. Chemical characteriza�on of the oligosaccharides in Brydeʼs whale (Balaenoptera edeni) andSei whale (Balaenoptera borealis) milk. Comparative
Biochemistry and Physiology, Part B 146: 153–159.
Watanabe, H., Tateno, H., Kusakabe, H., Kamiguchi, Y.,
Fujise, Y., Ishikawa, H., Ohsumi, S. and Fukui, Y. 2007.
Fer�lizability and chromosomal integrity of frozenthawed Brydeʼs whale (Balaenoptera edeni) spermatozoa intracytoplasmically injected into mouse oocytes.
Reproduction, Fertility and Development 19 (1): 306.
Watanabe, H., Tateno, H., Kusakabe, H., Matsuoka, T.,
Kamiguchi, Y., Fujise, Y., Ishikawa, H., Ohsumi, S. andFukui, Y. 2007. Fer�lizability and chromosomal integrityof frozen-thawed Brydeʼs whale (Balaenoptera edeni)
spermatozoa intracytoplasmically injected into mouse
oocytes. Zygote 15 (1): 9–14.
2008 (3)Birukawa, N., Ando, H., Goto, M., Kanda, N., Pastene,
L.A. and Urano, A. 2008. Molecular cloning of urea
transporters from the kidneys of baleen and toothed
whales. Comparative Biochemistry and Physiology,
Part B 149: 227–235.
Ishikawa, H. and Shigemune, H. 2008. Compara�ve experiment of whaling grenades in the Japanese Whale
Research Program under Special Permit (JARPA and
JARPN). Jpn. J. Zoo Wildl. Med. 13 (1): 21–28.
Ohishi, K., Fujise, Y. and Maruyama, T. 2008. Brucella spp.
in the western North Paciﬁc and Antarc�c cetaceans: areview. J. Cetacean Res. Manage. 10 (1): 67–72.

List of peer-reviewed publications based on the Institute of Cetacean Research (ICR)ʼs surveys up to 20202009 (9)Bhuiyan, M.M.U., Suzuki, Y., Watanabe, H., Hirayama, H.,
Matsuoka, K., Fujise, Y., Ishikawa, H., Ohsumi, S. andFukui, Y. 2009. A�empts at in vitro fer�liza�on andculture of in vitro matured oocytes in sei (Balaenoptera
borealis) and Brydeʼs (B. edeni) whales. Zygote 17 (1):19–28.
Fujita, H., Honda, K., Hamada, N., Yasunaga, G and Fujise,
Y. 2009. Valida�on of high-throughput measurement
system with microwave-assisted extrac�on, fully automated sample prepara�on device, and gas chromatography-electron capture detector for determina�onof polychlorinated biphenyls in whale blubber. Chemosphere 74: 1069–1078.
Hiwasa, M., Suzuki, Y., Watanabe, H., Bhuiyan, M.M.U.,
Matsuoka, K., Fujise, Y., Ishikawa, H., Ohsumi, S. andFukui, Y. 2009. Eﬀects of semen extenders and storage temperatures on characteris�cs of frozen-thawed
Brydeʼs (Balaenoptera edeni) whale spermatozoa. Journal of Reproduction and Development 55: 599–606.
Konishi, K., Tamura T., Isoda, T. Okamoto, R., Hakamada,T., Kiwada, H. and Matsuoka, K. 2009. Feeding strategies and prey consump�on of three baleen whale species within the Kushiro-Current Extension. J. Northw.
Atl. Fish. Sci. 42: 27–40.
Lee, E., Bhuiyan, M.M.U., Watanabe, H., Matsuoka, K.,
Fujise, Y., Ishikawa, H. and Fukui. Y. 2009. Produc�on ofcloned sei whale (Balaenoptera borealis) embryos byinterspecies soma�c cell nuclear transfer using enucleated pig oocytes. J. Vet. Sci. 10 (4): 285–292.
Matsukura, R., Yasuma, H., Murase, H., Yonezaki, S., Funamoto, T., Honda, S. and Miyashita, K. 2009. Measurements of density contrast and sound-speed contrast
for target strength es�ma�on of Neocalanus copepods(Neocalanus cristatus) in the North Paciﬁc Ocean. FishSci 75 (6): 1377–1387.
Murase, H., Ichihara, M., Yasuma, H., Watanabe, H., Yonezaki, S., Nagashima, H., Kawahara, S. and Miyashita, T.
2009. Acous�c characteriza�on of biological backscatterings in the Kuroshio-Oyashio inter-frontal zone andsubarc�c waters of the western North Paciﬁc in spring.
Fish. Oceanogr. 18 (6): 386–401.
Murase, H., Nagashima, H., Yonezaki, S., Matsukura, R.
and Kitakado, T. 2009. Applica�on of a generalized addi�ve model (GAM) to reveal rela�onships between
environmental factors and distribu�ons of pelagic ﬁsh
and krill: a case study in Sendai Bay, Japan. ICES Journalof Marine Science 66 (6): 1417–1424.
Tsuki, K., Sato, A., Kaneko, H., Yasunaga, G., Fujise, Y. and
Nomata, H. 2009. Comparison of physiologically sig-

