Baker et al. SC/60/BC2 1 of 8
FOR CONSIDERATION BY THE SCIENTIFIC COMMITTEE OF
THE INTERNATIONAL WHALING COMMISSION
SANTIAGO, CHILE, JUNE 2008
Market surveys of whale meat in Japan, 2007 – 2008, with
reference to the number of fin whales for sale
C.S. BAKER1, N. FUNAHASHI2, D. STEEL1
1Marine Mammal Institute, Oregon State University, Newport, Oregon 97365 USA
2 International Fund for Animal Welfare, 5-28-601 Higahsihoncho, HigashiKurume-shi, Tokyo 203-0014,
Japan
ABSTRACT
We report on species identification of whale-meat products purchased directly and via
the Internet from commercial markets of Japan from mid September 2007 to late
March 2008. A total of 99 products included seven species of baleen whale:
humpback (n=5), fin (n=39), Bryde’s (n=13), sei (n=21), North Pacific minke (n=15)
and Antarctic minke whales (n=5). The individual identity of market fin whales was
considered by comparison to products purchased since scientific hunting of fin whales
in the Antarctic was initiated in the austral season of 2005/06 as part of the JARPA II
programme. Although only 13 fin whales have been reported in the JARPA II catch in
the 2005/06 and 2006/07 seasons, a minimum of 15 individual fin whales were
represented by products on the market during this time. It is difficult to explain the
presence of these additional individuals given the small number of fin whales in
official reports of bycatch. To improve control of commercial whalemeat markets and
estimation of illegal, unreported or undocumented (IUU) takes, we recommend that
information from the Japanese DNA register be made available through the data
availability procedure of the IWC Scientific Committee.
INTRODUCTION
A diverse range of products originating from protected whales and unprotected dolphins continues to
be widely available on commercial markets throughout Japan and some coastal cities of the Republic
of (South) Korea more than 20 years after the 1986 international moratorium on commercial whaling
(e.g., Chan et al. 1995; Baker et al. 1996a, b, Grohman et al. 1999, Baker et al. 2000a, Baker et al.
2000b, Baker et al. 2002; Baker et al. 2006, Baker et al. 2007). In 2000, Japan increased the species
diversity of its legal market in whale products by expanding its scientific whaling programme in the
North Pacific (JARPN II) and the Antarctic (JARPA II). In addition, Japanese regulations were
changed from July 2001 to permit the killing and selling of whales that had been accidentally caught
in coastal fishing set nets (Anon. 2001). Finally, products from reported stockpiles of frozen products
from species killed before the 1986 moratorium, or taken in subsequent scientific hunting by Iceland
until 1989 may also be sold on the Japanese market.
Molecular species identification of cetacean products has become an important tool for monitoring the
species composition of market samples. We, and others, have conducted surveys of commercial
markets in Japan since 1993, and in South Korea since 1994 (e.g., Baker and Palumbi 1994, Baker et
al. 1996a,b, Baker et al. 2000, Congdon et al. 1999, Lento et al. 1997, 1998, 2000, 2001, Lavery et al.
2002). Here we report on the species identification of market products purchased in Japan between
September and November 2006. The results of these recent surveys highlight persistent uncertainties
regarding stock definitions for some protected species and stocks, and possible infractions of
international agreements with regard to the likely origins of some products. Previously (Baker et al.
2007), we presented evidence from products purchased in 2006 (referred to as the ‘2006 market
survey’) suggesting that some fin whale products sold in Japan prior to the start of the scientific hunt
of this species in the Antarctic in 2005/06 did not originate from the Icelandic scientific hunt in the
Baker et al. SC/60/BC2 2 of 8
North Atlantic, the last legal source of fin whale. Here we present results of market surveys during
2007 and early 2008 (referred to as the ‘2007 market survey’). Of particular interest was the minimum
number of individual fin whales available on the market, relative to the expected number from the
reported scientific hunt and coastal bycatch, and the likely oceanic origin of these individuals.
