TEREP‐ lCR「 CR

TECHNICAL REpORTS OF THE
INSTITUTE OF CETACEAN RESEARCHTEREP-ICR is published annually by the Institute of Cetacean Research.
ISSN: 2433-6084
Copyright© 2022, The Institute of Cetacean Research {ICR).

Citation
Entire issue:
Institute of Cetacean Research. 2022. Technical Reports of the Institute of Cetacean Research (TEREP-ICR) No. 6. The Institute of Cetacean Research, Tokyo,」apan, 86 pp.
Individual reports:'Author Name'. 2022.'Report title'. Technical Reports of the Institute of Cetacean Research (TEREP­/CR} No. 6:'pp'-'pp'.

Editorial correspondence should be sent to:
Editor, TEREP-ICR
The Institute 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 �TEREP-ICR.html

Cover photo: Fin whales against the background of the Hokkaido coast (top and middle); a com­
mon minke whale near the Japanese coast (bottom).

Copyright© 2022, The Institute of Cetacean Research (ICR).

TECHNICAL REPORTS OF THE
INSTITUTE OF CETACEAN RESEARCHTEREP-ICR
No. 6

The Ins�tute of Cetacean Research (ICR)
Tokyo, 2022

Technical Reports of the Ins�tute of Cetacean Research (2022)

p. i

Foreword
It is a pleasure for me to introduce the sixth issue of the Technical Reports of the Ins�tute of Cetacean Research (TEREPICR-6). Consistent with its stated objec�ves, TEREP-ICR describes and reports on the process, progress, and results oftechnical or scien�ﬁc research, as well as the state of technical or scien�ﬁc research programs conducted by the ICR,
including those commenced recently as is the case of the Japanese Abundance and Stock structure Surveys in the Antarc�c (JASS-A).
The Covid-19 pandemic has con�nued aﬀec�ng the ways of conduc�ng scien�ﬁc research at the ICR. Despite the diﬃcul�es, inconveniences and limita�ons derived from the pandemic, during 2022 the ICR has been able to safely and successfully conducts ﬁeld surveys, conducts laboratory works and analyses, and par�cipates in na�onal and interna�onal
mee�ngs. These eﬀorts enabled the ICR to make signiﬁcant progress on several research works during 2022 as shownin this issue of TEREP-ICR.
Similar as in previous TEREP-ICR issues, TEREP-ICR-5 was widely distributed in Japan and in foreign countries. Notwithstanding the challenges imposed by the pandemic, TEREP-ICR is on course towards achieving its objec�ves. At thesame �me, it has been a good opportunity for our scien�sts to compile and summarize their research conducted over
the years, as a precursor to submi�ng their works for peer review publica�ons.
I sincerely hope that this sixth issue of the TEREP-ICR will contribute further to an increased understanding amongna�onal and interna�onal scien�ﬁc communi�es of the technical and research ac�vi�es conducted by the ICR.
Dr. Yoshihiro Fujise
Director GeneralIns�tute of Cetacean Research
Tokyo, December 2022iTechnical Reports of the Ins�tute of Cetacean Research (2022)

p. iii

Editorial
Welcome to the sixth issue of the Technical Reports of the Ins�tute of Cetacean Research (TEREP-ICR-6).
This issue contains seven technical reports and one commentary ar�cle. In TEREP-ICR-6, we started with the series ofreports summarizing the research ﬁndings on whales and the ecosystem in the western North Paciﬁc Ocean.
Regarding the research ﬁndings in the western North Paciﬁc, Taguchi and colleagues (Part 1) focused on the studies
on stock structure in large baleen whale species. These studies have been possible because the ICR holds one of largest
collec�on of gene�c samples in this oceanic basin, which derived from both past whale research programs under special
permit and biopsy samples collected during dedicated sigh�ng surveys. Takahashi and colleagues (Part 2) focused on the
results of abundance es�mates of baleen whale species based on sigh�ng data collected systema�cally during dedicatedsigh�ng surveys. Bando and colleagues (Part 3) focused on the results of biological parameters in baleen whale species
based on samples and data collected during past whale research programs under special permits.
Results of three important dedicated sigh�ng surveys are summarized in this issue: Isoda and colleagues summarized
the results of the 2021/22 austral summer season survey of the Japanese Abundance and Stock structure Surveys in the
Antarc�c (JASS-A); Katsumata and Matsuoka summarized the results of surveys conducted in the North Paciﬁc Ocean
in 2021; and Murase summarized the results of the 2021 Interna�onal Whaling Commission-Paciﬁc Ocean Whale andEcosystem Research (IWC-POWER) survey.
Konishi and Isoda summarized the results of the long-term track movement of Antarc�c minke whales revealed bysatellite-monitored tagging experiments conducted by the ICR, and examined these preliminary results in the context of
the available stock structure hypothesis derived from gene�c analyses.
In the commentary ar�cle, Gómez Díaz presented his view on the development and relevance of the ICR in the context
of recent whaling events in Japan and the world.
TEREP-ICR-6 issue also includes sec�ons that outline the contribu�on of ICR scien�sts to interna�onal and na�onal
mee�ngs in 2022, as well as their contribu�on in terms of peer-reviewed publica�ons up to December 2022.
We trust that you will ﬁnd this sixth issue informa�ve and useful.
Dr. Luis A. Pastene
Dr. Satoko Inoue
Editorial Team, TEREP-ICR
Tokyo, December 2022

iii

Technical Reports of the Ins�tute of Cetacean Research (2022)

p. v

Contents
Forewordi

Editorialiii

Contentsv

Technical Reports
Taguchi, M., Goto, M. and Pastene, L.A. What do we know about whales and ecosystem in thewestern North Paciﬁc Ocean? Part 1: summary of results on stock structure in baleen whale
species

1

Takahashi, M., Hakamada, T. and Matsuoka, K. What do we know about whales and ecosystem in thewestern North Paciﬁc Ocean? Part 2: summary of results on abundance es�mates in baleen whale
species
11
Bando, T., Maeda, H., Inoue, S. and Kuno, T. What do we know about whales and ecosystem in thewestern North Paciﬁc Ocean? Part 3: summary of results on biological parameters in baleen whale
19
species
Isoda, T., Katsumata, T. and Matsuoka, K. Results of the dedicated sigh�ng survey under the JapaneseAbundance and Stock structure Surveys in the Antarc�c (JASS-A) in the eastern part of Area VI inthe 2021/22 austral summer season
25
Katsumata, T. and Matsuoka, K. Report and highlights of the dedicated sigh�ng surveys in the western North Paciﬁc Ocean in 2021
36
Murase, H. Results of the IWC-Paciﬁc Ocean Whale and Ecosystem Research (IWC-POWER) dedicated sigh�ng survey in 2021—An overview—
46
Konishi, K. and Isoda, T. Long-term track movement of Antarc�c minke whales revealed by satellitemonitored tag experiments conducted by the Ins�tute of Cetacean Research
52

CommentaryGómez Díaz, Gabriel. Some thoughts about the Ins�tute of Cetacean Research

56

Na�onal mee�ngs

59

Interna�onal mee�ngs

61

Peer-reviewed publica�ons

66v

Technical Reports of the Ins�tute of Cetacean Research (2022)

pp. 1–10

Technical Report

What do we know about whales and ecosystem in the western North
Paciﬁc Ocean? Part 1: summary of results on stock structure in baleen
whale species
Mioko Taguchi*, Mutsuo Goto and Luis A. Pastene
Institute of Cetacean Research, 4–5 Toyomi-cho, Chuo-ku, Tokyo 104–0055, Japan*Contact e-mail: taguchi@cetacean.jp

ABSTRACT

Iden�ﬁca�on of biological stocks within a species is fundamental to appropriately interpret demographic parameters. Informa�on on the temporal-spa�al stock structure is crucial in establishing eﬀec�ve management
and conserva�on strategies of marine living resources. This paper summarizes the recent gene�c studies on
stock structure of four baleen whale species in the North Paciﬁc: right, sei, Brydeʼs and common minke whales.
These studies used a large number of gene�c samples collected mainly in surveys conducted by the Ins�tuteof Cetacean Research since 1994. Results of the studies revealed novel stock structures, and this informa�on
is contribu�ng to stock assessments and conserva�on of each whale species in the North Paciﬁc.

INTRODUCTION
Diﬀerent biological stocks could have diﬀerent demographic characteris�cs and respond in diﬀerent ways toenvironmental stress. For this reason, the iden�ﬁca�onof stocks in a species is fundamental for the appropriate
interpreta�on of abundance es�ma�on and other demographic parameters. Informa�on on the temporal-spa�alstock structure is thus crucial in establishing eﬀec�vemanagement and conserva�on strategies. In this regard, Moritz (1994) deﬁned management units (MUs) as
historically isolated sets of popula�ons and suggested
that gene�c markers be used to deﬁne management
units within species: popula�ons with a signiﬁcant divergence of allele frequencies at nuclear or mitochondrial
loci, regardless of the phylogene�c dis�nc�veness of thealleles.
This paper describes the outline of the recent studies
which examine gene�c diversi�es and stock structure
based on the MUs deﬁned by Moritz (1994) for the four
North Paciﬁc whale species: right (Eubalaena japonica),sei (Balaenoptera borealis), Brydeʼs (B. brydei) and common minke whales (B. acutorostrata).

BACKGROUND ON GENETIC STUDIES ON STOCKSTRUCTURE IN NORTH PACIFIC BALEEN WHALES
North Paciﬁc right whales
LeDuc et al. (2012) was the ﬁrst study that focused on
North Paciﬁc right whaleʼs stock structure and gene�c di-versity. The study using the mitochondrial DNA (mtDNA)
control region and six microsatellite DNA (msDNA) loci,mainly showed the poten�al parentage and gene�c diversity in 24 individual whales (23 from the eastern North
Paciﬁc and one from the western North Paciﬁc). Theirresults inferred that North Paciﬁc right whales could be
isolated to some degree between the western and eastern regions based on paternity analysis. However, it wasnoted that the rela�onship of the eastern North Paciﬁcright whales to those in the western North Paciﬁc waslargely unexamined.
Sei whales
A pioneering work on stock structure of North Paciﬁc sei
whales was carried out on the feeding grounds by Wada
and Numachi (1991) using three polymorphic allozyme
loci, which suggested a single stock in the area eastof 160°E. The most comprehensive gene�c study was
performed by Kanda et al. (2009c) using msDNA genotypes at 17 loci and mtDNA control region sequencesfrom a total of 790 specimens collected on the feeding
grounds between 140°E and 135°W during 1972–1973
and 2002–2007. The study also showed no evidence ofgene�c heterogeneity in the research area. Kanda et al.
(2013) subsequently examined gene�c diﬀerences of thisspecies both spa�ally and temporally. The results of thestudy were consistent with those found by Kanda et al.
(2009). In addi�on, Kanda et al. (2015) examined spa�algene�c diﬀeren�a�on using genotypes at 16 msDNA
1

Technical Reports of the Ins�tute of Cetacean Research (2022)

loci by only using samples collected during the summers
in 2010, 2011 and 2012 in order to eliminate temporalnega�ve biases. The study again did not ﬁnd evidence ofmul�ple stocks of sei whales in the North Paciﬁc.
Brydeʼs whales
A series of allozyme studies (Wada and Numachi, 1991;
Wada, 1996) were conducted using Brydeʼs whale
samples collected between 150°E and 160°W, on the
North Paciﬁc feeding grounds. These studies did not ﬁnd
evidence of gene�c heterogeneity in the research area.
A subsequent RFLP analyses inves�gated the pa�ern ofgene�c varia�ons of Brydeʼs whales at intra- and interoceanic levels for mtDNA control region in specimensfrom four loca�ons: western North Paciﬁc, westernSouth Paciﬁc, eastern South Paciﬁc and eastern Indian
Oceans (Pastene et al., 1997). The study showed signiﬁcant gene�c diﬀerences among the four regions, but did
not reveal any signiﬁcant gene�c diﬀerences within thewestern North Paciﬁc Ocean. The most recent popula�ongene�c study was performed by Kanda et al. (2007), using
msDNA genotypes at 17 loci and mtDNA control region
sequences in a sample set used by Pastene et al. (1997),
as well as addi�onal samples from the western North
Paciﬁc feeding grounds. The study conﬁrmed signiﬁcantgene�c diﬀeren�a�ons among oceanic regions, but notwithin the western North Paciﬁc.
Common minke whales
At least two stocks of the common minke whales havebeen historically recognized in the western North Paciﬁc,namely the Okhotsk Sea–West Paciﬁc (known as the Ostock) and the Sea of Japan–Yellow Sea–East China Sea(known as the J-stock). These two stocks are diﬀeren�at-

ed morphologically, reproduc�vely (Omura and Sakiura,
1956; Kato, 1992) and gene�cally (Wada and Numachi,
1991; Goto and Pastene, 1997), which suggests their reproduc�ve isola�on.
Kanda et al. (2009a; b) examined gene�c varia�ons at
16 msDNA loci in a total of 2,542 gene�c samples collected in the waters around Japan and pelagic areas of
the western North Paciﬁc during 1994–2007, and succeeded in the assignment of individual to stocks using a
Bayesian clustering approach, i.e. STRUCTURE analysis.
The study presented the highest likelihood probability at
K=2, which indicated that the samples came from twogene�cally dis�nct popula�ons, i.e. O- and J-stocks.
Pastene et al. (2016a) updated these studies by increasing sample sizes, and showed a ﬁner spa�al distribu�on of each stock (Figure 1): J-stock whales mainly
distributed in the Sea of Japan (sub-areas 6E and 10E), Ostock whales mainly distributed in the oﬀshore North Paciﬁc (east of sub-area 7WR). Both stocks occur along the
Paciﬁc coast of Japan (sub-areas 7CN, 7CS and 2C) and in
the southern Okhotsk Sea (sub-area 11) but sub-area 2C
is mainly occupied by the J-stock whales. Furthermore,
the study showed the temporal distribu�on of the O- andJ-stock whales on the Paciﬁc side of Japan (sub-areas 2C,
7CN and 7CS) as shown in Figure 2: J-stock whales are
predominant throughout the year in sub-area 2C, while
the propor�on of the J-stock whales increases in autumn/
winter and decreases in spring/summer in sub-areas 7CS
and 7CN. The reverse is true for O-stock.
A subsequent study by Pastene et al. (2016b) inves�gated the possibility of addi�onal structure within
O-stock using hypothesis tes�ng and the discriminant
analysis of principal component (DAPC) approach. A
simula�on exercise showed that the sta�s�cal power of

Figure 1. Management sub-areas deﬁned by the IWC for the western North Paciﬁc common minke whales (le�), andspa�al occurrence of O- and J-stocks in sub-areas (right: BC2, BC6, BC7CS, BC7CN, BC10, BC11=bycatches fromthe respec�ve areas; K7CN=coastal JARPNII surveys at Kushiro; S7CS=coastal JARPNII surveys at Sanriku; 7CS,
7CN, 7WR, 7E, 8, 9 and 11=oﬀshore JARPN/JARPNII surveys). Sample sizes are at the top of each bar. Color
indicates stock assignment by STRUCTURE analysis (gray=O-stock, dark gray=J-stock, light gray=unassigned).
2

What do we know about whales and ecosystem in the western North Pacific Ocean? Part 1

Figure 2.

Monthly occurrence of J- and O-stocks in sub-areas 2C (le�), 7CN (middle) and 7CS (le�). Each bar is expressed as three months moving average. Sample sizes are on the top of each bar. Samples from sub-area
2C and subareas 7CN and 7CS were collected during 2001–2014 and 1994–2014, respec�vely. Color indicates stock assignment by STRUCTURE analysis (dark gray=J-stock, gray=O-stock, light gray=unassigned).

the hypothesis tes�ng was high. Both analyses showedno evidence of sub-structuring within O-stock.
Tiedemann et al. (2017) developed a new maximum
likelihood-based approach to infer Parent–Oﬀspring(P–O) rela�onships using full msDNA data at 16 loci ina total of 4,554 western North Paciﬁc common minke
whales including fetuses. The study iden�ﬁed a total of
49 validated P–O pairs (excluding mother-fetus pairs)
across sub-areas, with J-stock pairs on both sides of Japan
closer to the coast, while O-stock pairs were mostly located to the east of Japan, both close to the coast as well as
far oﬀshore. The study provided no evidence for further
stock structure other than O- and J-stocks.

RECENT GENETIC STUDIES ON STOCK STRUCTURE
IN THE NORTH PACIFIC
Basic laboratory procedures
DNA extraction
Total genomic DNA was extracted from 0.05 g of a �ssue
sample using either the standard phenol-chloroform
method (Sambrook et al., 1989) or Gentra Puregene kits
(QIAGEN) and stored in TE buﬀer (10 mM Tris-HCl, 1 mM
EDTA, pH 8.0).
mtDNA
Approximately 500 base pairs of a variable part of the
mtDNA control region were ampliﬁed by PCR using
primers MT4 (Árnason et al., 1993) and Dlp5R (5ʹ-CCA
TCG AGA TGT CTT ATT TAA GGG GAA C-3ʹ; Pastene
et al., 2022). PCR ampliﬁca�on was carried out in 25 µl
reac�on mixtures containing 10–100 ng of template DNA,0.1 µM of each primer, 0.5 units of Ex Taq DNA polymerase (Takara Bio Inc., Shiga, Japan), 0.2 mM of dNTPs,
and 1× Ex Taq buﬀer. Each reac�on was performed withan ini�al denatura�on step at 95°C for 5 min, followed
by 30 cycles of 30 s at 94°C, 30 s at 50°C and 30 s at 72°C,
with a ﬁnal extension step at 72°C for 10 min. PCR products were puriﬁed using MicroSpin S-400HR Columns(GE Healthcare, Illinois, USA). Cycle sequencing was

performed using BigDye terminator cycle sequence Kit(Applied Biosystems Inc., Massachuse�s, USA), following
the manufacturerʼs instruc�ons. The cycle sequencing
products were puriﬁed using AutoSeq G-50 spin Columns(GE Healthcare). The labeled sequencing fragments wereresolved by ABI gene�c analyzers (Applied Biosystems
Inc.) for Sanger sequencing by capillary electrophoresis.
Both strands of all samples were sequenced using theprimers used for ampliﬁca�on. The nucleo�de sequences
obtained were aligned by using the program MUSCLE
implemented in MEGA ver. 10.0.5 (Kumar et al., 2018) or
the program Sequence Navigator.
msDNA
Diﬀerent nuclear msDNA loci were genotyped according
to the whale species (Table 1). The polymerase chain reac�on (PCR) ampliﬁca�ons were performed in 15 µl reac�on mixtures containing 10–100 ng of DNA, 0.25 µM ofeach primer, 0.35 units of Ex Taq DNA polymerase (Takara
Bio Inc.), 0.2 mM of dNTPs, and 1× Ex Taq buﬀer, with
94°C for 2 min, followed by 30 cycles at 94°C for 20 s/54–
61°C for 45 s/72°C for 1 min, and a post-cycling extension
at 72°C for 10 min. Nega�ve and posi�ve control samples,one of each, were run in parallel with all PCR ampliﬁca�ons. The PCR products were run on a BaseSta�on100
DNA fragment analyzer (Bio-Rad, California, USA) withan internal size standard (GENESCAN400HD, Applied Biosystems Inc.), or on an ABI 3500 DNA Analyzer (Applied
Biosystems Inc.) with a 600 LIZ size standard (Applied Biosystems Inc.). In the former pla�orm, alleles were visualized using Cartographer so�ware speciﬁcally designed for
the BaseSta�on, and the fragment sizes were determinedmanually in rela�on to the internal size standard and aposi�ve control sample of known size that were run oneach gel. In the other pla�orm, each allele was determined by automated binning and assigned to allelic bins
predeﬁned in GeneMapper v. 4.0 (Applied Biosystems
Inc.). Any misassigned scores were corrected by hand.
If msDNA scores generated from both pla�orms were3

Technical Reports of the Ins�tute of Cetacean Research (2022)
Table 1
msDNA markers used in the recent studies on four North Paciﬁc baleen whales.
Norht Paciﬁc right whale

Sei whale

Brydeʼs whale

Common minke whale

EV1
EV14
EV21
EV37
EV94
GT023
GT211
GT310
GATA028
DlrFCB17
TR3G2
TR2G5
TR3F2

EV94
EV1
GT211
EV14
EV21
GGAA520
GATA53GT23
GT575
GATA417GT310
EV104
GATA28
GT271
GT011
DlrFCB17
GATA98

DlrFCB14
EV1
EV104
EV14
EV21
EV94
GATA28GATA417
GATA53
GATA98GGAA520
GT011
GT23
GT310
GT575
TAA31

EV1
EV14
EV21
EV37
EV94
GATA28GATA98GATA417
TAA31
GT23
GT195
GT211
GT310
GT509
GT575
DlrFCB14

Table 2Gene�c analyses used in the recent studies on four North Paciﬁc baleen whales.
Analysis
Gene�c diversity indices
HWE test
Hypothesis test
FST es�mates
G'ST es�mates
AMOVA
Mantel test
STRUCTUREDAPCsPCAGENELANDBAPS
TESS
Phylogene�c analyses
PO analysis

North Paciﬁc right whale

Sei whale

Common minke whale

mtDNA

msDNA

mtDNA

msDNA

mtDNA

msDNA

mtDNA

msDNA

●
●
●
●

●
●

●
●
●
●

●
●
●
●

●
●
●
●

●
●
●
●

●
●
●
●

●

●

●

●
●
●
●
●
●
●

●
●

●

contained in a single dataset, then all msDNA scores generated by the ABI 3500 DNA Analyzer were standardized
to those by the BaseSta�on100 DNA fragment analyzer.
Basic analy�cal procedures
The analy�cal procedures used in the recent gene�c studies on popula�on structure are shown in Table 2 for eachwhale species.
This sec�on brieﬂy explains these analy�cal procedures.
4

Brydeʼ s whale

●
●
●
●
●
●

●
●

Genetic diversity indices
mtDNA haplotype (h) and nucleo�de (π) diversi�es(Nei, 1987) were es�mated with sample standard devia�ons. The number of alleles (A), inbreeding coeﬃcient(FIS) and expected heterozygosity (HE) were es�mated per
msDNA locus and across loci. The departure from Hardy–
Weinberg equilibrium (HWE) was also tested for each
msDNA locus and across loci.

What do we know about whales and ecosystem in the western North Pacific Ocean? Part 1

Genetic diﬀerentiation
The gene�c diﬀerence between popula�ons was examined by several sta�s�cal tests for diﬀerences in allele/
haplotype frequencies between popula�ons, as well as
by pairwise FST es�mates and GʼST values (Hedrick, 2005).
The Analysis of Molecular Variance (AMOVA) (Excoﬃer
et al., 1992) was used to examine the hierarchical gene�cstructure. A Mantel test was performed between a matrixof pairwise Edwardʼs gene�c distances and a matrix of
Euclidian geographic distances to examine the pa�ern of
isola�on by distance (IBD).
Genetic clustering and stock assignment approaches
Es�ma�on of the most likely number of clusters and/or
individual-based stock assignment were performed usinga Bayesian clustering analysis with an admixture model
assuming correlated allele frequencies implemented in
STRUCTURE program (Pritchard et al., 2000).
To iden�fy and describe clusters of gene�cally related
individuals, a Discriminant Analysis of Principal Components (DAPC; Jombart et al., 2010), which relies on datatransforma�on using Principal Components Analysis as aprior step to discriminant analysis, was performed with orwithout a priori group assignments for msDNA genotype
set. A priori group was determined based on geographic
sampling site.
The spa�ally explicit clustering approaches, e.g. spa�al Analysis of Principal Components (sPCA; Jombart
et al., 2008), GENELAND (Guillot et al., 2005), TESS(François et al., 2006; Chen et al., 2007) and BAPS (Corander et al., 2008), have been also used to inves�gate
gene�c structure and stock assignment.
Phylogeographic analysis
A sta�s�cal parsimony network (Clement et al., 2000)
was generated to infer the phylogene�c rela�onships
among mtDNA haplotypes.
Parent–oﬀspring analysis
Parent–Oﬀspring (P–O) rela�onships were inferred
based on a maximum-likelihood approach (Tiedemannet al., 2017). mtDNA haplotypes and biological informa�on, e.g. sex, sexual maturity and sampling posi�on,were used to interpret the inferred P–O pairs. The geographical distribu�on of the pairs was discussed from theview point of stock structure.

Main results
North Paciﬁc right whales
Dataset
Pastene et al. (2022) summarized and analyzed all available gene�c data for North Paciﬁc right whales. A�erremoving all duplicates (i.e. samples that were collectedfrom the same individual) and one sample from eachmother and calf pair, a single dataset was generated forthe mtDNA sta�s�cal analyses, which consisted of 30
samples each in the western North Paciﬁc (WNP: 27 newsamples, one from LeDuc et al. [2012] and two historicalbaleen samples) and the eastern North Paciﬁc (ENP: 21
samples from LeDuc et al. [2012], three historical baleensamples and six new samples), respec�vely (Figure 3);
the sample size for the msDNA analysis was 19 whales
for WNP.
Main results and discussion
The study found a total of 16 haplotypes among 60
individual whales. Only four haplotypes were shared between WNP and ENP right whales. The FST between WNP
and ENP right whales was high (0.128) and sta�s�callysigniﬁcant, and the heterogeneity test resulted in sta�s�-

Figure 3.