niﬁcant imidazole dipep�des in cetaceans sampled inJapanese whale research (short paper). Sci. Rep. Hokkaido Fish. Exp. Stn. 74: 25–28 (in Japanese).
2010 (7)Bhuiyan, M.M.U., Suzuki, Y., Watanabe, H., Lee, E.,
Hirayama, H., Matsuoka, K., Fujise, Y., Ishikawa, H.,
Ohsumi, S. and Fukui, Y. 2010. Produc�on of sei whale(Balaenoptera borealis) cloned embryos by inter- andintra-species soma�c cell nuclear transfer. Journal of
Reproduction and Development 56: 131–139.
Funakasa, N., Yoshioka, M. and Fujise, Y. 2010. Featuresof the Ocular Harderian Gland in three Balaenopterid
species based on anatomical, histological and histochemical observa�ons. Mammal Study 35 (1): 9–15.
Ogawa, G., Ishida, M., Kato, H., Fujise, H. and Urano, N.
2010. Iden�ﬁca�on of faculta�ve anaerobic bacteria
isolated from the intes�ne of the minke whale Balaenoptera acutorostrata by 16S rRNA sequencing analysis. Fish Sci 76 (2): 177–181.
Pastene, L.A., Fujise, Y. and Hatanaka, H. 2010. JARPNII: achallenging mul�discipline research program focusedon whales in the western North Paciﬁc. Ottar (TromsøMuseum) 280: 23–28 (in Norwegian)．
Shishido, R., Ohishi, K., Suzuki, R., Takishita, K., Ohtsu,
D., Okutsu, K., Tokutake, K., Katsumata, E., Bando, T.,
Fujise, Y., Murayama, T. and Maruyama, T. 2010．Cetacean Toll-like receptor 4 and myeloid diﬀeren�a�on
factor 2, and possible cetacean-speciﬁc responses
against Gram-nega�ve bacteria．Comp. Immun. Microbiol. Infect. Dis. 33 (6): 89–98．
Suzuki, Y., Umatani, M., Bhuiyan, M.M.U., Watanabe,H., Mogoe, T., Matsuoka, K., Fujise, Y., Ishikawa, H.,
Ohsumi, S., Sasaki, M. and Fukui, Y. 2010. Eﬀects ofequilibra�on steps, type of sugars and addi�on ofwhale follicular ﬂuid on viability and in vitro matura�on of vitriﬁed whale oocytes．Jpn. J. Zoo Wildl. Med
15 (2): 65–72．
Tojo, N., Matsukura, R., Yasuma, H., Yonezaki, S., Watanabe, H., Kawahara, S., Murase, H. and Miyashita, K.
2010．Spa�al analysis of isada krill (Euphausia paciﬁca)
distribu�on in frontal environments in the North Pacific Ocean．GIS/spatial analyses in ﬁsheries and aquatic
science．4．Interna�onal Fishery GIS Society: 115–138.
2011 (20)
Hakamada, T. and Matsuoka, K. 2011. Abundance es�mate for common minke, Brydeʼs and sei whales in theoﬀshore region of the western North Paciﬁc. Bull. Jpn.
Soc. Fish. Oceanogr. 75 (3): 173–174 (in Japanese).
73Technical Reports of the Ins�tute of Cetacean Research (2020)

Kanda, N., Goto, M. and Pastene, L. A. 2011. Gene�c stock
structure of baleen whales. Bull. Jpn. Soc. Fish. Oceanogr. 75 (3): 191 (in Japanese).
Kato, H. and Fujise, Y. 2011. General discussion: Symposium on “Marine ecosystem of the western North Paciﬁc with special interest in cetaceans.” Bull. Jpn. Soc.
Fish. Oceanogr. 75 (3): 194–195 (in Japanese).
Kishiro, T., Yoshida, H., Tamura, T., Konishi, K., Kanda, N.,
Okamoto, R. and Kato, H. 2011. Rela�onship between
maturity and feeding ecology of common minke whales
oﬀ Kushiro in autumn. Bull. Jpn. Soc. Fish. Oceanogr. 75(3): 175–176 (in Japanese).
Maehara, T., Aoki, I., Yonezaki, S., Watanabe, H., Murase,
H. and Yamaguchi, A. 2011. Feeding ecology of theJapanese anchovy in the oﬀshore area east of Honshu,
Japan in summer. Bull. Jpn. Soc. Fish. Oceanogr. 75 (3):187 (in Japanese).
Matsuoka, K., Kiwada, H., Fujise, Y. and Miyashita, T. 2011.
Distribu�on of baleen whales in the western North Paciﬁc based on the JARPN and JARPN II sigh�ng surveys(1994 to 2007). Bull. Jpn. Soc. Fish. Oceanogr. 75 (3):178–179 (in Japanese).
Miyashita, K., Matsukura, R., Tojo, N., Minami, K.,
Yasuma, H., Nagashima, H., Onodera, K., Watanabe, H.,
Yonezaki, S. and Murase, H. 2011. Monitoring of preyof cetaceans by using quan�ta�ve echosounders. Bull.
Jpn. Soc. Fish. Oceanogr. 75 (3): 183–184 (in Japanese).
Mori, M., Watanabe, H., Hakamada, T., Tamura, T., Konishi, K., Murase, H. and Matsuoka, K. 2011. Development of an ecosystem model of western North Paciﬁcusing Ecopath with Ecosim. Bull. Jpn. Soc. Fish. Oceanogr. 75 (3): 189 (in Japanese).
Murase, H., Kawabata, A., Kubota, H., Nakagami, M.,
Amakasu, K., Abe, K., Miyashita, K. and Oozeki, Y. 2011.
Eﬀect of depth-dependent target strength on biomasses�ma�on of Japanese anchovy. Journal of Marine
Science and Technology 19: 267–272.
Murase, H., Tamura, T., Isoda, T., Okamoto, R., Kato, H.,
Yonezaki, S., Watanabe, H., Tojo, N. Matsukura, R.,
Miyashita, K., Kiwada, H., Matsuoka, K., Nishiwaki, S.,
Inagake, D., Okazaki, M., Okamura, H. and Fujise, Y.
2011. Prey selec�on of common minke, Brydeʼs andsei whales in the oﬀshore region of the western North
Paciﬁc. Bull. Jpn. Soc. Fish. Oceanogr. 75 (3): 180–181(in Japanese).
Nishiwaki, S., Otani, S. and Tamura, T. 2011. The satellite
tracking of the western North Paciﬁc. Bull. Jpn. Soc.
Fish. Oceanogr. 75 (3): 193–194 (in Japanese).
Onodera, K., Nagashima, H., Matsukura, R., Miyashita,K., Yonezaki, S. and Murase, H. 2011. Cetacean prey74

