IMMUNOGENETIC AND MARKING APPROACHES TO
IDENTIFYING SUBPOPULATIONS OF THE
NORTH PACIFIC WHALEsn
KAZUO FUJINO
To identify local subpopulations of whales, various methods have been
applied. These include indirect methods basing upon oceanographic information, seasonal change of localization of whaling grounds and catch
statistics, parasitological analysis, and direct methods of morphologicaland marking investigations. Recently another method using ecologicalcharacters was adopted to analyse mingling of southern humpback populations (Chittleborough, 1959).
Morphological and marking investigations are most frequently utilized
among these. Most morphological characteristics are controlled withpolygenic system and are apt to be affected by non-hereditary factors
during growth of individuals, so that discrimination of different populations may sometimes run into difficulties. On the other hand, marking
investigation gives the most definite informations about movements ofmarked whales. To deduce, however, distribution of general population
to which marked whales are belonged, effects of various conditions whenmarked or recaptured and of ecological factors should be discussed.
From these reasons, another method of population genetics might bringmore useful informations. In this method, some characters which arecontrolled with comparatively simple allele system, such as blood groups
in human (Boyd, 1950) (Race & Sanger, 1954), in cattle (Owen, Stormont& Irwin, 1947), in fowl (Brile et al., 1958) and pigmentations in insect(Komai, Chino & Hoshino, 1956) are used as genetic marker.
Recently serological characters have been utilized as 'genetic marker'
to identify intraspecific subpopulations of marine animals commerciallyvaluable in fisheries. These include studies on tuna (Cushing, 1956;
Suzuki et al., 1959), salmon (Takahashi & Suyehiro, 1957; Ridgway,
Cushing & Durall, 1958), Pacific sardine (Sprague, 1959), Atlantic herring(Sinderman, 1959), fur-seal (Fujino & Cushing, 1960) and whales (Fujino,
1956; Cushing, Fujino & Takahashi, 1959).
In the present paper serological basic studies on whales, carried out
since 1952, are summarized and analysis of raciation of the North Pacificfinback whales by means of population genetics with blood group and
1> Supported in part by the grants of the Japanese Fisheries Agency during from years
1955 to 1958. Outlines were read at International Oceanographic Congress, New York,
Sept., 1959.

85

86

K. FUJINO

marking investigations are discussed with attention to complemental
relationships between these two methods.
BASIC PROBLEMS IN SEROLOGY

To identify whale populations by means of blood typing investigation,
not only analysis of serological characteristics of themselves but also
standardization of experimental methods and materials should be undertaken. Results of discussion on these technical problems are describedin the following section. Other general methods used in successive
sections are same as those in the literatures cited there.
Technical problems, especially preserving methods of materials andblood typing with preserved materialsa) Glycerol-freezing preservation technique of intact erythrocytes.
Race and Sanger (1954) stated about studies on glycerol-freezing
technique, 'There seems general agreement that antigens are wellpreserved at least up to a year and probably much longer. The method
has been of great use in keeping available samples of blood representing rare genotypes'. Cushing et al. (1959) reported an application of thistechnique on marine animals of which sterile precautions are not available.
After collecting blood samples are added equal volume of a mixture offour parts of glycerol and six parts of five percent sodium citrate solution(additional antimicrobial of guanofuracin was used at a concentration of
200 mg per liter against accidental warming), and then the mixture are
kept frozen.
After transportation to laboratory, frozen samples are thawed under
running water and are dialysed against 1.5 percent saline for two hours,
and are washed several times with saline. Then the cells available fortest will be obtained in good yield. All blood samples of finback andhumpback whales, collected in 1958 and 1959, were analysed throughthis technique.
b) Blood typing of dried erythrocytes with 1311-labelled antibody.
Fujino & Cushing (1959) developed another blood typing technique using
dried erythrocytes and radioactive antibodies. Outlines of this techniqueare as follows. After being diluted one in ten with saline, one drop of
fresh blood sample is dried on stripe of filter paper and is preserved indesiccator. Then 1311-labelled antibody is diffused on the filter paperwith solvent of four per cent ammonium sulfate through the erythrocytes.
Specific radioactivity on blood spot shows positive reaction between
antibody and corresponding erythrocytes. According to their conclusive
description, this technique will be useful in blood-typing of dried erythrocytes, but there are still general problems about preparation of

SUBPOPULATIONS OF THE NORTH PACIFIC WHALES

87

samples before applying to large scale of investigation.
c) Preservation of blood typing reagents. To undertake blood typing
investigation of marine organisms in relation to fisheries problems, large
number of sample should be tested for a long period. For this purpose
it is desirable to use same standard reagents during a series of investigation. In past time many workers used formalin, merthiolate, carbolic
acid and sodium azide as preservative of serum. While Race & Sanger(1954) stated that reagents for human blood typing can be preserved
for long time by freezing at a temperature of -20°C without preservative. As blood typing of marine animals, however, is sometimes
carried out in field laboratory, it is required to add preservative toserums against warming during transportation. In practical, after
inactivation serum reagents are added 1/10 volume of five per cent ofcarbolic acid or one per cent of sodium-azide, and are kept frozen at
temperature -15 or -20°C. Another series of reagents divided fromoriginal serums have been kept frozen at same temperature without
preservative for use in central laboratory. No significant differences in
specificities and titers has been recognized between both series of reagents even after three years' preservation so far.
Classification and genetics of blood group
Agglutination or hemolysis reaction with immune antibody, isohemagglutinin and natural antibody are used for detecting blood groupantigens. Summarized results of blood typing on four species of baleen
whales and three species of toothed whales detected so far are shown in
Table 1. Details by species will be described in following paragraphs.
a) Blue-white dolphin, Stenella caeruleo-albus1 ). Two kinds of antigens are detected by agglutination and hemolysis reactions with rabbit
immune serums, and isoagglutinins specific to these two antigens arefound in low frequency (Yamaguchi & Fujino, 1953). Parenthesized
figures in Table 2 show expected frequencies of occurrence of phenotypes
calculated through hypothetical two allele system on population in HardyWeinberg's equilibrium.
It is not clear at present whether discrepancies between expected
figures and observed those are caused by scantiness of samples tested
or in compatibility of the allele system adopted.
Another kind of agglutinogen was detected in S. 23 dolphin cells, andoccurred one out of ten individuals tested. No information on genetics
on this antigen has not been obtained so far (Cushing, Fujino & Takahashi, 1959; Fujino & Cushing, 1959).
1 > More recently these common name and scientific name are used for this species
which blood typing were reported by Yamaguchi and Fujino (1953).

Baleenwhale

B. musculus

Megaptera nodosa

Humpback whale

B. physalus

Fin whale

Blue whale

Balaenoptera edeni

Bryde whale

Physeter catodon\ Sperm whale-J

Yamaguchi & Fujino (1952)

Literature

Agglutination reaction withl
immune and natural
J Fujino (1956)
antibodies\Fujino (1953)
Ditto and isoagglutination J\

Agglu~inatio~an~ hemolysislFujino (1954)
reactions with immune J
antibodies

Detecting method of antigens\ 1, 2, 3, 4Ifinback Ju !mm;ine andnatural ant1bod1es

Agglutinatio~ with anti-} Cushing, Fujino &Takahashi (1959)

Bm1+, Bm1- and Bm2+, Bm2-\Agglutination and hemolysis}
with immune antibodies Fujino (1953)
JMn1+, Mn 1- and Mn 2+, Mn2 and isoagglutination

Ju1Ju2, Jui, Ju2(Bp1+, Bpi- and Bp2+, Bp2-

Bb1+, Bb1- and Bb2+, Bb2-

{ Pc1Pc 2, Pc1, Pc2
Sp+, Sp-{ Br1Br2, Bri, Br2
Pb+, Pb-

Berardius bairdii

Phenotype

Toothed Baird-beaked whale
whale

Scientific name

Blood groups

BLOOD GROUPS IN VARIOUS SPECIES OF WHALES

Stenella caeruleo-albus Dc1Dc2, Dc1, Dc2

Common name

Whale species

TABLE 1.

Blue-white dolphin

Classificationofwhales

6

r
d
';::l

co
co

89

SUBPOPULATIONS OF THE NORTH PACIFIC WHALESTABLE 2.

FREQUENCY OF OCCURRENCE OF BLOOD TYPES OF
BLUE-WHITE DOLPHIN
Blood typeSum
Male
FemaleDc1
Dc1Dc2
Dc2
No. of whales observed

.200
.500
.300
20

.187
.688
.125
16

.194 (.235).583 (.500).223 (.265)36b) Baired-beaked whale, Berardius bairdii. Three kinds of antigens
Br11 Br2 and Pb are found by agglutination and hemolysis reactions withimmune rabbit serums. According to phenotypic relationships Pb antigen
seems to occur independently to Br1 and Br2 antigens. Frequencies ofoccurrence of phenotypes are shown in Table 3. Parentages of blood
types are tested on two pairs of cow and fetus.
TABLE 3.

Br system

Pb system

FREQUENCY OF OCCURRENCE OF BLOOD TYPES OFBAIRD-BEAKED WHALE

Blood type

Male

FemaleSum

f Br1 2

.820
.103
.077
39
.125
.875
32

.750
.083
.167
12
.OOO
1.000
8

.804
.098
.098
51
.100
.900
40

l
lf

BriBr
Br2
No. of whales observed
Pb present
Pb absent
No. of whales observed

After being tested with agglutination-inhibition reaction, blood types
both of male f etus 8 feet 3 inch. long taken from 37 feet cow and offemale fetus 8 feet 7 inch. long taken from 36 feet cow were identified
as Br1 Pb- which are same as those of their cow (Fujino, 1954).
Informations on isoagglutinin and allele system of three antigens havenot been obtained yet.
c) Sperm whale, Physeter catodon. Three kinds of antigens of Pc11
Pc2 and Sp are found by agglutination and hemolysis reactions withTABLE 4.

FREQUENCY OF OCCURRENCE OF BLOOD TYPES OF
SPERM WHALES

Blood typefPei
Pc system ( Pc 1Pc 2
Pc2
Sp system { Sp present
Sp absent
No. of whales observed

Male

Female

Sum

.400
.300
.300
.400
.600
10

.313
.313
.374
.500
.500
16

.346
.308
.346
.462
.538
26

immune rabbit and fowl serums. According to phenotypic relationships,
antigen Sp belongs into different allele system from those which antigens
Pei and Pc2 are concerned. Frequency of occurrence of phenotypes isshown in Table 4.

90

K. FUJINO

Natural antibody specific to Sp antigen is detected in the serums of sei
whales, Balaenoptera borealis and agglutinates Sp positive cells up to
dilution of one in eight or one in sixteen (Fujino, 1954). No available
data about isoagglutinin specific to known antigens and allele system has
been obtained yet.
d) Bryde whale (Ogasawara whale), Balaenoptera edeni1 ). Twokinds of blood group antigens Bb1 and Bb2 are detected by immune
rabbit agglutinins and hemolysins in the Bryde whale taken from Bonin
Islands' area, and occur as shown in Table 5 (Fujino, 1953). Sufficient
data discussing on allele system of both antigens are not obtained yet.
According to phenotypic relationships, however, antigens Bb1 and Bb2
seem to be controlled by different allele systems each other. No agglutinin, which has sufficient high titer and specificity to identify the
known antigens, has been positively detected yet.
TABLE 5.

FREQUENCY OF OCCURRENCE OF BLOOD TYPEANTIGENS OF BRYDE WHALEo

Present or absent
of antigen·/ Bb1 present
Bbi antigen \ Bbi absent
f Bb2 presentn \ Bb2 absent
No. of whales observed

Bb2 antige

Male

Female

Sum

.364
.636
.273
.727
22

.500
.500
.300
.700

.407
.593
.281
.719
32

10

e) Finback whale, Balaenoptera physalus. At first two kinds ofantigens Bp1 and Bp2 were found by immune rabbit agglutinins andhemolysins (Fujino, 1953). According to phenotypic relationships these
TABLE 6.

TWELVE PHENOTYPES OF FINBACK BLOOD GROUPS
CLASSIFIED WITH Bp AND Ju ANTIGENS*
Ju antigen
Bp antigen
JuiJU2Ju2
Jui
Bpi and Bp2 present
BpiBP2 Ju2
Bp1BP2 Jui
Bp1BP2 JuiJU2
Bp1 present
Bpi JuiJU2
Bpi Ju2
Bp1 Ju.i
Bp2 JuiJU2
Bp2 Ju2Bp2 present
Bp2 Jui00
Bpi and Bp2 absent
0
JuiJu2Ju2
Jui* cited from Fujino (1956)

two antigens seem to be controlled by different allele system each other.
Additional two antigens belong to Ju system were detected independently
to the known two antigens by immune rabbit and fowl agglutinins. In
consequence twelve phenotypes can be possible as shown in Table 6.
o Sei whales taken from Bonin Islands' area (Fujino, 1953) are identified as different
species from Balaenoptera borealis (Omura, Nishimoto & Fujino, 1952). More recently,
these common name and scientific name are adopted to this species by Omura et al. (1957)
and Omura (1959).

91

SUBPOPULATIONS OF THE NORTH PACIFIC WHALES

Another new antigen X were detected positively in Bp2J u 1 and OJ u 1
type cells, but relations to the known antigens have not been studied
enough yet (Fujino, 1956).
Cushing et al (1959) stated that homozygous and heterozygous types,
which belong to Ju system, reveal dosage effects against anti-Ju2 antibody. This was confirmed with additonal experiments using materials
TABLE 7.

DOSAGE EFFECT IN HETEROZYGOUS TYPE OF
FINBACK Ju BLOOD GROUP

Anti-Ju2 no. 50 immune rabbit serum (absorbed)
Dilution
Cells
503
!181
Ju1Ju2 \ 455

20

40

80

160

320

640

1280 2560 5120 10240

tit
tit
tit

tit

tit

tit

tit

*tit tit* tit+

Ju1

Ju2

504

* *tit tit+
tit

tit

+

+

256

512

* *

+

Anti-Ju 2 no. 34 immune rabbit serum (absorbed)
Dilution
Cells

Jui

10

20

40

80

tit
tit
tit

tit
tit
tit

tit
tit
tit

+H-

503

Ju1JU2 I 181
\ 455
504
Ju2

160

320

*tit tittit+ tit*+

No. 2 horse serum (unabsorbed)

640

1280

+

*

+

Dilution
Cells
1
503
{181
\ 455
504

+
+

tit

+

4

8

16

32

64

128

* tittittit
*

tit
tit
tit

tit

tit

*
tit

+

2

tit

Anti-Ju1 no. 47 immune fowl serum (absorbed)

+

* tit

tit

Dilution
Cells
Ju1

503

Ju1JU2 { i~§
Ju2

504

~

10

20

40

tit
tit
tit

tit
tit
tit

**

*

80

+
+
+

which were collected by glycerol-freezing technique in 1959. Conversely,
this phenomena may serve to distinguish heterozygous type from homozygous type.
Moreover another two types which revealed dosage effect against
anti-Ju 2 serum were found among Ju heterozygous types (Ju1 Ju2) asshown in Table 7. In this table results of reaction against anti-Ju1

92h. FUJINO

serum show that there is no difference in antigenic activity between
nos. 455 and 181 heterozygous erythrocytes after preservation. Large
number of investigation about this point has not been carried out, but
the results of Table 7 show that heterozygous type (Ju1Ju2) may be
subdivided into two groups, that is, stronger one and weaker one.
As shown in Table 8 natural antibodies specific to Ju antigen aredetected positively in the serums of fowl, rabbit, pig, horse and cattle.
As already reported in part by Cushing et al. (1959) no difference has
been recognized on anti-Ju 2 specificity between immune and natural
antibodies so far. Additional advantage, which these natural antibody
occur in high frequency, will be potentially useful in large scale ofblood typing investigations hereafter.
TABLE 8. ANTI-FINBACK Ju NATURAL ANTIBODIES FROMSERUMS OF VARIOUS SPECIES OF ANIMALSO
Animal Specificity ofantibody
fanti-Ju1
Fowl\anti-Ju2

Agglutinin
titer2J

1:2
1:4

Frequency ofoccurrence
one out of threeditto

Rabbitanti-Ju 2

1:8-1:64

three out of fivePig

anti-Ju2

1:256

only one tested

Horse

anti-Ju2

1:64-1:2560

all of eleven

Remarkdetected from a) same individual

Five out of eleven

rpossess finback species
1 ~pecific antibody

lm common.
1:32-1:40960
all of thirteen
1) partly reported already by Fujino (1958) and Cushing, Fujino & Takahashi (1959).
Cattle2)anti-Ju 2

agglutinin titer against homozygous type Ju 2 in room temperature.
TABLE 9. BLOOD TYPE ANTIGENS OF FINBACK WHALES USED FOR
LARGE SCALE OF INVESTIGATIONS IN VARIOUS AREAS
Year

1952
54
55
56
57
58
59

East China Kamchatka
Sea
off
Bp
Bp and Ju
Bp and Ju
Ju
Ju
Ju
Ju

South of east
Aleutian

North of east Off Navarin and
Aleutian
Olyutorskiy

Bp and Ju

Bp and Ju
Bp and Ju

Ju
Ju

Ju
Ju

Ju

While Fujino (1958) stated that immune animal which has natural
antibody specific to Ju antigen can produce more excellent antibody in
specificity and titer against corresponding Ju antigen than other animals.
Basing upon the above stated knowledges have been undertaken large
number of blood typing of finback whales from various areas of thenorth Pacific.
Fujino (1956) stated that in large scale of investigation blood typing

64
129

429

.744

.256

.851

.149

sum

438

348
786

.109 .114.034 .042

121

.025

.025

99

.051

.030

.950 .919

220

.036

.027

.937

male female sum,---------'~

13

.538

.462

.005

.984

810

821

.012 .011

.007

.981

.024 .057
41

.012
1631

35

.024 .029

76

.040

.026

z

5

13

'°
(;.)