Approximate core distribu�on of North Paciﬁc rightwhales in summer (July–August) based on 19th
century whaling records and 20th century sigh�ng
and whaling catch data compiled by Clapham et al.
(2004). The pa�ern of distribu�on shows high density in the western and eastern sides of the North
Paciﬁc (purple) and low density in the central area
(white). The monthly plots of right whale sigh�ngs
and catches by Clapham et al. (2004) suggesteda pa�ern of south–north migratory movement inspring (March–May) to summer (June–August)on both sides of the North Paciﬁc. The ﬁgure also
shows the distribu�on of gene�c samples examinedin the recent study (Pastene et al., 2022). Mostof the samples from the eastern side are from
LeDuc et al. (2012). Symbol color indicates sexes(red=female, blue=male, black=uniden�ﬁed).
5Technical Reports of the Ins�tute of Cetacean Research (2022)

cally signiﬁcant diﬀerences between whales on both sidesof the North Paciﬁc; however, there was no concordance
between geography and haplotype network pa�ern.
The number of msDNA alleles per locus ranged from
2 to 7 (4.08 on average), and the HE ranged from 0.285
to 0.787 (0.593 on average). The FIS in each locus ranged
from 0.045 to 0.350 with 0.032 on average. The study
found no signiﬁcant devia�ons from HWE across loci.
The mtDNA results, sugges�ng some degree of popula�on structuring, were consistent with the pa�ern of catchand sigh�ng data showing higher densi�es on either sideof the North Paciﬁc, but li�le in between as shown in Figure 3 (Clapham et al., 2004). These ﬁndings support the
hypothesis of diﬀerent popula�ons occurring in the eastern and western sides of the North Paciﬁc. The authors
also suggested the possibility of incomplete lineage sorting of ancestral polymorphisms from a lack of structure in
haplotype network.
An alterna�ve interpreta�on on these results was that
there is a single interbreeding popula�on in the North
Paciﬁc that is exhibi�ng mtDNA structuring as a result of
matrilineally driven seasonal site ﬁdelity. Although theauthors could not rule out the alterna�ve interpreta�on,they considered, as the more plausible, the hypothesisof two discrete breeding popula�ons of right whales onboth sides of the North Paciﬁc since such hypothesis is
supported also by the pa�ern of catches and sigh�ng
distribu�on, and diﬀerent catch and recovery histories(Brownell et al., 2001; Clapham et al., 2004).
Sei whales
DatasetA single dataset consis�ng of mtDNA control region
sequences (n=1,733) and msDNA genotypes at 17 loci(n=1,729) was generated from specimens of sei whales
collected from diﬀerent sources (Figure 3: Appendix 7
in Tamura et al., 2019). The gene�c data generated fromthese samples were divided into three areas for the purpose of data analyses (Figure 4).
Main results and discussion
Pairwise FST es�mates between areas showed noevidence of gene�c heterogeneity in North Paciﬁc sei
whales for both gene�c markers. These results were also
supported by AMOVA and STRUCTURE analyses, whichshowed a lack of gene�c structuring of this species in theoceanic areas of the North Paciﬁc. These ﬁndings werealso consistent with results of comparable gene�c diversi�es among areas as well as the results from the tests of
HWE and FIS es�mates. Overall, the recent reﬁned analyses supported a single stock hypothesis as suggested
6

Figure 4.

Sampling posi�on of sei whales used in the recent gene�c analyses (Appendix 7 in Tamura
et al., 2019). Color indicates sample source; yellow:
historical commercial whaling during 1972–1973,
red: JARPNII during 2002–2016, and blue: POWER
during 2010–2013. Solid red line shows subareas.
The gene�c data was stra�ﬁed according to thethree areas, i.e. western, central and eastern, fordata analyses.

Figure 5.

Sampling posi�on of Brydeʼs whales used in thegene�c analysis in Taguchi et al. (in press). Color
indicates sample source; yellow: historical commercial whaling operated in 1979 and 1983–1984,
red: JARPNII during 2000–2016, blue: POWER during 2013–2016, green: dedicated sigh�ng survey
in 2012 and 2014, and gray: bycatch in 2010. Areas
enclosed by solid and dashed lines are sample
stra�ﬁca�on according to management areas forthe western North Paciﬁc Brydeʼs whales deﬁned
by the IWC scien�ﬁc commi�ee (IWC, 2008; 2018).
Mul�ple gene�c samples that were collected outside of these areas were assigned to the longitudinally closest area, i.e. area 2.

previously.
Brydeʼs whales
DatasetIn the most recent study (Taguchi et al., in press), a total of
1,182 msDNA genotype sets at 17 loci and 1,195 mtDNA
control region sequences were analyzed (Figure 5). These
data were divided into the three management areas forthe purpose of data analyses (Figure 5).

What do we know about whales and ecosystem in the western North Pacific Ocean? Part 1

Main results and discussion
According to Taguchi et al. (in press), gene�c diversi�eswere similar among areas and a haplotype network did
not show any geographic structure. However, an AMOVA
found evidence of gene�c structure in this species. Pairwise FST and GʼST es�mates, as well as heterogeneity tests,a�ributed this gene�c structure to weak, but signiﬁcantdiﬀeren�a�on between areas 1W and 2. Furthermore,
a Mantel test and a high-resolu�on analysis of gene�cdiversity sta�s�cs showed a weak spa�al cline of gene�c
diﬀeren�a�on.
Given that this study focused on whales in their feeding grounds, these ﬁndings could be reconciled by two
possible scenarios: (1) the occurrence of a single popula�on with kin-associa�on being responsible for feeding
ground preference, and (2) the occurrence of two popula�ons, with feeding ground preference for either area 1Wor area 2; both popula�ons mix geographically througharea 1E. The results from the DAPC were not inconsistent
with these scenarios. Given an es�mated dispersal rateof less than 2% between areas 1W and 2, which indicatessome demographic independency among whales in these
two areas, both scenarios should be considered based on
the precau�onary principle in stock assessments.
Common minke whales
Recent gene�c works have focused on reﬁning the twostock hypothesis, O- and J-stocks, as well as inves�ga�ngwhether addi�onal structure exists within each stock.
DatasetRecent studies have been based on mtDNA control region
sequences (n=4,706) and msDNA genotype sets at 16 loci(n=4,707). The gene�c samples were collected during theJapanese Whale Research Programs under Special Per-

Figure 6.

mit in the western North Paciﬁc, Phases I and II (JARPN/
JARPNII) conducted systema�cally in the western North
Paciﬁc during spring–summer from 1994 to 2016, as well
as from bycatches which occurred along the Japanese
coast from 2001 to 2016.
Main results and discussion
P–O inferences and their geographical distribu�on
Goto et al. (2019) updated the analysis by Tiedemannet al. (2017), and iden�ﬁed a total of 40 and 13 P–O
pairs for O- and J-stocks, respec�vely. The O-stock pairs
were found between coastal and oﬀshore waters, while
J-stock pairs were distributed within and between the Sea
of Japan and the Paciﬁc side of Japan (Figure 6). These
results provided no evidence for further stock structureother than O- and J-stocks.
Spa�ally explicit clustering analyses
The results of sPCA in conjunc�on with those of DPAC
suggested the occurrence of the O- and J-stocks but
provided no evidence for addi�onal structure in eachstock (Taguchi et al., 2019a), which was also supportedby the mtDNA analyses. These ﬁndings indicated a low
possibility that mul�ple stocks exist (other than the J- andO-stocks) in the study area with overlapping geographic
ranges.
A study by de Jong and Hoelzel (2019) using threeother spa�ally explicit clustering tools including GENELAND, TESS and BAPS showed diﬀerent pa�erns of clustering, and the authors stated that the most informa�ve
approach was GENELAND using the mixture model withcorrelated allele frequency model which supported K=4.
However, an addi�onal work subsequently undertaken by
Taguchi et al. (2019b) examined the correspondence ofthe four clusters by GENELAND with the available gene�cand non-gene�c informa�on, and concluded that theDistribu�on of parent–oﬀspring pairs of O- (le�) and J-stocks (right) in western North Paciﬁc common
minke whale (blue square: parent, orange circle: mature oﬀspring, red cross: immature or unknown oﬀspring).
7

Technical Reports of the Ins�tute of Cetacean Research (2022)

most plausible scenario was for two popula�ons (O- andJ-stocks) with complex spa�al and temporal mixing along
the Paciﬁc coast of Japan. de Jong and Hoelzel (2019)
further noted that some of the analyses conducted wereconsistent with a scenario of coastal areas containing gene�cally admixed individuals, and recommended further
analyses under the GENELAND as well under the TESS
and BAPS.

CONCLUDING REMARKSOver the last two decades, many gene�c studies focusedon stock iden�ﬁca�on and structure in western North
Paciﬁc baleen whales have been undertaken. The driving force behind these analyses was to obtain informa�on to assist in determining eﬀec�ve conserva�on andmanagement. Of necessity, these studies were based ongene�c samples collected on feeding grounds and migratory corridors. In this context, they are based on theconcept of MUs described by Moritz (1994). The recent
studies summarized in this paper used several increasingly sophis�cated clustering approaches for the purposeof iden�fying stocks, which showed sta�s�cal results
consistent with this criterion for deﬁning stocks and contributed to their stock assessment. It is also important tomake eﬀort to inves�gate the stock structure of baleen
whales distributed in unsurveyed areas including breeding grounds, and the gene�c rela�onship of them withwhales distributed around Japan.

FUTURE WORKS
Gene�c samples for other baleen whale species in thewestern North Paciﬁc (i.e. humpback whales, ﬁn whales
and blue whales) are also available at the ICR. The gene�cdata generated from these samples will be analyzed in
the near future to inves�gate their stock structure.
The development of a new gene�c marker, which isless error-prone than msDNA, i.e. SNP (Single Nucleo�dePolymorphism) is on-going at the ICR. This marker will be
used to perform the gene�c analyses for all whale species
of interest. Such analyses will aid in a be�er understanding of stock structure of baleen whales in the western
North Paciﬁc.

ACKNOWLEDGEMENTS
We thank all of the whalers and researchers who contributed to the collec�on of samples. We are grateful to
all technicians for their laboratory works at the ICR. We
would also like to thank Naohisa Kanda who organized
most of the msDNA data used in the studies summarized
in this paper. We thank the Editorial Team of TEREP-ICR
8

for the editorial work on this manuscript.

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Technical Reports of the Ins�tute of Cetacean Research (2022)

pp. 11–18

Technical Report

What do we know about whales and ecosystem in the western North
Paciﬁc Ocean? Part 2: summary of results on abundance es�mates inbaleen whale species
Megumi Takahashi*, Takashi Hakamada and Koji Matsuoka
Institute of Cetacean Research, 4–5 Toyomi-cho, Chuo-ku, Tokyo 104–0055, Japan*Contact e-mail: takahashi@cetacean.jp

ABSTRACTThe Ins�tute of Cetacean Research (ICR) has been conduc�ng whale sigh�ng surveys in the western North
Paciﬁc Ocean for more than 25 years. The main purpose of these surveys is the collec�on of sigh�ng datarequired for abundance es�ma�on of baleen whale species in this oceanic basin. Results of the analyses ofthe data collected have revealed the pa�ern of distribu�on and abundance es�mates of each whale species in
diﬀerent years and in diﬀerent periods within a year. Abundance es�mates have been used for management
purposes (e.g. calcula�on of catch limits for sustainable commercial whaling) in the case of some species. This
paper presents a brief outline of the survey design and analy�cal procedures used, and of the main results
obtained so far.

INTRODUCTION
Informa�on regarding the distribu�on and abundance ofwhale species is essen�al in understanding the role andimpact of whales in the ecosystem. Abundance is also an
important piece of informa�on required for establishingpolicy of management of whales. For example, a �meseries of es�mates of absolute abundance derived fromsigh�ng surveys is required under the Revised Management Procedure (RMP) for se�ng safe catch limitsof commercial whaling. The RMP is the management
procedure developed by the Interna�onal Whaling Commissionʼs Scien�ﬁc Commi�ee (IWC SC) for baleen whale
species.
One of the most common approaches for collec�ngsigh�ng data for abundance es�ma�on purposes is thevessel-based survey using the Line Transect (LT) method.
Japan through the Ins�tute of Cetacean Research (ICR)
has conducted systema�c research on whales in the western North Paciﬁc for more than 25 years. The ﬁrst one
was the Japanese Whale Research Program under Special
Permit in the western North Paciﬁc (JARPN, 1994–1999),
which was followed by the second phase of this program(JARPNII, 2000–2016). Subsequently, the New Scien�ﬁcWhale Research Program in the western North Paciﬁc(NEWREP-NP) was designed and implemented between
2017–2019. All these three research programs (JARPN,
JARPNII, NEWREP-NP) had an important sigh�ng survey

component based on the LT method. Abundance es�mates based on sigh�ng data collected by this survey
component have been crucial in the context of some ofthe objec�ves of those programs, e.g. the es�ma�on of
prey consump�on by whale species. As noted above,abundance es�mates are key informa�on for calcula�ng
catch limit for commercial whaling on some species.
The objec�ve of this paper is to presents a brief outlineof the survey design and analy�cal procedures used by
the ICR sigh�ng surveys in the western North Paciﬁc, andto present the main results obtained so far.

RESEARCH AREA
The research area for the sigh�ng surveys comprises the
Paciﬁc side of Japan, north of 35°N, the Sea of Japan, andOkhotsk Sea (see Figure 1). Each of these research areas
were divided into smaller strata, and the surveys wereconducted in spring (mainly May to June) and summer(mainly August to September) seasons to inves�gate
whale distribu�on and abundance in each season.

VESSEL USED
Specialized vessels which have three observa�on platforms were used in the surveys (Figure 2). Observers collected whale sigh�ng data from the pla�orms using binoculars in the �me period between sunrise and sunset.

11Technical Reports of the Ins�tute of Cetacean Research (2022)

SURVEY METHODOLOGY
Sigh�ng surveys were conducted by the LT method. The
survey protocols followed the IWC SCʼs Requirements andGuidelines for Conduc�ng Surveys and Analyzing Data
within the Revised Management Scheme (IWC, 2012a).
Under the LT method, vessels move on pre-determinedzigzag track lines, which cover the en�re or a part of amanagement area (Figure 3A). The start point is randomly
selected for each survey. The design of track lines does

not follow physical features such as isobaths which may
be correlated with whale distribu�on and their migra�on.
Observers search for whales from each pla�orm on the
research vessels. When a whale school is detected, thedistance between observers and whales and the angle
from the track line to whales were recorded (Figure 3B)
together with the posi�on of the research vessel and �meof the sigh�ng.

ANALYTICAL PROCEDURES
Abundance es�ma�on using sigh�ng data is made using
distance sampling calcula�on (Buckland et al., 2001).
The methodology involves two main components. The
ﬁrst component involves ﬁ�ng the detec�on func�on tosigh�ng data. Ini�ally it is assumed that whales are uniformly distributed and that they have the same probability to be detected. However, searching ac�vity is aﬀectedby environmental factors (e.g. glare, fog, sea state), andwhales occurring far from the vessel are more diﬃcult
to detect (Figure 4). Therefore, detec�on func�on (e.g.
Half-normal func�on and Hazard-rate func�on) should
be examined, and the best model is selected using the
Akaike Informa�on Criterion (AIC) values.

Figure 1.

12

Research area on the Paciﬁc side of Japan, southern
Okhotsk Sea and Sea of Japan, covered by Japanesesigh�ng surveys.

Figure 2.

Specialized research vessels used for sigh�ng surveys (le�: Kaiyo-Maru No.7, right: Yushin-Maru No.2). The
ﬁgures show the observa�on pla�orms (taken from Takahashi [2020] and Isoda et al. [2021]).

Figure 3.

Diagrams showing the zigzag track lines for a research area under the LT survey (A); and the whale observa�on from diﬀerent pla�orms (B).

What do we know about whales and ecosystem in the western North Pacific Ocean? Part 2

Figure 4.

Diagram showing the detec�on probability assuming g(0)=1. The probability of detec�on decreases
with distance from the vessel.

 x2 
Half-normal: f ( x, z ) = exp  − 2  2σ 
 − x  − b 
Hazard-rate: f ( x, z ) = 1 − exp  ,
 aσ  
where x is perpendicular distance, z is vector of several
covariates; and σ represents scale parameter for incorpora�ng the covariates in the detec�on func�on and a andb are parameters of each key func�on.
The second component is for es�ma�ng abundance
using the detec�on probability derived from the detec�on func�on. The Horvitz–Thompson-like es�mator is
one of the most frequently used methods. The principleof the es�mator is that the total number of whales present are es�mated from the propor�on of the abundancein the area surveyed (the covered area) to the en�re survey area. To do this the abundance in the area surveyed is
calculated using the detec�on probability. The detec�onfunc�on models are denoted f(x; _) where x is perpendicular distance (from the track line to whales) and θ is avector of parameters of the key func�on to be es�mated.
Nˆ covered =

n


i=1

w 1

 w
0

si
, zi ; θ )dx
f ( x

,where w is trunca�on distance from the track line, si issize of ith detected cluster (I=1, …, n).
Then, it is scaled up to the full study area by mul�plying it by the ra�o of covered area to study area. Here,
A/2wL represents propor�on of en�re research area toarea searched:

A 
Nˆ =
N
2wL covered
In reality, because of diving behavior, not all animals
are available for detec�on even when they are near the

Figure 5.

The searching eﬀort by each Lat. 1°×Long. 1° gridsquare covered by JARPN and JARPNII surveys during
1994 to 2014 (Matsuoka and Hakamada, 2019). The
gray scale shows the degree of searching eﬀort.

vessel and/or on the track line. Therefore, this bias should
also be considered in the abundance es�ma�on process.
This bias is known as availability bias and is diﬀerent frompercep�on bias caused by human error under diﬀerent
environmental condi�ons.

RESULTS ON DISTRIBUTION AND ABUNDANCE
Figure 5 shows the main research area and the total primary searching eﬀort by each Lat. 1°×Long. 1° grid square
during 1994 to 2014 under JARPN and JARPNII surveys.
Table 1 shows the abundance es�mates for each whale
species. The numbers of whales distributed in the JARPNII
survey area were es�mated in the early and late seasons(see Hakamada and Matsuoka, 2016a; 2016b).
Blue whales
The main distribu�on area for blue whales moved northward from 35°N–40°N to north of 40°N during the spring(May to June) to summer (August to September) seasons(Matsuoka et al., 2016) (Figure 6). In the late 20th century, they had never been observed in this area in June,
but this could have been caused by the diﬀerence inregula�ons of the whaling opera�ons for coastal and oﬀshore whaling. In the late season, blue whales were more
frequently sighted north of 40°N (Figure 6), and the abundance es�mate was 958 animals (CV= 0.461) (Table 1).
Fin whales
Fin whales were observed mainly in the Paciﬁc side ofJapan but also occurred in the Okhotsk Sea and Sea ofJapan (Matsuoka et al., 2016). The main distribu�onarea was north of 37°N, and the highest density area wasnorth of 45°N. Fin whales were widely distributed acrossthe western North Paciﬁc in the spring season, but they13

Technical Reports of the Ins�tute of Cetacean Research (2022)
Table 1Abundance es�mates for several large baleen whale species. All es�mates assumed g(0)=1, except for common minke whales inwhich g(0)=0.798 (CV=0.134).
Spring season

SpeciesBlue whale
Fin whale
Sei whale
Bryde's whale
Humpback whale
North Paciﬁc right whale
Common minke whale

Summer season

20092011+20122008

38 (0.977)
413 (0.569)
4,734 (0.177)
2,957 (0.394)
1,136 (0.438)
0 (—)
3,629 (0.586)

161 (0.474)
1,369 (0.295)
2,988 (0.304)
1,851 (0.413)
1,921 (0.318)
1,147 (0.434)
2,122 (0.371)

958 (0.461)
3,958 (0.425)
5,086 (0.378)
13,306 (0.251)
392 (0.877)
416 (0.653)
3,080 (0.677)

References
Hakamada and Matsuoka (2016b)
Hakamada and Matsuoka (2016b)
Hakamada and Matsuoka (2016a)
Hakamada and Matsuoka (2016a)
Hakamada and Matsuoka (2016b)
Hakamada and Matsuoka (2016b)
Hakamada and Matsuoka (2016a)

Spring season: May to June, Summer season: August to September.

Figure 6.

Blue whales observed during the surveys (top). Sigh�ng posi�ons of blue whales (blue circles) on the tracklines, by survey period (bo�om).

were more frequently sighted north of 40°N in the summer season (Figure 7), and the abundance was es�matedat 3,958 animals (CV= 0.425) in this season (Table 1).
Humpback whales
The main distribu�on area of humpback whales was
north of 37°N from May to September (Matsuoka
et al., 2016). The highest density area was observed northof 35°N (west of 158°E) and north of 45°N (east of 158°E).
They moved northward from 37°N to 43°N in the inshore
area of Japan during May to June, and to further northof 45°N during July to August (Figure 8). The abundancees�mates were 1,136 (CV= 0.438) to 1,921 (CV= 0.318) inthe early seasons. Despite the surveys covering the en�rewestern North Paciﬁc in the summer season, sigh�ngs of
14

this species were not frequent in this season (Table 1).
North Paciﬁc right whales
During the survey programs, 48 schools (68 individuals)
were observed north of 37°N from May to September(Matsuoka et al., 2016) (Figure 9). The total abundance
was es�mated to be 1,147 animals (CV= 0.434) basedon the JARPNII data in the spring season of 2011/12(Table 1).
Sei whales
The main distribu�on area of sei whales shi�ed northward during spring to summer seasons (Murase et al.,2016; Figure 10). The abundance es�mate in the summer season was higher than that in the spring season

What do we know about whales and ecosystem in the western North Pacific Ocean? Part 2

Figure 7.

Fin whales observed during the surveys (top). Sigh�ng posi�ons of ﬁn whales (red circles) on the tracklines, by survey period (bo�om).

Figure 8.

Humpback whales observed during the surveys (top). Sigh�ng posi�ons of humpback whales (light blue
circles) on the track lines, by survey period (bo�om).

(Table 1). The es�mates should be considered as a part of
the total stock number of this species (see Taguchi et al.,this issue).
Brydeʼs whales
The main distribu�on of Brydeʼs whale shi�ed northward during the spring to summer season (Hakamada
et al., 2009). Their distribu�on was in lower la�tudes
compared to other large baleen whale species (Figure 9). Table 1 shows the abundance es�mates for
Brydeʼs whales in the spring and summer (July to September) seasons in this JARPNII area.

Common minke whales
Common minke whales are known to migrate further
north of the JARPNII survey area through early season tolate season (Hakamada et al., 2009) (Figure 12). Common
minke whales distribute primarily in the Sea of Okhotsk
and in the waters east of the Kamchatka Peninsula andthe Kuril Islands in August and September (Buckland et al,
1992; Miyashita, 2019). The es�mated numbers distributed in the survey area were 3,629 (CV= 0.586) and 2,122(CV= 0.371) in the spring season and 3,080 (CV= 0.677) inthe summer season (Table 1).

15

Technical Reports of the Ins�tute of Cetacean Research (2022)

Figure 9.

North Paciﬁc right whales observed during the surveys (top). Sigh�ng posi�ons of North Paciﬁc rightwhales (purple circles) on the track lines, by survey period (bo�om).

Figure 10.

Sei whales observed during the surveys (top). Sigh�ng posi�ons of sei whales (orange circles) on the tracklines, by survey period (bo�om).

USE OF THE ABUNDANCE ESTIMATES FOR MANAGEMENT PURPOSES
Japan resumed commercial whaling of North Paciﬁc sei,
Brydeʼs and western North Paciﬁc common minke whales
in July 2019. Catch limits for each species were set inline with the RMP (IWC, 2012b). The core component of
the RMP is the Catch Limit Algorithm (CLA) (Aldrin andHuseby, 2007; Aldrin et al., 2008), which is a feedback
16

control algorithm that sets baleen whale harvest levels
based on catch histories and a �me series of es�matesof absolute abundance derived from sigh�ng surveys.
Implementation Simulation Trials (ISTs) are conducted toassess uncertain�es for several parameters in the case ofmul�-stock scenarios.
Abundance es�mates from ICR sigh�ng surveys havebeen fundamental in the process of applying the CLA
and ISTs to the three North Paciﬁc baleen whale species.

What do we know about whales and ecosystem in the western North Pacific Ocean? Part 2

Figure 11.

Brydeʼs whales observed during the surveys (top). Sigh�ng posi�ons of Brydeʼs whales (green circles) on
the track lines, by survey period (bo�om).

Figure 12.

Common minke whales observed during the surveys (top). Sigh�ng posi�ons of common minke whales(purple circles) on the track lines, by survey period (bo�om).

For species like the sei and Brydeʼs whales, total stock
abundances have been obtained by combining the ICR es�mates in the western North Paciﬁc with es�mates fromother surveys in the central and eastern North Paciﬁc.
See details of the process for catch limit calcula�on in
Japanʼs RMP Team (2019).

FINAL REMARKSThe ICR has conducted sigh�ng surveys for more than
25 years, in collabora�on with other ins�tu�ons, usinga systema�c survey procedure. The surveys revealed the
distribu�on and abundance of several large baleen whale

species in the western North Paciﬁc Ocean. Moreover,abundance es�mates from the ICR surveys contributed
to the process for calcula�ng catch limits for sustainable
commercial whaling of sei, Brydeʼs and common minke
whales. There are several aspects that could improve theabundance es�mates in the future. Among them are thees�mates of g(0) for all species, evalua�on of addi�onal
variance for mul�-year surveys and the use of modelbased approaches.

ACKNOWLEDGEMENTS
We thank all researchers and crew members who par17

Technical Reports of the Ins�tute of Cetacean Research (2022)

�cipated in the ICR sigh�ng surveys for their hard work incollec�ng data in the ﬁeld. Our thanks are also extended
to all researchers who assisted in the analyses. We are
grateful to the Fisheries Agency of Japan for providing
funds for the surveys. Our gra�tude to Luis A. Pastene(ICR) for his assistance in preparing this manuscript. We
also thank the Editorial Team of TEREP-ICR for the editorial work on this manuscript.