species and oceanographic surveys in Sendai Bay. Bull.
Jpn. Soc. Fish. Oceanogr. 75 (3): 184–185 (in Japanese).
Pastene, L.A., Hatanaka, H., Fujise, Y., Kanda, N., Murase,
H., Tamura, T., Miyashita, T. and Kato, H. 2011. Whale
research program under special permit in the western
North Paciﬁc—Phase II (JARPNII): origin and objec�ves. Bull. Jpn. Soc. Fish. Oceanogr. 75 (3): 172–173 (inJapanese).
Pastene, L.A., Hatanaka, H., Fujise, Y., Kanda, N., Murase,
H., Tamura, T., Yasunaga, G., Kishiro, T., Yoshida, H.,
Miyashita, T. and Kato, H. 2011. The Japanese Whale
Research Program under Special Permit in the western
North Paciﬁc Phase-II (JARPNII): origin, objec�ves andresearch progress made in the ﬁrst six years period(2002–2007), including scien�ﬁc considera�ons for the
next research period. Aquabiology 33 (2): 171–185 (inJapanese).
Sasaki, Y., Saito, S., Matsuoka, K., Murase, H. and Kiwada.,
H. 2011. Spa�al distribu�on pa�erns of sei and Brydeʼswhales. Bull. Jpn. Soc. Fish. Oceanogr. 75 (3): 179–180(in Japanese).
Tamura, T., Konishi, K., Isoda, T., Hakamada, T., Kiwada, H.,
Okamoto, R. and Matsuoka, K. 2011. The food habitat,rela�on of habitat factor and prey consump�on of
three baleen whale species in the oﬀshore region of
the western North Paciﬁc. Bull. Jpn. Soc. Fish. Oceanogr. 75 (3): 174–175 (in Japanese).
Watanabe, H., Kiwada, H., Kumagai, S., Yonezaki, S.,
Kishiro, T., Konishi., K. and Yoshida, H. 2011. Habitatand prey selec�on of common minke whale oﬀ Kushiro
in autumn—Es�ma�on of distribu�on and migra�onof common minke whale using environmental informa�on—Bull. Jpn. Soc. Fish. Oceanogr. 75 (3): 182–183 (inJapanese).
Yamaguchi, A., Hanamiya, Y., Murase, H. and Watanabe,
H. 2011. Ver�cal ﬂux of materials by diel ver�cal migra�on of macrozooplankton. Bull. Jpn. Soc. Fish. Oceanogr. 75 (3): 185–186 (in Japanese).
Yasunaga, G. and Fujise, Y. 2011. Monitoring of pollutants in the baleen whales and their prey species in thewestern North Paciﬁc. Bull. Jpn. Soc. Fish. Oceanogr. 75(3): 190 (in Japanese).
Yoshida, H., Kishiro, T., Goto, M., Bando, T., Tamura, T.,
Konishi, K., Okamoto, R. and Kato, H. 2011. Feeding
ecology of common minke whales in coastal waters oﬀ
Sanriku in spring. Bull. Jpn. Soc. Fish. Oceanogr. 75 (3):
177–178 (in Japanese).
2012 (4)Maeda, H. and Kato, H. 2012. Technical improvements