Ul

;i:..
t"'
M

~
::i::

.OOO .OOO .OOO
80

.OOO .OOO .OOO

" Ij

;i:..
0

--

'"ti

::i::

Olyutorskiy,--------'-----------

z

M

::i::

....,

"Ij

0

Ul

0

....,

;i:..

ct"'

'"ti

0

'"ti

cco

Ul

0
?::!
....,

1427

.781

.219

1.000

.OOO

male female sum

701

.779

.221

1.000

.OOO

sum

.952 .914 .934 1.000 1.000 1.000

male female sum,--------''-----------.._

Navarin

726

.782

.218

1.000

.OOO

female~North of E. Aleut.

male

.006

.982

male female sum

~

32

.500

.500

1.000

.OOO

.OOO
1.000

sum

femaleNorth of E. Aleut.

19

.474

.526

1.000

.OOO

male

~

South of E. Aleut.

1) Include data of Fujino (1953), Fujino (1956), Cushing, Fujino & Takahashi (1959) and additional those in 1959.
2) Figures in 1952 and 1954 & 1955 are noted separately here, because same reagents were not used in these years.

65

.119

.200 .125

Ju2

No. of whales tested.048.108 .094 .101

Ju1JU2

.844

.833

.736

.692 .781

Jui

.857

male female sum

male female sum

.163

197

.736

.264

.852

.148

female

Off Kamchatka South of E. Aleut.

232

.750

.250

.849

.151

,--------'~

East China Sea

200

.770

.230

.715

.285

,--------'~Ju blood group

100

100

No. of whales tested

.270

.190

{present

.730

.740

.690

absent

.810

.260

.310

{present

absent

BP2

Bp,

male

~

sum

~

female1954 & 55

1952

male

Bp antigen

Off Kamchatka2>

TABLE 10. FREQUENCY OF OCCURRENCE OF Bp AND Ju BLOOD GROUP ANTIGENS OF FINBACK WHALES11

94

K. FUJINO

in Ju system can give higher precision than in Bp1 and Bp2 antigens
because of some misclassification resulting from unknown antigen-antibody
reactions involved. Therefore, Ju system alone is used for practicalinvestigation ever since. Blood groups, which were adopted for investigations in various areas in each year, are shown in Table 9. Table
10 shows their frequency of occurrence obtained. In this table theresults in 1952 are described separately from those in 1954 and 1955,
because in 1952 no discussion was performed on the effect of X antigen
which was found subsequently. It is clear from this table that BP1
antigen which occurs in the area off Kamchatka does not occur in thearea of east Aleutian and that frequencies of Ju2 antigen show geographical gradient descending from western to eastern sides of thenorth Pacific. Problems on identification and mingling of intraspecificsubpopulations will be discussed later.
TABLE 11. SIZE DISTRIBUTION OF FIN WHALE FETUSES OF
WHICH BLOOD GROUPS WERE IDENTIFIED
Male
FemaleBody length
in feet
Ju1Ju2
Ju2
Ju2
Ju1
Ju1
Ju1JU2
4
1
1
1
3
5
6
4
1
6
7
1
2
1
7
8
6
1
7
1
9
3
2
10
3
1
11
2
1
3
12
3
2
13
2
2
1
14
1
1
1
Sum
32
6
27
2
2

Parentages of blood types between cows and their f etuses taken andpopulation genetics using relative frequencies of occurrence of eachphenotype were used for finding allele system of blood types. Blood
samples taken from umbilical cords of fetuses were used for analysis.
Embryological problems of blood types in fetal stages were taken into
consideration. After discussing developments of human blood types infetuses by means of agglutination and agglutination-inhibition reactions,Mizu (1931) and Hirasawa (1935) stated that phenotypes in ABO andMN systems can be already identified approximately three months after
fertilization. Therefore, data on whale fetuses, of which blood types
were definitely identified with above stated two reactions alone are used
for discussion of parentages of blood types in whales.
Table 11 shows body length distribution of fetuses of which blood

SUBPOPULATIONS OF THE NORTH PACIFIC WHALES

95

tpyes were able to be identified. Judging from averaged growth curve
estimated by Nozawa et al (Nozawa et al. 1956, pp. 44-55) and Ohsumi
et al. (1958) it may be assumed that fetal blood types of the north Pacificfinbacks can be identified already five or six months after fertilization.
The parentages of blood types observed are shown in Table 12. According to this table Ju blood type seems to be hereditarily controlled
by two allele system, but one exception of parentage has been observed.
That is, Ju1 type fetus was taken from Ju2 type cowl). This does not
fit hypothesis in two allele system. Therefore, further discussions will
be taken up by means of population genetics using relative frequencyof occurrence of phenotypes as follows.
TABLE 12.

PARENTAGES OF Ju BLOOD GROUPS OF FINBACKSll

Mother whale1>

no. of tested
blood group
Ju1
56
JuiJu2
7
7
Ju2
Sum
70
Results in the years 1954, 55 & 59.

Blood groups of fetus

Ju1
53
5
1
59

JuiJU2
3

Ju2
0

1
4
8

2
3

1

Phenotypic relationships of Ju blood types noted in Table 6 are
discussed as follows from view points of presence or absence of antigens.
i) Erythrocytes lacking Ju 1 antigen are certain to have Ju 2 antigen, and erythrocytes lacking Ju 2 antigen are certain to have Jui antigen. That is, no erythrocytes which has neither
antigen Jui nor Ju2 has been observed.
ii) Erythrocytes possessing both antigens Jui and Ju 2 exists.

Probable hypothetical two allele and three allele systems were discussed
by means of population genetics. At first relations of dominancy among
alleles were defined as follows. Parenthesized symbols mean frequencyof occurrence of each gene. In two allele system, there is no relationof dominancy between j 1 (p) and j2 (q) genes. In three allele A
system, j~ (q) gene is completely dominant against j 2 (r) gene and no
relation of dominancy exist between j2 and j 1 (p) genes. In threeallele B system, j~ (r) gene is completely ressesive against j 1 (p) geneand j 2 (q) gene is equivalent to j 1 gene. Then genotype-phenotype
relations are given in Table 13. In two allele system one phenotype
corresponds to each one genotype. In three allele A system Ju1 phenotype
includes four genotypes and other two phenotypes have one genotype
each. In three allele B system phenotypes Jui. Ju1Ju2 and Ju2 include
two, one and three genotypes respectively.
tl According to the log supervised by Mr. K. Nasu, Whales Research Institute, these are no.
1542 cow 70 feet long and male fetus 8 feet and 6 inch. long taken in the area of the northof east Aleutian (54°27 1N, 168°321W) in Aug. 4, 1959.

K. FUJINO96
TABLE 13.

PHENOTYPE-GENOTYPE RELATIONSHIPS IN THREEKINDS OF ALLELE SYSTEMSGenotype

Phenotypetwo allele systemthree allele A systemthree allele B system

id1
iii2
iziz

idi. id1a, i1aha, hai2
id2
iziz

i1h, id2r
id2
iziz, izizr' izrizr

In a population at Hardy-Weinbergs' equilibrium, following relations
between frequencies of occurrence of phenotypes and those of genotypes
can be formulated. That is, in two allele system,[Jui]=pz[Ju1Ju2]=2pq[Ju2]=q2

(1)

p+q=l

, in three allele A system[Jui]=(p+q)2+2qr )l]uilu2]=2pr( 2)[Ju2]=r 2

p+q+r=l, and in three allele B system[Ju1]=p2+2pr[Ju1Ju2]=2pq[Ju2]=(1-p) 2( 3)

p+q+r=l, where [Ju1], [Ju1Ju2] and [Ju2] mean frequency of occurrence of eachphenotype and a relation of [Ju1]+[Ju1Ju2]+[Ju2]=l is provided.
TABLE 14.

FREQUENCY OF OCCURRENCE OF PHENOTYPES AND GENES OF Ju
BLOOD GROUPS OF FINBACK WHALESSouth of east North of east
Freq. of occurrence
East China Sea Kamchatka (A)Aleutian (B)Aleutian (C)of phenotypes,.---------.__,_and genes
act. no. freq.
act. no. freq.
act. no. freq.
act. no. freq.
.731
.844
663
.937
1602
.982
95
206
flJuil
.042
.027
13
.101
33
.006
Observed
10
6l [Ju1Ju2]
.163
90
.114
21
19
.012
8
.036
[Ju2]
sum
129
786
220
1631
.787
.865
.951
fp
.985
Two-allele
.213
.135
\q
.049
.015
.620
588
.748
system, exp. {[Ju1l80199.1 .905
1582.4 .9702
184.234
value [Ju1Ju2]
43.335
20.5 .093
48.3 .0296
.045
14
.018
6
[Ju2]
0.4 .002
0.3 .0002
.125
.062
.071
.027
.471
.600
.739
.863
Three.338
.404
.190
lr
.110
allele
.662
.596
.810
.890
system*
.032
.085
.017
.003
.306
.319
.173
.107
lr
* Calculated for convenience of comparison under a hypothesis which population in eacharea is in Hardy-Weinberg's equilibrium.
~

rAf~

lBf~

~'-----.

97

SUBPOPULATIONS OF THE NORTH PACIFIC WHALESTABLE 15. EXPECTED FREQUENCY OF OCCURRENCE OF BLOOD TYPES
OF CALVES FROM VARIOUS MATINGS IN HYPOTHETICAL
THREE ALLELE SYSTEMS
1) In A system
Expected blood type of calves
Mating~genotype
phenotype_.,,---_
phenotypegenotype
1
jdi<l h<lji<l h' h jih
JuiJu2 Ju2
Jui
jdi
hh
p4
p4
[Jui]
x[Jui]
hh Xhh
p3q
p3q
2p3q
xjdia
p2q2
p2q2
x ji<lji<l
Xji<ljz
p2qr p2qr
p2qr
p2qr
p2q2 2p2q2 p2q2
4p2q2
hh<l Xjiji<l
pq3
pq3
2pq3
Xji<lh<l
Xji<ljzpq2r pq2r pq2r pq2r
3pq 2r pq 2rq4
q4
ji<lji<l x ji<lh<l
qsr
qsr
2q 3r
Xji<lfa
q2r2
q2r2 2q2r2
q2r2
3q2r2
ji'ij2 Xjiahpsrpsr
psr psr[Jui]
x [JuiJu2] hh Xjifa
p2qr p2qr
p2qr p2qr
3p2qr p 2qr
hh" Xhh2pq2'Y'pq2r
pq2r
hahaxhhpqr2
pqr2 pqr2 pqr2
2pqr2 pqr2 pqr2
ji<lfa Xhh
p2q2
p2q2
[Jui]
x[Ju2]
jdi Xfahpqr2 pqr2
pqr2 pqr2
jdia Xfaj2
q2r2
q2r2
ji<lji<1Xj2fa
qrs
qrs
qrs
qrs
ji<ljz Xhh
p2r2
p2r2 2p2r2 p2r2
2p2r2 p2r2
[Ju1Ju2] X [Ju1Ju2] jih Xj1fa
prs
pra
prs
pra
[Ju1Ju2] X [Ju2]hh Xfajz
r4
r4
[Ju2]
x[Ju2]
j2h Xhh2)

In B system
Expected blood type of calves
Matingphenotypegenotype
phenotype[Jui]

X[Ju1][Jui]

x [Ju1Ju2][Jui]

x[Ju2][Ju1Ju2] x [Ju1Ju2][Ju1Ju2] X [Ju2][Ju2]x[Ju2]

genotype
jd1 Xjd1
Xjd2r
jd2r Xhhr
jd1 Xj1h
jifar Xjd2
jd1 Xhh
Xfafar
xfarjzr
hhr Xfah
Xjd2r
xfarhrhh Xhhjd2 Xj2h
Xhh"
Xj2"hrj2h Xhh
Xhh"
Xj2"hrhhr Xjd2"xhrhr
j 2rj 2r.)( jzrfar_______..,~

jd1
p4
psr
p2r2
psr
p2qr

p2q2

jihr

hh

hh

par
2p2r2

hhr

jzrjzrp2r2

psr
p2qr p2qr
p2q2
p2qr p2qr
p2q2
pq2r
pqr2 pqr2
prB
2p2q2 p2q2
pq8
pq3
pq2r pq2r pq2r
pqr2
q4
qsr
q2r2

p2qr

pq2r
pqr2 pqr2
prs

pq2r
pqr2
qsr
q2r2
2q2r2 q2r2
qrs
qrs
r4

Jui
Ju1Ju2 Ju2
p4
2p 3r
p2r2
3p2r2
pBr
psr
2p2qr p 2qr p2qr
p2q2
p2qr p2qr
p2q2
pq2r pq2r
pqr2 pqr2 2pqr2
pr3
prs
p2q2 2p2q2 p2q2
pq3
pq3
pq2r pq2r 2pq 2r
pqr2
pqr2
q'
2q 8r
q2r2
4q2r2
2qr 3
r4

K. FUJINO

98

Table 14 shows gene frequencies calculated from Table 10 throughthese formulas. Results of statistical tests shows that Ju blood typedoes not fit hypothetical two allele system. This confirm incompatibility of two allele system which was previously noted in parentages.
Therefore, analysis of raciation and mingling of different populations
will be discussed through three allele A and B systems.
Frequency of expected blood groups in fetuses from various matings
can be noted in Table 15 through two ways of A and B systems.
Dosage effect in heterozygous types noted in Table 7 would propose
existence of subtypes to which an additional gene will concern. However,informations sufficient to statistical treatment have not been obtained yet.
f) Blue whale, Balaenoptera musculus. Two kinds of antigens Bm1
and Bm 2 are found in the erythrocytes of north Pacific blue whale withrabbit immune agglutinins and hemolysins. Frequencies of occurrenceof antigens are shown in Table 16. Isoagglutinins specific to known
TABLE 16.