REFERENCES
Aldrin, M. and Huseby, R.B. 2007. Simula�on trials 2007 for a retuned Catch Limit Algorithm. Paper SC/59/RMP4 presented
to the IWC Scien�ﬁc Commi�ee, May 2007 (unpublished).
143 pp. [Available from the IWC Secretariat].
Aldrin, M., Huseby, R.B. and Schweder, T. 2008. A note on tuning
the Catch Limit Algorithm for commercial baleen whaling. J.
Cetacean Res. Manage. 10 (3): 191–194.
Buckland, S.T., Ca�anash, K.L. and Miyashita, T. 1992. Minke
whale abundance in the northwest Paciﬁc and the Okhotsk
Sea, es�mated from 1989 and 1990 sigh�ng surveys. Rep. int.
Whal. Commn 42: 387–392.
Buckland, S.T., Anderson, D.R., Burnham, K.P., Laake, J.L.,
Borchers, D.L. and Thomas, L. 2001. Introduction to DistanceSampling. Oxford University Press, Oxford. 595 pp.
Hakamada, T., Matsuoka, K. and Miyashita, T. 2009. Distribu�on
and the number of western North Paciﬁc common minke,
Brydeʼs, sei and sperm whales distributed in JARPNII Oﬀshore
component survey area. Paper SC/J09/JR15 presented to the
JARPNII Review Mee�ng, January 2009 (unpublished). 18 pp.
[Available from the IWC Secretariat].
Hakamada, T. and Matsuoka, K. 2016a. The number of western
North Paciﬁc common minke, Brydeʼs and sei whales distributed in JARPNII Oﬀshore survey area. Paper SC/F16/JR12
presented to the JARPNII Review Mee�ng, February 2016
(unpublished). 14 pp. [Available from the IWC Secretariat].
Hakamada, T. and Matsuoka, K. 2016b. The number of blue,
ﬁn, humpback, and North Paciﬁc right whales in the western North Paciﬁc in the JARPNII Oﬀshore survey area. Paper
SC/F16/JR13 presented to the JARPNII Review Mee�ng,
February 2016 (unpublished). 14 pp. [Available from the IWC
Secretariat].
Matsuoka, K. and Hakamada, T. 2019. Distribu�on of blue,18

ﬁn, humpback and North Paciﬁc right whales in the western North Paciﬁc based on JARPN and JARPNII surveys(1994–2014). Technical Reports of the Institute of Cetacean
Research (TEREP-ICR) No.3: 41–51.
Interna�onal Whaling Commission. 2012a. Requirements andguidelines for conduc�ng surveys and analysing data within
the revised management scheme. J. Cetacean Res. Manage.
(Suppl.) 13: 509–517.
Interna�onal Whaling Commission. 2012b. Requirements
and guidelines for Implementations under the Revised
Management Procedure (RMP). J. Cetacean Res. Manage.
(Suppl.) 13: 497–505.
Isoda, T., Katsumata, T. and Matsuoka, K. 2021. Results of thededicated sigh�ng survey under the Japanese Abundance
and Stock structure Surveys in the Antarc�c (JASS-A) in thewestern part of Area III in the 2020/21 austral summer season. Technical Reports of the Institute of Cetacean Research(TEREP-ICR) No.5: 7–15.
Japanʼs RMP Team. 2019. Catch limits for western North Paciﬁcsei, Brydeʼs and common minke whales calculated in linewith the IWCʼs Revised Management Procedure (RMP).
h�ps://www.jfa.maﬀ.go.jp/j/whale/a�ach/pdf/index-63.pdf.
Matsuoka, K., Hakamada, T. and Miyashita, T. 2016. Distribu�onof blue (Balaenoptera musculus), ﬁn (B. physalus), humpback(Megaptera novaeangliae) and north paciﬁc right (Eubalaena
japonica) whales in the western North Paciﬁc based on
JARPN and JARPNII surveys (1994 to 2014). Paper SC/F16/JR9
presented to the JARPNII Review Mee�ng, February 2016
(unpublished). 12 pp. [Available from the IWC Secretariat].
Miyashita, T. 2019. Abundance es�mate of common minke
whales in sub-areas 11, 10E and 7CN in 2014. Paper SC/68A/
ASI15 presented to the IWC Scien�ﬁc Commi�ee, May 2019
(unpublished). 3 pp. [Available from the IWC Secretariat].
Murase, H., Hakamada, T., Sasaki, H., Matsuoka, K. and Kitakado,
T. 2016. Seasonal spa�al distribu�ons of common minke, sei
and Brydeʼs whales in the JARPNII survey area from 2002 to
2013. Paper SC/F16/JR7 presented to the JARPNII Review
Mee�ng, February 2016 (unpublished). 17 pp. [Available
from the IWC Secretariat].
Takahashi, M. 2020. A note on g(0) es�mates derived from vessel-based sigh�ng surveys. Technical Reports of the Instituteof Cetacean Research (TEREP-ICR) No.3: 14–20.

Technical Reports of the Ins�tute of Cetacean Research (2022)

pp. 19–24

Technical Report (not peer reviewed)

What do we know about whales and ecosystem in the western North
Paciﬁc Ocean? Part 3: summary of results on biological parameters inbaleen whale species
Takeharu Bando1, *, Hikari Maeda2, Satoko Inoue1 and Tomoe Kuno1
1

Institute of Cetacean Research, 4–5 Toyomi-cho, Chuo-ku, Tokyo 104–0055, Japan
Fisheries Resources Institute, 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

ABSTRACT

Three baleen whale species, sei, Brydeʼs and common minke whales were surveyed during the former Japanese scien�ﬁc whaling programs in the western North Paciﬁc Ocean conducted from 1994 to 2019. This paper
summarizes some life history-related biological parameters es�mated for the three species, based on dataand samples collected during the research programs. These biological parameters are important as input data
for models aimed at studying the popula�on dynamics of the species. Results therefore will contribute to the
management of these whale species in the western North Paciﬁc.

INTRODUCTION
Three baleen whale species, sei (Balaenoptera borealis),
Brydeʼs (B. edeni brydei) and common minke (B. acutorostrata scammoni) whales were surveyed by former
Japanese scien�ﬁc whaling program in the western
North Paciﬁc conducted by the Ins�tute of Cetacean Research (ICR). The target species for the Japanese Whale
Research Program under Special Permit in the North Paciﬁc (JARPN) (1994–1999) was the common minke whales(GOJ, 1994). The second phase of this program (JARPNII)(2000–2016) targeted Brydeʼs, common minke and sei
whales. The la�er was added as a target species in 2002
(GOJ, 2000; 2002).
A�er the comple�on of the JARPNII, the New Scien�ﬁcWhale Research Program in the western North Paciﬁc(NEWREP-NP) commenced in 2017 and lasted un�l 2019
(GOJ, 2017). The target species were sei and common
minke whales. Figure 1 shows the baleen whale species
targeted by the former Japanese scien�ﬁc whaling programs in the western North Paciﬁc.
During these research programs, systema�c biologicalsampling was conducted on the whales taken. These included earplugs for age determina�on, testes and ovaries
for determina�on of sexual maturity and es�ma�on of
reproduc�ve history, fetus for reproduc�ve studies andbody length for growth studies. Some life history-related
biological parameters, which are important as input datafor popula�on dynamics models, were es�mated for thethree whale species based on data and samples collectedduring the research programs.
This paper summarizes the results of biologicalparameter es�ma�ons in sei, Brydeʼs and common
minke whales in the western North Paciﬁc. It follows
more detailed analyses presented previously in Bando(2017; 2018), Bando and Maeda (2020) and Maeda et al.
(2017).

SUMMARY OF BIOLOGICAL PARAMETERS IN
THREE BALEEN WHALE SPECIESLaboratory and analy�cal procedures to es�mate biological parameters in baleen whales were explained in previous ar�cles (Bando, 2017; 2018; Bando and Maeda, 2020;
Maeda et al., 2017). Details of these approaches are notrepeated here for brevity. The results of biological parameter es�ma�ons for the three species in the western
North Paciﬁc are presented in this sec�on.
Earplug age readability
In many baleen whale species, ages are es�mated bycoun�ng the number of growth layers accumulated in theearplugs which are formed in the external auditory meatus (Lockyer, 1984a). During the scien�ﬁc whaling programs, earplug age readability had improved comparedto the reading based on earplug collected from pastcommercial whaling. This was due to careful sampling
19

Technical Reports of the Ins�tute of Cetacean Research (2022)

Figure 1.

Baleen whale species targeted by the former Japanese scien�ﬁc whaling programs in the western North
Paciﬁc: sei whale (upper le�); Brydeʼs whale (upper right); and common minke whale (lower).

of earplugs by experienced researchers and introduc�onof a new gela�nized extrac�on method during the scien�ﬁc whaling programs (Bando, 2021; Bando and Maeda,
2020; Maeda, et al., 2013).
Sei whales
Bando and Maeda (2020) reported earplug age readability of immature sei whales as 55.9% and 48.9% for males
and females, respec�vely. Readability in sexually mature
animals was higher. The values were 72.5% and 67.4% formales and females, respec�vely.
Brydeʼs whales
Similar readability values were reported for Brydeʼswhales. They were 43.8% and 40.8% for immature males
and females, respec�vely, and 74.0% and 76.4% for mature males and females, respec�vely (Bando, 2021).
Common minke whales
In the case of common minke whale, the readability
was somewhat lower. They were 35.4% and 34.7% for
immature males and females, respec�vely, and 49.2%
and 59.0% for mature males and females, respec�vely(Maeda et al., 2016).
Growth curve
The growth curve of each whale species was es�mated by
20

ﬁ�ng the von Bertalanﬀy growth model to body lengthand age.
Sei whales
The growth curve of sei whales was es�mated as Lt =14.14(1−e−0.174(t+6.650)) for males, and Lt =15.17(1−e−0.150(t+7.407))
for females (Bando and Maeda, 2020).
Brydeʼs whales
The asympto�c length of Brydeʼs whales was slightly
smaller than that of sei whale. The growth curve was es�mated as Lt =12.65(1−e−0.189(t+5.250)) for males, and Lt =13.30(1−e−0.170(t+4.929)) for females (Bando, 2021).
Common minke whales
The asympto�c length of common minke whales wasmuch smaller than the other two species. The growth
curve was es�mated as Lt =7.49(1−e−0.41(t+0.90)) for males,and Lt =8.66(1−e−0.11(t+7.60)) for females (Maeda, 2012).
In the three whale species, the asympto�c length was
greater in females than in males.
Maximum life span
Sei whales
Bando and Maeda (2020) reported the maximum age ofsei whales (es�mated from earplug reading) as 54 years
for males and 43 years for females.

What do we know about whales and ecosystem in the western North Pacific Ocean? Part 3

Brydeʼs whales
Bando (2021) reported the maximum age of Brydeʼswhales as 51 and 56 years for males and females, respec�vely.
Common minke whales
Maeda et al. (2016) reported maximum age of maleand female common minke whales as 49 and 41 years,
respec�vely.
Age at sexual maturity
There are several methods to es�mate age at sexual
maturity of baleen whales (Lockyer, 1984b). Age at 50%
sexual maturity (tm50%) is es�mated by calcula�ng thera�o of immature and mature whales in each age. Age at
ﬁrst ovula�on (tmov) is es�mated by mean age of whales
with one corpus luteum and no corpus albicans in bothovaries. It is generally known that the transi�on phase inearplugs of large baleen whales indicates sexual maturity(Lockyer, 1972; Masaki, 1979; Kato, 1983; Ohsumi, 1986)
and age at sexual maturity (tmp) is es�mated by mean
age of whales when the transi�on phase is formed.
Sei whales
Bando and Maeda (2020) es�mated tm50% of sei whales
as 6.7 (SE= 0.29) years for males and 6.9 (SE= 0.27) years
for females, respec�vely. Tmov was es�mated as 8.6 (SD=3.14) years (Bando, 2017).
Brydeʼs whales
For Brydeʼs whales, tm50% were es�mated as 7.72 (SE=0.49) and 8.56 (SE= 0.39) years for males and females,
respec�vely (Bando, 2021). Tmov was es�mated as 10.1
(SD= 1.13) years (Bando, 2017).
Common minke whales
Tmp was es�mated as 7.68 (SD= 1.81) years for males and7.82 (SD= 1.58) years for females (Maeda et al, 2017).
Tmov was es�mated as 8.75 (SD= 1.5) years (Maeda etal, 2017).
Body length at sexual maturity
Body length at 50% sexual maturity (Lm50%) and ﬁrst
ovula�on (Lmov) are es�mated in the same manner as
the age at 50% sexual maturity (tm50%) and age at ﬁrst
ovula�on (tmov). Lm50% is es�mated by calcula�ng thera�o of immature and mature whales in each body lengthclass. Lmov is es�mated by mean body length of whales
with one corpus luteum and no corpus albicans in bothovaries.

Sei whales
Lm50% of sei whales was es�mated as 12.72 m (SE= 0.12)
for males and 13.31 m (SE= 0.12) for females (Bando and
Maeda, 2020). Lmov was es�mated as 13.70 m (SD= 0.47)(Bando, 2017).
Brydeʼs whales
Lm50% of Brydeʼs whales was slightly smaller than that ofsei whale, 11.41 m (SE= 0.25) for males and 11.75 m (SE=0.23) for females (Bando, 2021). Lmov was es�mated as
12.13 m (SD= 0.62) (Bando, 2017).
Common minke whales
There are no available es�mates of Lm50% and Lmov forcommon minke whales. Common minke whales are known
to segregate in the western North Paciﬁc by sex and sexual
maturity status (Zenitani et al., 2000a; 2000b). This fact will
bias the es�mate of length at sexual maturity.
Pregnancy rate
Pregnancy rates are usually calculated as the propor�onof pregnant females among captured mature females.
However, in some cases, this value may diﬀer from thetrue pregnancy rate due to biases such as segrega�on andthe eﬀect of mother and calf pairʼs occurrence.
Sei and Brydeʼs whales
Most sei and Brydeʼs whales are thought to give birth
every two or more years. Females accompanied by calf
are considered to have a low pregnancy rate because
they were pregnant in the previous year. If females withcalf were excluded from samples, pregnancy rate may beoveres�mated. Bando (2017) es�mated pregnancy rateof sei and Brydeʼs whales with correc�on of the eﬀect ofmother and calf pair (which were not targeted for sampling in most years) as 0.650 and 0.491, respec�vely.
Common minke whales
Pregnancy rates of common minke whale have not beenes�mated due to the small number of mature female
samples. Most of the mature females distribute outsideof the research area (Hatanaka and Miyashita, 1997;
Wada, 1989).
Concep�on Date
In baleen whales, growth of fetus a�er the early stage ofdevelopment is considered to be linear with �me, andmethods for es�ma�ng concep�on date from foetal lengthand date of capture has been developed (Hugge� andWiddas, 1951; Kato and Miyashita, 1991; Lockyer, 1984b).
21

22
72.5% (n=466)
68.4% (n=618)

MatureTotal

tmp
13.70 m0.650

13.31 mReferences

Bando (2017)
Bando and Maeda (2020)

1.007

0.481

Li�er size

0.744/year
0.519

Annual ovula�on rate

Peak from December to February

12.72 m—Foetal sex ra�oConcep�on datePregnancy rate

Reproductive characteristics

Lm50%

—

8.6 years
6.9 years

tm50%

43 years

Lt =15.17(1−e−0.150(t+7.407))

63.0% (n=736)

67.4% (n=562)

48.9% (n=174)

Femaletmov
6.7 years

54 years

Lt =14.14(1−e−0.174(t+6.650))

55.9% (n=152)

Immature

Body length at sexual maturity Lmov

Age at sexual maturity

Sexual maturity

Maximum life span

Growth curve

GrowthEarplug readability

Age reading

Male

Sei whale

0.491

11.75 m12.13 m—8.56 years

10.1 years

56 years

Lt =13.30(1−e−0.170(t+4.929))

66.1% (n=416)

76.4% (n=296)

40.8% (n=120)

Female

1.011

0.467

0.526/year

Bando (2017); (2018); (2021)

0.533

Year round with peak from October to May

11.41 m—7.72 years

51 years

Lt =12.65(1−e−0.189(t+5.250))

63.9% (n=313)

74.0% (n=208)

43.8% (n=105)

Male

Brydeʼs whale

Table 1
Summary of biological parameters of three baleen whale species in the western North Paciﬁc.

—

—

—

7.82 years—

8.75 years

41 years

Lt =8.66(1−e−0.11(t+7.60))

41.2% (n=684)

59.0% (n=183)

34.7% (n=501)

Female———Bando et al. (2010)
Goto et al. (2021)
Maeda (2012)
Maeda et al. (2016); (2017)—

Peak from autum to winter (J-stock)
One peak in winter (O-stock)——

7.68 years—

49 years

Lt =7.49(1−e−0.41(t+0.90))

45.2% (n=1,888)

49.2% (n=1,340)

35.4% (n=548)

MaleCommon minke whale

Technical Reports of the Ins�tute of Cetacean Research (2022)

What do we know about whales and ecosystem in the western North Pacific Ocean? Part 3

Sei and Brydeʼs whales
Bando (2017) calculated concep�on dates based on theanalysis of fetuses from 414 sei and 177 Brydeʼs whales.
He reported that the concep�on season of sei whales
lasts from November to March, with a peak from December to February, and that of Brydeʼs whales is year-round,with a moderate peak from October to May.

Common minke whales
There are no es�mates available for common minke
whales.
Table 1 summarizes the es�mates of life history-related
biological parameters in sei, Brydeʼs and common minke
whales in the western North Paciﬁc.

Common minke whales
As for minke whales, two biological stocks are known
to exist in the western North Paciﬁc, the J and O stocks(see reviews in Goto et al., 2017; 2021). Both stocks arereported to have diﬀerent concep�on seasons, the Ostock from November to June with a single winter peak,
and the J-stock from August to March with a peak fromautumn to winter (Bando et al., 2010, Goto et al., 2021).

Several biological parameters of three baleen whale
species were es�mated from the samples collected byscien�ﬁc whaling programs conducted by the ICR. During these surveys, a wide range of whale samples, fromyoung to old, were collected by random sampling method
which led to a more accurate es�ma�on of biologicalparameters than that es�mated from past commercial
whaling data. Some biological parameters respond tochanges in their environment such as food availabilityor fecundity, which will aﬀect the status of whale stocks.
Biological parameters are also important as input data
for models aimed at studying the popula�on dynamics
of the species. The es�mated biological parameters willcontribute to the management of these whale species in
the western North Paciﬁc Ocean.

Annual ovula�on rate
Annual ovula�on rate was es�mated from regression ofage and number of corpora in the ovaries.
Sei and Brydeʼs whales
Annual ovula�on rate was es�mated as 0.744/year for sei
whales and 0.526/year for Brydeʼs whales, respec�vely(Bando and Maeda, 2020; Bando, 2021).
Common minke whales
There is no es�mate available for common minke whales.
Foetal sex ra�o
Sei and Brydeʼs whales
Bando (2017) reported foetal sex ra�o (male ra�o) of sei
whales as 51.9% (n=418) and that of Brydesʼs whales as
53.3% (n=169). No signiﬁcant diﬀerence from parity was
detected for sei and Brydeʼs whales.
Common minke whales
There are no es�mates available for common minke
whales.
Li�er size
In baleen whale species, the number of fetuses is usually
one. However, mul�ple fetuses (twins in most cases) are
observed occasionally (Kimura, 1957; Lockyer, 1984b).
Sei and Brydeʼs whalesLi�er sizes were es�mated as 1.007 for sei whales (n=420),
and 1.011 for Brydeʼs whale (n=177) (Bando, 2017).

CONCLUDING REMARKSACKNOWLEDGEMENTS
Our sincere thanks to all researchers and crews who collected data and samples during the JARPN, JARPNII and
NEWREP-NP surveys. We thank the members of the Ins�tute of Cetacean Research (ICR) and Japan Fisheries Research and Educa�on Agency for their useful sugges�ons
and assistance in the laboratory work. We also thank Luis
A. Pastene (ICR) for his assistance in preparing this report.
We also thank the Editorial Team of TEREP-ICR for theeditorial work on this manuscript.

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thesis, Tokyo University of Marine Science and Technology.
147 pp. (in Japanese).
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based on samples collected by JARPNII. Technical Reports of
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Bando, T. 2021. Improved es�mates of some life-history parameters of the pelagic subspecies of Brydeʼs whale in the western North Paciﬁc. Cetacean Population Studies 3: 239–245.
Bando, T. and Maeda, H. 2020. Progress in the research on earplug-based age determina�on and biological parameters ofNorth Paciﬁc sei whales at the Ins�tute of Cetacean Research.
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557–564.
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306–317.
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Kato, H. and Miyashita, T. 1991. Migra�on strategy of southernminke whales in rela�on to reproduc�ve cycle es�mated24

from foetal length. Rep. int. Whal. Commn 41: 363–369.
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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
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[Available from the IWC Secretariat].
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minke whale as an indicator of age at sexual maturity. Open
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Masaki, Y. 1979. Yearly change of the biological parameters
for the Antarc�c minke whale. Rep. int. Whal. Commn 29:
375–395.
Ohsumi, S. 1986. Earplug transi�on phase as an indicator ofsexual maturity in female Antarc�c minke whales. Sci. Rep.
Whales Res. Inst. 37: 17–30.
Wada, S. 1989. La�tudinal segrega�on of the Okhotsk Sea –West Paciﬁc stock of minke whales. Rep. int. Whal. Commn
39: 229–233.
Zenitani, R., Fujise, Y. and Kato, H. 2000a. Some analyses on
biological parameters of western North Paciﬁc minke whales,from a viewpoint of stock iden�ﬁca�on. Paper SC/F2K/J13
presented to the IWC Scien�ﬁc Commi�ee Workshop to
Review the Japanese Whale Research Programme underSpecial Permit for North Paciﬁc minke whales (JARPN),
February 2000 (unpublished). 18 pp. [Available from the IWC
Secretariat].
Zenitani, R., Fujise, Y., Okamura, H. and Kato, H. 2000b. Further
examina�on of the distribu�on of western North Paciﬁcminke whales applying a logis�c regression analysis for reproduc�ve data collected by the JARPN surveys. Paper SC/52/
RMP17 presented to the IWC Scien�ﬁc Commi�ee, May 2000
(unpublished). 8 pp. [Available from the IWC Secretariat].

Technical Reports of the Ins�tute of Cetacean Research (2022)

pp. 25–35

Technical Report (not peer reviewed)

Results of the dedicated sigh�ng survey under the JapaneseAbundance and Stock structure Surveys in the Antarc�c (JASS-A) inthe eastern part of Area VI in the 2021/2022 austral summer season
Tatsuya Isoda*, Taiki Katsumata and Koji Matsuoka
Institute of Cetacean Research, 4–5 Toyomi-cho, Chuo-ku, Tokyo 104–0055, Japan*Contact e-mail: isoda@cetacean.jp

ABSTRACTThe results of the sigh�ng survey of the Japanese Abundance and Stock structure Surveys in the Antarc�c(JASS-A) in the 2021/2022 austral summer season are reported. A dedicated sigh�ng vessel was engaged in
the line transect method survey in the eastern part of Antarc�c Area VI East (130°W–120°W) for 33 days, from
11 January to 12 February 2022. For the survey, the research area was divided into northern and southern
strata. In addi�on, surveys were conducted successfully in coastal ice-free waters, south of 72°S, an area thatis normally covered by pack-ice and diﬃcult for vessels to access. The total searching distance in the researcharea was 1,333.5 n.miles (2,469.6 km). Four baleen whale species and at least four toothed whale species weresighted in the research area. Other research ac�vi�es such as biopsy sampling, photo-ID, satellite tagging, andoceanographic observa�ons were also conducted. The data and samples collected are required for the mainand secondary research objec�ves of JASS-A.

Long-term systema�c surveys on whales and the ecosystem in the Antarc�c, such as the JARPA/JARPAII1, NEWREPA2, and IDCR/SOWER3, obtained important data pertaining
to the study of abundance and abundance trends of large
whales and their biology as well as the role of whales in
the Antarc�c ecosystem. All these research programs havebeen terminated. The last NEWREP-A survey was carriedout in the 2018/2019 austral summer season.
The Japanese Abundance and Stock structure Surveys
in the Antarc�c (JASS-A) commenced in the 2019/2020
austral summer season because it was considered important to con�nue with the whale and ecosystem surveys
in the Indo-Paciﬁc region of the Antarc�c Ocean throughdedicated sigh�ng surveys and other non-lethal researchtechniques. JASS-A has two main research objec�ves,i) the study of the abundance and abundance trends oflarge whale species, and ii) the study of the distribu�on,movement, and stock structure of large whale species.
JASS-A also has several secondary research objec�ves
related to oceanography, marine debris, gene�c data toes�mate abundance, whale biology, and study on the u�lity of Unmanned Aerial Vehicle (UAV). The JASS-A program
was presented to the 2019 mee�ng of IWC SC4 (GOJ,
2019a), the 2019 mee�ng of CCAMLR-EMM5 (GOJ, 2019b),
and the 2019 mee�ng of NAMMCO SC6 (GOJ, 2019c).
The approach of JASS-A is systema�c vessel-basedsigh�ng surveys u�lizing the ʻline transect methodʼ. Surveys are designed and conducted following the protocols
included in the ʻRequirements and Guidelines for Conduc�ng Surveys and Analysing Data within the Revised
Management Schemeʼ (IWC, 2012). Sigh�ng protocols
are the same as those used in the former IDCR/SOWER
surveys (Matsuoka et al., 2003). The JASS-A surveys are
conducted alterna�vely in IWC Management Areas III, IV,
V, and VI by one or two specialized vessels, over a tenta�ve period of eight austral summer seasons.

1

4

INTRODUCTION

Japanese Whale Research Programs under Special Permit in
the Antarc�c, Phases I and II
2
New Scien�ﬁc Whale Research Program in the Antarc�c
Ocean
3
Interna�onal Decade for Cetacean Research/Southern Ocean
Whale and Ecosystem ResearchInterna�onal Whaling Commission-Scien�ﬁc Commi�eeCommission for the Conserva�on of Antarc�c Marine Living
Resources-Working Group on Ecosystem Monitoring andManagement
6
North Atlan�c Marine Mammal Commission-Scien�ﬁc Commi�ee
525

Technical Reports of the Ins�tute of Cetacean Research (2022)

The ﬁrst and second JASS-A surveys were carried out inthe 2019/2020 and 2020/2021 austral summer seasonsand covered the sector 000°–035°E of Antarc�c Area
III West. The third survey was carried out during the2021/2022 season and covered the sector 130°W–120°W
of Antarc�c Area VI East. This paper presents a summaryof the 2021/2022 JASS-A survey results. The Appendix
shows photographs of whales, experiments and Antarc�cenvironment taken during the 2021/2022 survey.