List of peer-reviewed publications based on the Institute of Cetacean Research (ICR)ʼs surveys up to 2020of the age character (earplug) collec�ons in common
minke whales. Bull. Jpn. Soc. Fish. Oceanogr. 76 (2):59–65 (in Japanese).
Murase, H., Kawabata, A., Kubota, H., Nakagami, M.,
Amakasu, K., Abe, K., Miyashita, K. and Oozeki, Y. 2012.
Basin-scale distribu�on pa�erns and biomass es�ma�on of Japanese anchovy Engraulis japonicus in thewestern North Paciﬁc. Fish Sci 78 (4): 761–773.
Nakamura, G., Kato, H. and Fujise, Y. 2012. Rela�ve
growth of the skull of the common minke whale Balaenoptera acutorostrata from the North Paciﬁc in comparison to other Balaenoptera species. Mammal Study
37 (2): 105–112.
Watanabe, H, Okazaki, M., Tamura, T., Konishi, K., Inagake, D., Bando, T., Kiwada, H. and Miyashita, T. 2012.
Habitat and prey selec�on of common minke, sei, andBrydeʼs whale in mesoscale during summer in the
sub-arc�c and transi�on regions of the western North
Paciﬁc. Fish Sci 78 (3): 557–567.
2013 (5)*Kershaw, F., Leslie, M.S., Collins, T., Mansur, R.M., Smith,B.D., Minton, G., Baldwin, R., LeDuc, R.G., Anderson,
R.C., Brownell, R.L. and Rosenbaum, H.C. 2013. Popula�on diﬀeren�a�on of 2 forms of Brydeʼs whales in the
Indian and Paciﬁc Oceans. Journal of Heredity 104 (6):755–764.
Maeda, H., Kawamoto, T. and Kato, H. 2013. A study on
the improvement of age es�ma�on in common minke
whales using the method of gela�nized extrac�on of
earplug. NAMMCO Scientiﬁc Publications 10: 17pp.
Matsukura, R., Sawada, K., Abe, K., Minami, K., Nagashima, H., Yonezaki, S., Murase, H. and Miyashita,
K. 2013. Comparison of measurements and model
calcula�ons of target strength of juvenile sandeel in
Sendai Bay. Nippon Suisan Gakkaishi 79 (4): 638–648(in Japanese).
Sasaki, H., Murase, H., Kiwada, H., Matsuoka, K., Mitani, Y.
and Saitoh, S. 2013. Habitat diﬀeren�a�on between sei(Balaenoptera borealis) and Brydeʼs whales (B. brydei)
in the western North Paciﬁc. Fish. Oceanogr. 22 (6):496–508.
Shimizu, Y., Ohishi, K., Suzuki, R., Tajima, Y., Yamada, T.,
Kakizoe, Y., Bando, T., Fujise, Y., Taru, H., Murayama, T.
and Maruyama, T. 2013. Amino acid sequence varia�ons of signaling lymphocyte ac�va�on molecule andmortality caused by morbillivirus infec�on in cetaceans. Microbiol Immunol 57: 624–632.

2014 (6)Inoue, S., Kishiro, T., Fujise, Y., Nakamura, G. and Kato,
H. 2014. Seasonal changes in the tes�s of the North
Paciﬁc common minke whale. Nippon Suisan Gakkaishi
80 (2): 185–190 (in Japanese).
Murase, H., Hakamada, T., Matsuoka, K., Nishiwaki, S.,
Inagake, D., Okazaki, M., Tojo, N. and Kitakado, T. 2014.
Distribu�on of sei whales (Balaenoptera borealis) in
the subarc�c–subtropical transi�on area of the western North Paciﬁc in rela�on to oceanic fronts. Deep
Sea Research Part II: Topical Studies in Oceanography
107: 22–28.
Nakamura, G., Kadowaki, I., Nagatsuka, S., Fujise, Y.,
Kishiro, T. and Kato, H. 2014. Varia�on in a color pattern of white patch on the ﬂippers of North Paciﬁc
common minke whales: poten�al applica�on for theirinteroceanic diﬀerence. La mer 52: 31–47.
Nakamura, G. and Kato, H. 2014. Developmental changes
in the skull morphology of common minke whales
Balaenoptera acutorostrata. Journal of Morphology
275: 1113–1121.
Niimi, S., Imoto, M., Kunisue, T., Watanabe, M.X., Kim,
E.Y., Nakayama, K., Yasunaga, G., Fujise, Y., Tanabe, S.
and Iwata, H. 2014. Eﬀects of persistent organochlorine exposure on the liver transcriptome of the common minke whale (Balaenoptera acutorostrata) fromthe North Paciﬁc. Ecotoxicology and Environmental
Safety 108: 95–105.
*Rosel, P.E. and Wilcox, L.A. 2014. Gene�c evidence reveals a unique lineage of Brydeʼs whales in the northern Gulf of Mexico. Endang Species Res. 25 : 19–34.
2015 (1)Kitayama, C., Sasaki, M., Ishikawa, H., Mogoe, T., Ohsumi,
S., Fukui, Y., Budipitojo, T., Kondoh, D. and Kitamura, N.
2015. Structure and func�ons of the placenta in common minke (Balaenoptera acutorostrata), Brydeʼs (B.
brydei) and sei (B. borealis) whales. Journal of Reproduction and Development 61 (5): 415–421.
2016 (6)Hirako, S., Suzuki, M., Kim, H., Iizuka, Y., Matsumoto,A., Wada, N., Okabe, M., Takenoya, F., Yasunaga, G.,
Yamanaka, S., Miyazaki, A. and Shioda, S. 2016. Hepa�clipid accumula�on is ameliorated in obese KK mice by
dietary sei whale oil. Fish Sci 82 (3): 509–517.
Miyakawa, N., Kishiro, T., Fujise, Y., Nakamura, G. andKato, H. 2016. Sexual Dimorphism in Pelvic Bone
Shape of the North Paciﬁc Common Minke Whales(Balaenoptera acutorostrata). Open Journal of Animal
75

Technical Reports of the Ins�tute of Cetacean Research (2020)