MATERIALS AND METHODS
Market product surveys
For the 2007 survey of Japan, products were purchased via the Internet from shops located in ten
prefectures on the islands of Honshu (Iwate, Chiba, Tokyo, Shiga, Osaka, Wakayama, Tottori, and
Yamaguchi), Shikoku (Kochi) and Kyushu (Nagasaki) from September 2007 to March 2008. Shops
were searched by key words, “whalemeat” and “whale” in Japanese, and one to four products were
purchased from those shops that specified whale species. Additionally, three prefectures (Ishikawa,
Osaka, and Wakayama) were visited in November. In an effort to evaluate changes to species
availability resulting from the JARPAII hunt, fin whale products were purchased from all shops
listing this species by name.
Field Amplification and Isolation of DNA
As in all previous surveys of commercials markets, DNA extractions from cetacean tissue and
subsequent amplification via the Polymerase Chain Reaction (PCR) were conducted on site. The field
analyses in the current surveys were performed using a Biometra thermocycler. Tissue from each
product was prepared for amplification using Chelex resin (BioRad Laboratories) using the methods
of Walsh et al. (1991), as described in Baker et al. (1996b). By using biotin-labelled primers and
streptavidin-coated plates (see below), all amplified (synthetic) products were isolated and washed
free of the original DNA template before transport to our home laboratory for direct DNA sequence
analysis. This is in accordance with the regulations of the Convention on International Trade in
Endangered Species (CITES 1973; Bowen and Avise 1994; Jones 1994).
Portions of two mitochondrial (mt) DNA loci, the control region (~ 800 base pairs, bp) and
cytochrome b gene (~ 500 bp), were amplified in the field for primary species identification. Control
region amplifications were performed using the primers Dlp1.5-L (modified by a 5’-M13 forward
primer extension to facilitate initiation of the subsequent sequencing reaction) and Dlp8G-H (see
Lento et al. 1997, Figure 1). Cytochrome b amplifications used the primers GLUDG-L and CB2-H
(Palumbi 1996). To enable purification of the amplified DNA, the Dlp1.5-L and CB2 primers used in
the field were labelled with biotin. PCR amplifications were transferred to tubes coated with
streptavidin (Nunc or ABgene streptavidin-coated 96-well plates), and incubated overnight to allow
binding of the biotin-labelled synthetic DNA. Native template DNA (which lacks the biotin label) was
removed by repeated high-stringency washing in the field. Re-amplification of the control region
fragment in the laboratory was performed using the primer, M13Dlp1.5-L and internal primers,
Dlp5-H (resulting in a re-amplified fragment ~ 450 -550 bp in length). Re-amplification of the
cytochrome b fragment used the original primers. Reactions were prepared for cycle sequencing using
SAP/ExoI treatment (incubation with shrimp alkaline phosphatase and exonuclease I; Werle et al.
1994). Cycle sequencing reactions used an ABI Prism Big-Dye Terminator Ready Reaction Kit (v3.1,
Applied Biosystems, Inc) and DNA sequencing was performed on an ABI3730 Automated DNA
Sequencer (Applied Biosystems, Inc).
Phylogenetic identification of species.
Species identification analysis was performed using the phylogenetic methods described by Baker and
Palumbi (1994) and Baker et al. (1996a) as reviewed by Dizon et al (2000), Briefly, identification of
‘test’ sequences was performed using phylogenetic reconstruction methods based on maximum
parsimony (MP) and neighbour joining (NJ) genetic distance algorithms available in the computer
program PAUP* (Swofford 1999). Each test sequence was compared with a suite of ‘type’ or
‘reference’ sequences to make an initial assignment to family, and in some cases, genus. Test
sequences identified to the same family were then grouped for secondary phylogenetic comparison to
additional type sequences of the same or related species and appropriate outgroups. The consistency
of species identification was estimated using 1,000 bootstrap simulations for both primary and
Baker et al. SC/60/BC2 3 of 8
secondary phylogenetic analyses. The phylogenetic methods of species identification used in this and
previous surveys have been implemented in the web-based programme, DNA Surveillance,
(www.DNA-surveillance.auckland.ac.nz; Ross et al. 2003).