SURVEY DESIGN
Research area
The research area of JASS-A is comprised of IWC Management Areas III, IV, V, and VI, south of 60°S (Figure 1). The
research area in the 2021/2022 season was the easternpart of Antarc�c Area VI East (130°W–120°W), south of
60°S (Figure 1). The area was divided into northern andsouthern strata. The boundary between these strata wasdeﬁned by a line 45 n.miles from the northern edge of the
pack-ice (Figure 2). In addi�on, an area of ice-free waters(polynya) was formed between the southern edge of the
pack-ice and the ice shelf, south of 72°S in early January(Figure 2). Details of the ice conﬁgura�on are shown in
Figure 3. The ice-free waters (in the range of 125°W to
110°W) became accessible to the vessel in early February.

in the survey. They had experience in conduc�ng linetransect surveys, biopsy sampling, photo-iden�ﬁca�on(photo-ID), satellite tagging, and oceanographic survey
in the Antarc�c through the previous JARPA/JARPAII and
NEWREP-A programs.
Sigh�ng procedures, mode and experiments
See Isoda et al. (2022) for details of the procedures used
for sigh�ng surveys and other research ac�vi�es such assigh�ng distance and angle experiment, photo-ID, biopsy
sampling, satellite tagging, oceanographic survey, marine
debris observa�on, and survey using UAV.

RESULTS OF THE SURVEY
Narra�ve of the survey
Table 1 shows the i�nerary of the survey. The dura�onof this cruise was 109 days. The YS2 departed Shiogama,
Japan on 3 December 2021, arriving back at the homeport on 20 December 2021. The YS2 started the sighting survey in Antarc�c Area VI East at 68°37ʹS; 120°00ʹW
on 11 January. The survey was completed at 73°51ʹS;
120°00ʹW on 12 February. The YS2 arrived back at thehome port on 1 March and in Japan on 21 March.

Research vessel
The dedicated sigh�ng vessel Yushin-Maru No. 2 (YS2)
was engaged in the survey. Its speciﬁca�ons are indicatedin Figure 4. Three Japanese researchers par�cipatedFigure 2.

Figure 1.

26

Research area of JASS-A. The shaded area
(130°W–120°W) indicates the surveyed area in the2021/2022 austral summer season.

Research area (130°W–120°W) indica�ng northernand southern strata, and searching eﬀorts (blacklines) of the JASS-A survey in 2021/2022 australsummer season. The research commenced at
68°37ʹS; 120°00ʹW and ended at 73°51ʹS; 120°00ʹW.
Note the ice-free waters south of 72°S.

Results of the dedicated sighting survey under the Japanese Abundance and Stock structure Surveys in the Antarctic (JASS-A)

Figure 3.

Maps of the pack-ice distribu�ons in the research area for dates 5 January (upper le�), 15 January(lower le�), 25 January (upper right) and 5 February (lower right) 2022, constructed by Japan Aerospace
Explora�on Agency (JAXA), based on observa�onal data acquired by the Advanced Microwave ScanningRadiometer 2 (AMSR2). Note that the ice-free waters became accessible to the vessel in early February.

Figure 4.

The dedicated sigh�ng vessel Yushin-Maru No. 2 and its three observa�on pla�orms: top barrel pla�orm(TOP), Independent Observer Pla�orm (IOP) and Upper Bridge Pla�orm (UBP).

Research eﬀort in the research area
Table 2 shows a summary of the searching eﬀort spent
during the survey. The YS2 was engaged in the researchfor 33 days, from 11 January to 12 February 2022. The
total searching eﬀort was 1,333.5 n.miles (2,469.6 km);659.0 n.miles in NSP mode during 60 hour 27 minutes ofresearch and 674.4 n.miles in IO mode during 62 hour 21
minutes of research. In the northern stratum, the total
searching eﬀort was 731.2 n.miles (NSP: 322.7 n.miles;
IO: 408.5 n.miles), and the searching eﬀort coverage was62%. In the southern stratum, the total searching eﬀort

was 254.6 n.miles (NSP: 145.8 n.miles; IO: 108.8 n.miles),and the searching eﬀort coverage was 92%. In the coastal
ice-free waters, the total searching eﬀort was 347.7 n.
miles (NSP: 190.5 n.miles; IO: 157.2 n.miles), and thesearching eﬀort coverage was 83%.
Therefore, a good distribu�on of eﬀort within both
strata and survey mode was achieved. The total experimental �me for photo-ID, biopsy sampling, tagging, anddistance and angle experiment was 22 hours 00 minutes.

27

Technical Reports of the Ins�tute of Cetacean Research (2022)
Table 1
I�nerary of the 2021/2022 JASS-A dedicated sigh�ng survey.
Date (y/m/d)

Event2021/11/16
2021/12/2
2021/12/3
2021/12/13
2021/12/20
2022/1/11
2022/2/12
2022/3/1
2022/3/9
2022/3/21

Planning mee�ng at Tokyo, Japan
Pre-cruise mee�ng in Shiogama, JapanYS2 departed Shiogama, Japan
Transit survey started at 0°00ʹ; 163°00ʹE
YS2 arrive at the home port
Transit survey ﬁnished and survey started in the research area at 68°37ʹS; 120°00ʹW
Survey completed in the research area (33 days) and transit survey start at 73°51ʹS; 120°00ʹW
YS2 arrive at the home port
Transit survey completed at 0°00ʹ; 163°00ʹE
YS2 arrived in Japan and post cruise mee�ng carried out in Shiogama, Japan

Figure 5(a) Geographical distribu�on of primary sigh�ngs of Antarc�c minke, ﬁn, humpback, and Antarc�c blue
whales. (b) The distribu�on of Antarc�c minke whales (pink circle) in the ice-free waters. Rolling 32 days average sea surface temperature (le�) and chlorophyll concentra�on (right) from 17 January to 17 February 2022
were observed by MODIS-Aqua and MODIS-Terra (Original data: Ocean colour web, from h�ps://oceancolor.
gsfc.nasa.gov/ (Accessed 2022-7-30)). Note that sea surface temperature was lower and chlorophyll concentra�on was higher in the western part, where the density of Antarc�c minke whales was high.

Whale sigh�ngs in the research area
Four baleen whale species and at least four toothed
whale species were sighted in the research area. The
dominant whale species in the research area was the Antarc�c minke whale (83 schools/142 individuals) followed
28

by the ﬁn whale (44/64). Sigh�ngs of other species wereas follows: humpback (26/36), Antarc�c blue (5/6), sperm(9/10), Arnouxʼs beaked (1/8), southern bo�lenose (1/1),
killer (5/71, include Types B, C and undetermined type)
and Ziphiidae (4/6) whales (Table 3).

Results of the dedicated sighting survey under the Japanese Abundance and Stock structure Surveys in the Antarctic (JASS-A)
Table 2
Summary of searching eﬀort (�me and distance) and �me (hours) spent during the 2021/2022 JASS-A survey.
Searching eﬀort
Date and �me

Experiment �me(distance [n.miles][hours:minutes:seconds]

and �me [hours:minutes:seconds])
Survey Sec�ons

Es�mated

Photo-ID,
Start

End

NSP

Transit survey

2021/12/13

2021/12/18(0°00–Entering foreign countries EEZ)

14:59

18:00

Transit survey

2021/12/26

2022/1/11(Leaving foreign countries EEZ–Research area)

8:35

9:02

Research area

2022/1/11

2022/2/1(Area VIE 130°W–120°W)

9:02

18:08

Naviga�on/transit survey

2022/2/1

2022/2/8

to the coastal ice-free waters, south of 72°S

18:08

14:06

Coastal ice-free waters, south of 72°S

2022/2/8

2022/2/12(130°W–120°W)

14:06

10:03

Transit survey

2022/2/12

2022/2/22(Research area–Entering foreign countries EEZ)

10:03

17:05

Transit survey

2022/3/8

2022/3/9(Leaving foreign countries EEZ–0°00)

6:30

12:05

2021/12/13

2022/3/9

14:59

12:05

Total

IO

Biopsy,angle andSatellite tag

distance training/

experimentexperiment124.2

10:44:29——

0:14:08—579.9

50:46:32——

2:55:24

1:40:57

468.5

43:18:40

517.2

47:36:56

13:18:08

3:44:27

—

—

—

—

3:56:13—190.5

17:08:29

157.2

14:44:45

1:02:08—782.2

67:39:37——

6:20:12—177.9

14:45:22

—

—

—

—

2323.2

204:23:09

674.4

62:21:41

27:46:13

5:25:24

Table 3
Number of sigh�ngs made during the 2021/22 JASS-A survey in the research area, by stratum and species.
Eastern part of Area VIE (130°W–120°W)
Southern stratum

Northern stratum

Prim.

Prim.

SpeciesSecond.

Sch.

Ind.

Sch.

Ind.

Antarc�c blue whale

0

0

0

0

Fin whale

0

0

0

0

Like ﬁn

0

0

0

0

Antarc�c minke whale

4

6

0

0

Like minke

0

0

0

0

Humpback whale

0

0

0

0

Like humpback

0

0

0

0

Baleen whale

1

1

0

0

Sperm whale

5

5

0

Southern bo�lenose whale

1

1

Arnouxʼs beaked whale

00Killer whale (undetermined type)

0

Killer whale (type B)
Killer whale (type C)

Sch.

Sub-total

Coastal ice-free waters,

Totalsouth of 72°S

Second.

Prim.

Second.

Ind.

Sch.

Ind.

Sch.

Ind.

Sch.

Ind.

4

5

1

1

0

0

0

0

41

60

3

4

0

0

0

0

1

1

0

0

0

0

0

0

11

13

1

1

64

119

3

3

0

0

0

0

2

2

1

2

18

24

8

12

0

0

0

0

2

2

0

0

0

0

0

0

2

2

0

0

0

0

0

0

0

3

4

1

1

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

1

8

0

0

0

0

0

0

0

3

1

21

0

0

0

0

0

0

0

0

0

0

0

0

0

0

Ziphiidae

1

1

0

0

2

3

1

Uniden�ﬁed whale

0

0

0

0

3

3

0

Prim.
Sch.

Second.

Ind.

Sch.

Ind.

Sch.

Ind.

4

5

1

1

5

6

41

60

3

4

44

64

1

1

0

0

1

1

79

138

4

4

83

142

2

2

1

2

3

4

18

24

8

12

26

36

2

2

0

0

2

2

3

3

0

0

3

3

0

8

9

1

1

9

10

0

0

1

1

0

0

1

1

0

0

1

8

0

0

1

8

44

0

0

3

44

0

0

3

44

0

0

0

0

1

21

0

0

1

21

1

6

0

0

1

6

0

0

1

6

2

0

0

0

0

3

4

1

2

4

6

0

0

0

0

0

3

3

0

0

3

3

Prim.: primary sigh�ng, Second.: secondary sigh�ng, Sch.: school, Ind.: individual.

29

Technical Reports of the Ins�tute of Cetacean Research (2022)
Table 4
Summary of the results of experiments conducted during the 2021/2022 JASS-A survey.
Experiments

Results and descrip�ons

Sigh�ng distance and angle experiment

128 trials completed by 30 January

Photo-ID

Obtained from 7 Antarc�c blue, 9 humpback and 10 killer whales

Biopsy sampling

Collected from 2 Antarc�c blue, 12 ﬁn, 15 Antarc�c minke, 11 humpback, 1 Brydeʼswhales and 2 killer whales

Satellite tagging

Deployed on 9 ﬁn and 15 Antarc�c minke whales

Oceanographic survey

116 XCTD casts

Marine debris observa�on

A ﬁshing buoy was observed in the research area

UAV

Aerial images collected from 2 Antarc�c blue, 6 Antarc�c minke and 3 humpback whales

Antarctic minke whales
Antarc�c minke whales were widely distributed in the
research area (Figure 5a). It was the only species sighted
in the ice-free waters with high density. This observa�on
is consistent with the interpreta�on of Fujise and Pastene(2021) that larger numbers of this species are being distributed in polynyas within the pack-ice in recent years,
possibly in search of alterna�ve feeding areas in responseto the increase in abundance and geographical expansion of other large whale species (e.g. humpback andﬁn whales). A sigh�ng survey using an icebreaker vessel(Ainley et al., 2007) found no Antarc�c minke whales
within coastal polynyas in an area similar to that surveyed
in the present survey. However, this earlier survey was
conducted in 1994 and the distribu�on of this species inpolynyas is postulated for recent years.
The density of Antarc�c minke whales in ice-free waters was generally high. However, the number of sigh�ngs
east of 125°W (eastern side) was no�ceably lower. The
sea surface temperature (SST) map shows that the SST in
the eastern side was rela�vely high and the chlorophyll
concentra�on was rela�vely low compared to the western side (Figure 5b). In addi�on, the water colour in theeastern side was clear, while the colour in the westernside was brownish and muddy sugges�ng waters withhigher produc�vity. Therefore, a possible explana�on forthe lower whale density on the eastern side is that waters
on this side are less produc�ve, as suggested by low chlorophyll concentra�on, which could be translated into less
phytoplankton and less krill for the whales to feed. Futuredetailed analysis of the XCTD observa�on data will lead toa be�er understanding of environmental factors aﬀec�ng
the Antarc�c minke whale distribu�on in ice-free waters.
Fin whales
Fin whales were widely distributed only in the northern
30

stratum (Figure 5a). The density was higher than that in
the previous 2000/2001 SOWER survey in the same sector (Ensor et al., 2001). The abundance of this species
appears to be increasing, as suggested for Antarc�c Areas
V+ VI West (Matsuoka and Hakamada, 2014).
Humpback whales
Humpback whales were widely distributed in the northern stratum with higher concentra�ons observed in thesouthern part of this stratum (Figure 5a). The density was
higher than that in the 2000/2001 SOWER cruise in thesame sector (Ensor et al., 2001), conﬁrming the view that
this species is increasing in this area.
Antarctic blue whales
All Antarc�c blue whales were observed in the northern
stratum (Figure 5a). The density was a li�le higher than in
the 2000/2001 SOWER cruise in the same sector (Ensor
et al., 2001).
Duplicate sightings
Duplicates sigh�ngs were those sigh�ngs made concurrently by both the IOP and TOP barrel observers during
the IO mode survey. These data will be used to es�mate
g(0), which in turn will be used to adjust es�mates ofabundance. There was a total of 31 duplicates involving
several whale species.
Other research ac�vi�es
Table 4 shows a summary of results of diﬀerent experiments.
Sighting distance and angle experiment
The sigh�ng distance and angle experiment was conducted in order to evaluate the accuracy of sigh�ng distanceand angle provided by primary observers. The results ofResults of the dedicated sighting survey under the Japanese Abundance and Stock structure Surveys in the Antarctic (JASS-A)

this experiment will be used for the calcula�on of abundance es�mates. A training for this experiment was conducted on 7 January. The actual experiments, comprising
128 trials, were successfully completed on 30 January.
Photo-ID
Photo-ID data is used for individual matching exercise toinves�gate distribu�on and movement of large whales. A
total of 7 Antarc�c blue, 9 humpback and 10 killer whales
were successfully photo-iden�ﬁed during the en�resurvey. These data will be registered into the Ins�tuteof Cetacean Research (ICR) database (see Matsuoka andPastene, 2014).
Biopsy sampling for large whales
Biopsy samples are used for gene�c studies on stock
structure of large whales and for other feasibility studies
related to the speciﬁc objec�ves of the JASS-A. For theen�re survey, a total of 43 biopsy samples were collectedfrom 2 Antarc�c blue, 12 ﬁn, 15 Antarc�c minke, 11
humpback, 1 Brydeʼs (in transit area), and 2 killer whales,
using the Larsen system (Larsen, 1998). Biopsy samples
were stored at −20°C.
Satellite tagging
Satellite tagging is used for the study of movement,
distribu�on and stock structure of whales. The satellitemonitored tags (SPOT and SPLASH-types, Wildlife Computers, Redmond, Washington, USA) were deployedwith the Air Rocket Transmi�er System (ARTS) (LK-ARTS,
Skutvik, Norway). The detail of deployment system andprotocols, and research results to date were described in
Konishi et al. (2020). In the research area, 9 and 15 satel-

Figure 6.

Oceanographic sta�ons (XCTD cas�ng points). Red
circle: research area; black circle: transit.

lite tags were deployed on ﬁn and Antarc�c minke whales
respec�vely.
Oceanographic survey
Oceanographic observa�ons are important to understandthe rela�onship of whales and the physical environment.
The ver�cal distribu�on of water temperature and salinity were recorded from sea surface to 1,850 m water
depth using XCTD system (eXpendable Conduc�vity,
Temperature and Depth proﬁler, Tsurumi-Seiki Co., Ltd.,
Yokohama, Japan; probe type: XCTD-4 and XCTD-4N)
with Digital Converter MK-150P at 116 sta�ons along thesurvey track lines (Figure 6). Oceanographic data will beanalysed to study the oceanographic structure of the research area and the rela�onship with whale distribu�on.
Marine debris observation
Studies on marine debris in the Antarc�c are very scarce.
It is therefore important to con�nue with this kind of survey in order to monitor future trends in the occurrence ofmarine debris. One ﬁshing buoy was observed in the research area. These data will be registered into the ICR database and reported in the future (e.g. Isoda et al., 2021).
Feasibility study on the utility of UAVThis technique will be used to reﬁne observa�ons relatedwith whale abundance and distribu�on, e.g. determine
the number of individuals in the schools. The technique
can be used also for photogrammetry studies. Aerial images were collected for a total of two Antarc�c blue, six
Antarc�c minke and three humpback whales using small
UAV, Inspire 2 Pro and DJI phantom 4 Pro (video clips
can be accessed at h�ps://www.youtube.com/channel/
UCz3c9IIMiQPVeryAogmJIig). These data will be registered into the photo-ID catalogue of ICR.
Sigh�ng survey in low-middle la�tude area
Sigh�ng survey was conducted between the equator andthe research area (includes transit to the star�ng point
within the research area), excluding areas of the foreigncountriesʼ EEZs. The searching eﬀort was 704.1 n.miles andthe total sigh�ngs included ﬁn (4/4), Antarc�c minke (4/5),
and humpback (1/2), Brydeʼs (1/1) and sperm (4/4) whales.
During the transit survey from the Antarc�c researcharea to the equator (includes transit from the ending
point within the research area), the searching eﬀort was
960.1 n.miles and the total sigh�ngs included ﬁn (10/24),
Antarc�c minke (2/2), humpback (8/15), sperm (1/1) andArnouxʼs beaked (1/3) whales.
During the transit surveys, biopsy samples were col31

Technical Reports of the Ins�tute of Cetacean Research (2022)

lected from ﬁve ﬁn, two Antarc�c minke, seven humpback, and one Brydeʼs whales.

HIGHLIGHTS OF THE SURVEY
The 2021/2022 JASS-A survey covered the eastern part ofArea VI East (130°W–120°W) and succeeded in collec�ngsigh�ng data necessary for the abundance es�ma�on of
cetaceans in this area. Of par�cular importance was thesurvey conducted in ice-free waters south of 72°S. Several
other data necessary for understanding stock structure,
movement and the environment of whales were collected during the survey. The data collected throughthe JASS-A will be analysed in conjunc�on with the datacollected by the previous JARPA/JARPA, NEWREP-A and
IDCR/SOWER surveys in the same region so that theanalyses can be based on a long and consistent data set.

ACKNOWLEDGEMENTS
We thank the captain, crew members and researchers
who par�cipated in the 2021/2022 JASS-A survey for
their eﬀort in the collec�on of data and samples. We also
thank the Fisheries Agency of Japan for the funding andlogis�c support for the JASS-A program. We appreciate
Yoshihiro Fujise, Tsutomu Tamura (ICR), Hiroto Murase(Tokyo University of Marine Science and Technology),
Takashi Hakamada, Mioko Taguchi (ICR) and the steering group of JASS-A for their guidance in the design andimplementa�on of the research. Kyodo Senpaku Co. Ltd.
is acknowledged for their arrangements and support forthis cruise. We thank Kenji Konishi (ICR) for preparing thesatellite tagging, Takashi Yoshida (ICR) for compiling thesea ice informa�on, and Toshihiro Mogoe (ICR) for drawing the ﬁgures for the pack-ice distribu�on. We are grateful to Luis A. Pastene (ICR) for his assistance in preparing
this report. We also thank the Editorial Team of TEREP-ICR
for the editorial work on this manuscript. Finally, we acknowledge all of those who helped during the implementa�on of the survey despite the very diﬃcult situa�ons
caused by the COVID-19 pandemic.

REFERENCES
Ainley, D.G., Dugger, K.M., Toniolo, V. and Gaﬀney, I. 2007.
Cetacean occurrence pa�erns in the Amundsen and southern
Bellingshausen Sea sector, Southern Ocean. Marine Mammal
Science 23 (2): 287–305.
Ensor, P., Matsuoka, K., Marques, F., Miura, T., Murase, H.,
Pitman, R., Sakai, K. and Van Waerebeek, K. 2001. 2000–
2001 Interna�onal Whaling Commission-Southern Ocean
Whale and Ecosystem Research (IWC-SOWER) circumpolarcruise, Areas V, VI and I. Paper SC/53/IA5 presented to the32

IWC Scien�ﬁc Commi�ee, July 2001 (unpublished). 56 pp.
[Available from the IWC Secretariat].
Fujise, Y. and Pastene, L.A. 2021. Whales as indicators of historical and current changes in the marine ecosystem of
the Indo-Paciﬁc sector of the Antarc�c. pp. 84–104. In: M.
Kanao, D. Godone and N. Dema�eis (eds.). Glaciers and Polar
Environment. IntechOpen. 200 pp.
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.
Interna�onal Whaling Commission. 2012. Requirements andguidelines for conduc�ng surveys and analysing data within
the Revised Management Scheme. J. Cetacean Res. Manage.
(Suppl.) 13: 509–517.
Isoda, T., Matsuoka, K., Tamura, T. and Pastene, L.A. 2021.
Spa�al and temporal distribu�on of ﬂoa�ng marine macrodebris in the Indo-Paciﬁc region of the Antarc�c. Cetacean
Population Studies 3: 198–211.
Isoda, T., Katsumata, T., Ueta, E., Kasai, H. and Matsuoka, K.
2022. Results of the Japanese Abundance and Stock structure
Survey in the Antarc�c (JASS-A) during the 2021/2022 australsummer season. Paper SC/68D/ASI/10 presented to the IWC
Scien�ﬁc Commi�ee, April–May 2022 (unpublished). 28 pp.
[Available from the IWC Secretariat].
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.
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

Results of the dedicated sighting survey under the Japanese Abundance and Stock structure Surveys in the Antarctic (JASS-A)
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 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 IWC Scien�ﬁc Commi�ee,

February 2014 (unpublished). 27 pp. [Available from the IWC
Secretariat].
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
(unpublished). 11 pp. [Available from the IWC Secretariat].

33

Technical Reports of the Ins�tute of Cetacean Research (2022)

Appendix 1.
Photographs from 2021/2022 JASS-A survey

34

Photo 1.

Antarc�c minke whale.

Photo 2.

Antarc�c minke whale within the pack ice.

Photo 3.

Antarc�c blue whale.

Photo 4.

Fin whale.

Photo 5.

Humpback whale.

Photo 6.

Killer whale.

Results of the dedicated sighting survey under the Japanese Abundance and Stock structure Surveys in the Antarctic (JASS-A)

Photo 7.

Naviga�ng within the pack-ice.

Photo 8.

Sigh�ng ac�vity at the Upper Bridge Pla�orm.

Photo 9.

Sigh�ng ac�vity near an iceberg.

Photo 10.

Buoy used in the angle and distance experiment.

Photo 11.

Oceanographic observa�on using XCTD.

Photo 12.

Biopsy sampling on humpback whale.
35

Technical Reports of the Ins�tute of Cetacean Research (2022)

pp. 36–45

Technical Report (not peer reviewed)

Report and highlights of the dedicated sigh�ng surveys in thewestern North Paciﬁc Ocean in 2021
Taiki Katsumata* and Koji Matsuoka
Institute of Cetacean Research, 4–5 Toyomi-cho, Chuo-ku, Tokyo 104–0055, Japan*Contact e-mail: katsumata@cetacean.jp

ABSTRACT

Vessel-based sigh�ng surveys were conducted in 2021 by the Ins�tute of Cetacean Research to examine the
distribu�on and abundance of whales in the western North Paciﬁc Ocean. The research area was between
30°N–53°N and 150°E–155°W. The surveys were conducted between 4 August and 30 September, and two
research vessels, Yushin-Maru and Kaiyo-Maru No.7, were engaged in the surveys. A total of 3,713.8 n.miles
was searched by the passing mode in the research area. Coverage of the searching eﬀorts on the planned
cruise track line was 70.4%. In total, seven large whale species, including blue (17 schools/20 individuals), ﬁn(78/124), sei (102/156), Brydeʼs (57/72), common minke (1/1), humpback (13/16) and sperm (94/193) whales
were sighted during the research period. Photo-ID images were collected from blue (12 individuals), humpback (5 individuals) and sperm (1 individual) whales. Biopsy skin samples using a Larsen system were collectedfrom blue (n=1), ﬁn (n=5), sei (n=15) and humpback (n=3) whales. The density indices of blue and ﬁn whales
have increased notably since the last survey in the same research area 10 years ago. Furthermore, Bryde's andhumpback whales, which were not detected in the previous surveys, were sighted in the 2021 surveys.

INTRODUCTION
Dedicated cetacean sigh�ng surveys in the western North
Paciﬁc have been conducted in the summer season since
1995, as a part of the Japanese Whale Research Program
under Special Permit in the western North Paciﬁc (JARPN/
JARPNII) and the New Scien�ﬁc Whale Research Programin the western North Paciﬁc (NEWREP-NP), based on thesurvey procedures of the Interna�onal Whaling Commission/Southern Ocean Whale and Ecosystem Research(IWC/SOWER) (Anon, 2008). Based on the data collectedfrom those surveys, the distribu�on pa�erns of large
whales such as blue, ﬁn, sei, Bryde's, common minke,
humpback, North Paciﬁc right and sperm whales, andabundance es�mates of common minke, sei, and Bryde's
whales were inves�gated and reported to the IWC Scien�ﬁc Commi�ee (SC) (IWC, 2001; 2010; 2016).
The Fisheries Resources Ins�tute (FRI) has also conducted dedicated sigh�ng surveys for cetaceans in the
North Paciﬁc Ocean since the 1980s (Buckland et al.,
1992; Miyashita et al., 1995; Miyashita and Kato, 2004;
Shimada, 2004; Kanaji et al., 2012). In 2019, the Government of Japan decided to con�nue the sigh�ng surveys
in the North Paciﬁc (IWC, 2019) under the ra�onale thatthe collec�on of sigh�ng data to es�mate abundance
36

and biopsy/photo-iden�ﬁca�on data to examine stock
structure have contributed in the past to the work on the
management and conserva�on of large whales by the
IWC SC (IWC, 2016).
This paper reports the results of the latest Japanese
dedicated sigh�ng surveys conducted by the Ins�tute of
Cetacean Research (ICR) between 4 August and 30 September 2021.