Sciences 6: 131–136.
Murase, H., Tamura, T., Otani, S. and Nishiwaki, S. 2016.
Satellite tracking of Brydeʼs whales Balaenoptera edeni
in the oﬀshore western North Paciﬁc in summer 2006
and 2008. Fish Sci 82 (1): 35–45.
Nakamura, G., Kadowaki, I., Nagatsuka, S., Hayashi, R.,
Kanda, N., Goto, M., Pastene, L.A. and Kato, H. 2016.
White patch on the fore-ﬂipper of common minke
whale, as a poten�al morphological index to iden�fy
stocks. Open Journal of Animal Sciences 6: 116–122.
Nakamura, G., Takahashi, M., Hirose, A., Hayashi, R.,
Fujise, Y., Miyakawa, N., Inoue, S. and Kato, H. 2016.
Presence of an interparietal bone and morphologicalvaria�on in the vertex of the skull in North Paciﬁc common minke whale. La mer 54 (1): 1–10.
Ohishi, K., Bando, T., Abe, E., Kawai, Y., Fujise, Y. andMaruyama, T. 2016. Long-term and large-scale epidemiology of Brucella infec�on in baleen whales andsperm whales in the western North Paciﬁc and Antarc�c Oceans. J. Vet. Med. Sci. 78 (9): 1457–1464.
2017 (2)Bando, T., Nakamura, G., Fujise, Y. and Kato, H. 2017.
Developmental changes in the morphology of western
North Paciﬁc Brydeʼs whales (Balaenoptera edeni).
Open Journal of Animal Sciences 7: 344–355.
Ishii, M., Murase, H., Fukuda, Y., Sawada, K., Sasakura,T., Tamura, T., Bando, T., Matsuoka, K., Shinohara, A.,
Nakatsuka, S., Katsumata, N., Okazaki, M., Miyashita,
K. and Mitani, Y. 2017. Diving behavior of sei whales
Balaenoptera borealis rela�ve to the ver�cal distribu�on of their poten�al prey. Mammal Study 42 (4):
191–199.
2018 (3)Hirose, A., Kishida, T. and Nakamura, G. 2018. Nasal mucosa resembling an olfactory system in the common
minke whale (Balaenoptera acutorostrata). Cetacean
Population Studies 1: 25–28.
Huijser, L.A.E., Bérubé, M., Cabrera, A.A., Prieto, R., Silva,
M.A., Robbins, J., Kanda, N., Pastene, L.A., Goto, M.,
Yoshida, H., Vikingsson, G.A. and Palsbøll, P.J. 2018.
Popula�on structure of North Atlan�c and North Paciﬁc sei whales (Balaenoptera borealis) inferred frommitochondrial control region DNA sequences and microsatellite genotypes. Conserv Genet 19: 1007–1024.
Nakamura, G., Ryeng, K.A., Kadowaki, I., Hayashi, R.,
Nagatsuka, S., Hirose, A., Fujise, Y. and Haug, T. 2018.
Comparison of shapes of the white ﬂipper patch
between two sub-species of common minke whales
76(Balaenoptera acutorostrata). Cetacean Population
Studies 1: 15–24.
2019 (1)
Watari, S., Murase, H., Yonezaki, S., Okazaki, M., Kiyofuji,
H., Tamura, T., Hakamada, T., Kanaji, Y. and Kitakado, T.
2019. Ecosystem modeling in the western North Paciﬁcusing Ecopath, with a focus on small pelagic ﬁshes. Mar
Ecol Prog Ser 617–618: 295–305.
2020 (1)Yasunaga, G. and Fujise, Y. 2020. Concentra�on of persistent organic pollutants (POPs) in three species ofbaleen whales in the western North Paciﬁc. Cetacean
Population Studies 2: 39–53.

Other surveys
1993 (1)Fujise, Y., Ishikawa, H., Saino, S. and Kawasaki, M. 1993.
Catch and struck-and-lost rate in the Japanese Dallʼs
porpoise hand-harpoon ﬁshery. Rep. int. Whal. Commn
43: 453–457.
1995 (1)Shimada, H. and Pastene, L.A. 1995. Report of a sightings survey oﬀ the Solomon Islands with comments on
Brydeʼs whale distribu�on. Rep. int. Whal. Commn 45:
413–418.
1997 (4)Kimura, T., Ozawa, T. and Pastene, L.A. 1997. Sample
prepara�on and analysis of mitochondrial DNA fromwhale baleen plates. Marine Mammal Science 13 (3):495–498.
Ljungblad, D.K., Staﬀord, K.M., Shimada, H. and Matsuoka, K. 1997. Sounds a�ributed to blue whales recorded
oﬀ the southwest coast of Australia in December 1995.
Rep. int. Whal. Commn 47: 435–439.
Ohmagari, K. and Berkes, F. 1997. Transmission of indigenous knowledge and bush skills among the WesternJames Bay Cree Women of Subarc�c Canada. Human
Ecology 25 (2): 197–222.
Shimamura, M., Yasue, H., Ohshima, K., Abe, H., Kato, H.,
Kishiro, T., Goto, M., Munechika, I. and Okada, N. 1997.
Molecular evidence from retroposons that whales
form a clade within even-toed ungulates. Nature 388:
666–670.
1998 (2)Miyazaki, N., Fujise, Y. and Iwata, K. 1998. BiologicalList of peer-reviewed publications based on the Institute of Cetacean Research (ICR)ʼs surveys up to 2020