FREQUENCY OF OCCURRENCE OF BLOOD TYPEANTIGENS OF BLUE WHALES

Presence or absenceof antigen
Bm1 antigen f Bm1 present\ Bm1 absent
Bm 2 antigen I Bm2 present\ Bm2 absent
No. of whales observed

Male

Female

Sum

.033
.267
.133
.567
30

.059
.411
.059
.471
17

.043
.319
.106
.532
47

antigens occur irregularly. Titers of anti-Bm 1 reach up to 1: 4 or 1: 64
and those of anti-Bm 2 up to 1: 2 or 1: 64 (Fujino, 1953). According
to phenotypic relationships, antigens Bm 1 and Bm 2 seem to be controlled by genes belong to different allele systems each other, though
informations enough to discuss has not been obtained.
g) Humpback whale, Megaptera nodosa. Two kinds of blood groupantigens Mn 1 and Mn 2 are found in the north Pacific humpback whales
TABLE 17.

FREQUENCY OF OCCURRENCE OF BLOOD TYPEANTIGENS OF HUMPBACK WHALES

Presence or absenceof antigen
Mn1 antigen f Mn1 present\ Mn1 absent
Mn2 antigen f Mn2 present\ Mn2 absent
No. of whales observed

Male

Female

Sum

.111
.278
.056
.555
18

.118
.235
.118
.529
17

.114
.257
.086
.543
35

with immune rabbit agglutinins and hemolysins. Frequency of occurrence of each antigen is shown in Table 17. Isoagglutinins specific toknown antigens were detected positively at high frequency. Out of ten

99

SUBPOPULATIONS OF THE NORTH PACIFIC WHALES

samples tested, three have anti-Mn11 three have anti-Mn 2 , two have bothand two have none of agglutinins. Titers of anti-Mn1 and anti-Mn 2 agglutinins reach up to 1: 2 or 1: 128 and 1: 4 or 1: 128 respectively.
Table 18 shows results of isoagglutination (Fujino, 1953).
TABLE 18. ISOHEMAGGLUTINATION IN Mn BLOOD TYPEANTIGENS OF HUMPBACK WHALES
Erythrocytes
Serums
142

143

142
143
151
180
184
191
192
203
204
206

151

184

180

192

191

203

204

206

*
*
*
*

*
*+
*
*
*

+ft
+ft
+ft
+ft

Additional four kinds of blood types were . temporarily classified on
humpbacks taken from Okinawa waters with anti-finback Ju agglutinins(immune rabbit and fowl serums and natural antibodies from cattle and
TABLE 19. NEW BLOOD TYPES OF HUMPBACK WHALES TEMPORARILY
CLASSIFIED BY ABSORBING TESTS OF ANTI-FINBACK Ju SERUMS
Blood types of erythrocytes used as agglutinogen

Blood types ofabsorbingerythrocytes
1
2
3

4

1

2

3

+
+
+

+
+

+

4

TABLE 20. ISOHEMAGGLUTINATION IN HUMPBACK WHALE BLOOD
TYPES CLASSIFIED BY ANTI-FINBACK Ju SPECIFICITIESBlood types of erythrocytes
Serums
K 9
K24
RIO
R29
K 4
R38

1
K9

+ft
+ft
+ft
+ft
+ft

2
K24

3
RIO

3
R29

+

+

4
K4

4
R38

+

*

+

horse) as shown m Table 19. Isoagglutinins specific to these antigens
were found. Especially agglutinin specific to type 1 erythrocytes werepositively detected in all samples excepting type 1 individual out of

K. FUJINO

100

thirty-three those tested. Table 20 shows summarized results of theisoagglutination.
Any informations to discuss relationships between these four types andMn antigens have not been obtained. Phenotypic relations noted inTable 19 would suggest existence of sub-blood types, but additional data
should be collected to conclude on allele system definitely. Frequenciesof occurrence of these four phenotypes of humpbacks taken from Okinawa
area are shown in Table 21 (Cushing et al., 1959).
TABLE 21.

FREQUENCY OF OCCURRENCE OF HUMPBACKJu BLOOD TYPES

Blood types
1

2

3

4

No. ofwhalesobserved

.010

.040

.071

.879

99

Locality
taken
Okinawa

Serological constitution of whales from view points of human ABO blood
types
Natural antibodies specific to human ABO blood types were detected
on whale serums by means of agglutination reactions. As shown in
Table 22, anti-human A (a') agglutinin was detected positively in all
species tested, but anti-human B (fi') agglutinin was found in finbackand blue whales only. Agglutinin titers of anti-A reached up to 1 : 2
or 1 : 4 and those of anti-B up to 1 : 8 (Yamaguchi & Fujino, 1952;
Fujino, 1953).
TABLE 22. FREQUENCY OF OCCURRENCE OF ANTI-HUMAN A(a')
AND ANTI-HUMAN B(f'l') NATURAL ANTIBODIES IN
SERUMS OF VARIOUS SPECIES OF ANIMALS
Natural antibody
Whale species
Toothed f blue-white dolphin
whale \ sperm whaler

bryde whale
Baleen fin whale
whale blue whale
humpback whalela'f'!'

a'

I'!'

o'

.OOO
.OOO
.OOO

.160
.154
.188
.065
.297
.400

.OOO
.OOO
.OOO

.840
.846
.812
.774
.514
.600

.096
.189

.OOO

.065

.OOO
.OOO

No. ofwhalestestedLocality
taken

25
13
16
31
37
15

east side of Izu
Penninsula
Antarctic
Bonin Islands
northern part of( North Pacificl)Friedenreich and With (1933) and Iseki and Murakami (1940) state thatanti-human B and anti-human A natural antibodies in serums of animal
does not have uniform construction in specificity and can be analysed
into partial antibodies specific to corresponding partial antigens of human
B and A blood groups. While Owen (1954) describes on heterogeneityof antibodies. However no analysis on these points of natural antibodyof whales has been undertaken yet.
According to many studies it has been clarified that human blood type

SUBPOPULATIONS OF THE NORTH PACIFIC WHALES

101

antigens of A, B and 0 have mosaic constructions of partial antigens
and that these partial antigens distribute in erythrocytes and body
fluids of animals. Terashima (1942) reported that human A antigen
can be analysed into four partial antigens A1 , An, Am and A1v. Friedenreich and With (1933) subdivide human B antigen into three partial
antigens Bi. B2 and B3 • Recently Furuhata (1957) proposed to substitute
Bv Bu and Bm for three symbols noted above to avoid confusion withthose of sub-blood types. Inoue (1943) reported that human 0 erythrocytes have a mosaic constitution which consists of three partial
antigens 0 1, Ou and Orn.
After analysing B antigen by this method, Fujino (1958) reported thatfinback erythrocytes have B;v partial antigen of human B which has
simpler structure serologically than Brn in guinea pig. Summarized
results including additional species are shown in Table 23. Materials
and methods in experiments are same as those noted by Fujino (1958).
No analysis on blood group substances A and 0 has been undertaken.
TABLE 23.

DISTRIBUTION OF PARTIAL ANTIGENS OF HUMAN B
BLOOD TYPE SUBSTANCES
Whale species
Partial antigen
No. of whale, testedfblue-white dolphin
Toothed whale 1 sperm whale\ baird-beaked whale
fin whale
Baleen whale f\ seiwhale

Bn1 and B1v'
ditto
ditto
Brv'
ditto

10
25
8
32
7

Schiff and Sasaki (1932) showed that secretion or non-secretion ofblood type substance in human saliva is a hereditary character. Thereafter it has been clarified by many workers that this serologicalcharacter can be identified with other body fluid than saliva. After
analysing saliva of rabbit by the use of antibody specific to human
A blood type, Koshino (1938 and 1939) stated that secreter-nonsecretertypes can be distinguished in rabbit and that this serological property isclosely related to producing ability of anti-human A antibody when used
as immune animal. The author tried to analyse secreter-nonsecretertypes in sperm whales using mucous membrane of second stomach as
material. Human B blood type substance was used as indicator for
analysis. After being washed, the mucous membrane of second stomach
was ground down and was extracted with equal weight of 1.5 percent
saline in refrigerator for 24 hours. Original sample for test were prepared by addition of 1/10 volume of five percent carbolic acid to extracts
after boiling for thirty minutes. After being inactivated, anti-human
B type immune rabbit serum (agglutinin titer of anti-B1 = 1: 2560,
Fujino, 1958) wai;; diluted 1 : 20 with i;;aline and wai;; added half volume

K. FUJINO

102

of human A type erythrocytes to absorb anti-human species-specificand anti-C agglutinin common to A and B cells.
Series of extracts prepared by successive dilution of original sample
with saline was mixed with equal volume of absorbed anti-B1 serum.
After two hours' absorption in room temperature, results of agglutination inhibition tests were observed by mixing of human B type erythrocytes as shown in Table 24. It can be seen from this table thatextracts of sample nos. 2, 3, 5, 6, 8 and 9 inhibit agglutination ofanti-B1 antibody significantly up to 1 : 4 dilution but others inhibit
TABLE 24.

No. ofextracts
1
2
3
4
5
6
7
8

ANTI-HUMAN B AGGLUTININ-INHIBITING TESTS BY SALINEEXTRACTS OF STOMACH OF SPERM WHALES
Dilution of extracts
1:1
1:2
1:4
1:8
1:16
1:32
1:64

+

*

*

-!It

+

*

-!It

-!It

-!It

+

Control

-!It

-!It

+

*+

±
-!It
±
±
-!It

+

9
10

-!It

*-!It

-!It
-!It

-!It
-!It

-!It

+
+
-!It

+

*

-!It
-!It

-!It
-!It

*
*
*
-!It
*
*-!It
-!It

-!It

-!It
-!It
-!It
-!It
-!It
-!It
-!It
-!It
-!It
-!It
-!It

slightly. These facts seem to show existence of secreter-nonsecreter typein sperm whales. This serological property will serve for analysing ofsubspecific population in future. Relationship between these charactersand known blood types of sperm whales has not been studied yet.
POPULATION ANALYSIS OF THE NORTH PACIFICFINBACK WHALESAs regards raciation of the North Pacific fin whales several studies havebeen published as follows. After discussing seasonal change of localization of whaling grounds, catch statistics and oceanographic informations,
Omura (1950) and Kasahara (1950) assumed that in the east (Pacific) andwest (Japan sea) sides of Japan distribute different local populations
which have different migratory courses each other. Basing upon geographical difference of frequency of occurrence of blood type, Omura (1955)
and Fujino (1956) stated that fin whales distribute in areas around Aleutian Islands do not belong to one uniform population, but rather different
local populations migrate to area off Kamchatka and area of north ofeast Aleutian. While Kawakami & Ichihara (1958) stated from results

SUBPOPULATIONS OF THE NORTH PACIFIC WHALES

103_/

of marking investigation that both populations in American and Asiansides migrate to same feeding area around Aleutian, but it is not clear
whether these populations mingle in this area or are kept isolated eachother. After comparing external dimensions of whale body, Ichihara(1957) found significant difference between fin whales from area around
Aleutian and those from East China Sea, but does not discuss whether
whales taken from these two areas belong to different populations ornot. In the present report racial study of whales from these various
areas will be discussed by means of population genetics with bloodgroups and marking investigations.
Distribution of finback whales in the North PacificFin whales in the North Pacific have very widely geographical distribution, winter in lower latitude areas for breeding or calving and insummer season migrate northwards for feeding. Whaling grounds areformed in the waters adjacent to California, British Columbia, Alaska,
Aleutian Chain, Kuril Islands and Japan during from June to October.
Number of finback whales taken in the years 1949 to 1959 are shownin Table 25. In the areas of Okhotsk and Pacific sides of Japan number of catch has remarkably decreased in recent years. Whaling in the
East China Sea has been resumed since 1955 after several years' suspension after World War II.
Population genetics with blood types
As already stated, several kinds of blood group antigens have beenfound in finback whales. Four antigens out of these are used for large
scale of investigations as markers. Fujino (1956) stated that higher
precision in blood typing can be expected in Ju system than in Bp system because of purification of typing reagents and phenotypic relationships. Therefore, Ju system alone has been used since 1956.
Cushing et al. (1959) stated that dosage effects were recognized between homozygous and heterozygous types in Ju system. Thereafter
additional dosage effect was observed among heterozygous types as already shown in Table 7. On the other hand original anti-Ju 2 immune
serums used in the years 1954 to 1959 have been always kept in highagglutinin titers and were used at dilutions 1 : 10 or 1 : 20 in actual
typing. Therefore weaker heterozygous types seem to have beenscarecely misclassified as Ju1 homozygous type. From above-statedreasons Bp antigens were used for only qualitative comparison between
populations and Ju antigens were used for qualitative and quantitative
analysis of populations from different localities.
Geographical difference of distributions of Bp and Ju blood type antigens, which were shown in Table 11, will suggest existence of local
populations of the North Pacific finback whales. After comparing by

K. FUJINO

104

year frequencies of occurrence in various areas, yearly fluctuation canbe scarecely seen in the area of north of east Aleutian (area C) but canbe recognized in the areas off Kamchatka (area A) and of south of east
Aleutian (area B) (see Fig. 2). It can be assumed from these facts thatwhales which migrate to area C in these successive years, belong to
the identical population, and that different populations migrate to areas
A and B in different proportions by year. Concerning to this point,

Fig. 1. Whaling grounds in the North Pacific Fin whale.
TABLE 25.

NUMBER OF FINBACK WHALES CAUGHT IN VARIOUS AREAS
OF THE NORTH PACIFIC, 1949-59
Year

1949 1950 1951 1952 1953 1954 1955 1956 1957 1958 1959 Sum
01>
0
0
0
Bonin
0
2 > 277
3>
225
972
80
22
0
184
171
0
0
12
J
{West
1
apan North-east 208 141 209 209 234 197 12970
50
47 1570
76
Kuril
53 106 156 241 179 266 219 241 173 328 223 2185
Kamchatka & Be- 64 92 90 447 615 1554 1439 1480 1486 1370 1582 10219
ring (Pelagic)
British Columbia 105 150 216 240 181 150 120 168 284 573 369 2556
258
California
1093
22 109 105Sum
441 489 680 1159 1221 2167 2132 2239 2225 2601 2406 17760
Area1) Whaling has ceased since 1953.
2) Whaling in the East China Sea has been operated since 1955.
3) Includes thirty-five Whales caught from Tsushima Strait, 1949-53.

Fujino (1953) stated that remarkable seasonal fluctuations of frequencyof occurrence of Bp antigens were seen in the area off Kamchatka (areaA) in 1952. This fact was confirmed with the results in Ju system
obtained in successive years as shown in Table 26.

105

SUBPOPULATIONS OF THE NORTH PACIFIC WHALES

From these things it can be thought that migratory ranges of different local populations cover this area off Kamchatka and that relative
size of these different stocks which actually migrate there fluctuatesfrom year to year.
Number of whales tested in the East China Sea is small, but no remarkable yearly or seasonal fluctuations seem to be recognized judging
from daily occurrence of blood types shown in Fig. 3.
south of east Aleut.

off Kamchatka

ND<1>li212£JEWU&WiZZIJlllWd
ND

r-

u
c

~
:J

~ g

g

lD

1954

ND

WW

ND

ND

•

Ju2
Ju I Ju2
Ju11959 1954

1959 1954
Fig. 2.

north of east Aleut.