RESULTS
Species identification of market products
Amplification of the two mtDNA fragments was attempted from total of 100 products, of which 99
produced reliable nucleotide sequences for either the mtDNA control region or cytochrome b gene, or
both, for identification to the species level (Table 1).
Sei whales. 21 products were identified as sei whale (16 females, 3 males and 2 unknowns). Based
on the number of unique mtDNA haplotypes and sex, the 21 products represent at least 12 individual
sei whales.
Bryde’s whales. 13 products were identified as Bryde’s whales (6 females, 6 males and 1 unknown).
Based on the number of unique mtDNA haplotypes and sex, the 13 products represent a minimum of
9 individual Bryde’s whales.
Humpback whales. 5 products were identified as humpback whales. All were male and all shared a
single mtDNA haplotype, common in some North Pacific populations.
North Pacific minke whales. 15 products were identified as North Pacific minke whales. These
represented at least 11 different individuals. Using the informative nucleotide sequence-based system
of stock identification (allowing identification of haplotypes characteristic of J-and O-stock animals –
‘haplogroups’) developed in previous surveys (Baker et al 2000; Goto and Pastene, 1998), 8 of the
products (representing at least 6 individuals) were identified as ‘J’ type likely to have originated from
the East Sea/Sea of Japan and perhaps other coastal waters of Japan.
Antarctic minke whales. 6 products were identified as Antarctic minke whales, two males, one
female and three unknowns. Based on sex and unique mtDNA sequences these 6 products represent a
minimum of 6 individuals.
Fin whales. 39 products were identified as fin whales, 21 females, 16 males and 2 unknowns. Based
on haplotype variation and sex, at least 12 individuals were represented in the 39 products.
Expected and minimum number of fin whale market individuals
To determine the expected number of individual fin whales on the market during the two surveys
years, we reviewed annual progress reports to the IWC and the Marine Mammal Stranding Data Base
(The National Science Museum/The Institute of Cetacean Research, http://svrsh1.kahaku.go.jp/). The
JARPA II programme reported killing 10 fin whales (4 males: 6 females) during the 2005/06 season
and 3 fin whales during the 2006/07 season (1 males: 2 females). Three strandings of fin whales
during 2005 and 2006 seemed to have been disposed of rather than released to market (22nd Jan 2005,
Okinawa, 2nd Feb 2005, Akita, 19th Mar 2006, Nagasaki). One fin whale was entangled alive and
then died (or was killed) on 17th Dec 2007, in Iwate. Although products from this individual were
reportedly sold to market, only one fin whale products in our survey was purchased after this date:
J08-03 purchased on 21 January 2008. The haplotype of this product matched several products
purchased before this date and so did not contribute an additional market individual. Given that it has
been more than 15 years since the last reported importation of fin whale from Iceland (in 1991 from
whales killed in 1989), we did not consider the North Atlantic as a likely source of market individuals
(see below). Consequently, we expected to find no more than 13 market individuals in our combined
survey of 2006 and 2007.
Contrary to this expectation, at least 15 fin whale market individuals were represented in the market
surveys of 2006 and 2007, based on haplotype variation alone. Of the 12 individual fin whales
represented in the 2007 survey, 5 matched to the 8 individuals represented in the 2006 survey, giving
Baker et al. SC/60/BC2 4 of 8
a minimum census of 15 market individuals (5 males: 10 females). Interestingly, none of the 15
haplotypes from the 2006-2007 surveys matched to haplotypes found on the market prior to 2006.