analysis of a mass stranding of melon-headed whales(Peponocephala electra) at Aoshima. Japan. Bull. Natn.
Sci. Mus., Tokyo, Ser. A 24 (1): 31–60.
Otani, S., Naito, Y., Kawamura, A., Kawasaki, M., Nishiwaki, S. and Kato, A. 1998. Diving behavior and performance of harbor porpoises, Phocoena phocoena,
in Fukuoka bay, Hokkaido, Japan. Marine Mammal
Science 14 (2): 209–220.
1999 (4)Iino, Y. 1999. Treaty Implementa�on: the 1946 Interna�onal Conven�on for the Regula�on of Whaling.
System Compliance in Ocean Management. Southeast
Asian Programme in Ocean Law. Policy and Management: 189–193.
Pastene, L.A. and Shimada, H. 1999. Report of a sigh�ng
survey in Chileʼs Exclusive Economic Zone with comments on sei whale distribu�on. Anales Instituto Patagonia, Serie Cs. Nat. 27: 51–62.
Shimamura, M., Abe, H., Nikaido, M., Ohshima, K. andOkada, N. 1999. Genealogy of families of SINEs in cetaceans and ar�odactyls: the presence of a huge superfamily of tRNAGlu-derived families of SINEs. Mol. Biol.
Evol. 16 (8): 1046–1060.
Yoshida, H. and Kato, H. 1999. Phylogene�c rela�onshipsof Brydeʼs whales in the western North Paciﬁc andadjacent waters inferred from mitochondrial DNA sequences. Marine Mammal Science 15 (4): 1269–1286.
2000 (3)Goto, M. and Ohsumi, S. 2000. Response to “Scien�ﬁc
Whaling Source of illegal Products for Market” By Baker
et al. Science 290: 1695–1696.
Otani, S., Naito, Y., Kato, A. and Kawamura, A. 2000. Diving behavior and swimming speed of a free-ranging
harbor porpoise, Phocoena phocoena. Marine Mammal Science 16 (4): 811–814.
Takahashi, S., Le, L.T.H., Saeki, H., Nakatani, N., Tanabe,S., Miyazaki, N. and Fujise, Y. 2000. Accumula�on ofbutyl�n compounds and total �n in marine mammals.
Water Science and Technology 42 (7–8): 97–108.
2001 (2)Abe, H., Goto, M., Pastene, L.A., Dewa, K. and Naito, E.
2001. Prac�cal use of mul�plex ﬂuorescent PCR for
cetacean sex iden�ﬁca�on. Marine Mammal Science
17 (3): 657–664.
Yoshida, H., Yoshioka, M., Shirakihara, M. and Chow, S.
2001. Popula�on structure of ﬁnless porpoises (Neophocaena phocaenoides) in coastal waters of Japan

based on mitochondrial DNA sequences. Journal of
Mammalogy 82 (1): 123–130.
2002 (1)Konishi, K. and Sakurai, Y. 2002. Geographical varia�ons ininfec�on by larval Anisakis simplex and Contracaecum
osculatum (Nematoda, Anisakidae) in walleye pollock
Theragra chalcogramma stocks oﬀ Hokkaido, Japan.
Fish Sci 68 (3): 534–542.
2003 (1)Matsuoka, K., Ensor, P., Hakamada, T., Shimada, H., Nishiwaki, S., Kasamatsu, F. and Kato, H. 2003. Overview
of the minke whale sigh�ng survey in IWC/IDCR and
SOWER cruises from 1978/79 to 2000/01. J. Cetacean
Res. Manage. 5 (2): 173–201.
2004 (5)Branch, T.A., Matsuoka, K. and Miyashita, T. 2004. Evidence for increases in Antarc�c blue whales based on
Bayesian modelling. Marine Mammal Science 20 (4):726–754.
Escorza-Trevino, S., Pastene, L.A. and Dizon, A.E. 2004.
Molecular analyses of the Truei and Dalli morphotypes of Dallʼs porpoise (Phocoenoides dalli). Journal of
Mammalogy 85 (2): 347–355.
Ikemoto, T., Kunito, T., Watanabe, I., Yasunaga, G., Baba,N., Miyazaki, N., Petrov, E.A. and Tanabe, S. 2004.
Comparison of trace element accumula�on in Baikal
seals (Pusa sibirica), Caspian seals (Pusa caspica) andnorthern fur seals (Callorhinus ursinus). Environmental
Pollution 127: 83–97.
Murase, H., Matsuoka, K., Nishiwaki, S., Hakamada, T. andMori, M. 2004. Eﬀect of observed covariates (school
size, sigh�ng cue, la�tude and sea state) on the Antarc�c minke whale abundance es�ma�on parameters in
the IWC/IDCR-SOWER surveys. J. Cetacean Res. Manage. 6 (3): 283–292.
Yanagimoto, T. and Konishi, K. 2004. Acanthochondria priacanthi (Copepoda: Chondracanthidae) as a biologicalindicator for stock iden�ﬁca�on of sandﬁsh Arctoscopus japonicus (Steindachner). Fish Sci 70 (2): 336–338.
2005 (5)Agusa, T., Matsumoto, T., Ikemoto, T., Anan, Y., Kubota, R.,
Yasunaga, G., Kunito, T., Tanabe, S., Ogi, H. and Shibata,Y. 2005. Body distribu�on of trace elements in blacktailed gulls from Rishiri Island, Japan: age-dependentaccumula�on and transfer to feathers and eggs. Environ. Toxicol. Chem. 24 (9): 2107–2120.
77

Technical Reports of the Ins�tute of Cetacean Research (2020)