1959___..Year
Yearly fluctuation of frequency of occurrence of Ju blood groups.
ND: No available data

TABLE 26. NON-RANDOM DISTRIBUTION OF BLOOD GROUPS BY YEAROR BY SEASON IN THE WHALES TAKEN FROM AREAS OFF
KAMCHATKA AND SOUTH OF EAST ALEUTIAN.

Jui
.646
.765
.858
.989
.821
.929

Ju1Ju2
.098
. 068
.027

1958/a\b
1959

No. ofwhalesobserv.
82
234
113
91
28
238

.036
.021

Ju2
.256
.167
.115
.011
.143
.050

1955

32

.907

.031

.062

1958la
\b
1959

140
37

.943
.946
.909

.021
.027
.091

.036
.027

Year andseasonArea

1954/a\b
1955
OffKamchatka \lSouth of east!
Aleutian

11

Blood groups.OOO

Remark.OOO

June
July-Aug .
May-June
May 23-June 9
July 24-Aug 1
May 27-June 27
June (13), July (2) &Aug. (17)June 10-July 2
July 14-July 23
July 20-29

Even in case which yearly fluctuations are seen as stated above, it
can be discussed by use of averaged figures of gene frequency whether
two populations from areas of both side out of geographically neighbouring three areas mingle in the middle area or not. For this purpose
it should be discussed whether continued ratios of corresponding genefrequencies from three areas are proportionate in three alleles or not.
Gene frequencies are given in Table 13. Results of calculation in fourK. FUJINO

106

Fig. 3.

Occurrence of Ju blood groups in daily catch in East China Sea.

TABLE 27. CORRELATION BETWEEN POPULATIONS TAKEN FROMNEIGHBOURING THREE AREAS REVEALED BY GENE FREQUENCIES, in middle area
Combination ofneighbouring threeareas

Ratio of mingl.ll Freq. of phenotype
Gene freq.
Allele:x2
system obs.
exp.2>
obs. exp.2>
1-bb663 684.6.28.72 Ju,p .062.087
38.5 13.14
33.65
A. q .600.630.35 Ju1JU2
62.9
90.23 Ju2.77\. r .338.283*average
.43
786
.57
644.5
.77
.23 Jui.662.686
63.7 17.24
.34
.032
.059
.66 Ju1JU2
77.8.94.06 Ju2.254\ r .306*average.32.68
663 693.3
.062
.090
-.17
1.17 Jui
37.7 24.18
.52
33
.600
.649
.48 Ju1JU2
55.0
.69
90
.338
.265
.31 Ju2.65
786.35*average
642.9
.69
.662
.679
.31 Jui
62.1
15.26
.22
.032
.058
. 78 Ju1Ju2
81.0.91.09 Ju2
.306
.262
.39
.61
*average663 683.0.36.64 Jui.062.086
38.5 11.45
33
.67
.600
.628
.33 Ju1JU2
64.5
90
.78
.22 Ju2
.338
.286
.40
786
*average
.60
644.5
.687
.78
.22 Juir p .662
64.5
18.11
.35
.65 Ju1JU2
B q .032 .060
77.0
.94
.06 Ju2
r .306 .253
.31
*average
.69
198.5
1.26
206
- .26 Jui
(p .071 .051
3.99
6
5.9
.47
.682
.53 Ju1Ju2
A I q .739
15.6
.35
8
.65 Ju2'r .190 .267
.31
220
.69
*averageIp .801
.35
206.4
.801
.65 Jui
BJ q .017
4.8
0.37
.48
.014
.52 Ju1JU2
8.8.33.67 Ju2
.173
.185
\ r
.61
*average.39,------'~f

East China Sea

fKamchatkaa)

I East Aleutian\(Sum of north& south)East China Sea

fKamchatkab)

I South of east\

Aleut.

East China Sea
Kamchatka
North of east
Aleut.

c)1 Kamchatka
J South of eastd)
Aleut.
North of east
Aleut.

l

B(~

Ratio of mingling in the middle area of populations from western and eastern areas.
Calculated from figures of populations from areas of both side and averaged ratio ofmingling.

1)
2)

SUBPOPULATIONS OF THE NORTH PACIFIC WHALES

107

combinations of neighbouring three areas among which gene frequencies
reveal successive gradients are shown in Table 26. These include fourcases of a) East China Sea: area A: sum of areas B and C, b) East
China Sea: area A: area B, c) East China Sea: area A: area C, andd) area A: area B: area C.
From Table 26 it was denied that populations from both sides' areasmingle in middle area in any three cases a), b), and c) out of four throughA or B allele systems. While in case of d) possibility of mingling seemto be assumed through B allele system only. This will be discussed later
more in detail together with results of marking investigations. As resultsof emigration of marked whales show that some degree of minglings occur
between west and east areas around Aleutian, it can be concluded thatwhales from East China Sea belong to a local populations which nearly
isolated from populations that migrate to areas around Aleutian Chains.
Results from areas off Navarin and Olyutorskiy Peninsulas are excluded
from calculations because of scantiness of data available.
Ratios of mingling of different populations calculated from blood typegene frequencies and rate of recapture of marked whales
A. Rate of emigration of populations calculated from marking return.
A part of marking investigations, which has been undertaken in thenorth Pacific since 1949, was already reported by Omura & Kawakami(1956) and Kawakami & Ichihara (1958). As shown in Table 28, numberof finback whales recorded as ' Hit' reach up to 14 in the areas off
TABLE 28.

NUMBER OF FINBACK WHALES MARKED IN VARIOUS AREAS
OF THE NORTH PACIFIC, 1949~1959*
Year

Area
1949 1950 1951 1952 1953 1954 1955 1956 1957 1958 1959 Sum
1
Bonin
1
Japan, Pacific
7
1
13
5
42)
221)
Kamchatka
4
1
2
61
28
l83l 15
10
8
10
77
16
East
{South344l
210Bl 32
165 l 377
Aleutian North
57
28
North of Pribilof
31
913
and Navarin
0
1
52
74
Sum
7
55
64
26 542
1
27 235* Includes whales recorded as 'Hit' only. 1), 2), 3), 4) and 5) include each one whale
which was not recorded as ' Hit' but was recovered in successive years. 6) includes same
type of three whales as above-noted.

Bonin Islands and Pacific side of Japan and to 528 in the area around
Aleutian Islands. Seventy whales out of these have been recaptured in
the areas around Aleutians, but no recovery has been reported fromareas of East China Sea, north-east side of Japan, around Kuril Islands,
off British Columbia and California in which whalings have been operated.

20

.so\

60N

120

120E

150

\
<[

'' \

M,,}
~

150

-

--- --

D-----@ --NORTH PACIFIC

180

1-•,,on f·

180

I

No. of whales moved to different areas

No. of whales marked

No. of whales recovered in the some area during marking year
one year after marking
two years after marking

'~-

120W

Fig. 4. Summarized results of movements of marked finback whales in the North Pacific (1953-1959).

/ J\

150

40

ISO

60N

100

......

~

d
._

!"1

@

SUBPOPULATIONS OF THE NORTH PACIFIC WHALES

109

Fig. 4 outlines interrelations among various whaling areas revealed byresults of movement of marked whales. It can be thought to suggest
existence of local populations that in spite of fairly large number ofwhales are taken from various areas noted above in every year no recovery of marker has been reported from these areas excepting thosearound Aleutians. Especially regarding to relations between populations
in East China Sea and those around Aleutians, above-stated facts could
be interpreted to confirm that the former is isolated from the latter as
already concluded from blood typing investigations.
Relations among various areas around Aleutians will be analysed infollowing part. As shown in Fig. 4, 5 and 1 out of 61 whales marked
in area A were recaptured in the same area and area C respectively.
44, 3, 2 and 4 out of 377 whales marked in area Chas been recaptured
in the same, A, north of Pribilof and off Navarin, and B areas respectively. Out of 77, 2 and 3 whales marked in area B have been recaptured in the same and C areas. Additional recaptures of one andthree whales from those marked in areas C and B respectively were reported. As their markers, however, were recovered from freezing shipor oil-cooker of factory ship after treatment of whale body it is
not clear whether these whales were recaptured from areas B or C.
Judging from non-random geographical distribution in blood type genefrequency noted already, it can be thought that population which migrates to area C belong to different those migrate to area A, but abovestated results of marked whales show that mingling occurs in part
between east and west populations. When two populations of west andeast sides are temporarily called as I and II, above-stated situations
may be interpreted as follows. These populations I and II are kept
isolated each other in wintering breeding season and their feeding
migratory ranges overlap in part in the areas around Aleutian Islandsin summer season.
As Japanese pelagic whaling for baleen whales in the North Pacifichas been operated by one company's whaling fleet, conditions at recovery
of markers can be thought to be approximately identical by year and
by area. Therefore problems of mingling between populations I and II
in the areas A and C could be discussed quantitatively basing upon
results of emigration of marked whales. Relative size of populations I
and II which actually migrate to areas A and C can be calculated as
average for periods of investigation from blood type gene frequencies
and rate of emigration which is obtained as relative figures of rate ofrecapture against total catch in each area. As number of whales recaptured in the year of marking is remarkably effected by the season ofmarking and whaling operation, they are excluded from calculation.

110

K.FUJINO

Symbols listed in 'A Standard Terminology and Notation for Fishery
Dynamics' (Holt et al., 1959) are applied to calculating results of marking investigation, and some other additional terms and symbols are
adopted.
If average rate of recapture of whales marked in area A against catchin areas A and C in successive years after marking is given as ri andr~, and those of whales marked in area C against catch in area A and C
is given as ri and r:, following representation will be possible.
1

r1

Rate of return of whales to area A: - 1
1
r1+r2

r12

rate of emigration of whales from areas A to C:

rI+r~

rate of return of whales to area C:
r21

and rate of emigration of whales from areas C to A:

ri+~

·

Whales against which marking investigations have been actually carried out in the various areas around Aleutian Islands should be thought
to consist of not pure populations but rather parts of both populations
I and II. If marking investigations in areas A and C, however, could
be thought to have been undertaken against unmingled populations I andII respectively for convenience to calculation, ratios of mingling of both
populations in areas A and C can be given by following formulas,ratio of mingling in area A:(4)ratio of mingling in area C:, where N 1 and N 2 are relative sizes of parts of populations I and II
respectively which actually migrate to areas A and C.
Figures of ri, r~, ri, and r~ can be calculated as follows. At first various
symbols necessary for formulation are given. Term of u means averagerate of recapture of whales, marked in area C in a year, against numberof catch in area C in successive years. Number of catch, of whales
recaptured, and of survivor in area C in successive years after marking
are represented as follows.
Lapse of years after marking
f no. of whales recaptured
In area C, \.
f . ..
no. of survivor o m1tlal stock in catch0

1

2 ... xRo

Ri

R2···R,,

Co' Ci' C/ · · · C,/

As it can be assumed that total number of whales recaptured up tox years after marking proportionates to total number of survivor of

111

SUBPOPULATIONS OF THE NORTH PACIFIC WHALES

initial stock, when marked, in catch, u will be given as rate of theformer against the latter which is averaged for the figures of successive
years as follows.
(5)

After averaging figures of u for different years with weight,
formula 4 will be obtained through formula 5.
2g - ( -Uo- · x+-U1
Ux-1
r 22 = · ( x-1+ ···+
· 1)x(x+l) T0 -R~
T 1-R;
Tx-1-R~_ 1
.

r~

in

(6)

, where T means number of whales 'hit' in area C, R' means total
number of whales recaptured in the year of marking, that is, includes
whales recaptured after emigrating to different areas, f means rate ofrecovery of marker from whales actually recaptured, and g means
rate of confirming whales reported as 'Hit' against number of whales
effectively marked. Figures of f and g are given as constant by year
and by area. When various figures of area A are adopted to T, C'
and R' in formulas 5 and 6, ri can be calculated by the same way asstated-above. To calculate r7, figures of area C are adopted to T and
those of area A are used for R' and C' in the same formulas.
Various figures used for these calculations such as number of whales
hit, of those recaptured and of catch are shown in Tables 29 and 30.
Calculation of number of survivor C' from number of catch C will be
discussed later. Number of marked whales in Table 29 include whales
reported as ' Hit ' only and does not include finback whales recaptured
from those reported as 'No verdict', 'Miss' or 'misconceived as different species' when marked. As remarked in the table, three and
one whales marked in areas B and C respectively are excluded from'number of whales recaptured' in this table, because their markers
were recovered from cooker or refrigerated ship and their localities anddates of recapture are indefinite.
Figure of f was estimated from the experimental results on recovery
of markers reported by Kawakami & Nasu (1956) as shown in Table 31.
They stated that significant difference of rate of recovery was seen between standard type dart and dart with streamer in their experiments.
In actual case, however, most streamer is fallen off from dart when recovered. Therefore f is calculated for all kinds of dart (1/f correction)from number of dart recovered on the treating deck of factory ship
among standard type dart shot into finback whales, that is, f=17/35=0.485.
In actual marking investigation, whales recaptured can be divided into
two cases in which their markers were recovered from whales re-

112

K. FUJINOTABLE 29. NUMBER OF FIN WHALES MARKED AND RECAPTURED
Recaptured in the areas:
Off Kamchatka
Marked in
the areas:

Off
Kamchatka

South of east
Aleutian

North of east
Aleutian

North of
Pribilof
and off
NavarinSouth of east Aleut.

North of east Aleut.

1953 54 55 56 57 58 59 1953 54 55 56 57 58 59 1953 54 55 56 57 58 59--- 1--1953 - - - 1 - - 54
- - - 2
55
56
57
1 1
58
59
1953
54
--1--55-1--2- - - 56
-

57
58
59
1953
54
55
56
57
58
59

-

1 1 -

-

-

1-

-----

-

-

1 -

7 7 2 1 1 3

1

2- 1-

1 1 1 3
6 3
1 1
4

2 1 1

7 9 5 2 11 14-

2 -J 1953
54

l

55
57

56
58

59

Sum

2 2 1 32a) Includes whales recorded as ' Hit' only. There are one additional recapture fromb) A marker (no. 4452), which no information on marking was recorded, is excluded fromc), d) Marker no. 6860 was recovered in the year 1959. Markers of nos. 6859 and 7269
TABLE 30. NUMBER OF FINBACK WHALES TAKEN BY JAPANESE FLEETIN VARIOUS AREAS AROUND ALEUTIAN ISLANDS AND BERING SEA
Area
Year

Off
Kamchatka

1952
53
54
1955
56
57
58
59
Sum

213
470
564
148
595
280
269
694
3233

East Aleutian
~

South

168
35
46
500
442
53
1244

North

584
1177
774
286
298
703
3822

North of
Pribilof

Off
Off
Navarin Olyutorskiy

158

174
275

6
47

158

449

53

Sum
213
470
1316
1360
1415
1404
1331
1450
8959

SUBPOPULATIONS OF THE NORTH PACIFIC WHALES

113

IN THE VARIOUS AREAS AROUND ALEUTIAN ISLANDS, 1953-1959

North of Pribilof and
off Navarin

Number of whales
markedSum ofrecapture

1953 54 55 56 57 58 59------------

2·

-----

2

------

2------------

-----

1
3

---

1

----

-------------

1- ----1- -

22
9
4
12
2
4

------------- - 1- ----

1

1- -

1

4

66

Remarkrecordedcorrected
as 'Hit', Ti•) Tix l/g
211>
26.2
4
5.0
32)
3.7
1
1.2
2
2.5
28
34.9
0
0.0
0
0.0
173 >
21.2
15
18.7 (Ono whal• (mack no. 16149) ,..
captured from areas B or C in
16
20.0
1958, isnot included in no. ofrecapture.
10
12.5{Two markers of no. 6860 &
8
10.0
6859, recovered from cooker, are10
12.5
not included in no. of recoveryc).
0
0.0
2066)
256.6
32
39.9
57
71.0 {One marker of no. 7269, re334 >41.0
covered from refrigerated ship, is28
34.9
not included in no. of recoveryd).
15 5 )
18.7
0
0.0
3
3.8
1
1.2
0
0.0
9
11.2
0
0.0
0
0.0
519b)
646.7

whales not recorded as ' Hit' in each of 1), 2), 3), 4), 5) and four those in 6) respectively.
number of marking and recapture.
were recovered in the same year of marking.
TABLE 31.