Oceanic origins of fin whale market individuals
As described in Baker et al. (2007), reference sequences of the mtDNA control region (>400 bp in
length, as reported in Hatch et al. 2006) representing fin whales of known origin were assembled to
investigate the oceanic origins of fin whale sold on the Japanese markets. We expected that sequences
of all fin whale products purchased prior to the 2005/06 JARPA II hunt (and assumed to have
originated from the 1991 importation of Icelandic scientific hunt) should group with reference
sequences from the North Atlantic and should be distinct from products purchased after the 2005/06
JARPA II (and assumed to have originated from the Southern Hemisphere).
The observed phylogenetic relationship of market and reference haplotypes was not consistent with
this expectation (Figure 1). The phylogenetic reconstructions (neighbour joining and maximum
parsimony) showed that most haplotypes grouped into one of three clades supported in a strict
consensus of all equally parsimonious trees (but with only weakly supported by bootstrap
simulations). The first clade consisted only of North Atlantic reference samples and 15 haplotypes
representing market products purchased between 1993 and 2003 (mostly before 1998). The second
clade consisted of two North Pacific reference haplotypes (from Alaska) and all 15 haplotypes
representing market products purchased in 2006 and 2007, as well as 10 haplotypes representing
market products purchased from 1993 to 2004. Two haplotypes, one purchased before JARPA II
(J97.012) and one after (J0720&), were placed intermediate between the first and second clade. The
third clade consisted only of reference sequences from the North Pacific, including the coast of
California and Russia (SWFSC tissue archive, K. Robertson, pers. comm.).
The close genetic relationship of haplotypes in the second clade suggests that some of the products
purchased before the 2005/06 JARPA II hunt might have originated from IUU takes in the North
Pacific or Southern Hemisphere, rather than from long-term storage of the Icelandic scientific hunt (as
previously assumed). Unfortunately, we have no access to reference sequences from Southern
Hemisphere fin whales, limiting the power of this test of expectations.
CONCLUSIONS
Market survey results have previously raised concerns for Illegal, Unreported or Undocumented
(IUU) takes from stocks of at least five species of whales: sei, Bryde’s, humpback, gray and North
Pacific minke whales. Here, we present additional evidence for IUU takes from a sixth species, the fin
whale. Unfortunately, in the absence of information presumably available in the Japanese DNA
register of whaling, we cannot confirm the identity of whales in scientific hunting and coastal
bycatch, or the source of the potential IUU takes. Based on the limited availability of reference
sequences, however, we suggest that some IUU take of fin whales from the North Pacific or Southern
Hemisphere has been ongoing since the start of market surveys in 1993. To improve control of
commercial whalemeat markets and estimation of IUU takes, we recommend that information from
the Japanese DNA register be reported annually to the Scientific Committee or through the data
availability procedure of the IWC Scientific Committee.
ACKNOWLEDGMENTS
Funding for market surveys in Japan was provided by the International Fund for Animal Welfare and
the endowment of the Marine Mammal Institute of Oregon State University.
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Baker et al. SC/60/BC2 7 of 8
Table 1: Species origins of whale, dolphin and porpoise products purchased in commercial markets of Japan
between 1993 and 2007, as determined by phylogenetic reconstruction of sequences from the mtDNA control
region. All identifications of species or family were supported by high bootstrap values >95% using the methods
described previously (Baker et al. 1996b; Baker and Palumbi 1994). Sample sizes indicate number of products
identified to species, not necessarily the number of unique individuals represented by the products (see text for
discussion).
Species Japan Japan Japan Japan Last legal source or year of
1993-03 2003/04 2006 2007 international protection
Mysticeti
Northern Pacific minke, 231 40 5 15 Japanese scientific whaling, ongoing
Balaenoptera acutorostrata since 1994
Antarctic minke, 466 20 2 6 Japanese scientific whaling, ongoing
B. bonaerensis since 1987/88
Common-form Bryde's whale, 23 4 - 13 1986 moratorium, or 1987 by Japan
B. brydei following Wada et al. under objection. Japanese scientific
2003 whaling (JARPNII) since 2000.