Agusa, T., Kunito, T., Yasunaga, G., Iwata, H., Subramanian, A., Ismail, A. and Tanabe, S. 2005. Concentra�onsof trace elements in marine ﬁsh and its risk assessment in Malaysia. Marine Pollution Bulletin 51 (8–12):
896–911.
Fukui, Y., Ishikawa, H. and Ohsumi, S. 2005. Diﬃcul�es
in publishing research results from scien�ﬁc whaling.
Marine Mammal Science 21 (4): 781–783.
Prudente, M., Tanabe, S., Kunisue, T., Yasunaga, G., Watanabe, I. and Miyazaki, N. 2005. Accumula�on of traceelements and persistent organochlorines in resident
and migratory waders from Calatagan Bay, Philippines.
Coastal Marine Science 29 (2): 133–140.
Sanino, G.P., Waerebeek, K.V., Van Bressem, M. andPastene, L.A. 2005. A preliminary note on popula�on
structure in eastern South Paciﬁc common bo�lenose
dolphins, Tursiops truncatus. J. Cetacean Res. Manage.
7 (1): 65–70.
2006 (6)Acevedo, J., Aguayo-Lobo, A. and Pastene, L.A. 2006.
Filopatria de la ballena jorobada (Megaptera novaeangliae Borowski, 1781), al area de alimentacion delEstrecho de Magallanes. Revista de Biologia Marina y
Oceanograﬁa 41 (1): 11–19.
Acevedo, J., Aguayo-Lobo, A., Acuna, P. and Pastene, L.A.
2006. A note on the ﬁrst record of the dwarf minke
whale (Balaenoptera acutorostrata) in Chilean waters.
J. Cetacean Res. Manage. 8 (3): 293–296.
Hayashi, K., Yoshida, H., Nishida, S., Goto, M., Pastene,
L.A., Kanda, N., Baba, H. and Koike, H. 2006. Gene�c
Varia�on of the MHC DQB locus in the ﬁnless porpoise(Neophocaena phocaenoides). Zoological Science 23(2): 147–153.
Mori, M. and Bu�erworth, D.S. 2006. A ﬁrst step towards
modelling the krill–predator dynamics of the Antarc�cecosystem. CCAMLR Science 13: 217–277.
Sekiguchi, K., Olavarria, C., Morse, P., Olson, P., Ensor,
P., Matsuoka, K., Pitman, R., Findlay, K. and Gorter, U.
2006. The spectacled porpoise (Phocoena dioptrica)
in Antarc�c waters. J. Cetacean Res. Manage. 8 (3):265–271.
Takahashi, Y., Ohwada, S., Watanabe, K., Ropert-Coudert,Y., Zenitani, R., Naito, Y. and Yamaguchi, T. 2006. Doeselas�n contribute to the persistence of corpora albican�a in the ovary of the common dolphin (Delphinusdelphis). Marine Mammal Science 22 (4): 819–830.
2007 (2)Acevedo, J., Rasmussen, K., Felix, F., Castro, C., Llano, M.,
78Secchi, E., Saborio, M.T., Aguayo-Lobo, A., Haase, B.,
Scheidat, M., Dalla-Rosa, L., Olavarria, C., Fosestell, P.,
Acuna, P., Kaufman, G. and Pastene, L.A. 2007. Migratory des�na�ons of humpback whales from the Magellan Strait feeding ground, Southeast Paciﬁc. Marine
Mammal Science 23 (2): 453–463.
Branch, T.A., Staﬀord, K.M., Palacios, D.M., Allison, C.,
Bannister, J.L., Burton, C.L.K., Cabrera, E., Carlson, C.A.,
Galle� Vernazzani, B., Gill, P.C., Hucke-Gaete, R., Jenner, K.C.S., Jenner, Mn. M., Matsuoka, K., Mikhalev,
Y.A., Miyashita, T., Morrice, M.G., Nishiwaki, S., Sturrock, V.J., Tormosov, D., Anderson, R.C., Baker, A.N.,
Best, P.B., Borsa, P., Brownell Jr, R.L., Childerhouse,
S., Findlay, K.P., Gerrode�e, T., Ilangakoon, A.D., Joergensen, M., Kahn, B., Ljungblad, D.K., Maughan, B.,
Mccauley, R.D., Mckay, S., Norris, T.F., Oman Whaleand Dolphin Research Group, Rankin, S., Samaran, F.,
Thiele, D., Van Waerebeek, K. and Warneke, R.M. 2007.
Past and present distribu�on, densi�es and movements of blue whales in the Southern Hemisphere andadjacent waters. Mammal Rev. 37 (2): 116–175.
2009 (1)Nambu, H., Tokutake, K., Ishikawa, H., Ohta, M., Fujita, K.
and Yamada, T.K. 2009. Observa�on of a gray whale,
Eschrichtius robustus, in Tokyo Bay in 2005. Japan Cetology 19: 17–22 (in Japanese).
2010 (3)Nambu, H., Ishikawa, H. and Yamada, T. 2010. Recordsof the western gray whale Eschrichtius robustus: its
distribu�on and migra�on. Japan Cetology 20: 21–29(in Japanese).
Pastene, L.A. and Quiroz, D. 2010. An Outline of the History of Whaling in Chile. pp. 73–98. In: Interna�onal
Center for Folk Culture Studies (Ed). Human Culture
from the Perspective of Traditional Maritime Communities, International Symposium Report No. 1. Kanagawa
Shimbun Press, Kanagawa. 199pp.
Taguchi, M., Ishikawa, H. and Matsuishi, T. 2010. Seasonal
distribu�on of harbour porpoise (Phocoena phocoena)
in Japanese waters inferred from stranding and bycatchrecords．Mammal Study 35 (2) : 133–138．
2011 (2)Acevedo, J., Olavarría, C., Plana, J., Aguayo-Lobo, A., Larrea, A. and Pastene, L.A. 2011. Occurrence of dwarf
minke whales (Balaenoptera acutorostrata subsp.)
around the Antarc�c Peninsula. Polar Biol 34 (2):313–318.