RATE OF RECOVERY OF MARKERS*

Fin whale

Humpback whale~~

Sperm whale~

no. of hit no. of recovery no. of hit no. of recovery no. of hit no. of recovery
11
With streamer 21
2
1
10
15
Without
182 )
2111
20
3
3
35
streamer
1) Includes four markers recovered from refrigerated ship.
2) Includes two markers recovered from refrigerated ship.
* Cited from Kawakami & Nasu (1956).

corded as 'Hit' and from those not so recorded. Then g was calculated
from Table 31 as a rate of number of whales recaptured from theformer against total number of recapture, that is, g

66-9
6+4+l
6

57
71=0.803(l/g correction). This denominator includes four whales of which localities

K. FUJINO

114

recaptured are indefinite and one whale recaptured from those on which
no information is recorded at the time of marking.
C' in formula 5 was calculated from number of catch C in each year
which were given in Table 30. While initial stock at the year of marking gradually decreases with lapse of years through natural mortality
and catch, it is recruited from younger generation in population. If
stock number, number of recruitment and number of survivor of initialstock at the year of marking are represented as follows, stock in eachyear consists of survivor and recruit, and rate of recruit rand survival
rate S have constant figures through full period of investigation, thenformulas 7, 8 and 9 can be introduced.
Lapse of year after marking
Stock number
Number of annual recruitment
Number of survivor of initial stock
when marked

0

1

2 ... xNo

Ni

N2 ··· N,,
R2 ···Rx
N/ ··· N,/

Ri
Ni'

R2- ... -R,,r -Ri--- Ni - N2 -

S

Nz-R2

Ni-Ri
No

N,,-R,,
Nx-i '

Ni

( 8)

(9)

N;=No·S".

From formulas 7 and 8, N,,=No( 1 ~r)".

( 7)- N,, '

Eliminating N 0 from this rela-tion and formula 9,
N~=N,,(1-r)".

(10)

If survival rates m sample of catch represent those of whole population,
C'

N'

C,,

N,,

--"'=-"' =(1-r)".

When stock is m a stabilized state, S+r=l.
formula 11,(11)

From this relation and(12)

Survival rates S used for formula 12 are estimated from frequency
distributions of number of corpus albicans in pregnant females taken
and from those of number of lamination of ear-plug in each area.
Data on ovaries used include major part of pregnant females taken
during from 1952 to 1959, and data on ear-plug used base on the
results collected in the three years 1957, 58 and 59 as shown in Table
32 and 34. At first it was discussed whether figures of survival rate

SUBPOPULATIONS OF THE NORTH PACIFIC WHALES

115

show significant differences or not during periods from 1952 to 1959 inwhich Japanese pelagic whalings have been operated. As regards
averaged number of ovulation in a breeding cycle, Laws & Purves(1956) offered figure of approximate 2.8 for Atlantic finbacks, Purves &Mountford (1959) obtained approximately same figure for Antarctic finbacks. While, Nishiwaki et al. (1958) reported figures of 1.6 and 1.8
for the North Pacific and Antarctic finbacks respectively, so that significant discrepancy between results reported by both authors for Antarctic
finbacks. In relation to this problem Laws (1960) states, 'There is stillsome uncertainty about the interpretation of the lamination of ear-plug
at present.' Therefore age composition are obtained from frequency
distribution of number of corpus albicans shown in Table 32 for twocases inwhich averaged number of ovulation is 1.6 or 2.8, and total survival rate S is calculated from these age compositions in two cases asrate of sum of five years' and older survivors against sum of four years'
and older survivors as shown in Table 33. In Figure 5 survival curves
for whole areas around Aleutian Islands against averaged numberof ovulation 1.6 are drawn separately- for three successive periods ofyears 1952-55, 1956-57, and 1958-59. Total survival rates for thesethree periods reveal gradual increase as shown by figures 0.8783, 0.9008
and 0.9095. Even if selective catching larger whales would effect upon
the above-stated trend because of steadiness in whaling operation inrecent years, it might be impossible to conclude that the north Pacificfinback stock have depleted remarkably at present. After discussing
separately by areas A, B and C, same trends can be clearly recognized.
Of course these facts can be seen also in another case that averaged
number of ovulation is 2.8 as shown in Table 35. In consequence, it
may be possible for convenience of calculation to regard that the northPacific finback stocks are nearly in a stabilized state. Then, C~ in formula
5 will be calculated by formula 12, and total survival rates S can begiven as averaged figures through whole periods for various areas.
Total survival rates calculated from frequency distribution of numberof lamination of ear-plug are shown in Table 35. It can be seen fromthis table that these figures are mostly consistent with those calculated
from ovarian data in a case of averaged number of ovulation 1.6. According to Baranov (1918) mortality rates of a population may be calculated from the slope of the right limb of a frequency curve of ages.
But frequencies of occurrence at peak and successive older several age
groups in right limb of this curve drawn by ear-plug data may be assumed to be not representative of population because of size limitationin regulation of whaling. Therefore, number of survivors in these agegroups were corrected by geometrical mean of survival rate calculated

116

K. FUJINO

from successive older three age groups. Total survival rate S was obtained as rate of survivors of age group tc+i and older against those ofminimum catchable age tc and older. No significant difference in figure
TABLE 32. FREQUENCY DISTRIBUTION OF NUMBER OF CORPUS ALBICANS
Off Kamchatka (A)No. of corpora
albicans
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44

1952~54 1955~56 1957~59

23
26
30 25.6
28 25.0
21 24.0
20 21.4
21 19.4
17 18.0
18 16.2
14 13.6
11 10.8
8 7.6
3 5.6
2 3.8
4 2.4
2 2.2
1 1.8
2 1.0
0.6
0.4
0.2
0.2
1 0.2
0.2
0.2

13
9
9 10.27 10.413 10.2
14 11.8
8 10.817 9.8
2 8.8
8 8.4
9 5.6
6 5.8
3 4.6
3 3.6
2 3.2
4 2.4
2 2.0
1 1.6
1 1.0
0.8
1 0.6
1 0.4
0.4
1 0.4
0.2
0.2
0.2
0.2

0.2
0.2
0.2
0.2
0.2
0.2
0.2
1 0.2
0.2
0.2

25
20
21 22.6
27 22.8
20 21.2
16 20.4
12 18.6
17 20.0
18 18.0
27 18.2
16 17.6
13 15.8
14 11.6
9 9.8
6 8.0
7 5.8
4 4.8
3 3.6
4 2.4
1.6
1 1.2
0.4
1 0.4
0.2
0.2

South of east Aleutian (B)
Sum
61
55
60 58.4
62 58.2
54 55.4
60 53.6
41 48.8
51 47.8
38 43.0
49 40.2
36 34.0
27 28.6
20 21.8
14 17 .2
12 13.6
13 10.4
7 8.6
6 6.2
5 4.0
2.8
2 1.8
1 1.0
1 1.0
1 0.8
0.6
1 0.4
0.2
0.2

0.2
0.2
1 0.2
0.2
0.2

1954~55 1956~57 1958~59

6
11
4
3
2
5
3
4
5
1
2
3
1
1
1

1

5.2
5.0
3.4
3.4
3.8
3.6
3.0
3.0
2.2
1.2
1.2
0.4
0.4
0.6
0.6
0.4
0.4
0.4
0.2
0.2
0.2
0.2

Sum

12
10
28
16
36
9
16 12.4 7 8.6 27 26.2
16 12.4 4 8.2 23 25.6
9 12.4 6 7.4 17 23.2
12 11.6 8 7.2 25 22.2
9 10.0 12 7.0 24 20.8
12 9.6 6 6.8 22 20.0
8 8.8 3 5.8 16 17.6
7 8.6 5 4.0 13 15.6
8 7.6 3 3.8 13 13.6
8 6.4 3 3.8 14 11.4
7 5.2 5 3.0 12 9.4
2 5.0 3 2.6 5 7.0
1 2.6 1 2.0 3 5.0
2 1.6 1 1.0 3 3.2
0.4 2 2.2
1 1.2
0.2 3 1.8
2 1.2
1.0
1.4
1 1.0
1 1.4
1 0.6
1 0.8
2 0.8
1 0.6
0.6
0.8
1 0.4
1 0.6
0.2
0.2
0.2
0.2

0.2
0.2
1 0.2
0.2
0.2

0.2
0.2
1 0.2
0.2
0.2

0.2
0.2
1 0.2
0.2
0.2

S was seen in two cases which three or four age were adopted as mm1mum catchable age tc. As total survival rates calculated from ear-plug's
data were consistent with one of two those calculated from ovarian
data, figures calculated from only ovarian data were used for actual calculation of formula 12.

117

SUBPOPULATIONS OF THE NORTH PACIFIC WHALESIn actual marking investigation in the ocean, whales smaller than
1 egal limit must be marked also. This may be assumed from figure 6
showing comparison between estimated size distribution of whales when
OF PREGNANT FINBACK FEMALES (OBSERVED AND SMOOTHED FIGURES)
North of east Aleutian (C)1954-55

1956-57

1958~59

56
63
37
51
30
36
23
20
9
9
6
6
2
2
1

26
28
34
21
11
14
9
6
5
3
5
6
3
3

34
20
21
15
18
10
13
13
9
8
1
7

47.4
43.4
35.4
32.0
23.6
19.4
13.4
10.0
6.4
5.0
3.0
2.2
1.2
0.8
1 0.4
0.6
0.4
2 0.4
0.6
0.6
1 0.2
0.2
0.2

3
1
2
1

22.8
21.6
17.8
12.2
9.0
7.4
5.6
5.0
4.4
4.0
3.4
3.0
1.8
1.4
1.2
1.2
0.6
0.8
0.6
0.2
0.2
0.2

1
2
1
2
1
1
1
1
2
1

1

1
1

0.2
0.2
0.2
0.4
0.4
0.2
0.2
0.2

1

21.6
16.8
15.4
13.8
12.0
10.6
8.8
7.6
5.0
3.4
2.2
2.0
0.8
1.2
1.2
1.0
1.0
1.0
0.8
1.0
0.8
0.8
0.8
0.6
0.2
0.2
0.2
0.2
0.2
0.2
0.2
0.2
0.2
0.2
0.2
0.2
0.2

Sum of three areas
Sum

116
111
92
87
59
50
45
39
23
20
12
19
5
6
3
3
2
3
3
3
2
2
2
1

1

1
1
1

91.8
81.8
68.6
58.0
45.2
37.4
27.8
22.6
15.8
12.4
8.6
7.2
3.8
3.4
2.8
2.8
2.0
2.2
2.0
1.8
1.2
1.2
1.0
0.6
0.2
0.2
0.2
0.2
0.2
0.2
0.2
0.4
0.4
0.4
0.6
0.6
0.4
0.2
0.2

1952-55

1956-57

1958-59

89
102
75
84
55
67
49
46
32
26
23
17
5
4
6
2
3
3

54
48
61
52
35
38
27
32
23
23
21
25
17
9
4
10
5
4
5
1
2
3

62
52
43
36
40
40
34
34
32
33
17
18
15
12
8
7
3
5
3
1
1
1
2
2

2
1
1
1

81.0
76.6
66.0
60.2
49.8
44.0
35.2
28.8
20.6
15.0
11.0
6.8
4.0
3.6
2.8
2.0
1.6
1.2
0.8
1.0
0.6
0.6
0.4
0.2

50.0
46.8
42.6
36.8
31.0
28.6
25.2
24.8
21.8
19.0
15.2
13.0
9.0
6.4
5.6
5.0
3.4
3.0
2.2
1.4
1.2
1 1.0
0.4
1 0.4
0.2
0.2

1
1
1
1
1

1

0.2
0.2
0.2
0.2
0.2

1

0.2
0.2
0.4
0.6
0.6
0.6
0.8
0.6
0.4
0.6
0.4
0.2
0.2
0.2

1
1
1

1

1
1

46.6
42.2
38.6
36.8
36.0
34.6
30.0
26.8
23.0
19.0
14.0
12.0
9.0
7.0
5.2
3.8
2.6
2.2
1.6
1.4
1.2
1.0
1.0
0.6
0.4
0.6
0.6
0.4
0.4
0.4
0.2
0.2
0.2
0.4
0.4
0.4
0.4
0.4
0.2

sum

205
202
179 177.6
172 165.6
130 147.2
145 133.8
110 116.8
112 107.2
87 90.4
82 80.4
61 65.4
60 53.037 40.2
25 31.8
18 22.0
19 17.0
11 13.6
12 10.8
8 7.6
4 6.4
3 4.6
5 3.8
3 3.0
4 2.6
1.8
1 1.2
1 0.6
0.8
1 0.6
1 0.6
0.6
1 0.8
0.8
2 0.8
1 0.8
1.2
1 1.0
2 0.8
1 1.0
0.8
1 0.6
0.4
1 0.4
0.2
0.2

marked and those of catches. Therefore correction (e-correction) on
number of catch and number of whales recaptured were discussed asfollows. If all age groups older than one year are marked and all age
groups older than four years in average are included in catchable stock,it can be assumed that all younger part of former which is smaller than

118

K. FUJINO

legal limit will recruit into catchable stock three years after marking.
Therefore it may be mostly enough to make correction on number ofcatch and number of whales recaptured m one and two years aftermarking. e1e2C~, e1e 2R1 and ep;, e2R 2 will be used in formula 5 instead
TABLE 33. NUMBER OF SURVIVOR AND DEATH OF FEMALENUMBER OF CORPUS ALBICANS
Off Kamchatka (A)Age

1952-54 1955-56 1957-59
Sum
....
....
....
....
0
0
0
0
t) .?:
t; .a
~.?:
t;.a
t;.a
~..d
i>
..., ~.?:i>..., ~.?:
i>
...,
i>
...,
• ....