SURVEY DESIGN
Research vessels
The surveys in 2021 were conducted by the research vessels Kaiyo-Maru No.7 (KY7) and Yushin-Maru (YS1). The
vessels were equipped with a top barrel pla�orm (TOP),
IO barrel pla�orm (IOP) and upper bridge. Speciﬁca�onsof these vessels are shown in Figure 1.
Research period and area
In 2021, surveys were conducted in late summer (August
to September), and the research areas were between
30°N–40°N and 150°E–180°for KY7, and between 40°N–
53°N and 180°–155°W for YS1 (Figure 2).
Track line design
The pre-determined track lines in the research area areReport and highlights of the dedicated sighting surveys in the western North Pacific Ocean in 2021

Figure 1.

Figure 2.

Research vessels Kaiyo-Maru No.7 (KY7) (le�) and Yushin-Maru (YS1) (right).

Research areas of the 2021 sigh�ng survey held from August to September. KY7 covered the green area,
and YS1 covered the blue area.

shown in Figure 2. The start point of the track lines were
decided randomly using the “Distance program ver. 7.0”
(Thomas et al., 2010), and the number of the line transects was decided by considering the research scheduleand based on the IWC survey guidelines (IWC, 2012).
Sigh�ng procedures
The sigh�ng survey was conducted using (1) Normal Passing mode (NSP) and (2) Passing with Independent Observer mode (IO). Data obtained under the IO mode areuseful for future es�ma�ons of g(0). Both survey modes
followed the protocol endorsed for the SOWER surveys(e.g. Matsuoka et al., 2003; Anon, 2008; IWC, 2012).
For NSP mode, there were two primary observers
on the top barrel pla�orm (TOP) and two (captain andhelmsman) on the upper bridge. All primary observers
engaged in the searching ac�vity for cetaceans by using
angle board and scaled binoculars (7×).
For IO mode, there were two primary observers on
the TOP and two on the IO barrel pla�orm (IOP). Theseobservers conducted the searching ac�vity for cetaceans

by using angle board and scaled binoculars (7×). There
was no open communica�on between the IOP and theTOP. The observers and researchers on the upper bridge
communicated to the TOP (or IOP) independently only toclarify informa�on, and did not distract the top-men fromtheir normal searching procedure. These primary observers report sigh�ng-informa�on to researchers and otherobservers on the upper bridge for data recording.
The sigh�ng survey eﬀort began 60 minutes a�er
sunrise and ended 60 minutes before sunset, with amaximum of 12 hours per day when the weather condi�ons were acceptable for observa�ons: visibility be�erthan 2.0 n.miles and wind speed less than 21 knots. The
searching speed was planned to be 10.5 to 11.5 knots
with slight adjustments to avoid vibra�on of the vessel.
Experiments
Distance and angle experiments were conducted in the
middle of the survey period. The experiment was conducted to evaluate measurement error, and followed theprotocol of the IWC/SOWER and IWC-POWER surveys
37

Technical Reports of the Ins�tute of Cetacean Research (2022)
Table 1
Summary of the survey periods and searching eﬀort in each research area.
Research period

Planned cruise
track(n.miles)Searching eﬀort
NSP(n.miles)Searching eﬀort
IO(n.miles)Searching eﬀort
Total(n.miles)Coverage of
eﬀort
(%)

2021/08/11–08/21

1,226.1

455.1

470.7

925.8

75.5

2021/08/30–09/09

1,392.5

533.1

534.4

1,067.5

76.7

YS1

2021/08/18–09/20

2,658.6

752.3

968.2

1,720.5

64.7

Total

–

5,277.2

1,740.5

1,973.3

3,713.8

70.4

VesselKY7Table 2
Numbers of sigh�ngs of large cetaceans, by species and research vessel in the 2021 surveys.
VesselSpeciesBlue whale
Fin whale
Sei whale
Brydeʼs whale
Common minke whale
Humpback whale
Sperm whale

Transit(Including both vessels)
Prim.

Second.

KY7(150°E–180°)
Prim.

YS1
(180°–155°W)

Second.

Prim.

Total

Second.

Prim.

Second.

Sch.

Ind.

Sch.

Ind.

Sch.

Ind.

Sch.

Ind.

Sch.

Ind.

Sch.

Ind.

Sch.

Ind.

Sch.

Ind.

0
8
1
12
0
0
32

0
13
1
20
0
0
72

0
0
0
1
0
0
3

0
0
0
1
0
0
5

0
1
0
20
0
0
34

0
1
0
25
0
0
90

0
0
0
0
0
0
0

0
0
0
0
0
0
0

17
66
98
24
1
11
24

20
107
146
26
1
13
25

0
3
3
0
0
2
1

0
3
9
0
0
3
1

17
75
99
56
1
11
90

20
121
147
71
1
13
187

0
3
3
1
0
2
4

0
3
9
1
0
3
6

Prim.: primary sigh�ng, Second.: secondary sigh�ng, Sch.: school, Ind.: individual.

(IWC, 2012).
When large cetaceans such as blue and humpback
whales were found, photo-ID images were obtained using
Canon EOS 7D Mark II (with 100–400 mm lens) from thebow or upper deck. Further, biopsy skin sampling using
the Larsen system (Larsen, 1998) was conducted when
blue, ﬁn, sei, humpback whales were sighted. A satellite
tagging experiment using the Air Rocket Transmi�er System (LK-ARTS) was also conducted for ﬁn and sei whales.

RESULTS
Brief narra�ve of the surveys
KY7 departed Shiogama, Miyagi, Japan on 4 August, andstarted the transit survey on 5 August. KY7 began the survey in the research area from the east on 11 August. KY7
paused the survey at 170°E on 21 August for a scheduled
port call, and entered Kushiro, Hokkaido, Japan, on 26
August for refueling and disembarka�on of researchers.
On 27 August, KY7 departed Kushiro, and reached theresume point at 170°E on 30 August. The survey was completed on 9 September. KY7 arrived in Shiogama, Miyagi
on 17 September.
YS1 departed Shimonoseki, Yamaguchi, Japan on 4
38

August, and started the transit survey on 6 August. On
18 August, YS1 reached the start point at 155°W, andcompleted the survey on 20 September. YS1 and arrived
in Shiogama, Miyagi, on 30 September.
Searching eﬀort
A summary of searching eﬀort and coverage by eachresearch vessel is shown in Table 1. A total of 3,713.8 n.
miles (6,878.0 km) were searched. The total searching effort by KY7 was 925.8 n.miles (1,714.6 km) (75.5% coverage) in the eastern part, and 1,067.5 n.miles (1,977.0 km)(76.7% coverage) in the western part. The total searchingeﬀort by YS1 was 1,720.5 n.miles (3,186.4 km) (64.7%
coverage).
Sigh�ngs
OverallSigh�ngs were summarized by each research vessel inTable 2 for large cetaceans and in Table 3 for small cetaceans.
There was a total of 349 schools sighted of seven large
whale species, involving 560 individual whales. The largest number of sigh�ngs was for sperm whales (in terms ofReport and highlights of the dedicated sighting surveys in the western North Pacific Ocean in 2021
Table 3
Numbers of sigh�ngs of small cetaceans, by species and research vessel in the 2021 surveys.
VesselSpecies
Killer whale
Melon-headed whale
Rissoʼs dolphin
Striped dolphin
Common dolphin
Paciﬁc white-sided dolphin
Northern right whale dolphin
True type Dallʼs porpoise
Dalli type Dallʼs porpoise
Unid. type Dallʼs porpoise
Ziphiidae
Mesoplodon spp.

Transit(Including both vessels)
Prim.

Second.

KY7(150°E–180°)
Prim.

YS1
(180°–155°W)

Second.

Prim.

Total

Second.

Prim.

Second.

Sch. Ind. Sch. Ind. Sch. Ind. Sch. Ind. Sch. Ind. Sch. Ind. Sch. Ind. Sch. Ind.
0
0
3
11
0
1
3
0
0
0
4
1

0
0
45
1,549
0
12
14
0
0
0
16
1

0
0
0
0
0
0
0
0
0
0
0
0

0
0
0
0
0
0
0
0
0
0
0
0

0
1
3
3
1
0
0
0
0
0
21
2

0
105
21
225
6
0
0
0
0
0
44
3

0
0
0
0
0
0
0
0
0
0
0
0

0
0
0
0
0
0
0
0
0
0
0
0

7
0
1
0
7
14
2
2
11
7
3
1

18
0
3
0
288
392
223
4
74
27
4
3

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

0
0
0
0
0
13
0
0
0
0
0
0

7
18 0
1
105 0
7
69 0
14 1,774 0
8
294 0
15
404 1
5
237 0
2
4 0
11
74 0
7
27 0
28
64 0
4
7 0

0
0
0
0
0
13
0
0
0
0
0
0

Prim.: primary sigh�ng, Second.: secondary sigh�ng, Sch.: school, Ind.: individual.

Figure 3.

The sigh�ng distribu�on of blue whales (blue circle) with the searching track of KY7 (blue line) and YS1(red line). Rolling 32 days average sea surface temperature from 13 August to 13 September 2021 was observed by MODIS-Aqua (Original data: Ocean color web, from h�ps://oceancolor.gsfc.nasa.gov/ (Accessed2022-7-18)).

number of individuals) and the smallest was for common
minke whales. The largest number of sigh�ngs in baleen
whales was for sei whales (Table 2).
In terms of small cetaceans, seven species were iden�ﬁed in the family Delphinidae, and one species wasiden�ﬁed in the family Phocoenidae. No species wereiden�ﬁed in the family Ziphiidae because of their elusive
behavior. The most common species sighted were striped
dolphins (14/1,774), followed by Paciﬁc white-sided dolphins (16/417) (Table 3).

Geographical distribution of sightings by cetacean species
Blue whale
The geographical distribu�on of the sigh�ngs is shown in
Figure 3 together with sea surface temperature. In total,
one mother and calf pair was sighted at posi�on 44°45ʹN,
174°43ʹW. The mean school size was 1.2 and the rangeof sea surface temperature (SST) at the sigh�ng posi�ons
was 11.5–19.7°C (mean SST: 16.3°C).
The density index (DI: schools sighted/100 n.miles
searching distance) based on primary sigh�ngs in the research area for YS1 was 0.99. The DIs were 0.15 and 0.48
39

Technical Reports of the Ins�tute of Cetacean Research (2022)

in the 2010 and 2011 IWC-POWER cruises, respec�vely,
which were conducted north of 40°N. The longitudinalranges of those cruises were 170°E–170°W and 170°W–
150°W, respec�vely (Matsuoka et al., 2011; 2012). It
should be noted that the survey period in 2021 was about

Figure 4.

40

A blue whale observed in the research area on 10
September 2021.

one month later than in the previous IWC-POWER surveys. The DIs suggest that the popula�on of blue whalesin the North Paciﬁc is increasing. A photograph of a blue
whale is shown in Figure 4.
Fin whale
The geographical distribu�on of the sigh�ngs is shown in
Figure 5 together with sea surface temperature. In total,
one mother and calf pair was sighted at posi�on 45°53ʹN,
174°00'W. Fin whales were distributed mainly throughout the research area surveyed by the YS1. The observed
mean school size was 1.6 and the range of SST at thesigh�ng posi�ons was 11.0–20.3°C (mean SST: 14.9°C).
The DI of YS1 was 3.84. The DI in the previous 2010 and2011 IWC-POWER surveys were 0.53 and 1.79, respec�vely (Matsuoka et al., 2011; 2012). As in the case of blue
whales, the DIs suggest that the ﬁn whale popula�on in
the North Paciﬁc is increasing.

Figure 5.

The sigh�ng distribu�on of ﬁn whales (red circle) with the searching track of KY7 (blue line) and YS1 (redline). Rolling 32 days average sea surface temperature from 13 August to 13 September 2021 was observedby MODIS-Aqua.

Figure 6.

The sigh�ng distribu�on of sei whales (orange circle) with the searching track of KY7 (blue line) and YS1(red line). Rolling 32 days average sea surface temperature from 13 August to 13 September 2021 wasobserved by MODIS-Aqua.

Report and highlights of the dedicated sighting surveys in the western North Pacific Ocean in 2021

Figure 7.

The sigh�ng distribu�on of Brydeʼs whales (green circle) with the searching track of KY7 (blue line) andYS1 (red line). Rolling 32 days average sea surface temperature from 13 August to 13 September 2021 wasobserved by MODIS-Aqua.

Figure 8.

The sigh�ng distribu�on of common minke whales (pink circle) with the searching track of KY7 (blue line)
and YS1 (red line). Rolling 32 days average sea surface temperature from 13 August to 13 September 2021
was observed by MODIS-Aqua.

Sei whale
The geographical distribu�on of the sigh�ngs is shownin Figure 6 together with sea surface temperature. This
species was sighted north of 42°N in the survey area of
the YS1. No mother and calf pairs were observed. Highdensity was observed in the la�tudinal band between
47°N and 49°N (Figure 6). The observed mean school size
was 1.5 and the range of SST at the sigh�ng posi�ons was
10.9–17.4°C (mean SST: 12.6°C).
The DI of YS1 was 5.70. The DIs of the 2010 and 2011
IWC-POWER surveys were 3.55 and 2.21, respec�vely(Matsuoka et al., 2011; 2012). These values were smaller
than those found in the 2021 surveys.
Bryde's whale
The geographical distribu�on of the sigh�ngs is shownin Figure 7 together with sea surface temperature. One
mother and calf pair was observed at posi�on 41°03ʹN,
178°18'W. Bryde's whales were widely distributed lon-

gitudinally north of 30°N, which is similar to the pa�ern
found in previous surveys (Shimada, 2004; Hakamada
et al., 2017). Based on sigh�ng data by KY7, the observed
mean school size was 1.3, and the range of SST at thesigh�ng posi�ons was 21.4–29.7°C (mean SST: 27.2°C).
Based on sigh�ng data by YS1, the observed mean school
size was 1.1, and the range of SST at the sigh�ng posi�ons
was 17.5–20.7°C (mean SST: 20.1°C).
The DIs were 1.00 and 1.39 by the KY7 and YS1, respec�vely. Brydeʼs whales were not sighted north of 40°N
in the previous IWC-POWER surveys (Matsuoka et al.,2011; 2012), however in the 2021 surveys some sigh�ngs
were made north of that la�tude (Figure 7) The northern
most sigh�ng posi�on in these surveys was 42°36ʹN.
Common minke whale
The geographical distribu�on of the sigh�ngs is shown in
Figure 8 together with sea surface temperature. Only one
school (one individual) was sighted at 48°07ʹN, 161°34'W
41

Technical Reports of the Ins�tute of Cetacean Research (2022)

Figure 9.

The sigh�ng distribu�on of humpback whales (black circle) with the searching track of KY7 (blue line) andYS1 (red line). Rolling 32 days average sea surface temperature from 13 August to 13 September 2021 wasobserved by MODIS-Aqua.

in Figure 11 together with sea surface temperature. This
species was distributed widely in the research areas (Figure 11). The observed mean school size diﬀered markedly
between the research areas covered by the two vessels:
2.6 in the area covered by KY7 and 1.0 by the area covered by YS1.

Figure 10.

A humpback whale sighted at 40°N on 18
September, 2021.

in calm weather condi�ons. The SST at the sigh�ng posi�on was 12.5°C. Common minke whales are hard to ﬁnd
under harsh sea surface condi�ons because their small
blows are diﬃcult to detect in such condi�ons.
Humpback whale
The geographical distribu�on of the sigh�ngs is shownin Figure 9 together with sea surface temperature. No
mother and calf pair of this species were sighted. Sightings were made north of 40°N only in the area surveyed
by YS1 (Figure 9). The range of SST at the sigh�ng posi�ons was 10.7–20.7°C (mean SST: 13.8°C).
The DI based on primary sigh�ngs of YS1 was 0.64. In
previous surveys, humpback whales were not sighted
south of 47°N (Matsuoka et al., 2011; 2012). In the 2021
surveys, two schools were sighted around 40°N, and one
individual was photographed (Figure 10). A biopsy sample
was collected from this individual.
Sperm whale
The geographical distribu�on of the sigh�ngs is shown
42

Small cetaceans
The geographical distribu�on of the sigh�ngs of small
cetaceans is shown in Figures 12a and 12b together withsea surface temperature. Killer whales, Paciﬁc whitesided dolphins and Dall's porpoises were mainly sighted
north of 42°N, while common dolphins were sighted
primarily in the 40°N–42°N la�tudinal band. Striped dolphins were not sighted north of 40°N. These distribu�onpa�erns of small cetaceans were consistent with thosefound in the previous IWC-POWER surveys conducted ten
years ago (Matsuoka et al., 2011; 2012).
Duplicate sightings
A total of 31 and 96 duplicates of large cetaceans wererecorded during IO mode surveys by KY7 and YS1, respec�vely.
Experiments
Sighting distance and angle experiment
The Es�mated Angle and Distance Training Exercise wasconducted during the surveys using a buoy that resembles
a blow with a reﬂector. During the exercise, the observers
familiarized themselves with distance es�mates from theTOP, IO and Upper Bridge. The Es�mated Angle and Distance Experiment was conducted on 23 September by YS1
on 14 August by KY7. The results of this experiment will be
used for the calcula�on of abundance es�mates.

Report and highlights of the dedicated sighting surveys in the western North Pacific Ocean in 2021

Figure 11.

The sigh�ng distribu�on of sperm whales (inverted brown triangle) with the searching track of KY7 (blueline) and YS1 (red line). Rolling 32 days average sea surface temperature from 13 August to 13 September
2021 was observed by MODIS-Aqua.

Figure 12.

(a) The sigh�ng distribu�on of killer whales, Dallʼs porpoise, uniden�ﬁed Mesoplodon spp., and Ziphiidae
with the searching track of KY7 (blue line) and YS1 (red line). Rolling 32 days average sea surface temperature from 13 August to 13 September 2021 was observed by MODIS-Aqua. (b) The sigh�ng distribu�onof striped, common, Paciﬁc white-sided, northern right whale, Rissoʼs dolphin and melon-headed whalewith the searching track of KY7 (blue line) and YS1 (red line). Rolling 32 days average sea surface temperature from 13 August to 13 September 2021 was observed by MODIS-Aqua.

43

Technical Reports of the Ins�tute of Cetacean Research (2022)

Photo-ID
Photographs were taken of blue (n=12), humpback (n=5)
and sperm (n=1) whales. All photographs were stored in
the ICR catalog and will be used for inves�ga�ng stock
structure and movement of these species in the future.
Biopsy sampling
A total of 24 biopsy samples were collected from blue(n=1), ﬁn (n=5), sei (n=15) and humpback (n=3) whales.
All samples were stored at the ICR laboratory and will be
used in gene�c analyses to inves�gate stock structure ofthese species in the future.
Satellite tagging
Two ﬁn and nine sei whales were successfully tagged andtracked. Results of this experiment are shown in Konishi
and Isoda (2022).

HIGHLIGHTS OF THE SURVEY
The sigh�ng surveys conducted in 2021 were completed
successfully. They provided unique data for summer
months, as informa�on on cetacean distribu�on andabundance during summer have been very scarce. Somemain characteris�cs of the surveys are summarized
below.
The surveys in August–September covered the northern part of the central Paciﬁc, and provided importantsummer sigh�ng data for, ﬁn, sei, Brydeʼs and sperm
whales. These data will be used for the abundance es�ma�on of those species. At the same �me, it was conﬁrmed that blue, humpback and common minke whales
were also distributed in these areas, although in small
numbers.
The DI of blue, ﬁn and sei whales increased signiﬁcantly compared with previous IWC-POWER surveys conducted ten years ago. The distribu�on pa�ern of thesethree species overlapped in terms of the la�tudinal band
and SSTs (Figures 3, 5 and 6). The distribu�on pa�ern ofthese three species were consistent with those in theprevious surveys from ten years ago. On the other hand,the distribu�on of Brydeʼs and humpback whales were
diﬀerent. Ten years ago, Brydeʼs whales were not sighted
and humpback whales were sighted only north of 47°N.
The result of the 2021 surveys showed that Brydeʼs whale
distribu�on either expanded or shi�ed northward, andhumpback whale distribu�on expanded or shi�ed southward.
As with the previous surveys, the 2021 surveys collected data on small cetaceans in the same way as large
cetaceans. The analyses of these data will provide valu44

able informa�on on the distribu�on and abundance ofsmall cetaceans in diﬀerent seasons.

ACKNOWLEDGEMENTS
We thank Captains Chikamasa Okoshi, Yasuaki Sasaki,
and their oﬃcers and crew of the Yushin-Maru and KaiyoMaru No.7 for their hard work and dedica�on that led
to the successful implementa�on of these surveys. We
express our gra�tude to Takashi Hakamada and Megumi
Takahashi (ICR) for their assistance in survey design. We
thank Yoshihiro Fujise and Tsutomu Tamura and the staﬀ
of the ICR and Kyodo Senpaku Co. Ltd. for their arrangements and support for the survey. We also thank Kenji
Konishi (ICR) for satellite tagging and Mioko Taguchi (ICR)
for the management of biopsy samples. We thank Luis A.
Pastene (ICR) for his assistance in preparing this report.
We also thank the Editorial Team of TEREP-ICR for theeditorial work on this manuscript.

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whale abundance in the northwest Paciﬁc and the Okhotsk
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Hakamada, T., Takahashi, M., Matsuoka, K. and Miyashita, T.
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sigh�ng surveys. Paper SC/M17/RMP2 presented to theWorkshop on the Implementa�on Review of western North
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Paciﬁc Japanese Special Permit Programme (JARPNII). Paper
SC/66b/REP/06 presented to the IWC Scien�ﬁc Commi�ee,June 2016 (unpublished). 96 pp. [Available from the IWC
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Interna�onal Whaling Commission. 2019. Report of theReport and highlights of the dedicated sighting surveys in the western North Pacific Ocean in 2021
Scien�ﬁc Commi�ee. Annex Q. J. Cetacean Res. Manage.
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Kanaji, Y., Iwasaki, T., Kishiro, T. and Miyashita, T. 2012. Cruise
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Konishi, K. and Isoda, T. 2022. Results of the Japanese dedicatedcetacean sigh�ng survey in the North Paciﬁc in 2021 summer season. Appendix B. Results of the satellite-monitored
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2021. Paper SC/68D/ASI06 presented to the IWC Scien�ﬁcCommi�ee, April–May 2022 (unpublished). 2 pp. [Available
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IWC Scien�ﬁc Commi�ee, May 1998 (unpublished). 8 pp.
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Matsuoka, K., Ensor, P., Hakamada, T., Shimada, H., Nishiwaki,S., Kasamatsu, F. and Kato, H. 2003. Overview of minke
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Scien�ﬁc Commi�ee, May–June 2011 (unpublished). 40 pp.
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Research (IWC-POWER). Paper SC/64/IA5 presented to the
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45

Technical Reports of the Ins�tute of Cetacean Research (2022)

pp. 46–51

Technical Report (not peer reviewed)

Results of the IWC-Paciﬁc Ocean Whale and Ecosystem Research(IWC-POWER) dedicated sigh�ng survey in 2021—An overview—
Hiroto Murase*
Tokyo University of Marine Science and Technology, 4–5–7 Konan, Minato-ku, Tokyo 108–8477, Japan*Contact e-mail: hmuras0@kaiyodai.ac.jp

ABSTRACTThis paper outlines the main results of the 2021 dedicated sigh�ng survey of the Interna�onal Whaling Commission-Paciﬁc Ocean Whale and Ecosystem Research (IWC-POWER). The IWC-POWER surveys are designed
and implemented by the IWC Scien�ﬁc Commi�ee, in special partnership with the Government of Japan. The
surveys have been conducted since 2010 as the ﬁrst phase with the long-term objec�ve: ʻ(to) provide informa�on to allow determina�on of the status of popula�ons (and thus stock structure is inherently important) oflarge whales that are found in the North Paciﬁc waters and provide the necessary scien�ﬁc background forappropriate conserva�on and management ac�onsʼ. The 2021 survey was conducted successfully between 2
August and 30 September 2021 in the eastern North Paciﬁc by the Japanese R/V Yushin-Maru No. 2. The following whale species were sighted in the survey area: blue (6 schools / 7 individuals), ﬁn (77/113), sei (23/37),
Brydeʼs (20/22), sperm (14/14) and killer (1/4) whales. Photo-iden�ﬁca�on data were collected from 7 blue,
31 ﬁn, 15 sei 13 Brydeʼs and 3 killer whales. A total of 19 biopsy samples were collected from 3 blue, 9 ﬁn, 4sei, 2 Brydeʼs and 1 killer whales. A total of 88 objects of marine debris were observed and recorded. Data collected during this survey will be used mainly for abundance es�ma�on and stock structure purposes.

INTRODUCTION
The Interna�onal Whaling Commission-Paciﬁc Ocean
Whale and Ecosystem Research (IWC-POWER) programis an interna�onal research eﬀort in the North Paciﬁccoordinated by the IWC and designed by the IWC Scien�ﬁc Commi�ee (SC) in special partnership with theGovernment of Japan. Scien�sts from the Ins�tute of Cetacean Research and coopera�ng ins�tutes such as Tokyo
University of Marine Science and Technology par�cipate
regularly in the IWC-POWER program, both in designingand implemen�ng the surveys. The IWC-POWER surveys
in the North Paciﬁc follow the series of IWC Interna�onal
Decade for Cetacean Research/Southern Ocean Whaleand Ecosystem Research (IDCR/SOWER) surveys 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 statusof 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�ons.ʼ The ﬁrst survey of this program was conducted
in 2010 and the most recent one in 2021 as the ﬁrst phase
46(IWC, 2021).
The IWC SC is close to comple�ng the ﬁrst phase of the
IWC-POWER, which is related to its short-term priori�es.
The IWC SC is preparing for the second phase related tomedium-term priori�es, based on the results of the ﬁrst
phase (see Matsuoka, 2020).
Originally, the 2021 IWC-POWER was planned to be
conducted in the region east of the Kuril archipelago
within the Russian EEZ, and Japan submi�ed the research
applica�on to the Russian Federa�on. However, Russia
sent a note verbale to Japan sta�ng that the government
could not accept the applica�on. No reason was given. As
a result, the 2021 IWC-POWER was conducted following
the back-up plan in the eastern North Paciﬁc waters.
The objec�ve of this document is to present an overview of the results of the 2021 IWC-POWER survey.
Details can be found in Murase et al. (2022). For generalbackground of the IWC-POWER including objec�ves, research area and general methodology, see Matsuoka(2020).