List of peer-reviewed publications based on the Institute of Cetacean Research (ICR)ʼs surveys up to 2020

Acevedo, J., Plana, J., Aguayo-Lobo, A. and Pastene, L.A.
2011. Surface feeding behavior of humpback whalesin the Magellan Strait. Revista de Biologia Marina y
Oceanograﬁa. 46 (3): 483–490.
2013 (4)Acevedo J., Haro D., Dalla R.L., Aguayo-Lobo A., HuckeGaete R., Secchi E., Plana J. and Pastene L.A. 2013.
Evidence of spa�al structuring of eastern South Paciﬁchumpback whale feeding grounds. Endang Species Res
22 (1): 33–38.
Kitamura, S., Matsuishi, T., Yamada, K., Tajima, Y., Ishikawa, H., Tanabe, S., Nakagawa, H., Uni, Y. and Abe, S.
2013. Two gene�cally dis�nct stocks in Bairdʼs beaked
whale (Cetacea: Ziphiidae). Marine Mammal Science
29 (4): 755–766.
Murase, H., Temoai, I., Kirata, T., Finkaso, S., Yasunaga, G.
and Pastene, L.A. 2013. A note on cetaceans oﬀ Kiriba�
and Tuvalu from a research cruise in October 2010. J.
Cetacean Res. Manage. 13 (2): 153–158.
Sugino, T., Yasunaga, G. and Fukuda, M. 2013. Eﬀect ofwhale meat extract on fa�gue induced by physical load
and by daily ac�vi�es in humans. Jpn. Pharmacol. Ther
41 (9): 879–893 (in Japanese).
2014 (3)Pastene, L.A. 2014. Macaya Hermanos y Compania.
Notas sobre las ac�vidades de la empresa ballenerade más larga historia en Chile. Colección Mocha Dick
pp 255–274.
Quiroz, D. and Pastene, L.A. 2014. Norwegian whaling inChile in the 20th Century and its inﬂuence on Chilean
local communi�es. pp. 61–70. In: Jan Erik Ringstad(Ed). Whaling and History IV. Commander Chr. Christensenʼs Whaling Museum, part of Ves�oldmuseene
IKA, Sande�ord. 192pp.
Quintela, M., Skaug, H.J., Øien, N., Haug, T., Seliussen,
B.B., Solvang, H.K., Pampoulie, C., Kanda, N., Pastene,
L.A. and Glover, K.A. 2014. Inves�ga�ng popula�on gene�c structure in a highly mobile marine organism: theminke whale Balaenoptera acutorostrata acutorostrata in the North East Atlan�c. PLoS ONE 9 (9): e108640.
2015 (3)Funasaka, N., Kirihata, T., Kato, H. and Ohsumi, S. 2015.
The ﬁrst record of a true albino common bo�lenose
dolphin Tursiops truncatus from Japan. Mammal Study

40 (1): 19–22.
Pastene, L.A., Acevedo, J., Siciliano, S., Sholl, T.G.C., de
Moura, J.F., O�, P.H. and Aguayo-Lobo, Anelio. 2015.
Popula�on gene�c structure of the South American
Brydeʼs whale. Revista de Biologia Marina y Oceanograﬁa 50 (3): 453–464.
Sakamoto, M., Itai, T., Yasutake, A., Iwasaki, T., Yasunaga,G., Fujise, Y., Nakamura, M., Murata, K., Chan, H.M.,
Domingog, J.L. and Marumoto, M. 2015. Mercury
specia�on and selenium in toothed-whale muscles.
Environmental Research 143: 55–61.
2017 (3)Funasaka, N., Kirihata, T., Hosono, M., Kato, K. andOhsumi, S. 2017. Three Cases of Anomalously White
Rissoʼs Dolphins Grampus griseus in Japan. Mammal
Study 42 (3): 173–178.
Hakamada, T., Matsuoka, K., Murase, H. and Kitakado, T.
2017. Es�ma�on of the abundance of the sei whale
Balaenoptera borealis in the central and eastern North
Paciﬁc in summer using sigh�ng data from 2010 to2012. Fish Sci 83 (6): 887–895.
Malde, K., Seliussen, B.B., Quintela, M., Dahle, G., Besnier,F., Skaug, H.J., Øien, N., Solvang, H.K., Haug, T., SkernMauritzen, R., Kanda, N., Pastene, L.A., Jonassen, I. andGlover, K.A. 2017. Whole genome resequencing reveals diagnos�c markers for inves�ga�ng global migra�on and hybridiza�on between minke whale species.
BMC Genomics 18: 76.
2019 (1)Takahashi, M., Kato, H. and Kitakado, T. 2019. Sex predic�on by machine learning methods for small sample
sizes: a case study of cranial measurements of killer
whales (Orcinus orca). Mammalian Science 59 (1):67–78 (in Japanese).
2020 (2)Kawasaki, K., Mikami, M., Goto, M., Shindo, J., Amano,
M. and Ishiyama, M. 2020. The evolu�on of unusuallysmall amelogenin genes in cetaceans; pseudogeniza�on, X–Y gene conversion, and feeding strategy. Journal of Molecular Evolution 88 (2): 122–135.
Pastene, L.A., Acevedo, J. and Branch, T.A. 2020. Morphometric analysis of Chilean blue whales and implica�ons
for their taxonomy. Marine Mammal Science 36 (1):
116–135.

79

TEREP-ICR

The InsƟtute of Cetacean Research,
4-5 Toyomi-cho, Chuo-ku, Tokyo 104-0055, Japanwww.icrwhale.org/TEREP-ICR.html