0 ;:li:: rn

4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24

25
26
27
28
29
30
31
32
33
34
35
36
37

South of east Aleutian (B)

0

• ro

v

i:l"O

• '""'

0 ;:li:: rn

0

• ro
v

i:l"O

• l-1

0

::l

i:: rn0• Cl:!

vi:l"Cl• '""'

0 ;:li:: rn

0

• ro
v

i::"O

284 12 146 7 246 9 626 18
272 13 139 5 237 8 608 23
259 14 134 8 229 8 585 15
245 22 126 7 221 7 570 16
223 5 119 6 214 6 554 19
218 14 113 4 208 6 535 23
204 15 109 10 202 5 512 24
189 16
99 7 197 5 488 26
173 17
92 8 192 6 462 28
156 1984 7 186 7 434 30
137 2077 8 179 8 404 42
69 9 171 9 362 48
117 2493 21
60 8 162 15 314 49
52 8 147 28 265 57
72 18
54 14
44 5 119 20 208 39
99 15 169 27
40 9
39 6
84 16 142 22
31 10
33 5
68 12 120 19
21 4
28 6
56 11 101 17
17 4
22 3
45 9
13 2.5 19 4
84 17
10.5 2.0 15 3
36 9
67 14
8.5 2.0 12 2.5 27 653 20
6.5 2.0 9.5 1.5 21 6
33 11
4.5 0.5 8 1.5 15 4
22 7
4.0 0.5 6.5 1.5 11 4
15 4
3.5 0.5 5 0.5 7 33
11
3.0 0.5 4.5 1.0 4 2
8 2
2.5 0.3 3.5 0.5 2 1
6 2
2.2 0.2 3 0.2 1 0.5 4 1
2.0 0.4 2.8 0.3 0.5 0.3 3 1
1.6 0.2 2.5 0.5 0.2 0.2 2 1
1.4 0.4 2 0.2 0.0
1 o.5
1.0 1.0 1.8 1.8
0.5 0.5
0.0
0.0
0.0

1954-55 1956-57 1958-59
....
....
....
0
0
0

Sum
....
0

4-< i>
4-< i>
t; .::t) .a1) .?;
1' ..c::
i>
...,
i>
...,
i> ~-E.....i> ~-;J
.
....
• ro
• lo-I
• ro
• ro
• M
• ttl
0 ;:l ov
0 ;:l OV 0 ;:l ov 0 ;:l 0 v
o·~i:: rno·~i::"Cli:: rni:: "Cl

51.8 2.6
49.2 2.4
46.8 2.4
44.4 2.0
42.4 2.1
40.3 1.9
38.4 2.1
36.3 1.9
34.4 2.6
31.8 2. 7
29. l 3.4
25.7 5.5
20.2 6.6
13.6 4.2
9.4 1.9
7 .5 1.3
6.2 0.7
5.5 1.0
4.5 0.5
4.0 0.4
3.6 0.3
3.3 0.5
2.8 0.3
2.5 0.3
2.2 0.3
1.9 0.4
1.5 0.3
1.2 0.4
0.8 0.2
0.6 0.3
0.3 0.1
0.2 0.2
0.0

141
135
130
125
120
114
108
102
96
91
85
79

6
5
5
5
6
6
6
6
5
6
6
8
8
9

i:: rn

91.5
87.3
83.2
79.1
76.0
72.5
69.5
65.5
61.2
55.2
49.7
45.0
71
40.0
63
36.0
54 10 32.0
43 12 28.l
31 10 21.7
21 4 15.3
17 4
9.2
13 2
4.8
11 1
2.3
10 1.5 0.8
8.5 1
0.0
7.5 0.5
7 1
6 1
5 0.5
4.5 1
3.5 1
2.5 1
1.5 0.5
1 0.5
0.5 0.5
0.0

i:: "Cli:: rni:l"Cl4.2 283 12
4.1 271 12
4.1 259 12
3.1 247 12
3.5 235 11
3.0 224 11
4.0 213 10
4.3 203 13
6.0 190 14
5.5 176 15
4.7 161 17
5.0 144 17
4.0 127 18
4.0 109 17
3.9 92 18
6.4 74 17
6.4 57 16
6.1 41 11
4.4 30 8
2.5 22 4
1.5 18 3
0.8 15 2
13 2
11 1
10 1
9 1
8 1
7 2
5 1.5
3.5 1.3
2.2 0.2
2 1.0
1 1.0
0

of C~, Ri and c;, ~. where e1 an e2 can be obtained approximately as
follows.
1

ei

e2

total survival rate for age groups older than two years of the stock '1=~~~~~~~~~~~~~~~~~~~~~~~

total surviv;i,l rate for age groups older than three years of the stock '119

SUBPOPULATIONS OF THE NORTH PACIFIC WHALES

However, survival curve has not been obtained for these younger partof the stock from data observed. Therefore, adopting the figure ofdeath rate of 0.025, which was estimated arbitrarily for younger age
groups by Ottestad (1956), against two and three age groups, calculation
FINBACKS, ESTIMATED BY FREQUENCY DISTRIBUTION OF
OF PREGNANT FEMALES
North of east Aleutian (C)1954-55....
0t:.?:> t;..d...... ...."'

1956-57....
0->o·:; t:£i:: (/)

i:: (/)

0 :::l..... <1.l
0 :::l 0 . "'1958-59....
0->. ....> t:£
0. <1.lo·~

Sum of three areas
Sum
....

1952-55....

0t; .?: t; ..d

>

....

:::l
i:: (/)

i::'O"' ...... "'0t; .?; t: ..c:>....

......0 "'0 <1.l0 :::li:: rni:: 'O43 278 27 212 14 1010 60
45 251 28 198 16
950 9042 223 28 182 15
860 7046 195 33 167
790 908
48 162 33 159 18
700 100
46 129 26 141 13
600 9047 103 24 128 14
510 90420 7042
79 12 114 1243
9 102 13350 6067
290 60
36
58
7
89 14
51
230 50
25
5
75 14
46
180 4018561 16
140 30
41
4
45 10
16
11
37
110 20
5
35
9
4
8
32
26
90 20
7
28
19
70 20
5
3
7
4
23
3
16
3
50
8
9 2
20
4
13
42
4
1
16
3
12
38
7 1.5
8
1.5
5.5 0.5 13
2
10.5 0.5 30
2
5 0.5 113
10
0.8 284
4.5 0.5 8
2
1.5 9.2 0.2 24
4 0.3 6.5 1.5 92
0.0 2291
3.7 0.6 51
0.0 20
3.1 0.1 4
1
92
0.0 193 0.2 31
92
0.4 17
2.8 0.3 2
0.2 8.6 0.6 15
3
0.4 122
2.5 0.3 1.8 0.4 8
1
2.2 0.2 1.4 0.4 7.6 0.6 10
9
2.0 0.2 1
0.5 7
1.0
2
7
2
1.8 1.8 0.5 0.5 6
1.0
0.0
5
2
0.0
5
1.0
4
3
1
1.0
2
3
1
1.0
1
2
1
1.0
1
0
1.0
0

904
848
784
724
660
600
544
480
424
360
292
240
184
136
99
68
48
36
28
22
20
16
12

56
64
60
64
60
56
64
56
64
68
52
56
48
37
31
20
12
8
6
2
4
4
1
3
1
1
2
1
1
1

0

<1.l

i:: 'Oi:: 'O

0 :::li:: (/)

i:: 'O

0

536
493
448
406
360
312
266
219
177
134
98
73
55
39
28
20
13

<1.l

11

8
7
6
4
3
2
1
0

1956-57....
0t;.:=: t;..c:>...... ...."':::li:: (/)

0

564
528
488
452
414
372
335
306
282
260
238
218
198
177
156
132
110
88
70
57
47
40
33
27
23
18

14
11
8
6

5
4
3
2
1
0

0

1958-59
....
0

~ .?:

>

• 1-1

t: ..c:....
•. . . . . . C<l
0 :::l 0 <1.l

>

....

i:: rni:: 'O

1960
1840
1720
1600
1470
1360
1240
1140
1020
910
800
700
600
500
410
330
260
200
160
140
110
90
70
60
50
40
30
25
20
15
10
5
0

120
120
120
130
110
120
100
120
110
110
100
100
100
90
80
70
60
40
20
30
20
20
10
10
10
10
5
5
5
5
5
5

0 :::li:: rni:: 'O

36
40
36
38
42
37
29
24
22
22
20
20
21
21
24
22
22
18
13
10
7
7
6
4
5
4
3
3
2
1
1
1
1
1
1

506
475
448
414
388
372
361
349
330
304
270
233
205
172
142
120
100
80
63
50
38
30
23
18
16
14
13
12
10
9
8
6
5
4
2
1
0

31
27
34
26
16
11
12
19
26
34
37
28
33
30
22
20
20
17
13
12
8
7
5
2
2
1
1
2
1
1
2
1
1
2
1

0

will be given as follows.
Age
2
3
4
older than 5

Number ofsurvivor

az
as
a,
a;

Number of
death

az-aa
as-a•
\a,
f

Death rate
0.025
0.025
11-s
I

0

~.::: ~.d

cd
<1.l

<1.l

i::'O

Sum
....

K. FUJINO

120

TABLE 34. FREQUENCY DISTRIBUTION OF NUMBER OF LAMINATION OF EARPLUG OF THE NORTH PACIFIC FIN WHALES, SUM OF MALES AND FEMALES
TAKEN DURING FROM 1957 TO 1959
No. oflaminationof ear-plug
2- 3
4- 5
6- 7
8- 9
10- 11
12- 1314- 1516- 1718- 1920- 21
22- 2324- 2526- 2728- 2930- 31
32- 3334- 3536- 3738- 3940- 4142- 4344- 4546- 4748- 4950- 5152- 5354- 5556- 5758- 5960- 6162- 6364- 65
66- 67
68- 6970- 7172- 7374- 75
76- 7778- 79so- 81
82- 8384- 8586- 87
88- 8990- 91
92- 93
94- 95
96- 97
98- 99100-101
102-103104-105
106-107108-109110-111

Off Kamchatka (A)South of east Aleut. (B),.--"~

observ.

0
14
34
23
30
22
16
20
29
23
24
20
21
16

17
26
26
10
17
9
10
12
6
11
8
7
6
3
2
2
4
3
1
3
1
3
1
5
2
4
1
1
1
1
0
1
0
1
0
0
0
2

smooth.

24.6
25.0
22.2
23.4
22.0
22.4
23.2
23.6
21.0
19.8
20.2
21.4
19.0
19.2
12.6
14.4
11.6
10.8
9.6
9.4
8.8
7.6
7.0
5.2
4.0
3.4
2.8
2.4
2.6
2.4
2.2
1.8
2.6
2.6
3.2
2.8
2.8
2.0
1.6
0.8
0.8
0.6
0.6
0.4
0.4
0.2
0.6
0.4
0.4
0.4
0.4
0.0

,,....--'

observ.

0
5
13
15
23
25
15
18
12
16
22
15
12
13
6
16
12
11
8
8
8
5
4
3
2
5
2
3
5
2
1
1
0
0
2
0
0
0
2
2
0
1
0
0
1

--------smooth.
16.2
18.2
19.2
18.6
17.2
16.6
16.6
15.4
15.6
13.6
12.4
11.8
11.6
10.6
11.0
9.4
8.0
7.6
5.8
4.4
3.8
3.2
3.0
3.4
3.4
2.6
2.4
1.8
0.8
0.8
0.6
0.4
0.4
0.8
0.8
0.8
1.0
1.0
0.6
0.4
0.2
0.2
0.2
0.2
0.2

North of east Aleut. (C)~-----------

observ.

4
45
87
95
66
50
38
27
33
33
22
21
25
19
18
12
16
10
17
16
9
13
6
5
4
4
5
4
3
1
6
2
2
3
2
4
5
0
2
1
1
1
1
1
0
1
2
0
1
1

smooth.

68.6
67.2
55.2
42.8
36.2
30.6
27.2
26.8
24.0
21.0
19.0
18.0
15.0
14.6
14.2
13.6
13.0
12.2
9.8
7.4
6.4
4.8
4.4
4.0
3.4
3.8
3.0
2.6
2.6
2.8
2.4
3.0
2.8
2.6
2.4
1.8
1.2
1.2
1.0
0.8
0.8
1.0
0.8
0.8
0.8
0.8
0.4
0.4
0.2
0.2
0.0

121

SUBPOPULATIONS OF THE NORTH PACIFIC WHALES, then1-S1-S
a4=-- · a5.
8a5az = ----S- · (0.975) 2 'In consequence,
az+aa+a4+a5
aa+a4+a5

1.975-S
0.975

3.001-2.026·8
1.975-S(13)

Figures e1 and e2 will be given after adopting those of S in Table 35
to formula 13. As in this case figures of S in any areas A, B and C
give approximately same results to u in formula 5, averaged value by
area around Aleutians was used. In consequence figures of e1 e2=1.4,....--.

.... ---o

5

10

15

20

1952~55
1956~57
1958~59

25

30

35

age

,,40

Fig.5.
Age composition of the North Pacific Fin whales, estimated from distributionof frequency of corpus ablicans of pregnant females.
TABLE 35.

SURVIVAL RA TES (S) OF THE NORTH PACIFIC FINBACKS
East Aleutian
Off
Kamchatka (A)
.8261
.8490
.8729
.8573*
.9010
.9131
.9274
.9194*.9397

south (B)
.8386
.8562
.8609
.8481
.9079
.9175
.9108
.9133
.9140

north (C)
.7505
.7496
.8075
.7589*
.8564
.8534
.8891
.8714*.8730

Whole area

.7914
(Season 1.8248
Season 2.8440
Season 3.8192
Average.8783
JSeason 1.9008
II Season 2.9095
Season 3.8962
Average
Ill
Figures in columns I and II are calculated from freq. distribution of no. of corpus albicans of pregnant females as the average number of ovulation of 2.8 or 1.6 in one breeding season. Figures in column Ill are calculated from freq. distribution of no. of lamination of ear-plug.
Seasons 1, 2 and 3 correspond to three periods of 1952-54, 1955-56, and 1957-59 in area
A, those of 1954-55, 1956-57, and 1958-59 in areas B and C and those of 1952-55,
1956-57, and 1958-59 in whole area.
* These figures are used for calculation in formula 12 as survival rate (S).
I

l

122

K. FUJINO

e2=1.2 and eie2=1.2, e2=1.1 were obtained against two cases of averaged
number of ovulation 2.8 and 1.6 respectively. Thus various coefficients
necessary for calculation of formulas 5 and 6 were obtained.
It should be discussed here that whether marked whales show random
distribution in the stock, to which they belong, or not in various areas.
For this purpose whether rate of recapture of marked whales fluctuate
little or not from year to year were examined firstly on the data on 206
whales which were recorded as 'Hit' in area C in 1954. Table 36 shows

,,-.,

\

./I

f

--533 whales marked, estimated;e---.. 8324 whales caught, 1952~59

' I

*,
I

f

f

measured
I

~

~

~

~

~

~

~

~

m

~

body length in feetFig. 6. Comparison of size distributions of whales caught (measured)and marked (estimated).

number of whales recaptured, number of catch and rate of recapture
calculated from formula 5 for five years during from one year (1955) andfive years (1959) after marking. But no 1/f correction is made in those
calculations. It can be seen from this table that yearly fluctuations ofrate of recapture are very small as shown by figures of u = 0. 007300 ±0.000455 and 0.005950±0.000465 against two cases of averaged numberof ovulation 2.8 and 1.6. Number of whales recaptured from 1000 whalesof annual catch are given by formula 14. y means number of whales
recaptured in each year, and Y means total number of those, and I
II correspond to two cases of averaged number of ovulation 2.8 and 1.6.
y=7.30x0.7589"}I
Y=22.98(1-0.7589")

y=5.95x0.8714"'

Y =40.32(1-0.8714")

}
II

(14)*

* These formulas are given in general as follows.
y=lOOOu·S"
, and

u·S
Y =lOOO(u· S+u· sz+u· ss+. · · +u· 8")=1000--(1-Sx)
1-S

(15)

, where S means survival rate and u means rate of recapture of marked whales against annual catch. If marked whales distribute at random in the stock to which they belong,
u=T/N
(16)
, where T is number of whales marked effectively and N means stock number.