RESULTS OF THE 2021 IWC-POWER SURVEY
The main results of the 2021 IWC-POWER survey aresummarized here based on Murase et al. (2022).

Results of the IWC-Pacific Ocean Whale and Ecosystem Research (IWC-POWER) dedicated sighting survey in 2021—An overview—
Table 1The 2021 IWC-POWER survey i�nerary.
Date

Event

1 August 2021

Pre-cruise mee�ng in Shiogama

2 August

Vessel departs Shiogama

17 August

Vessel starts the survey in the researcharea

18 September

Vessel completes the survey in the research area

30 September

Vessel arrives in Shiogama

1 October

Post-cruise mee�ng in Shiogama

Table 2
Speciﬁca�ons of the R/V Yushin-Maru No. 2 used in the 2021
IWC-POWER survey.
Call sign

JPPV

Length overall [m]69.61

Molded breadth [m]11.5Gross tonnage (GT)

747

Barrel height [m]19.5

IO barrel height [m]13.5

Upper bridge height [m]11.5

Bow height [m]6.5

Engine power [PS/kW]

5303/3900

Research area
The research area was between 40°00ʹN and US andCanadian EEZ boundaries, between 135°00ʹW and
155°00ʹW, comprised en�rely of the high seas (Figure 1).
The areas between 150°00ʹW and 155°00ʹW, and between 135°00ʹW and 150°00ʹW were surveyed in the2011 and 2012 POWER surveys, respec�vely, althoughthe �ming of the surveys were diﬀerent from the 2021
POWER survey.
Research vessel and scien�ﬁc personnel
The R/V Yushin-Maru No. 2 was used for this survey. The
vessel is shown in Figure 2 and its speciﬁca�ons in Table 2.
Figure 1.

Figure 2.

Research area and survey track lines with start andend points for the 2021 IWC-POWER survey.

Photograph of the R/V Yushin-Maru No. 2 used in
the 2021 IWC-POWER survey.

I�nerary
The survey was conducted between 2 August and 30 September 2021 by the Japanese R/V Yushin-Maru No. 2. Thei�nerary is shown in Table 1.

Three interna�onal researchers were nominated by the
IWC SC for this survey:
Hiroto Murase (Japan)—Cruise Leader (CL)/Chief Scien�st
James W. Gilpatrick, Jr. (US)—Photo-ID
Isamu Yoshimura (Japan)—sigh�ng data, marine debris
and biopsy sample management
Searching eﬀort
Survey trackline coverage in the research area was 77.2%(1,562.5 n.miles of a planned distance of 2,022.4 n.miles),with a total of 833.7 n.miles in Passing with abeam closing
mode (NSP) and 728.8 n.miles in Independent Observer
passing mode (IO). The eﬀort spent in sigh�ng and several experiments are shown in Table 3.
Summary of the sigh�ngs
During the survey in the research area, the followingsigh�ngs were made: blue (6 schools / 7 individuals), ﬁn(77/113), sei (23/37), Brydeʼs (20/22), sperm (14/14) andkiller (1/4) whales (Table 4). These data will be used toes�mate abundance of several species.
47

Technical Reports of the Ins�tute of Cetacean Research (2022)
Table 3
Summary of the searching eﬀort (�me and distance) and experimental �me (hours) in the survey area of the 2021 IWC-POWER survey.
Leg
No.

Start

End

Start

Date

Date

End

Time

Time

Research Area (between
150°W and 135°W)

101

17-Aug. 11-Sep.

121

12:43

Research Area (between
155°W and 150°W)

121

11-Sep. 16-Sep.

127

14:27

NSP

IO

Photo-ID, Es�mated angle andBiopsy, TDR distance training/tagexperiment

NSP+IO

Area

14:27

13:54

Time

Dist.
(n.m.)

TimeDist.
(n.m.)

Dist.
(n.m.)

59:04:25

624.3052:33:36

552.83 111:38:01 1177.13 14:08:44

6:40:47

19:45:13

209.44

16:26:07

175.990:00:00

Time

36:11:20

Time

385.43

Time

0:24:46

Table 4
Number of sigh�ngs for all species observed in the research area during the 2021 IWC-POWER survey (original track lines), by effort mode. NSP: Normal Passing with abeam closing mode; IO: Independent Observer mode, OE: Top down (TD) and dri�ing (DR).
Numbers of Individuals include the number of calves.
Species

NSP

IO

Total

Sch.

Ind.

Calf

Sch.

Ind.

Calf

Sch.

Ind.

Calf

Sch.

Ind.

Calf

Blue whale

2

3

0

4

4

0

0

0

0

6

7

0

Fin whale

39

58

1

37

54

3

1

1

0

77

113

4

Like ﬁn whale

0

0

0

1

1

0

0

0

0

1

1

0

Sei whale

14

26

0

8

10

0

1

1

0

23

37

0

Like sei whale

1

2

0

0

0

0

0

0

0

1

2

0

Brydeʼs whale

11

13

1

9

9

0

0

0

0

20

22

1

Sperm whale

9

9

0

5

5

0

0

0

0

14

14

0

Bairdʼs beaked whale

1

3

0

0

0

0

0

0

0

1

3

0

Ziphiidae

1

2

0

3

6

0

0

0

0

4

8

0

Killer whale

1

4

0

0

0

0

0

0

0

1

4

0

Striped dolphin

0

0

0

1

74

0

0

0

0

1

74

0

Common dolphin

1

84

2

3

410

8

0

0

0

4

494

10

Paciﬁc white-sided dolphin

3

85

3

1

157

7

0

0

0

4

242

10

Northern right whale dolphin

1

107

4

0

0

0

0

0

0

1

107

4

Dalli type Dallʼs porpoise

10

55

0

5

42

0

0

0

0

15

97

0

Unid. type Dallʼs porpoise

3

11

0

2

2

0

0

0

0

5

13

0

Unid. dolphin

2

13

0

0

0

0

0

0

0

2

13

0

Unid. small cetacean

1

1

0

1

1

0

0

0

0

2

2

0

Unid. cetacean

2

2

0

3

3

0

0

0

0

5

5

0

Unid. large baleen whale

7

7

0

5

6

0

0

0

0

12

13

0

Geographical distribu�on by species
Blue whale (Balaenoptera musculus)
Blue whales were mainly distributed in the northern part48

OEof the research area (Figure 3). Sea surface temperaturesat the sigh�ng posi�ons were between 12.7°C and 13.6°C.

Results of the IWC-Pacific Ocean Whale and Ecosystem Research (IWC-POWER) dedicated sighting survey in 2021—An overview—

Figure 3.

The searching eﬀort (black line) and sigh�ng posi�ons (blue circles) of blue whales during the 2021
IWC-POWER survey.

Figure 6.

The searching eﬀort (black line) and sigh�ng posi�ons (green circles) of Brydeʼs whales during the2021 IWC-POWER survey.

Figure 4.

The searching eﬀort (black line) and sigh�ng posi�ons (red circles) of ﬁn whales during the 2021
IWC-POWER survey.

Figure 7.

The searching eﬀort (black line) and sigh�ng posi�ons of sperm and killer whales (brown circles
and black cross, respec�vely) during the 2021
IWC-POWER survey.

Fin whale (Balaenoptera physalus)
Fin whales were mainly distributed in the northern partof the research area (Figure 4). Sea surface temperaturesat the sigh�ng posi�ons were between 12.7°C and 19.7°C.
Sei whale (Balaenoptera borealis)
Sei whales were distributed in mid-la�tudes of the research area (Figure 5). Sea surface temperatures at thesigh�ng posi�ons were between 13.0°C and 16.1°C.

Figure 5.

The searching eﬀort (black line) and sigh�ng posi�ons (orange circles) of sei whales during the 2021
IWC-POWER survey.

Brydeʼs whale (Balaenoptera edeni brydei)
Brydeʼs whales were distributed in the southern part of
the research area (Figure 6). Sea surface temperatures atthe sigh�ng posi�ons were between 17.4°C and 21.3°C.

49Technical Reports of the Ins�tute of Cetacean Research (2022)
Table 5
Summary of the Photo-ID experiments for each species conducted in the 2021 IWC-POWER research area.
Photo-ID

Blue

Fin

Sei

Brydeʼs

Killer

Total

7
0

31
0

15
0

2
11

3
0

58
11

Between longitude 150°W and 135°W
Between longitude 155°W and 150°W

Table 6
Summary of the number of species-speciﬁc biopsy samples collected in the 2021 IWC-POWER research area.
Biopsy samples

Blue

Fin

Sei

Brydeʼs

Killer

Total

3
0

9
0

4
0

1
1

1
0

18
1

Between longitude 150°W and 135°W
Between longitude 155°W and 150°W

Figure 8.

Researchers and crew of the 2021 IWC-POWER survey with the Yushin-Maru No. 2 in the background. The
picture was taken at the end of the cruise in Shiogama.

Sperm (Physeter macrocephalus) and killer whales(Orcinus Orca)
Sperm whales were widely distributed in the researcharea (Figure 7). Sea surface temperatures at the sigh�ng
posi�ons were between 12.7°C and 20.0°C. A total of 1
school (4 individuals) of killer whales was sighted in thenorthern part of the research area where the sea surface
temperature was 13.1°C.
Iden�ﬁca�on of duplicated sigh�ngs
Resight data were recorded for a total of 108 sigh�ngs
during IO Mode involving several baleen whale species.
These data will be used to es�mate g(0), which in turnwill be used to adjust abundance es�mates.
Photo-ID experiments
Photo-ID data were obtained for a total of 69 whales:
blue (7 individuals), ﬁn (31), sei (15) Brydeʼs (13) and killer(3) whales (Table 5). Images collected during the survey
were uploaded to the IWC master photographic databasein Adobe Lightroom (LR) (Anon, 2020). Photo-ID data will50

be used to study movement, distribu�on and stock structure of the species involved.
Biopsy sampling
Biopsy samples were collected using the Larsen sampling system from 19 individual whales: 3 blue, 9 ﬁn, 4sei, 2 Brydeʼs and 1 killer whales (Table 6). Every biopsy
sampling was documented photographically. All biopsy
samples were catalogued and stored in cryo-vials frozen
at a temperature of −30°C on the vessel. These samples
will be used for molecular gene�cs analyses on stock
iden�ﬁca�on.
Marine macro debris observa�on
During the survey, a total of 88 marine macro debris objects were observed. All items were recorded ʻon eﬀortʼ
(i.e. during the ﬁrst 15 minutes of each hour).
Feasibility experiment of dive behavior tagging
During the survey, satellite linked dive behavior tags wereexperimentally deployed as a feasibility study at the dis-Results of the IWC-Pacific Ocean Whale and Ecosystem Research (IWC-POWER) dedicated sighting survey in 2021—An overview—cre�on of Japan. The tags were a�ached to 2 ﬁn and 3 sei
whales and the data were obtained via satellite. Detailed
analysis will be conducted by Japanese scien�sts and theresults will be reported to relevant scien�ﬁc communi�es.

HIGHLIGHTS OF THE SURVEY
It is concluded that the 2021 IWC-POWER survey wascompleted successfully by a group of interna�onal scien�sts (Figure 8) and that valuable data were collected forseveral cetacean species. Such data will allow for studies
on distribu�on, abundance and stock structure of large
cetaceans in this par�cular area of the North Paciﬁc.
There are two aspects of this survey that should behighlighted. The ﬁrst aspect is a seasonal change in the
distribu�on of baleen whales. Although the 2021 POWER
survey area was covered by the 2011 and 2012 POWER
surveys, the survey �ming in 2021 was approximately
a month later than the previous two surveys. From theresults, it appeared that the distribu�on of baleen whales
has shi�ed northward. For instance, the Brydeʼs whale, atropical species, was sighted in the southern part of the2021 survey area while none was sighted in the previous
two surveys. Such a change will be inves�gated in futurestudies.
The second aspect is the success of the feasibility
experiment of dive behavior tagging. This was the ﬁrst
a�empt of this kind of experiment in the IWC-POWER
surveys. The obtained data could be used to correct availability bias in future abundance es�ma�on work.
Although the IWC-POWER survey within the Russian
EEZ has not been completed, it is an�cipated that the
IWC-POWER program will move toward its second phase,
which will be designed a�er examining in details the results from the ﬁrst phase between 2010 and 2021.

ACKNOWLEDGEMENTS
I am grateful to the Governments of the United States(NOAA Fisheries) and Japan (Fisheries Agency of Japan)
for funding this cruise. I also acknowledge the Government of Japan for their assistance in arranging the research permit. I extend my thanks to Iain Staniland andthe Secretariat of the IWC for their support and funding
for this cruise. I appreciate the ongoing support by the
POWER Steering Group and the Technical Advisory Group(TAG). I thank Takashi Hakamada, Megumi Takahashi and
Mioko Taguchi from the Ins�tute of Cetacean Researchand the staﬀ of Kyodo Senpaku Co. Ltd. for making arrangements and suppor�ng the cruise. I also wish to
express my gra�tude to the crew of the R/V Yushin-Maru
No.2 for their hard work and dedica�on. I thank the
Editorial Team of TEREP-ICR for the editorial work on thismanuscript.

REFERENCES
Anon. 2020. IWCʼs IDCR/SOWER and POWER cruise photographic database manual. 21 pp. [Available from the IWC
Secretariat].
Interna�onal Whaling Commission. 2021. Report of theMee�ng of the IWC-POWER Technical Advisory Group(TAG) and 2021 Planning Mee�ng: November 2020. Paper
SC/68C/REP/01Rev01 presented to the IWC Scien�ﬁcCommi�ee, April–May 2021 (unpublished). 24 pp. [Available
from the IWC Secretariat].
Matsuoka, K. 2020. An outline of the IWC-Paciﬁc Ocean Whaleand Ecosystem Research (IWC-POWER) including resultsof the 2019 survey. Technical Reports of the Institute of
Cetacean Research (TEREP-ICR) No. 4: 23–34.
Murase, H., Gilpatrick, J.W. Jr., Yoshimura, I. and Eguchi, H.
2022. Cruise report of the 2021 IWC-Paciﬁc Ocean Whale andEcosystem Research (IWC-POWER). Paper SC/68D/ASI/03
presented to the IWC Scien�ﬁc Commi�ee, May 2022 (unpublished). 34 pp. [Available from the IWC Secretariat].

51

Technical Reports of the Ins�tute of Cetacean Research (2022)

pp. 52–55

Technical Report (not peer reviewed)

Long-term track movement of Antarc�c minke whales revealed bysatellite-monitored tag experiments conducted by the Ins�tute of
Cetacean Research
Kenji Konishi* and Tatsuya Isoda
Institute of Cetacean Research, 4–5 Toyomi-cho, Chuo-ku, Tokyo 104–0055, Japan*Contact e-mail: konishi@cetacean.jp

ABSTRACTThis paper presents progress of the work on long-term track movement of Antarc�c minke whales based onsatellite tagging experiments conducted by the Ins�tute of Cetacean Research. In par�cular, the experiments
focused on inves�ga�ng the migratory links between whales in the feeding and breeding areas. Five examplesof northward tracks from Antarc�c feeding areas in the Indo-Paciﬁc sector of the Antarc�c to lower la�tudes
areas are presented. The ﬁve tracks suggested some migra�on pa�erns in the Indian and Paciﬁc Oceans.
These results showed that tracking of Antarc�c minke whales by satellite-monitored tags provides precise anddirect evidence of migratory links. This work should be con�nued and the number of whales tagged should be
increased. This informa�on will greatly assist the interpreta�on of the stock structure hypothesis of Antarc�cminke whales derived from gene�c and sigh�ng data analyses.

INTRODUCTION
Understanding the stock structure of whales is crucial forassessment and management purposes. The Antarc�cminke whale Balaenoptera bonaerensis (herea�er AMW)
is one of the most abundant species in the Southern
Hemisphere, and its abundance exceeds 500,000 animals(IWC, 2012). As in the case of other rorqual whales, theAMW is assumed to migrate between winter tropical andsub-tropical breeding areas and summer Antarc�c feeding areas (Kasamatsu et al., 1995). However, evidences of
direct links are scarce. Only three Discovery tag-marked
AMWs were recovered at lower la�tude areas, showing movements from feeding to breeding areas. An
AMW marked at 35°W (east of Antarc�c Peninsula) andtwo AMWs marked at 19°E were recaptured oﬀ Brazil
at la�tude 6°S–7°S in the Atlan�c Ocean (Buckland andDuﬀ, 1989). These were the only cases showing direct evidence of links between breeding and feeding grounds forthis species.
The Ins�tute of Cetacean Research (ICR) has conducted
detailed research of the AMW and its ecosystem in the
Indo-Paciﬁc sector of the Antarc�c in austral summer,
through the former programs JARPA (Japanese Whale
Research Program under Special Permit in the Antarc�c),
JARPAII (the second phase of JARPA) and the NEWREP-A(New Scien�ﬁc Whale Research Program in the Antarc52

�c Ocean). A summary of the biological and ecologicalresearch outputs from these programs is available in
Murase et al. (2020). In the austral summer 2019/20
the ICR commenced a new non-lethal research programin the Indo-Paciﬁc sector of the Antarc�c called JASS-A(Japanese Abundance and Stock-structure Surveys in the
Antarc�c). The main objec�ve of the JASS-A is the es�ma�on of the abundance and abundance trends of the AMW
and other large whale species. Research on the stock
structure of large whales is the other main objec�ve of
the JASS-A, which is essen�al for appropriately interpreting the results of abundance es�mates.
A review of the stock structure studies on the AMW
in the Indo-Paciﬁc sector was provided by Murase et al.
(2020). The current stock structure hypothesis of the AMW
in the Indo-Paciﬁc sector of the Antarc�c involves the occurrence of at least two stocks, the eastern Indian Ocean
stock (I-stock) and the western South Paciﬁc stock (P-stock)(IWC, 2008; Pastene and Goto, 2016; Kitakado et al., 2014;
Murase et al., 2020). These stocks are presumed to berelated to the breeding areas for this species suggested by
Kasamatsu et al. (1995) based on sigh�ng density indices.
Figure 1 shows a schema�c representa�on of the distribu�on of the I- and P-stocks in the Antarc�c feeding grounds
and the geographical loca�ons of the low la�tude breedingareas suggested by Kasamatsu et al. (1995).
One of the important aspects to be inves�gated is theLong-term track movement of Antarctic minke whales revealed by satellite-monitored tag experiments conducted by the Institute

Figure 1.

Hypothesis on stock structure of Antarc�c minke whales in the feeding grounds of the Indo-Paciﬁc sector of
the Antarc�c derived from gene�c analyses (I- and P- stocks) (IWC, 2008; Pastene and Goto, 2016; Kitakado
et al., 2014) and loca�on of breeding areas of this species suggested by the sigh�ng records (Kasamatsu
et al., 1995). The ﬁgure shows the distribu�on of the eastern Indian Ocean stock (I-stock) and westernSouth Paciﬁc stock (P-stock) (Pastene and Goto, 2016), and an overlap sector (Kitakado et al., 2014). The
four breeding areas suggested by sigh�ng density indices (Kasamatsu et al., 1995) are also shown withdashes-line, and the highest density parts are marked with the le�er “H”. Ver�cal bars represent possible
boundaries (Kasamatsu et al., 1995).

links between the hypothesized stocks in the feedingareas and breeding areas. Satellite tracking of whales
can be useful for this purpose. Long-term tracking of
AMWs were previously reported (Lee et al., 2017; Konishi
et al., 2020). However only a few tracks showed evidenceof northward migra�on pa�ern from the Antarc�c Peninsula (Lee et al., 2017).
This paper presents the progress on the work of longterm track movement of AMWs based on satellite tagging
experiments conducted by the ICR. Informa�on on thelink between whales in the feeding and breeding areas
will greatly assist the interpreta�on of the stock structure hypothesis in the feeding ground based on gene�canalyses.

Figure 2.

SPOT6-type tags (Wildlife Computers Inc.) connected to ICRʼs ISOD-type anchor and LK-Carrier.

Figure 3.

Whale tagger LK-ARTS used during the NEWREP-A
and JASS-A tagging experiments.

TAGGING EXPERIMENTS ON ANTARCTIC MINKEWHALES BY THE ICR
In this report, tagging results using SPOT6-type tags,
which are designed to inves�gate horizontal movement,
are presented (Wildlife Computers Inc.; Figure 2). The
tags were deployed using an air compressed launcher
LK-ARTS (Figure 3), ﬁred from the research vessel YushinMaru No.2 (Konishi et al., 2020). Five tracking records
with clear northward migra�on movements were selected for this paper.
Long-term track movement of AMWs
The ﬁve northward tracking movements of AMWs areshown in Figure 4 and the relevant data are presented

in Table 1. The tag deployments were made in the sector
10°E–30°E (west of Indian Ocean Sector) and in the sector
170°W–140°W (Paciﬁc Sector), south of 60°S. These records indicated that the western and eastern movements
seem to be closely related to the direc�on of westward
Antarc�c Coastal Current and eastward Circumpolar Current, respec�vely (see Nicol et al., 2000).
Two AMWs in the Indian Ocean Sector were tagged in53Technical Reports of the Ins�tute of Cetacean Research (2022)

Figure 4.

Five cases of northward tracking movement of Antarc�c minke whales tagged during the NEWREP-A and
JASS-A programs. All deployments were conducted during the austral summer on the feeding ground. Twoformer whaling loca�ons in South Africa and Brazil are also shown.

Table 1
Relevant data of ﬁve satellite-tracked Antarc�c minke whales in Figure 4 during the NEWREP-A and JASS-A programs.
Tracking colour

Tracking periods

Loca�ons of deployments

Loca�on of last transmission

Green
Blue
Red
PurpleYellow

2 Feb. 2020–2 Sept. 2020
28 Jan. 2021–1 Jul. 2021
27 Feb. 2018–3 Jun. 2018
6 Feb. 2022–6 Aug. 2022
28 Jan. 2022–1 Jul. 2022

68.5°S; 14.6°E
69.2°S; 19.4°E
76.7°S; 163.9°W
71.5°S; 133.4°W
66.6°S; 129.8°W

30.5°S; 35.1°E
40.2°S; 67.5°E
46.7°S; 140.7°W
15.9°S; 109.2°W
59.9°S; 104.3°W

a similar longitudinal sector, however they showed different movement pa�erns. One animal moved eastward
and started migra�ng to the north around 70°E (blue in
Figure 4). The other animal showed a westward movement along the Antarc�c coast, then changed its course
to the east at 45°W in the oﬀshore area, then started
migra�ng in the northeast direc�on. This animal ﬁnallyarrived at the area oﬀ the coast of Durban, South Africa,in September (green in Figure 4).
In the Paciﬁc Sector, AMWs moved into the Ross Sea
at ﬁrst, then one whale started migra�ng to the east (yellow in Figure 4). Two others moved ﬁrst to the east andthen to the north (red and purple in Figure 4). The longest
tracked animal showed northward migra�on and ﬁnallyarrived at 16°S; 109°W in August.
In the previous study by Lee et al. (2017), two AMWs
tagged in the west of the Antarc�c Peninsula showednorthward migra�on at diﬀerent longitudinal bands in
the Paciﬁc Sector, while another one moved into the
Atlan�c Sector.
Interpreta�on of movements in the context of stock
structure
The ﬁve cases of tracking AMWs presented in this paper
demonstrated that their longitudinal range of movement
at the feeding area are wider than previously reported
54(up to 80°). On the other hand, the migratory corridors
to breeding areas involve a wide longitudinal range. In
addi�on, AMWs appear to move in the northeast direc�on when they start the migra�on to the breeding areas.
These la�tudinal movement pa�erns will assist the interpreta�on of the stock structure hypothesis of this species
in the Indo-Paciﬁc sector of the Antarc�c derived fromgene�c analyses.
As indicated earlier, the current hypothesis on stock
structure by gene�c evidence in the Indo-Paciﬁc sector of
the Antarc�c feeding grounds involves at least two stocks,
the I-stock and the P-stock (Pastene and Goto, 2016) (Figure 1). These authors suggested that the former could be
related to the breeding area in the eastern Indian Ocean
while the la�er could be related to the breeding area inthe western South Paciﬁc suggested by sigh�ng density
indices examined by Kasamatsu et al. (1995). It is important to note that no gene�c analysis has been conducted
on whales in the hypothesized breeding areas at lower
la�tudes and that the gene�c rela�onship among the
proposed breeding areas remains unknown.
Indian Ocean sector
One of the whales tagged in the Indian Ocean Sector of
the Antarc�c reached the coast of Durban, in South Africa(Figure 4). This loca�on corresponds to the western Indian

Long-term track movement of Antarctic minke whales revealed by satellite-monitored tag experiments conducted by the Institute

Ocean breeding area proposed by Kasamatsu et al. (1995)
and Best (1982). This is a cri�cal piece of evidence on
movement from the tagging program conducted by the
ICR. The migratory track of the second whale tagged in the
Indian Ocean sector of the Antarc�c is not complete therefore the migratory des�na�on of this whale in the Indian
Ocean is unknown. It should be noted that the two tracked
AMWs were both at 0–10°E in the feeding ground.
Paciﬁc sector
The migratory pa�erns of the three AMWs tagged in the
Paciﬁc sector of the Antarc�c (Ross Sea) to lower la�tudes
were diﬀerent from each other, however, they all moved
in a northeast direc�on (Figure 4). The whale with the longest tracking record arrived at la�tudes<20°S, in the eastern Paciﬁc Ocean breeding area proposed by Kasamatsu
et al. (1995). The ﬁnal migratory des�na�ons of the othertwo whales are unknown. However, the incomplete movement pa�ern, in par�cular that of the ʻyellowʼ whale,suggests that it is unlikely that they reached the westernSouth Paciﬁc breeding area (Figures 1 and 4).
The three whales were tagged ini�ally in the Antarc�cin a sector corresponding to the P-stock (Figure 1). Assuming that whales in the proposed breeding areas of thewestern and eastern South Paciﬁc Ocean are diﬀeren�ated gene�cally, the tagging data are consistent with theno�on that whales from gene�cally diﬀeren�ated breeding stocks mix spa�ally in the feeding grounds, in this casein the Ross Sea. Mark-recovery tags in the past showedthat AMWs marked in the Atlan�c feeding ground at
35°W were recovered in the breeding area oﬀ Cos�nha,
Brazil in the Atlan�c Ocean (Buckland and Duﬀ, 1989),
sugges�ng that whales from a breeding ground (Brazil)
could be distributed in a very distant feeding ground.
Future works
The number of tagged whales should be increased in the
Indo-Paciﬁc sector of the Antarc�c. In par�cular, Antarc�c minke whales in the core areas of distribu�on of the
hypothesized I- and P- stocks should be targeted in the
future. Technological improvement of the satellite tagging is also required in order to increase the dura�on of
tracking whales.