coastal-form Bryde's whale, B. 0 - - Species not formally recognised until
edeni following Wada et al. 2003 but presumably protected since
2003; also Baker et al. 1996b, 1986 moratorium, or 1987 by Japan
Yoshida and Kato, 1999) under objection
Sei, B. borealis 16 16 6 21 1986 moratorium, 1977/78 for Antarctic;
1977, Brazil; 1978, Peru; 1979, Chile,
1979; 1980, Spain; 1988 Iceland; 2002
for Japan (JARPNII)
Fin, B. physalus 35 2 15 39 1986 moratorium, or 1989 for Iceland
scientific whaling; Japanese scientific
whaling in the Antarctic (JARPAII)
since 2005/06
Blue/Fin hybrid* (confirmed in 2 - - - 1989, Iceland scientific whaling
2008) (Cipriano & Palumbi 1999a)
Humpback, Megaptera 7 1 3 5 1966
novaeangliae
Gray, Eschrichtius robustus 7 - - - 1937
Odontoceti
Sperm whale, Physeter 6 - - - 1986 moratorium, or 1988 by Japan
macrocephalus under objection. Japanese scientific
whaling (JARPNII) since 2000.
Dwarf Sperm, Kogia simus 1 - - - No international protection
Beaked whales, Berardius 56 3 - - No international protection
bairdii, Ziphius cavirostris,
Mesoplodon carlhubbsi, M.
densirostrus
Killer whales, Orcinus orca 1 - - - No international protection
False killer whale, Pseudorca 1 - - - No international protection
crassidens
Dolphins – including pilot whales 119 6 - - No international protection
and members of the
Stenella-Tursiops-Delphinus
complex etc (>6 species)
Finless porpoise, Neophocaena 0 - - - No international protection
phocoenoides
Dall’s porpoise, Phocoenoides 14 - - - No international protection
dalli
Other mammals
Sheep 2 - - - n/a
Horse 3 - - - n/a
Total 990 92 31 99 1504 (Grand Total)
Baker et al. SC/60/BC2 8 of 8
AY582748-NA&
AY822094_finB-NA&
AY822097 finE
100 AY822103 finK
100
JE93.B02(07)
100 J98.C17
BphICE27&BphUA&J98.054
BphMd04
BphMd08
J95b.073
J95b.079
J97.015
100 BphCAN North Atlantic
BphMd01&J95b.082
BphMd03 and market,
BphMd05&BphMd07 1993 - 2003
100 BphMd11
BphMd12
J00.001
100 100 J03.019
JS93.003&JS95.069
JS95.031
100 BphN1
J97.024
JE93.B17(38)
JS93.011
J97.012
J0720&21&22&23&
AY822093_finA-NP
AY822106_finN-NP
100 J00.028&J01.039
100 J01.040
100
100 J06.001&J06.022&
J02.125&J03.011
100 J99.095
100 100 J0740A&
100 100
J0708
J0753
J0767&J0724&
J00.034&J99.077 North Pacific
100 100 J03.078&J04.022
J06.028&J0718A and market,
100 J97.017 1993 - 2007,
JS95.076
100 J98.048 including
JS93.041 JARPA II
J0703A&J0705&
J0716&J0758&
J06.005&J06.021
J06.015
J06.018
J0711
J0737
J0739
AY822095_finC-NP
AY822096_finD-NP
AY822098_finF-NP
100 AY822099_finG-NP
AY822100_finH-NP North Pacific
Ay822101_finI-NP
AY822104_finL-NP
AY822105_finM-NP
100
100
100
Blue whale
outgroup
Fig. 1. Phylogenetic relationships among fin whale reference sequences (with Genbank numbers) and market
products purchased between 1993 and 2007, based on mtDNA control region sequences. Numbers above
branches indicate strict consensus of many equally parsimonious trees. Arrows point to the 15 haplotypes found
in the 2006 and/or 2007 market surveys. Single arrows point to haplotypes found in only one survey year;
double arrows point to products found in both survey years. Haplotypes representing multiple products are
indicated by an ‘&’.