SUBPOPULATIONS OF THE NORTH PACIFIC WHALES

123

Calculated figures of y and Y are shown in Table 37 and Fig. 7. In
this table observed values are obtained from Table 36 directly andexpected those are obtained from formula 14. Figure 7 shows that
number of whales recaptured in each year fluctuate along expectedcurve, but that observed values in total number of those are highly
consistent with expected values in general. Curve Yin averaged numberof ovulation 2.8, however, seem to have too low values in five (and may
be also in successive) years after making, and this should be discussed
TABLE 36. ANNUAL RECAPTURE FROM 206 WHALES MARKED
IN THE AREA OF NORTH OF EAST ALEUTIAN (C) IN 1954
Year

Ja)

r%5

IlbJ

r%5

56
57
58
59

56
57
58
59

x
1
2
3
4
5
1
2
3
4
5

Rx

C,,

_________,,.~

~

obs.

cor.dJ

obs.

cor.dJ

7
2
1
1
3
7
2
1
1
3

9.8
2.4
1
1
3
8.4
2.2
1
1
3

1177
774
286
298
703
1177
774
286
298
703

1648
929
286
298
703
1412
851
286
298
703

Sxc)

Cx'

J:R,,

J:C,/

I:Rx/I:C,/

.7589
.5759
.4371
.3317
.2517
.8714
.7593
.6617
.5766
.5024

1251
535
125
99
177
1230
646
189
172
353

9.8
12.2
13.2
14.2
17.2
8.4
10.6
11.6
12.6
15.6

1251
1786
1911
2010
2187
1230
1876
2065
2237
2590

.007834.006831.006907.007065.007865.006829.005650.005617.005633.006023***

Means & stand. deviat., * .007300±.000455, ** .005950±.000465.
a), b) In columns I and II, figures for correction of Rx, C., and survival rate S usedcorrespond to those in two cases that average no. of ovulation in a breeding season is 2. 8
and 1.6 respectively, that is, survival rate S=0.7589, eie 2 =l.4 and ez=l.2 for the former
and S=0.8714, e1e2 =1.2 and ez=l.1 for the latter.

with additional data hereafter. Thus, following conclusion can be introduced. Whales in area C in the year 1954, distribute uniformly in thestock which migrate to the same area in successive years after marking.
Gradual decrease in number of whales recaptured in successive vears arecaused by dilution of original stock in the year of marking with approximately constant rate of annual recruit from younger generation of
the stock, and dilution of stock by mingling with different populations
could be hardly supposed.
Table 38 shows rates of recapture against one thousand whales ofannual catch on whales marked in the years besides 1954. It canbe seen from this table that rate of recapture for years 1954, 55
and 56 are well consistent with each other. But figure for year 1958
is two times of the former and figure for year 1957 is very high. As
ri is obtained finally as averaged value of u with weight, it is little
effected by figure for the year 1958, but is remarkably effected by
figure for the year 1957. Detailed records on marking and recoveryof whales marked in this year are shown in Table 39. According tothis table, six whales (dart nos. J6815, J6801, J6802, J6837, J6829,

K. FUJINO

124

J6843) out of those marked in neighbouring area each other for several
days in this year are taken in neighbouring area again in the year 1958,
and this fact cause to make rate of recapture remarkably high. Therefore, it is more appropriate to remove data on whales marked in 1957
from calculation of ri and r~ through formula 6. Results of actual
calculation will be described later.
TABLE 37.

Year

1955
56
57
58
59

NUMBER OF RECAPTURE FROM 1000 WHALES OF ANNUAL CATCH
No. of annual recept., ySum. of recapt., yYears aftermarking, xobserv.

1
2
3
4
5

5.9
2.6
3.5
3.4
4.3

expect.

observ.

,,-----~

I
5.5
4.2
3.2
2.4
1.8

II
5.2
4.5
3.9
3.4
3.0

expect.
,--------~~

II

5.9
8.5
12.0
15.4
19.7

5.5
9.7
12.9
15.3
17.1

5.2
9.7
13.6
17.0
20.0

On the other hand rate of recapture of whales marked in area A show
remarkable annual fluctuations. This seems to be caused by not only
scantiness of number of whales marked but also non-random distribution of marked whales in the stocks which migrate to this area fromyear to year ; in other words, migratory ranges of different populations
cover this area. Practically, however, various causes and mechanisms
TABLE 38. RATE OF RECAPTUER OF WHALES MARKED AND RECAPTUREDIN THE AREA OF NORTH OF EAST ALEUTIAN FROM 1000 WHALESOF ANNUAL CATCHa)
Average value with weightYear,u,:fTx-R,,'
WeightT,,-R,/xUx
not exclude exclude exclude
marked1957 & 58 1957 1957 & 58
1
243.08.02447
1954
5.947
5
2
35.88
.02795455
1.003
3.02832
2.018
71.25
3
.08699.0283656.026594 17.379
37.13.46806257
532.94.0495758
1.633
1a) Figure corrected for f and g and calculated for average number of ovulation 1.6.

of yearly fluctuations can not be analysed in detail as undertaken on
those in area C because of scantiness of data, so that ri and r~ were
calculated from figures of u in each year through formula 6.
Calculated results of rate of recapture of whales marked in areas A,
B and C are shown in Tables 40, 41 and 42. Table 40 shows comparisonby two cases of averaged number of ovulation 1.6 and 2.8 in a breeding
season which were adopted for calculation of survival rate from frequency
distribution of number of corpus albicans of pregnant whales. It canbe seen from this table that individual values of rate of recapture in case

SUBPOPULATIONS OF THE NORTH PACIFIC WHALES

125

of averaged number of ovulation 2.8 are somewhat higher than thosein another case, but there is little difference in their relative values.
Table 41 show comparison by two cases before and after 1/g and 1/f
corrections. It may be seen in this comparison also that despite of

40._3_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __
40

--"'

.8

"'

u

II

<o::J
c

c

"'

'-'0
"1:J

c

30

15\
0

::J

--"'
_..,

Q)

c
0

E

......e23 0-------- - -

V>
Q)

---

<o
_c
!:
"1:J

-=-~...:-=-===-

---~-

20

Q)

~

<O

E

......0

c 15

Q)

>
0

u

\l!

......0

w 10

..1)

E
::J
c

5

--2

3

4

5

------------------lO

15

lapse of years after marking

Fig 7. Annual recovery of whales marked in 1954 (206 whales marked) in
the area C, north of east Aleutian, showing two cases I and II against
survival rates of 0.7589 and 0.8714 respectively.

tolerable difference in individual values of rate of recapture, there 1s
little difference in relative values. According to above-stated facts it
can be concluded that errors in estimated values of averaged number ofovulation, g and f hardly effect on the results of rates of emigration.

126

K. FUJINOIn Tables 40 and 41 localities of recapture of two whales out of thosemarked in area B are indefinite, because their markers were recoveredfrom oil-cooker or oil-separator of factory ship after treatment of whale
TABLE 39. REMARKS ON WHALES MARKED IN THE AREA OF NORTH OF
EAST ALEUTIAN IN 1957 AND RECAPTURED IN SUCCESSIVE YEARS, marked
No. of No. of Dart
Locality whales
1 > whales
no.

Date
June 10, 1957 (

"
"
"

11,
25,
29,

July

6,

Sept.

7,

"
"
"

8,

, recaptured

,,

"
"
"
"
"

9,

"

10,

"

55N
168-9W
( 54N
168W
( 54N
170W
( 55N
168W
( 56N
170W
( 54N
167W
( 54N
168W
( 54N
169W
( 54N
168W

4

1

7269

June~July,

1

1

7263

July

1

1

J 65062

1

J 67884 > June 30, 1959

"

( 54N
167W

30, 1959(

Area2>

Nis>

B or C6>

53N
168-169W

C7l

2Aug. 11, 1957

off
( 63N
NavarinB>
178W
(
53N
177-178W c

3
2
6

4

f'

J 6815July

4, 1958* J 6801

"
"
"
"
"

11,

,r

J 6802

J 679911,

1957

Locality

2

l*

"

Date

10

( )6837"
J 6831)* J 6829

11,
16,
3,
8,

"
"

"
"
"
"

1*J6843"8,J 6807"

9, 1959

J 6834May

27, 1958

l

( 54N
170W
( 54N
169W
( 54N
169W
f 50N
\170W
( 53N
169W
( 54N
166W
( 54N
166W
( 53N
169W
( 52N
171E

c
c
c
B

c
c
c
c
A

1) number of whales recorded as 'Hit' only when marked, 2) areas temporarily classified
as A (off Kamchatka), B (south of east Aleutian) and C (north of east Aleutian), 3) NI, nodetailed information obtained, 4) recovered from whales recorded as ' Possible hit' whenmarked, 5) two markers in one whale, 6) recovered from cooker, 7) 8) recovered from refrigerated ship.

body. One was marked in area B (54°42'N, 159°25'W) in July 18,
1956 (marker number J6149) and was recaptured in area B or C duringfrom June 29 to July 11, 1958. Another was marked in area B (50°37' N, 168°33' W) in June 17, 1958 (marker number 6860) but locality

SUBPOPULATIONS OF THE NORTH PACIFIC WHALES

127

of recapture is indefinite because its marker was recovered from oilseparator Aug. 17, 1959. Therefore, as shown in Tables 40 and 41 rateof recapture are calculated against four cases of combination which eachof these two whales is recaptured in area B or C. As rate of recapture
show remarkable difference by case, rate of emigration from areas B to
C is not able to be discussed quantitatively so far. Rate of emigrationfrom areas B to C, however, show tolerably high value in all four cases,
so that it is obvious that whales migrate to area B are closely related
to those migrate to area C and it is difficult at present to suppose
existence of the third different populations migrate to the middle area
besides east (II) and west (I) populations.
TABLE 40. COMPARISON OF RATES OF RECAPTURE OF MARKED WHALESPER 1000 OF ANNUAL CATCH BY CASE OF AVERAGE
NUMBER OF OVULATION 1.6 OR 2.8ll

Karnchatka~arkedm areaof1south of ~ >east
3
Aleut.
4n. of e. Aleut.

1

ri+r~

_•_i_. --5_r~+r~

for no. of ovul. 2.8:
recaptured f rorn area ofKarnchatka

s. of e.
Aleut.

n. of e.
Aleut.

Karnchatka

s. of e.
Aleut.

n. of e.
Aleut.

.13541.00000.18125.00753.17372.00000.02208.00307.05371.06742.06402.07773.08699.16106.00000.19465.00807.18658.00000.02797.00426.07374.08948.08648.10222.10392

2

~.-1_
ri+r~

for no. of ovul. 1.6:
recaptured from area of

ri+r~.00000.02404.00000.02663.978: .022

.974: .026

.217: .783

.204: .796

1) f, g corrected.
2) Figures in columns l, 2, 3 and 4 show those for four cases in which two whales recaptured from B, B and C, C and B and C areas respectively.

In Table 29 whaling grounds in the northern part of north Pacific aretemporarily divided into four areas. Among these, however, area off
Navarin and Pribilof is closely related to area of north of east Aleutian(c) as shown by results of emigration of whales in the year of marking,so that rates of recapture are calculated for two cases in which NavarinPribilof area is put together to area C or not. Little difference in rateof emigration was seen between two cases. Then figurse of ri, ri, r~
.
.
r} .
ri
ri . r~and r~ and rates of em1grat10n - 1 - 1 • - - and • were
2
2
2
2
1
1
~+~

~+~

~+~

~+~

calculated for two cases including data on whales marked in area C in
1957 or not. Though significant differences are seen between these twocases, but the latter figures were used for the successive calculations

128

K. FUJINO

because of the reasons noted already (p. 123, 1. 21-24). Figures of rateof emigration from areas A to C, 0.978: 0.022 in table 42 show thatwest sides' population hardly migrate to area C. This fact confirm theexistence of west and east sides' populations which was clarified byblood typing investigations already. In following paragraph gene frequencies and relative sizes of populations I and II will be calculated fromrates of emigration and gene frequencies in various areas through formula
4, and averaged ratios of mingling of populations I and II in areas A,
B and C will be given.
TABLE 41.

COMPARISON OF RATE OF RECAPTURE OF MARKED WHALESPER 1000 OF ANNUAL CATCH BEFORE AND AFTER
f AND g CORRECTIONs11

lKamobotb

1'.Iarked south of !~m area east
3ofAleut.
4not corrected for f and grecaptured from area ofcorrected for f and grecaptured from area ofKarnchatka

s. of e.
Aleut.

n. of e.
Aleut.

Karnchatka

s. of e.
Aleut .

n. of e.
Aleut.

.07617.00000.20646.00941.19704.00000.03151.00384.06468.08023.07757.09312.10473. 13541.00000.18125.00753.17372.00000.02208.00307.05371.06742.06402.07773.0869921.00000n. of e. Aleut.
r11 •
r12
r1+r1
.
r1+r1
1 2
1 2
r22 _
_ r2
_
1_. _ _

.02744

ri+r; • r~+r~.00000.02404.952: .048

.978: .022

.208: .792

.217: .783

1) Figures calculated for average number of ovulation 1.6 per breeding season.
2) Figures in columns 1, 2, 3 and 4 correspond to those in previous table.

B. Estimation of ratios of mingling of populations I and II by means ofrates of recapture of marked whales and blood type gene frequencies.
As stated already, various figures in Table 42 were calculated,for convenience, under hypothesis that marking in areas A and C
were carried out against unmingled populations I and II respectively. But actually whales in areas A and C should be regarded as
mingled those of populations I and II. Therefore figures of right and
1

1

r1+r2

r1+r2

left terms in ratio of ~: ~ show upper and lower limits inprobable values respectively, and figures of left and right terms in ratioof - 2-

r21

r2

-

2

r1 +r2

r11
- 1-

:

~
r1 +rzr12-1

r1 +r2: - 1-

r1 +rz1

show upper and lower limits respectively.

=.978:.022

Ratio ofin Table 42 may be substituted by 1.000: .OOO

without giving significant difference to final results of rates of emigration.

SUBPOPULATIONS OF THE NORTH PACIFIC WHALES129

TABLE 42. COMPARISON OF RATES OF RECAPTURE BETWEEN TWO CASES
IN WHICH RECAPTURES FROM WHALES, MARKED IN THE AREA OFNORTH OF EAST ALEUTIAN IN 1957, ARE EXCLUDED OR NOT!>
Recapture from whales marked in 1957 areincluded
recapt. from area of

not included
recapt. from area of~Marked in{Kamchatka
area of n. of e. Aleut.

r;
.
r~
r1+r1
. r1+r1
1
2
1
2

ri

.

r~

r2+r2
. r2+r2
12
12

Kamchatka

north of e.
Aleut.

Kamchatka

north of e.
Aleut.

.13541.02404.00307.08699.13541.01673.00307.02836.978: .022

.978: .022

.217: . 783.371 : .629

1) Figures corrected for f and g, and calculated for average number of ovulation 1.6
per breeding season.

On the other hand intact figure of ---1_ · _1_ can be used, because
r~ +r;· ri +r;

population I hardly migrates to area C. Schematic relationships ofmingling of populations I and II in areas A and C may be shown as
Fig. 8. Gene frequencies in both three allele A and B systems wereadopted to calculations. Relationships stated above will be formulated
as follows.
In area A, and m area Cr~

Po=-- - N1p1
11

r 1 +r 2

r~(16)+-- N2P2
2
2
r 1 +r 2

r; N1q1 +- r;

qo=1

r 1 +r 12r~

r o = -- - N1r1
1
1

r 1+r 2

- - N2q2
2

r 12 +r 2

r;+-- N2r2
2
2
r 1+r 2

, where N1 and N2 are averaged relative sizes of populations I and II which
actually migrate to areas A and C, P1 q1 ru p 2 q2 r 2 , PA qA rA and Pa q0 r 0
are blood type gene frequencies of populations I and II and those ofareas A and C respectively. Results of various figures on populations I

K. FUJINO

130

and II calculated from this formula and figures in Tables 13 and 42 areshown in Table 43. It is obvious from Table 43 that there is no difference in gene frequencies of population II between two cases including
marking data in 1957 or not, but that there is some difference in thoseof population I. Moreover relative sizes of populations I and II shownorth of east Aleut.

r'1
r~ + r~II

Fig. 8. Diagramatic correlation between west (I) and east (II) populations
estimated from the movements of marked whales. Ni and N 2 mean
relative size of each population actually migrate to areas A and C.

TABLE 43.