ACKNOWLEDGEMENTS
The authors would like to thank the vessel crews and ICR
members involved in the tagging experiments in the ﬁeld.
We also thank Mutsuo Goto and Mioko Taguchi (ICR)
for providing explana�on of the current stock structure
hypothesis of AMWs in the Indo-Paciﬁc sector of the Ant-arc�c. Our gra�tude to Luis A. Pastene (ICR) for his input
in the interpreta�on of the tagging data in the context
of the stock structure and for his assistance in preparing
this manuscript. Finally, we thank the Editorial Team of
TEREP-ICR for the editorial work on this manuscript

REFERENCES
Best, P. 1982. Seasonal abundance, feeding, reproduc�on,age and growth in minke whales oﬀ Durban. Rep. int. Whal.
Commn 32: 759–786.
Buckland, S.T. and Duﬀ, E.I. 1989. Analysis of the Southern
Hemisphere minke whale mark-recovery data. Rep. int. Whal.
Commn (special issue) 11: 121–143.
Interna�onal Whaling Commission. 2008. Report of theIntersessional Workshop to Review Data and Results fromSpecial Permit Research on Minke Whales in the Antarc�c,
Tokyo, 4–8 December 2006. J. Cetacean Res. Manage.
(Suppl.) 10: 411–445.
Interna�onal Whaling Commission. 2012. Report of the
Scien�ﬁc Commi�ee. J. Cetacean Res. Manage. (Suppl.) 14:
1–468.
Kasamatsu, F., Nishiwaki, S. and Ishikawa, H. 1995. Breedingareas and southbound migra�ons of southern minke whales
Balaenoptera acutorostrata. Mar. Ecol. Prog. Ser. 119: 1–10.
Kitakado, T., Schweder, T., Kanda, N., Pastene, L.A. and Walløe,
L. 2014. Dynamic popula�on segrega�on by gene�cs andmorphometrics in Antarc�c minke whales. Paper SC/F14/J29
presented to the IWC Scien�ﬁc Commi�ee JARPA review
mee�ng, February 2014 (unpublished). 20 pp. [Available
from the IWC Secretariat].
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: 343–357.
Lee, J.F., Friedlaender, A.S., Oliver, M.J. and DeLiberty, T.L.
2017. Behavior of satellite-tracked Antarc�c minke whales(Balaenoptera bonaerensis) in rela�on to environmental factors around the western Antarc�c Peninsula. Anim
Biotelemetry 5: 23.
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. and Kato, H. 2020.
Review of the assessment of two stocks of Antarc�c minke
whales (eastern Indian Ocean and western South Paciﬁc). J.
Cetacean Res. Manage. 21: 95–122.
Nicol, S., Pauly, T., Bindoﬀ, N.L., Wright, S., Thiele, D., Hosie,
G.W., Stru�on, P.G. and Woehler, E. 2000. Ocean circula�onoﬀ east Antarc�ca aﬀects ecosystem structure and sea-ice
extent. Nature 406: 504–507.
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.
55

Technical Reports of the Ins�tute of Cetacean Research (2022)

pp. 56–58

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

Some thoughts about the Ins�tute of Cetacean Research
Gabriel Gómez Díaz
Institute of Cetacean Research, 4–5 Toyomi-cho, Chuo-ku, Tokyo 104–0055, JapanA�er the establishment of the Interna�onal Whaling
Commission (IWC) moratorium on commercial whaling
and the following start of Japanʼs whale research programin the Antarc�c Ocean (the JARPA program), the Ins�tuteof Cetacean Research (ICR) became the focus of a�en�on
and repeated publicity stunts by protest extremist groups
reac�ng to the lethal component, one of many elements
forming part of the research program. Over the years,
that undeserved a�en�on mainly aimed to diminish theimportance and signiﬁcance of ICR research ac�vi�es, waso�en accompanied by seemingly endless hype directed
to an uninformed public and s�rred up in great part bywestern mainstream media. An o�en-seen cliché was theallega�on that, in spite of long years of research ac�vi�esin the Antarc�c, the ICR had only managed to produce asingle scien�ﬁc publica�on. Apparently, all that cri�cism
was just a ﬂank of a nefarious, systema�c and sustained
eﬀort deployed over the years by a huge variety of actors
to isolate and s�gma�ze Japan for its whaling policies, as
exposed in detail in a paper �tled “International pressure
and Japanese withdrawal from the International Whaling Commission: when shaming fails” (Kolmaš, 2020). In
fact, the ICRʼs research programs have produced a large
amount of scien�ﬁc informa�on required for conserva�on and management of large whales (see list of peerreviewed publica�ons in this issue).
Since the 2019 resump�on of commercial whaling
following its withdrawal from the Interna�onal Conven�on for the Regula�on of Whaling (ICRW), most of thatcacophony has subsided. For a brief �me soon a�er thisdecision was taken by the Government of Japan, the ICR
got a few calls and email queries from the general public
and the media wan�ng to know whether ICRʼs role andreason to exist had abruptly come to an end. Althoughsome pundits may disagree, the scien�ﬁc data produced
over the years through ICR research programs andJapanʼs contribu�on have been and con�nues to be ofparamount importance to part of the work and discussions of the IWC and other interna�onal organiza�ons.
Soon a�er Japanʼs expression of inten�on to withdraw
56

from the ICRW, the IWC unequivocally stated that ʻformany years Japan has played an ac�ve and integral rolein both the Commission and its Scien�ﬁc Commi�eeʼ(IWC, 2019). So, the answer to those queries is no, therole and mission of ICR presumably will con�nue as far
as Japan remains commi�ed to the realiza�on of sustainable use of whale resources.
Japan withdrew from the ICRW in 2019 a�er careful considering that the IWC had walked away from its
original objec�ve of promo�ng sustainable use of whale
resources, namely maintaining a commercial whaling
moratorium without any scien�ﬁc jus�ﬁca�on. Enacted
in 1972, the U.S. Marine Mammal Protec�on Act (MPA),
ʻis celebra�ng this year 50 years of marine mammal science, conserva�on, and recoveryʼ (NOAA, 2022a). The
MPA was coincidentally created the same year when,according to mainstream media, a recommenda�on
for a 10-year moratorium on commercial whaling wasapproved overwhelmingly at the United Na�ons Conference on the Human Environment in Stockholm (NewYork Times, 1972). Ten years later, the IWC decided that
there should be a pause (a zero-quota regula�on) in commercial hun�ng on all species and popula�ons of whales
from the 1985/1986 season onwards. This purportedly
temporary pause, which became to be known as the IWC
commercial whaling moratorium didnʼt last for 10 years
as originally intended, and, although it went remarkably
unno�ced by the media, in July 2022 a few an�-whaling
groups celebrated in Brighton, UK, its 40th anniversary.
Brighton was the venue of the 34th IWC Annual Mee�ng
where the principal stated goal of the United States was
to gain an indeﬁnite moratorium on commercial killing ofwhales.
This outcome was summarized as the conserva�onist
countries achieving an outstanding victory for whale protec�on, although they had yet to close the book on thehistory of commercial whaling (U.S. GPO, 1982). Another,less o�en quoted, major achievement by the UnitedStates at that mee�ng was the IWCʼs agreement to create
a new management scheme for aboriginal/subsistenceSome thoughts about the Institute of Cetacean Researchwhaling. This decision established a Technical Subcommi�ee to consider subsistence, nutri�onal and culturalrequirements of aboriginal peoples and advise the Commi�ee in much the same way as the Scien�ﬁc Commi�eedoes (U.S. GPO, 1982).
It is noteworthy that in spite of the 40-year-old moratorium, commercial whaling, an economic ac�vity not much
diﬀerent from other ﬁsheries, is now being conductedsustainably by two IWC member countries (Norway and
Iceland) and one ex-member country (Japan) (IWC, 2022).
Remarkably enough, consump�on for food of some 87
species of cetaceans and other marine mammals has
been reported in at least 114 countries (Robards andReeves, 2011), while the UN Organiza�on for Food andAgriculture (FAO) has yet to include marine mammals asa source of food in its sta�s�cs and reports (FAO, 2022).
Current sustainable whaling does not produce a nega�ve eﬀect on the exploited stocks. Meanwhile, due todiﬀerent causes other than whaling, at present two iconic
cetacean species, the North Atlan�c right whale (Eubalaena glacialis) (NOAA, 2022b) and the vaquita (Phocoena sinus) (CEMDA, 2021) are facing imminent ex�nc�on.
Further, more than 500,000 cetaceans are being killedworldwide annually, by ﬁshing gear bycatch and otherhuman ac�vi�es (FAO, 2021).
The ICR 1987 incep�on as a general incorporated founda�on came as a direct result of the moratorium establishment. However, the history of our ins�tu�on goes further back in �me: ICR is, in reality, older than the 40-year
IWC moratorium. The introduc�on of the moratorium
brought an abrupt halt to Japanʼs commercial coastal andhigh seas whaling. The 1987 reorganiza�on of our ins�tu�on was a direct result of the moratorium establishment
and the proac�ve measures by Japan to undertake a research program in the Antarc�c to systema�cally collectscien�ﬁc data necessary for the resump�on of that ac�vity, ideally under the aegis of the IWC.
Harking back to the �me when Japanʼs whaling in
the Antarc�c resumed a�er the WWII, a ﬁrsthand witness wrote the following: ʻA unique thing is happening
in Japan today. ( . . . ) Their islands do not have space todevote to growing food for domestic animals. Hence theirmeat must come largely from the sea. ( . . . ) When General MacArthur authorized two whaling ﬂeets to operate in Antarctic waters during 1946–47, he undoubtedly
expected objections to be raised by competitor whaling
nations. But his reason was partly humanitarian. Japanneeded the proteins and fats ( . . . ) of whale meat andblubber. ( . . . ) No one can deny that Japan utilizes thewhale more than other nations do––they actually eat themeat and blubber, which is more than anyone else does,
except the Eskimos.ʼ (McCracken, 1948). It seems several
countries protested then against Japanʼs vessels going to
the Antarc�c, because of the postwar turmoil and poli�cal situa�on. The ICRʼs origins go back to such �mes. The
Ins�tute was born at a �me of need for food, while theworldʼs whaling industry was compe�ng for increasingly
dwindling resources. Whaling was for oil as a resource,
but some countries, as is the case today, were pursuingthis ac�vity for food. Such was the case of Japan then, as
it is now.
What was then the need for establishing an ins�tu�on
specialized in whale research, several decades before themoratorium? The parent ins�tu�on, or the originator of
ICR was the Nakabe Ins�tute of Science, founded in 1941.
In 1942 it became an incorporated founda�on with theﬁrst approval from the Planning Agency. A�er the endof the war, the facili�es and researchers of the NakabeIns�tute of Science had not been dispersed and weres�ll available. According to one of the founders, if there
had been no war, the Nakabe Ins�tute of Science would
have con�nued to exist and perform the ac�vi�es of its
future successor, the Whales Research Ins�tute (WRI)(Maruyama, 1959).
In 1946, the year following the end of WWII, with themomentum of two of the three authorized whaling ﬂeets
actually going out to the Southern Ocean to help overcome the food shortage, a plan was made to establish anins�tu�on specializing in whale research, and on August
20, 1947, the Ministry of Agriculture and Forestry OrderNo. 22-4014 authorized the establishment of the WRI(Ikeda, 1988). Ever since, un�l the 1987 incep�on of thepresent ICR, our ins�tu�on was a rare, world-renowned
private biological research ins�tute in the ﬁshery industry, with an already long history of 40 years and outstanding scien�ﬁc achievements contribu�ng to cetology andwhale resource management worldwide (Ikeda, 1988).
Without taking account of this historical fact, cri�cism
against ICR scien�ﬁc ac�vi�es would lack solid ground.
The charter of the WRI emphasized the importance of
the whaling industry, the eﬃciency of whale ﬁshing andprocessing technology, and the necessity of thoroughu�liza�on of whale products. It called for the expansionof research and the implementa�on of systema�c andcomprehensive research on whale stocks, and the applica�on of the results for the public beneﬁt (Ikeda, 1988),
while the stated purpose of establishment of ICR is ʻtocontribute to the proper management and u�liza�on ofﬁshery resources by conduc�ng research on cetaceans
and other marine mammals, as well as studies on related57

Technical Reports of the Ins�tute of Cetacean Research (2022)

interna�onal ma�ers.ʼ
One may argue that the pres�ge gained then by the
WRI and later ICR, was due in part to the long-year
proliﬁc inven�ve and strenuous eﬀorts of its outstanding researchers, but it is also due to the fact that since
its origins, the ins�tute has had an established system
to reach out and publicize its ac�vi�es. The publica�onof the Geiken Tsushin newsle�er con�nues even today,
and was planned at the same �me as the publica�onof the English-language The Scientiﬁc Reports of the
Whales Research Institute (SRWRI), ﬁrst published in1948 (Maruyama, 1959). Although the SRWRI journal had
to be discon�nued when the moratorium introduc�onmade mandatory for all ICR research staﬀ to apply anddevote themselves to scien�ﬁc produc�on oriented to
discussions at the IWC Scien�ﬁc Commi�ee, the edi�onof the now ﬁve-year old Technical Reports of the Instituteof Cetacean Research (TEREP-ICR) may be consideredpart of these agelong eﬀorts to reach both to the public
and scholars.
While ins�tu�onal introspec�on is always necessary, inpar�cular for a highly specialized and small organiza�onas the ICR, perhaps a renewed appraisal about the ins�tuteʼs nature, its background, origins and achievements
thus far, is in order. Hopefully more and more people
would understand be�er Japanʼs a�tude towards whales
and whaling, and perhaps bear in mind the unchanging
importance, both domes�c and interna�onal, of the present and future role of our ins�tute.

REFERENCES
Centro Mexicano de Derecho Ambiental. 2021. Estrategias
de Conservación de la Vaquita Marina. Propuestas enla Administración Federal 2018–2024, Análisis de contexto, Julio, 2020. Centro Mexicano de Derecho Ambiental,
Cuauhtémoc, Mexico. 29 pp. h�ps://www.cemda.org.mx/wpcontent/uploads/2021/03/CEM-informe-vaquitamarina.pdf.
Food and Agriculture Organiza�on. 2021. Fishing Operations.
Guidelines to Prevent and Reduce Bycatch of Marine
Mammals in Capture Fisheries. FAO Technical Guidelines for
Responsible Fisheries No. 1, Suppl. 4. Food and Agriculture
Organiza�on, Rome, Italy. 118 pp. h�ps://doi.org/10.4060/
cb2887en.

58

Food and Agriculture Organiza�on. 2022. The State ofWorld Fisheries and Aquaculture 2022. Towards Blue
Transformation. Food and Agriculture Organiza�on, Rome,
Italy. 266 pp. h�ps://doi.org/10.4060/cc0461en.
Ikeda, I. 1988. Zaidan Houjin Nihon Geirui Kenkyusho no hossoku ni saishite. Geiken Tsushin 372: 1–5.
Interna�onal Whaling Commission. 2019. Statement on
Government of Japan withdrawal from the IWC. h�ps://iwc.int/resources/media-resources/news/statement-ongovernment-of-japan-withdrawal.
(Accessedon
22
August 2022).
Interna�onal Whaling Commission. 2022. Commercial whaling. h�ps://iwc.int/management-and-conserva�on/whaling/commercial. (Accessed on 31 August 2022).
Kolmaš M. 2020. Interna�onal pressure and the Japanese withdrawal from the Interna�onal Whaling Commission: Whenshaming fails. Australian Journal of International Aﬀairs 75(2): 197–216.
Maruyama, T. 1959. Geiken Tsushin no omoide. Geiken Tsushin
100: 1–3.
McCracken, D.R. 1948. Four Months on a Jap Whaler. First
Edi�on. Robert M. Mc Bride & Co., New York, USA. 228 pp.
Na�onal Oceanic and Atmospheric Administra�on. 2022a.
Celebra�ng 50 Years of the Marine Mammal Protec�on Act.
News, 30 July 2022. h�ps://www.ﬁsheries.noaa.gov/featurestory/celebra�ng-50-years-marine-mammal-protec�on-act.
(Accessed on 31 August 2022).
Na�onal Oceanic and Atmospheric Administra�on. 2022b. North
Atlan�c Right Whale. Species Directory, 16 September 2022.
https://www.fisheries.noaa.gov/species/north-atlanticright-whale#overview. (Accessed on 27 September 2022).
New York Times. 1972. Whaling halt urged in Stockholm.
The New York Times, 10 June 1972. h�ps://www.ny�mes.
com/1972/06/10/archives/whaling-halt-urged-in-stockholm.
html. (Accessed on 31 August 2022).
Robards, M.D. and Reeves, R.R. 2011. The global extent andcharacter of marine mammal consump�on by humans:
1970–2009. Biological Conservation 144: 2770–2786.
U.S. Government Prin�ng Oﬃce. 1982. Review of the 34th
International Whaling Commission Meeting: Hearing Before
the Subcommittee on Human Rights and International
Organizations of the Committee of Foreign Aﬀairs, Houseof Representatives, Ninety-seventh Congress, second session, September 16, 1982. U.S. Government Prin�ng Oﬃce,
Washington, USA. 100 pp.

Technical Reports of the Ins�tute of Cetacean Research (2022)

pp. 59–60

National meetingsPar�cipa�on of scien�sts from the Ins�tute of Cetacean Research inNa�onal Mee�ngs in 2022
Seminar of the research group on the North
Paciﬁc Marine Ecosystem, Mari�me Order and
Diploma�c Security System
The Kajima Ins�tute of Interna�onal Peace was foundedin July 1966. The purpose of this ins�tute is to promoteresearch and studies to promote interna�onal peace.
From its incep�on, the ins�tute has a�empted, on theini�a�ve of the founder, to design itself as a forum forexperienced public ﬁgures in poli�cs, civil service and thebusiness world, by providing them with opportuni�es toexchange views on important issues of interna�onal concern. This research group was started from 2019 as oneof their ac�vi�es.
The 5th mee�ng of this research group was held online
on 22 October 2021 and was focused on whales andecosystem. Two scien�sts from the Ins�tute of Cetacean
Research (ICR) (Pastene and Tamura) par�cipated in themee�ng by making a presenta�on en�tled ʻWhales andecosystem studies at the Ins�tute of Cetacean Research(ICR)ʼ.

The 2022 mee�ng of the Japanese Society ofFisheries Science (JSFS)
The Japanese Society of Fisheries Science (JSFS) wasestablished in 1932. It is a non-proﬁt, registered society
dedicated to the promo�on of all aspects of ﬁsheriesscience. The society fulﬁlls its global commitment by promo�ng science, striving to achieve sustainable development in the ﬁeld of ﬁsheries, while recognizing the crucial
need to preserve the natural aqua�c resources. It also
strives to forge rela�onships with the ﬁshing industry,comprising both capture and culture ﬁsheries. The main
events organized by the society are the biannual meetings held in spring and autumn in one of the main ci�esof Japan. This forum is where members present theirresearch ac�vi�es, exchange informa�on, and foster collabora�ve research in areas of common interest.
The 2022 spring mee�ng of the JSFS was held onlinefrom 26 to 29 March. Three scien�sts from the ICR (Katsumata, Yoshida and Matsuoka) contributed to the mee�ngwith the presenta�on en�tled ʻDeveloping new technology UAV – Development of VTOL-UAV ʻAsuka”.

The 2022 mee�ng of the Mammal Society ofJapan (MSJ)The Mammal Society of Japan (MSJ) was established in
1987 by uni�ng two academic organiza�ons, the Mammalogical Society of Japan and the Research Group of
Mammalogists, which were founded in 1949 and 1955,
respec�vely. The mission of the MSJ is not only to promote the global advance of mammalogy, but also to
encourage social interac�ons among its members. The
MSJ convenes an annual mee�ng, which consists of academic sessions (symposia, oral presenta�ons, and poster
presenta�ons). Members of the MSJ always give great
importance to social problem related with mammals.
The annual mee�ng of the MSJ was held online from26 to 29 August 2022. Taguchi, Katsumata and Goto fromthe ICR were co-authors of a presenta�on to the meeting en�tled ʻPreliminary popula�on gene�c analyses of
humpback whales from Hachijo Islandʼ.

The 165th mee�ng of the Japanese Society of
Veterinary Science (JSVS)
The Japanese Society of Veterinary Science (JSVS) has its
origin in the Greater Japan Veterinary Society, which was
founded in 1885. The focus of this society is veterinaryscience from basic research to applied science and clinical research. The JSVS held an annual academic mee�ng
in autumn.
The annual mee�ng was held at Azabu University,
Kanagawa from 6 to 8 September 2022. Tamura andBando from the ICR were co-authors of an oral presenta�on to the mee�ng en�tled ʻDiscovery of new molecules
as s�mulators for gonadotrophs in whale brains: possible
mechanisms for long term lifespan and fer�lity of whales
compared to bovinesʼ.

The 27th Annual Mee�ng of the Associa�on of
Wildlife and Human Society (AWHS)The Associa�on of Wildlife and Human Society (AWHS)
was established in 1994. The Society aims to serve as apla�orm for a wide range of academic ﬁelds that deal
with the diverse rela�onships between wildlife and humans. The role of the AWHS is to promptly present theresults of academic research, prac�cal knowledge anddiscussions to not only the academic world but to the so59

Technical Reports of the Ins�tute of Cetacean Research (2022)

ciety at large. The AWHS held annual academic mee�ngs.
The 27th Annual Mee�ng of the AWHS was held at
Rakuno Gakuen University, Hokkaido from 28 to 30 October 2022. Konishi from the ICR was a co-author of twooral presenta�ons to the mee�ng en�tled ʻHabitat use
pa�ern of ﬁn whales oﬀ Abashiri, Hokkaido, Japanʼ andʻPossibility of individual iden�ﬁca�on based on photoiden�ﬁca�on of areas around the dorsal ﬁn of ﬁn whales
oﬀ Abashiri, Hokkaidoʼ.

The 60th Anniversary Conference of the JapaneseSociety of Fisheries Oceanography (JSFO)
The Japanese Society of Fisheries Oceanography (JSFO)
was established in 1962. Its aim is to iden�fy scien�ﬁcissues of local ﬁsheries, and to promote studies in coopera�on with local scien�sts and ﬁshermen in order to address such issues. Recent ﬁeld research ac�vi�es in JSFO

60

include, for example, the study on the rela�onship between marine organism distribu�ons and oceanographic
condi�ons, acous�c assessment of ﬁsh abundance,
oceanographic approach using satellite image, popula�on dynamics, modeling approaches. The researchac�vi�es of the JSFO are within the framework of theﬁsheries oceanography deﬁni�on, which is ʻthe study ofoceanic processes aﬀec�ng marine ecosystems and therela�onship of these ecosystems to the abundance andavailability of ﬁshʼ.
The 60th Anniversary Conference of the JSFO was heldat the Yokohama ﬁeld sta�on of the Fisheries ResourcesIns�tute from 3 to 6 November 2022. Hakamada andMatsuoka from the ICR were co-authors of an oral presenta�on to the mee�ng en�tled ʻDevelopment of acomprehensive app for cetacean species distribu�onmodeling in the North Paciﬁc Oceanʼ.

Technical Reports of the Ins�tute of Cetacean Research (2022)

pp. 61–65

International meetingsPar�cipa�on of scien�sts from the Ins�tute of Cetacean Research inInterna�onal Mee�ngs in 2022
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, to ʻprovide for the proper conserva�on ofwhale stocks and thus make possible the orderly development of the whaling industryʼ. One of the important
subsidiary bodies of the IWC is its Scien�ﬁc Commi�ee(SC), which meets annually.
The 2022 mee�ng of the IWC SC was carried out
through a series of virtual sessions from 25 April to 13
May (SC68D). A total of twelve scien�sts from the In-

Observers from Japan par�cipa�ng in the online 2022 IWC
Scien�ﬁc Commi�ee mee�ng at the ICR mee�ng room.

s�tute of Cetacean Research (ICR) par�cipated in themee�ngs (Fujise, Kato, Pastene, Tamura, Matsuoka,
Hakamada, Taguchi, Konishi, Goto, Isoda, Takahashi and
Katsumata). They presented a total of 10 documents:
six documents at the Standing Working Group on Abundance Es�mates, Stock Status and Interna�onal Cruises(ASI), one document at the Working Group on StockDeﬁni�on and DNA tes�ng (SDDNA), one document at
the Sub-Commi�ee on Conserva�on Management Plans(CMP) and two general documents (O: PICES Observer
Report and Japanʼs Scien�ﬁc Progress Reports).
The report of the IWC SC mee�ng can be found on
the website of the IWC (h�ps://iwc.int/events-and-workshops/sc68d-2022).

IWCʼs Workshop of the Technical Adviser Group(TAG) and 2023 cruise planning mee�ng for the
IWC-POWER program
IWC-POWER (Paciﬁc Ocean Whale and Ecosystem Research) is a large interna�onal research program aimed
to study the abundance and abundance trend of large
whales in the North Paciﬁc through annual dedicatedsigh�ng surveys. The program is designated and implemented by the Scien�ﬁc Commi�ee of the IWC througha Steering Group, which in turn count with a TechnicalAdviser Group (TAG). The TAG usually meets once a year.
The IWC-POWER program has carried out surveys withthe par�cipa�on of scien�sts from Japan, USA, RussianFedera�on, Republic of Korea and Mexico.
The TAG and the 2023 cruise planning mee�ngs were

Scien�sts par�cipa�ng in the 2022 IWCʼs TAG and 2023 cruise planning mee�ng for the IWC-POWER program.
61

Technical Reports of the Ins�tute of Cetacean Research (2022)

2022 online mee�ng of the CCAMLR-EMM Working Group.

held from 6 to 10 September 2022 at the Fisheries Agencyʼs Crew house, Tokyo. The TAG mee�ng was co-chairedby Matsuoka of the ICR, who also chaired the planningmee�ng. In addi�on, Pastene, Taguchi and Isoda from the
ICR par�cipated in the mee�ngs. A total of 10 primary
documents were presented.
The report of the mee�ngs can be found on the website of the IWC (h�ps://iwc.int/events-and-workshops).

68th Mee�ng of the Interna�onal Whaling Commissionth

The plenary sessions of the 68 mee�ng of the IWC
was carried out between 17 and 21 October 2022 at theGrand Hotel Bernardin and Conference Centre, Portoroz,
Slovenia. Among the main items in the mee�ng agenda
were: the Scien�ﬁc Commi�ee presenta�on of four consecu�ve mee�ngs; discussion on ﬁnance and administra-

68th mee�ng of the Interna�onal Whaling Commission.
62

�ve issues; proposals to amend the Schedule; discussion
on proposed resolu�ons; and aboriginal subsistence
whaling. Fujise and Ohmagari from the ICR par�cipatedin the mee�ng. The next mee�ng of the IWC will be heldin Lima, Peru in 2024.

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 topreserve marine life and environmental integrity in andnear Antarc�ca. It was established in large part in response to concerns that an increase in krill catches in theSouthern 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. One of these is the Working Group on Ecosystem Monitoring and Management(EMM), which meets annually.
The online mee�ng of the EMM Working Group washeld from 4 to 11 July 2022. The main items on the meeting agenda were: krill management including risk analyses, spa�al management and climate change. Under therisk analyses approach, several papers on distribu�on,abundance and prey consump�on of whales in westernParticipation of scientists from the Institute of Cetacean Research in International Meetings in 2022

Antarc�c were presented. Three scien�sts from the ICR
par�cipated in the mee�ng (Tamura, Isoda and Katsumata).
The report of the mee�ng can be found on the websiteof the CCAMLR (h�ps://mee�ngs.ccamlr.org/en/wgemm-2022).

Annual mee�ng of the North Atlan�c Marine Mammal Commission (NAMMCO) Scien�ﬁcCommi�ee (SC)The North Atlan�c Marine Mammal Commission(NAMMCO) is an interna�onal body for coopera�on on
the conserva�on, management and study of marine
mammals in the North Atlan�c. The NAMMCO Agreement was signed in Nuuk, Greenland on 9 April 1992 by
Norway, Iceland, Greenland and the Faroe Islands, andentered into force on 8 July 1992. The agreement focuseson modern approaches to the study of the marine ecosystem as a whole, and to be�er understanding the roleof marine mammals in the ecosystem. NAMMCO has a
Scien�ﬁc Commi�ee (SC), which meets annually.
The 2022 NAMMCO SC (SC28) mee�ng was held onlinefrom 24 to 28 January 2022. Five scien�sts from the ICR
par�cipated in the mee�ng (Pastene, Tamura, Konishi,
Hakamada and Takahashi) as observers from Japan. They
presented the following documents: ʻ2020–2021 Japan
progress report on large cetacean researchʼ, ʻ2019–2020
Japan progress report on small cetacean researchʼ, ʻOutline of the process to calculate a catch limit for North
Paciﬁc common minke whaleʼ, and ʻ2020–2021 report on
satellite tagging experiments at the Ins�tute of Cetacean
Researchʼ.
The report of the mee�ng can be found on the website of NAMMCO (h�ps://nammco.no/wp-content/up
loads/2022/06/ﬁnal_report-sc28-2022.pdf).

Mee�ngs of the Steering Commi�ee of the
NAMMCO-Japan MINTAG project
The essence of the Miniature-Tag project (MINTAG project) is the development of smaller and lighter satellite
transmi�er tagging systems than the ones exis�ng on themarket today. The objec�ve is to develop a tag (MINTAG)
adapted to the study of the lesser-known fast swimming baleen whales such as ﬁn, common minke, sei and
Brydeʼs whales, as well as pilot whales. The project is acoopera�on between the four NAMMCO member countries (see above) and Japan. MINTAG is led by a SteeringGroup, which meets regularly. Pastene and Konishi fromthe ICR are members of the Steering Group. An outlineof the MINTAG project can be found at the NAMMCO
homepage.

Fourth online mee�ng of the Steering Group of the
NAMMCO-Japan MINTAG project.

In 2022 several mee�ngs of the Steering Group havebeen carried out. The fourth mee�ng of the SteeringGroup was held online on 9 September 2022. The main
objec�ves of the mee�ng were to review of summer testshoo�ng, tag design and mee�ng with Wildlife Computers, project website, plan and schedule for 2023, data
depository and budget.

29th Annual Mee�ng of the NAMMCO Council
The 29th NAMMCO Council mee�ng was held at the Grand
Hotel, Oslo, from 13 to 15 September 2022. Pastene fromthe ICR par�cipated in the mee�ng as member of the observer delega�on from Japan composed by the Fisheries
Agency and the Ministry of Foreign Aﬀairs (MOFA) members. The Council mee�ng was carried out concurrently
with the mee�ng of the Management Commi�ee.

Scien�sts par�cipa�ng in the 28th online mee�ng of the
NAMMCO Scien�ﬁc Commi�ee.
63Technical Reports of the Ins�tute of Cetacean Research (2022)

par�cipated in the mee�ng (Tamura) introducing the observer report of the 2022 IWC-SC mee�ng. The report of
the PICES mee�ng can be found on the website of PICES(h�ps://mee�ngs.pices.int/).

Workshop of the project Basin-scale Events to
Coastal Impacts (BECI)

Observers from Japan to the 29th Annual Mee�ng of the
NAMMCO Council.

Annual mee�ng of the North Paciﬁc Marine Science Organiza�on (PICES)The North Paciﬁc Marine Science Organiza�on (PICES) isan intergovernmental science organiza�on established in
1992. It aims to promote and coordinate marine scien�ﬁcresearch in the North Paciﬁc Ocean and its adjacent seas,
and to provide a mechanism for informa�on and dataexchange among scien�sts in its member countries. Itspresent members are Canada, Japan, People's Republic ofChina, Republic of Korea, the Russian Federa�on, and theUnited States of America.

Hotel Paradise Busan, the venue of the PICES-2022 mee�ng.

The 2022 mee�ng of the PICES was held at the HotelParadise Busan, Busan, Korea. The business mee�ngof the Marine Bird and Mammals (S-MBM) group washeld on 24 September 2022. One scien�st from the ICR
64

The Basin-scale Events to Coastal Impacts (BECI) is anapproved UN Decade of Ocean Science (UNDOS) project
supported by the North Paciﬁc Anadromous Fish Commission (NPAFC) and the North Paciﬁc Marine Science
Organiza�on (PICES). The objec�ve of BECI is, over thecourse of the UN Decade (2021–2030), to develop andimplement an interna�onal ocean intelligence system
of monitoring, research and analy�cal approaches thatwill provide �mely knowledge and advice to decision
makers regarding the impact of current and future ocean
condi�ons from the high seas to coastal socio-ecologicalsystems. In order to develop a high-level science plan for
BECI, a series of workshops are carried out to assemble
experts to advise on the state of climate and ocean models, knowledge of the current and projected condi�ons ofclimate and ocean in the North Paciﬁc and their likely effects on biological produc�on including ﬁshery resourcesand emerging tools and technology that can be used toeﬀec�vely monitor and study the connec�on between
ﬁsh and the ecosystem.
The Workshop 3 of the BECI (ʻTechnology and tools
for monitoring and synthesisʼ) was held online from 13
to 14 June 2022. Katsumata from the ICR par�cipated inthis workshop by presen�ng the study en�tled ʻProgressin the development of a long-range ﬂying VTOL-UAVʻASUKA”. Presenta�ons made at the Workshop 3 can befound at h�ps://beci.info/workshop_3/.

Presenta�on by an ICR scien�st at the Workshop 3 of the BECI.

24th Biennial Conference of the Society for Marine Mammalogy (SMM)
The Society for Marine Mammalogy (SMM) was foundedParticipation of scientists from the Institute of Cetacean Research in International Meetings in 2022

in 1981 and is the largest interna�onal associa�on ofmarine mammal scien�sts in the world. The mission of
the SMM is to promote the global advancement of marine mammal science and contribute to its relevance andimpact in educa�on, conserva�on and management. The
SMM held conferences every two years.
The 24th biennial conference of the SMM was held at
Palm Beach, Florida, USA from 1 to 5 August 2022. This
was the ﬁrst fully hybrid, live-virtual conference of theSMM. Kim, Bando, Fujise and Kato from the ICR were coauthors of a poster presenta�on en�tled ʻPrenatal development stages of the Antarc�c minke whaleʼ, presented
to the conference.

2022 Conference of the Spanish Cetacean Society(SEC)of 1999 during the XIII Annual Conference of the European Cetacean Society held in Valencia. The SEC was createdwith the global goals of promo�ng and coordina�ng theco-opera�on between people and ins�tu�ons dedicatedto research and conserva�on of marine mammals in
Spain and to reinforce the link between researchers andthe relevant authori�es in order to integrate research
eﬀorts in the framework of the regional, na�onal and interna�onal biodiversity conserva�on strategies. The SEC
held annual conferences.
The 2022 annual conference of the SEC was held in
Ibiza, Spain from 27 to 30 October. Pastene, Konishi andNakai from the ICR were co-authors of an oral presenta�on en�tled ʻLa distribu�on especial del epibionte
Xenobalanus globicipitis como indicador de diferencias
morfocinema�cas entre cetaceosʼ.

The Spanish Cetacean Society (SEC) was founded in April

65

Technical Reports of the Ins�tute of Cetacean Research (2022)

pp. 66–86

Peer-reviewed publications

List of peer-reviewed publica�ons based on the Ins�tute of
Cetacean Research (ICR)ʼs surveys up to 2022
This sec�on presents a list of peer-reviewed publica�ons
based on data collected by surveys conducted underformer special scien�ﬁc permit programs (JARPA/JARPAII/
NEWREP-A and JARPN/JARPNII/NEWREP-NP), including
both lethal and non-lethal techniques. Peer-reviewed
publica�ons based on these surveys are focused mainly
on topics related to assessment and management oflarge whales. However samples and data collected by
the surveys have also been useful to carry out studies ofa more academic-oriented nature. Publica�ons based onsuch 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
66

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 2022

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 (2): 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. and67

Technical Reports of the Ins�tute of Cetacean Research (2022)

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.
68

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 2022

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: A. Nishimura (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
69Technical Reports of the Ins�tute of Cetacean Research (2022)

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 (Balae70

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 2022

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).
71Technical Reports of the Ins�tute of Cetacean Research (2022)

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 Ant72

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 2022

�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 (4)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.
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. 2014. Areolaeof the placenta in the Antarc�c minke whale (Balaenoptera bonaerensis). Journal of Reproduction andDevelopment 60 (1): 62–67.
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.
73

Technical Reports of the Ins�tute of Cetacean Research (2022)

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.
2021 (13)Cunen, C., Walløe, L., Konishi, K. and Hjort, N.L. 2021.
Decline in body condi�on in the Antarc�c minke whale(Balaenoptera bonaerensis) in the Southern Ocean
during the 1990s. Polar Biol 44 (2): 259–273.
Fujise, Y. and Pastene, L.A. 2021. Whales as indicators ofhistorical and current changes in the marine ecosystemof the Indo-Paciﬁc sector of the Antarc�c. pp. 85–104.
In: M. Kanao, D. Godone and N. Dema�eis (Eds.). Glaciers and Polar Environment. IntechOpen. 200 pp.
Isoda, T., Matsuoka, K., Tamura, T. and Pastene, L.A. 2021.
Spa�al and temporal distribu�on of ﬂoa�ng marine
macro debris in the Indo-Paciﬁc region of the Antarc�c.
Cetacean Population Studies 3: 198–211.
Isoda, T., Tamura, T. and Pastene, L.A. 2021. Inges�on ofmarine debris and evidence of entanglements involving Antarc�c minke whales (Balaenoptera bonaerensis)
sampled in the Indo-Paciﬁc sector of the Antarc�c.
Cetacean Population Studies 3: 286–294.
Kato, H., Fujise, Y., Nakamura, G., Hakamada, T., Pastene,
L.A. and Best, P.B. 2021. Dwarf minke whales: morphology, growth and life history based on samples collectedfrom the higher la�tudes in the Southern Hemisphere.
Cetacean Population Studies 3: 93–128.
Kim, Y., Nishimura, F., Bando, T., Fujise, Y., Nakamura, G.,
Murase, H. and Kato, H. 2021. Fetal development in tailﬂukes of the Antarc�c minke whale. Cetacean Population Studies 3: 231–238.
Konishi, K. and Tamura, T. 2021. Preliminary use of nearinfrared spectroscopy to es�mate the biochemicalcomponents of the muscles of the Antarc�c minke
whale Balaenoptera bonaerensis. Cetacean Population
Studies 3: 281–285.
Nakamura, G., Zenitani, R., Bando, T., Fujise, Y., Yamamoto, R., Nishimura, F., Hirose, A., Kim, Y. and Kato, H.
2021. Skeletal measurements on some large cetacean
species done by scien�sts of TUMSAT and ICR. Cetacean Population Studies 3: 215–230.
Nishimura, F., Kim, Y., Bando, T., Fujise, Y., Nakamura,G., Murase, H. and Kato, H. 2021. Morphological differences in skulls and feeding apparatuses between
Antarc�c (Balaenoptera bonaerensis) and common(Balaenoptera acutorostrata) minke whales, and theimplica�on for their feeding ecology. Can. J. Zool. 99(12): 1067–1079.
74

Ohashi, Y., Goto, M., Taguchi, M., Pastene, L.A. and Kitakado, T. 2021. Evalua�on of a paternity method basedon microsatellite DNA genotypes for es�ma�ng theabundance of Antarc�c minke whales (Balaenoptera
bonaerensis) in the Indo-Paciﬁc region of the Antarc�c.
Cetacean Population Studies 3: 139–151.
Pastene, L.A., Goto, M., Taguchi, M. and Matsuoka, K.
2021. Gene�c matches of southern right whales in the
Indian sector of the Antarc�c: a contribu�on towards
understanding their movement and site-ﬁdelity. Cetacean Population Studies 3: 129–138.
Uchida, M., Suzuki, I., Ito, K., Ishizuka, M., Ikenaka, Y.,
Nakayama, S.M.M., Tamura, T., Konishi, K., Bando, T.
and Mitani, Y. 2021. Es�ma�on of the feeding recordof pregnant Antarc�c minke whales (Balaenoptera
bonaerensis) using carbon and nitrogen stable isotope
analysis of baleen plates. Polar Biol 44 (3): 621–629.
*Yamazaki, K., Aoki, S., Katsumata, K., Hirano, D. andNakayama, Y. 2021. Mul�decadal poleward shi� of thesouthern boundary of the Antarc�c Circumpolar Current oﬀ East Antarc�ca. Sci. Adv. 7 (24): eabf8755.
2022 (3)Ishikawa, H., Otsuki, M., Tamura, T., Konishi, K., Bando,
T., Ishizuka, M., Ikenaka, Y., Nakayama, S.M.M. and Mitani, Y. 2022. Foraging ecology of mature male Antarc�c minke whales (Balaenoptera bonaerensis) revealed
by stable isotope analysis of baleen plates. Polar Science 31: 100785.
Pastene, L.A., Goto, M., Taguchi, M. and Fujise, Y. 2022.
Phylogeny and popula�on gene�c structure of minke
whales worldwide: a review of recent studies. pp.
11–37. In: H.A.E. Kaoud (Ed.). Marine Mammals. IntechOpen. 126 pp.
Yasunaga, G., Inoue, S., Bando, T., Hakamada, T. and Fujise,
Y. 2022. Aspar�c acid enan�omer quan�ﬁca�on using
ultraperformance liquid chromatography–tandem
mass spectroscopy combined with deuterium-chloridehydrolysis to improve age es�ma�on in Antarc�cminke whale Balaenoptera bonaerensis. Marine Mammal Science h�ps://doi.org/10.1111/mms.12977.

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.

List of peer-reviewed publications based on the Institute of Cetacean Research (ICR)ʼs surveys up to 2022

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,
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
whale. pp. 109–113. In: A. Nishimura (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.
75

Technical Reports of the Ins�tute of Cetacean Research (2022)

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 Paciﬁc76

during 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 Physiology, 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

List of peer-reviewed publications based on the Institute of Cetacean Research (ICR)ʼs surveys up to 2022

(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
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.
2009 (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,77

Technical Reports of the Ins�tute of Cetacean Research (2022)

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 signiﬁcant 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
78(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).
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.,List of peer-reviewed publications based on the Institute of Cetacean Research (ICR)ʼs surveys up to 2022

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 preyspecies 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(in Japanese).
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 improvementsof 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.
79Technical Reports of the Ins�tute of Cetacean Research (2022)

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: 17 pp.
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 west80

ern 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
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List of peer-reviewed publications based on the Institute of Cetacean Research (ICR)ʼs surveys up to 2022

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 (3)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.
Maeda, H., Fujise, Y., Kishiro, T. and Kato, H. 2017. U�lityof the transi�on phase in earplugs of the North Paciﬁc
common minke whale as an indicator of age at sexual
maturity. Open Journal of Animal Sciences 7: 414–424.
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(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.
2021 (8)Bando, T. 2021. Improved es�mates of some life-history
parameters of the pelagic subspecies of Brydeʼs whalein the western North Paciﬁc. Cetacean Population
Studies 3: 239–245.
Gomes, T.L., Quiazon, K.M., Kotake, M., Fujise, Y., Ohizumi, H., Itoh, N. and Yoshinaga, T. 2021. Anisakis spp.
in toothed and baleen whales from Japanese waters
with notes on their poten�al role as biological tags.
Parasitology International 80: 102228.
Maeda, H. and Kato, H. 2021. Seasonal changes in theearplug germinal layers of North Paciﬁc common minke
whales. Cetacean Population Studies 3: 246–251.
Matsuoka, K., Hakamada, T. and Miyashita, T. 2021. A
note on recent surveys for right whales Eubalaena japonica in the western North Paciﬁc. Cetacean Population Studies 3: 252–257.
Nakamura, G., Zenitani, R., Bando, T., Fujise, Y., Yamamoto, R., Nishimura, F., Hirose, A., Kim, Y. and Kato, H.
2021. Skeletal measurements on some large cetacean
species done by scien�sts of TUMSAT and ICR. Cetacean Population Studies 3: 215–230.
Nishimura, F., Kim, Y., Bando, T., Fujise, Y., Nakamura,G., Murase, H. and Kato, H. 2021. Morphological differences in skulls and feeding apparatuses between
Antarc�c (Balaenoptera bonaerensis) and common(Balaenoptera acutorostrata) minke whales, and theimplica�on for their feeding ecology. Can. J. Zool. 99(12): 1067–1079.
Suyama, S., Yanagimoto, T., Nakai, K., Tamura, T., Shiozaki,
K., Ohshimo, S. and Chow, S. 2021. A taxonomic revision of Pennella Oken, 1815 based on morphology andgene�cs (Copepoda: Siphonostomatoida: Pennellidae).
Journal of Crustacean Biology 41 (3): ruab040.
Taguchi, M., Goto, M., Milmann, L., Siciliano, S., Tiedemann, R. and Pastene, L.A. 2021. New insights into thegene�c structure of sei whales (Balaenoptera borealis)at the inter-oceanic scale. Cetacean Population Studies
3: 152–163.
81

Technical Reports of the Ins�tute of Cetacean Research (2022)

2022 (4)Pastene, L.A., Goto, M., Taguchi, M. and Fujise, Y. 2022.
Phylogeny and popula�on gene�c structure of minke
whales worldwide: a review of recent studies. pp.
11–37. In: H.A.E. Kaoud (Ed.). Marine Mammals. IntechOpen. 126 pp.
Taguchi, M., Goto, M., Matsuoka, K., Tiedemann, R. andPastene, L.A. 2022. Popula�on gene�c structure of
Brydeʼs whales (Balaenoptera brydei) on the central
and western North Paciﬁc feeding grounds. Can. J. Fish.
Aquat. Sci (in press).
Takahashi, M., Tamura, T., Bando, T. and Konishi, K. 2022.
Feeding habits of Brydeʼs and sei whales in the western North Paciﬁc inferred from stomach contents andskin stable isotope ra�os. Journal of Sea Research 184:
102204.
Zhu, Y., Mizutani, K., Minami, K., Shirakawa, H., Kawauchi, Y., Shao, H., Tomiyasu, M., Iwahara, Y., Tamura,T., Ogawa, M., Tatsuyama, K. and Miyashita, K. 2022.
Target strength measurements of free-swimming
sandeel species, Ammodytes spp., in a large indoor
experimental aquarium. Journal of Marine Science andEngineering 10 (7): 966.

Other surveys
1993 (2)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.
Taga, Y. 1993. Generaliza�on of HT strategy and its
applica�on. Yokohama Mathematical Journal 40 (2):
163–173.
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 indig82

enous 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. Biologicalanalysis 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 (5)Corkeron, P.J., Ensor, P. and Matsuoka, K. 1999. Observa�ons of blue whales feeding in Antarc�c waters. Polar
Biol 22 (3): 213–215.
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 Mam-

List of peer-reviewed publications based on the Institute of Cetacean Research (ICR)ʼs surveys up to 2022

mal 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.
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�c
83

Technical Reports of the Ins�tute of Cetacean Research (2022)

ecosystem. 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.,
Secchi, 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
84

from the Perspective of Traditional Maritime Communities, International Symposium Report No. 1. Kanagawa
Shimbun Press, Kanagawa. 199 pp.
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.
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: J.E. Ringstad (Ed.).
Whaling and History IV. Commander Chr. Christensenʼs
Whaling Museum, part of Ves�oldmuseene IKA, Sande�ord. 192 pp.

List of peer-reviewed publications based on the Institute of Cetacean Research (ICR)ʼs surveys up to 2022

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.
2021 (4)Diallo, S.T., Sane, A., Nelson, T., Katsumata, T. and Hakamada, T. 2021. Cetaceans oﬀ Gabon based on a 2011sigh�ng survey, with a preliminary density es�mate of
the humpback whale Megaptera novaeangliae. Cetacean Population Studies 3: 258–264.
Katsumata, T., Hirose, A., Nakajo, K., Shibata, C., Murata,
H., Yamakoshi, T., Nakamura, G. and Kato, H. 2021.
Evidence of winter migra�on of humpback whales tothe Hachijo island, Izu Archipelago oﬀ the southerncoast of Tokyo, Japan. Cetacean Population Studies 3:
164–174.
Milmann, L., Taguchi, M., Siciliano, S., Baumgarten,
J.E., Oliveira, L.R., Valia�, V.H., Goto, M., O�, P.H.
and Pastene, L.A. 2021. New gene�c evidences fordis�nct popula�ons of the common minke whale(Balaenoptera acutorostrata) in the Southern Hemisphere. Polar Biol 44 (8): 1575–1589.
Takahashi, M., Nakamura, G. and Kato, H. 2021. Growthrelated changes in the cranium of killer whales in thewestern North Paciﬁc. Cetacean Population Studies 3:
175–188.
2022 (5)Cabrera, A.A., Schall, E., Bérubé, M., Anderwald, P.,
Bachmann, L., Berrow, S., Best, P.B., Clapham, P.J.,
Cunha, H.A., Dalla Rosa, L., Dias, C., Findlay, K.P., Haug,
T., Heide-Jørgensen, M.P., Hoelzel, R., Kovacs, K.M.,
Landry, S., Larsen, F., Moreira Lopes, X., Lydersen, C.,
Ma�la, D.K., Oos�ng, T., Pace III, R.M., Pape�, C., Paspa�, A., Pastene, L.A., Prieto, R., Ramp, C., Robbins, J.,
Sears, R., Secchi, E.R., Silva, M.A., Simon, M., Víkingsson, G., Wiig, Ø., Øien, N. and Palsbøll, P.J. 2022. Strong
and las�ng impacts of past global warming on baleen
whales and their prey. Global Change Biology 28 (8):2657–2677.
Hirose, A., Kodera, R., Uekusa, Y., Katsumata, H., Katsumata, E., Nakamura, G. and Kato, H. 2022. Compara�veanatomy of and around the posterior nasofrontal sacof a beluga whale. Marine Mammal Science 38 (3):1272–1285.
85

Technical Reports of the Ins�tute of Cetacean Research (2022)

Matsuoka, K., Crance, J.L., Taylor, J.K.D., Yoshimura, I.,James, A. and An, Y.-R. 2022. North Paciﬁc right whale(Eubalaena japonica) sigh�ngs in the Gulf of Alaska andthe Bering Sea during IWC-Paciﬁc Ocean Whale andEcosystem Research (IWC-POWER) surveys. Marine
Mammal Science 38 (2): 822–834.
Pastene, L.A., Taguchi, M., Lang, A., Goto, M. and Mat-86

suoka, K. 2022. Popula�on gene�c structure of North
Paciﬁc right whales. Marine Mammal Science 38 (3):1249–1261.
Ponganis, P.J., Fujise, Y. and Miyazaki, N. 2022. Morphology and physiology in some small pelagic cetaceans: is
Dallʼs porpoise a deep diver and a thoroughbred of thesea? Marine Mammal Science 38 (4): 1442–1469.