GENE FREQUENCIES OF Ju BLOOD GROUPS AND RELATIVESTOCK NUMBER OF POPULATIONS I AND II

A system{!
B system{!
Relative stock number

Include recapture fromwhales marked in 1957

Exclude recapture fromwhales marked in 1957

popul. I

popul. II

popul. I

popul. II

.075
.500
.425
.575
.043
.382
.74

.026
.869
.105
.896
.002
.102
1.26

.112
.222
.666
.334
.074
.592
.43

.026
.869
.105
.896
.002
.102
1.57

differences between both cases, and N 2 is always larger than Ni as shownby Nz/Ni=l.7 or 3.7. Ratios of mingling of populations I and II inareas A and C are calculated from figures of Ni and N 2 and those in
Table 43 through formula 4 and mingling rate in area Bis calculated fromgene frequencies of populations I and II (Table 42). These results areshown in Tables 44 and 45. It can be seen from the latter table thatthe results show remarkable difference by allele systems A or B.

131

SUBPOPULATIONS OF THE NORTH PACIFIC WHALES

Ratios of mingling band 1-b in three alleles does not proportionate eachother in A system, but approximately proportionate in B system. These
facts were confirmed with chi-square tests of observed values against
expected those in frequency of occurrence of phenotypes. Possibilityof mingling shown in Table 44 is denied in A system but is affirmed in
TABLE 44. RATIO OF MINGLING OF POPULATIONS I AND II IN THE AREAS
OFF KAMCHATKA, SOUTH AND NORTH OF EAST ALEUTIAN
S. of e. Aleut.
North of east
Kamchatka~
Aleut.
A system B system
Include recapture from { popul. I
whales marked in 1957 popul. II
Exclude recapture from { popul. I
whales marked in 1957 popul. II

.73
.27
.42
.58

.51
.49
.29
.71

.29
.71
.10

.90

.02
.98
.01
.99

TABLE 45. COMPARISON BETWEEN OBSERVED VALUE AND EXPECTEDVALUE OF BLOOD GROUP GENE FREQUENCIES IN THE AREA
OF SOUTH OF EAST ALEUTIAN
Ratio of mingI.1>
Gene frequency~~
Allele system
exp.zi
obs.
1-b
b
.071
.051
.92
.08
.65
.739
.681
.35
.73
.190
.268
.27
lndudo '<oaptu" Afrom whales.49
average.51
marked in 1957
{p
.73
.810
.803
.27
.64
B q
.017
.014
.36
.75
.173
.183
.25
r
average
.71
.29
.48
.071
.051
.52
.681
.20
.80
.739
Exclude recapturef A
.85
.268
.15
.190
from whales
average.29.71.89
marked in 1957 B
.810
.818
.11
.93
.017
.024
.07
.173
.158
.87
.13
average.90.101) ratio of mingling of populations I and II.
2) calculated from gene frequencies of populations I and II and averaged ratio of mingling.

I ff
{i

l ff

B system. This fact confirm that B allele system can better interpretthe results observed than A system as assumed already by quantitative
comparisons of blood type gene frequencies only. But additional basic
discussions on phenotype-genotype relationships should be undertaken
hereafter before reaching definite conclusions on allele system. In present paper, for convenience, three allele B system and rates of emigration
excluding marking data in 1957 are adopted finally. Then ratio of mingling in area Bis obtained as m1 ,~ 1 = .10: .90. According to the results
discussed above, it can be concluded that population I migrate to areas
A and B but does hardly to area C, and population II migrate not only
to area C but also to areas B and A, that is, migratory ranges of

132

K. FUJINO

Fig. 9. Relative positions of the local populations of the North PacificFinback whales in the triangular coordinates, expressed through the threeallele B system of Ju blood group.
E : East China Sea
S : South of east Aleut. (observed)
N: North of east Aleut.
K : Kamchatka
I : Population I
II : Population II
TABLE 46.
a)

SUMMARY OF VARIOUS FIGURES DISCUSSED IN THIS SECTION

Blood group gene frequenciesii

Gene
freq.

{i

Population I

Kamchatka

.334
.074
.592
.503
.053
.444

.662
.032
.306
.844
.042
.114

South of east Aleut.
obs.

exp.2iNorth ofeast
Aleut.

.810
.017
.173
.937
.027
.036

.818
.024
.158
.928
.039
.033

.890
.003
.107
.982
.006
.012

~

Population II

.896
.002
.102
.985
.004
.011

Freq. of { [Jui]pheno- (Ju1Ju2]
types
[Ju2]
1l in three allele B system only, 2) calculated from ratio of mingling.
b) Averaged ratio of mingling between populations I and II:
Area

Population { ~c)Kamchatka

South of east
Aleut.

North of east
Aleut .

.42

.10
.90

.01

.58

.99

Relative stock number of populations I and II actually migrate to the areas off Kamchatka and north of east Aleutian:N2=1.57SUBPOPULATIONS OF THE NORTH PACIFIC WHALES

133

populations I and II overlap in areas A and B. Relative size of both populations which actually migrate to area A fluctuate from year to year,and gene frequencies observed in area A fluctuate correspondingly. As
blood typing data, however, have not been obtained during full period
for marking investigation, there are assumed to be some bias in observed
values of gene frequency of population I, because they were obtained
as averaged figure by year. On the other hand, gene frequency of
population II is not affected by this and seems to include little bias.
Even if there is some bias in gene frequencies of population I, thesefigures will be available for discussing yearly fluctuation of ratio ofmingling between both populations in various areas. For this purpose
additional number of whales should be observed hereafter. Summarized
results of various figures in this paragraph are shown in Table 46.
DISCUSSION AND SUMMARY

In the present paper after analysing sub-populations of the north Pacificfinback whales, the author clarified that two different populations, which
their migratory ranges overlap each other in feeding season, distributein areas around Aleutian Chains, and that whales taken from East China
Sea belong to a different local population which is isolated from above
those. Various assumptions were rendered during these analysis, sothat it should be discussed here that how do they effect upon final
conclusions in the present report.

Problems in immunogenetics. Despite of somewhat lower precisionin blood typing Bp antigen, the facts that Bp1 antigen occurs in area
off Kamchatka (A) and while does not in area of north of east Aleutian(C) are sufficient evidence to show existence of local populations in these
two areas. On the other hand, it can be supposed that misclassificationof Ju blood types hardly has happened, because of obverse and reverse
correlations between Ju1 and Ju 2 antigens, dosage effect between homozygous and heterozygous types and purification of standard reagents.
Moreover, as specific agglutinin has sufficient high titer even after dilution
with saline for actual use, it can be hardy assumed that weaker heterozyous type was misclassified as homozygous type J u 1 • As regards allele
system of Ju blood type, after discussing discrepancies from HardyWeinberg's Equilibrium in frequency of occurrence of phenotypes andparentages between cows and fetuses, two allele system is denied andthen at least three alleles should be supposed. Then problems of identification and mingling of populations were investigated through two kindsof hypothetical three allele systems A and B. It was discussed whether
continued ratios of gene frequencies between neighbouring three areas

134

K. FUJINO

reveal approximately same figures in three alleles each other or not. Consequently it was clarified from discussions through both allele systems
A and B that whales taken from East China Sea belong to a local
population isolated from those around Aleutians. As regards correlations
between three areas off Kamchatka (A), south of east Aleutian (B), andnorth of east Aleutian (C), after above-stated discussion it was denied
through A system that populations from both side's areas mingle inmiddle area but it was affirmed through B system. On the other
hand, as emigration of marked whales between both side's areas, havebeen reported it is not probable that mingling in middle area is completely denied, so that A allele system seems to give results contradictory
to the facts observed. In other words, though direct evidences havenot been obtained yet, B allele system could better interpret the factsof observation than A system at present. As revealed by the rate ofemigration of marked whales, populations in the areas A and B aremingled those of populations I and II. But results of discussions on:figures in Table 27 and successive those, which were calculated throughformulas 2 and 3, can lead to final conclusions in the present paper.
According to the phenomena of dosage effect in Ju heterozygous types,
additional alleles related to possible sub-types can be supposed, butavailable data are too scanty to discuss this problem definitely. Existenceof this new fourth allele, however, might not effect upon analysing
results of racial study basing on three allele systems.

Ejfects of estimation of various figures in calculation of ratio ofmingling and some biological factors to the results discussed in thepresent paper. To calculate rate of emigration, it was presumed for convenience that marking in areas A and C were performed against unmingled
populations I and II. As population II, however, actually migrate toarea A, rate of emigration of population II to this area is obtained asa upper limit averaged for period of investigations. According to same
thinking it can be thought that major part of population I distribute toarea A and a part to area B, but it scarecely migrate to area C. This
fact is confirmed by the blood typing evidences that Bp1 antigen occurs
in area A but does not in area C at all.
Effects of errors in estimates of f, g, e1 , e2 and S in formulas 5 and
6 to ratios of mingling and other final results in present report will be
discussed here. At first g means rate of confirming number of whales
hit, against those effectively marked and was given as rate of numberof whales recaptured from those recorded as 'hit' when marked against
total number of recapture. Coefficient f means rate of recovery ofmarker from wba.les recaptured, and was estimated experimentally.

SUBPOPULATIONS OF THE NORTH PACIFIC WHALES

135

Actually one whale is shot by two or more darts in low frequency, so thatit could be approved that rate of recovery of marker f is equal to rateof confirming recaptured whales. As stated in discussions on Table 41,
figures f and g thus obtained fairly effect upon values of r}, r~, r~ andri, but hardly do to rates of emigration calculated as relative value ofthese. Figures of e1' e2 and S were calculated for two cases assuming
average number of ovualation per breeding season 2.8 or 1.6. Errorsof estimate of e1 and e2 effect upon only first and second terms ofnumerator and denominator in formulas 5 and 6. As regards estimateof survival rates S matters are as follows. For example populations I
and II migrate to area A and some geographical segregation by age ina population may be supposed. As being obvious from formula 12,
however, errors of survival rate S estimated from ovarian data of catchin each area effects a little upon only first term and hardly upon second
term and after as negligible minute more than two orders of each
denominator of formula 5 and will bring scarce difference on final
figures of rates of emigration. Estimates of e1' e2 and S show different
figures by two cases in which averaged number of ovulation is 2.8 or
1.6, and give some differences to individual figures of r:, r~, ri and r~, but
hardly bring difference to rates of emigration of their relative values.
Significant difference is seen in figure of r~ calculated from formula
6 for area C between two cases including marking data in 1957 or not(see Table 42). According to detailed records in marking logs, in spiteof scantiness of number of whales marked, those marked in closely
neighbouring areas at approximately same time are recaptured in closely
neighbouring areas again in a successive year. This makes figure ofu for this year higher, so that some bias is brought to figure of r~.
Therefore it is more appropriate that the data in this year is excluded.
Thus after being corrected with f and g coefficients and adopting figuresof ei. e2 and S in the case which average number of ovulation is 1.6,
rates of emigration was finally calculated from data of marking in~vestigation excluding these in 1957. On the other hand, regarding area
A it is assumed from yearly fluctuations of frequency of occurrence ofblood type that relative size of populations I and II which migrate tothis area vary from year to year. But analysis stated above for area
C could not be performed for area A because of scantiness of numberof whales marked and recaptured in each year, so that intact figuresof rl and r~ obtained from formulas 5 and 6 are used for calculation ofrates of emigration.
Next problems to be discussed are how do ecological factors during
migration of whales effect upon the results in previous discussions.
Regarding ecology of whales in general, some degrees of seasonal or

136

K. FUJINO

geographical segregations by sex, pregnancy and some other physiologicalfactors have been observed. In the north Pacific finback whales, however,slight segregations by pregnancy only have been observed. Sufficient
data to estimate quantitatively biases in rates of recapture which seemto be caused by individual factors have not been obtained yet. But thefact, that rates of recapture ri for area C against number of survivor
in catch keep approximately constant value from year to year, would
suggest that the above-stated various factors against biological segregations hardly effect upon rates of recapture finally. On the other hand,such analysis has not been performed for data on area A, so that it
must be kept in mind that some degree of biases in rates of recapture
are involved.

2

2

r1 +r2

r1 +r2This means that ratio of ~ : ~ includes somedegree of error. Errors in this ratio and in rates of emigration whichis caused by bias between periods for blood typing and those for marking
investigations, might bring somewhat effects upon gene frequency of
population I and figures of N 1 and N 2 • But these figures must havesufficient precision to discuss problems involved in present paper. Higher
precision could be expected through discussions with additional data
hereafter.
Other biological evidences suggesting existence of local populations.
After comparing morphological characteristics, following differences,which may confirm the existence of local populations, have been observed
between whales from East China Sea and those from areas around
Aleutian Islands. That is, whales from East China Sea 1) have lower
growth rate in body length, attain sexual maturity at averaged bodylength of 57 or 58 feet in male and of 60 or 61 feet in female approximately three feet smaller (Fujino, unpublished data), 2) are different
significantly in external body proportions (Ichihara, 1958) and in shapeof skull (Fujino and Ichihara, unpublished data), and 3) have shorter
baleen plates in same ages and in same body lengths than those fromareas around Aleutians. Full discussions on these points will be publishedlater (Fujino, to be published in next issue). Such significant difference
have not been recognized between whales from area off Kamchatka andthose from area of east Aleutions. Therefore it can be assumed thatdifferentiation between populations I and II is situated in earlier stage
than that between those and populations in East China Sea. In the areasof Okhotsk sea and off Pacific coast of Japan sufficient investigations havenot been carried out in recent years because of poor catch, but shapeof baleen plates from these whales reveal same characteristics as thosefrom Aleutian waters (Fujino, unpublished data).
Results of discussions in the present paper can be summarized as

SUBPOPULATIONS OF THE NORTH PACIFIC WHALES

137

follows.
1) After studying technical problems in large scale of blood typing
investigations on whales, standardization in basic methods has been
established as follows. Glycerol freezing technique has potential value inpreserving intact erythrocytes used for analysing blood type antigens,and standard reagents can be preserved by adding preservatives at concentration of 0.5% carbolic acid or 0.1% sodium azide and by being
kept frozen for several years in good condition available for testing.
Preservatives will serve against accidental warming during transportation
between central and field laboratories.
2) Blood type antigens and other serological constitutions were analysed in four species of baleen whales and three species of toothed whales
and allele systems in some species among these were discussed. Especially in Ju blood types of finback whales, two allele system was denied andtwo kinds of three allele systems were discussed.
3) Racial study on the north Pacific finback whales were performedby means of population genetics with blood types. As Ju antigens havehigher precision than Bp antigens in large scale of investigation, theformer was used for quantitative analysis.
4) Basing upon non-random geographical distribution in frequency ofoccurrence of blood types and rates of emigration of marked whales, it
became to be obvious that different local populations of finback whales
distribute in various areas of the North Pacific including East China Sea,
west and east sides' areas around Aleutian Islands.
5) According to comparison of continued ratios of blood type genefrequencies of populations from neighbouring three areas, finback whales
in East China Sea belong to a local population which is isolated fromthose in areas around Aleutians. Furthermore, this fact was confirmed
significantly by comparison of morphological characteristics such as
growth rate in body length, body length at which sexual maturity is
attained, external body proportions and shape of skull, shape and growthrate of baleen plate.
6) Rate of emigration which was calculated from results of marking
investigations strongly confirm the existence of the two different populations in the west (population I) and east (population II) sides' areas
around Aleutian Islands which was previously clarified by blood typing
investigations, and that population I hardly migrate to the area northof east Aleutian (area C), but population II distribute not only to area
C but also to area off Kamchatka (area A). In other words, migratory
ranges of populations I and II are overlap in the areas A and south ofeast Aleution (B), but relative sizes of both populations which actually
migrate to these areas fluctuates from year to year.

138

K. FUJINO

7) Gene frequencies of populations I and II are estimated by blood
type gene frequencies observed in various areas and rates of emigrationof marked whales through three allele B system. These includes P1:.334, q1: .074, r 1 : .592 in population I and p 2 : .896, q2: .002, r2: .102 in
population II. These figures will be useful for estimating annual ratioof mingling between populations I and II in various areas around AleutianIslands, although some bias may be involved in figures of population I.
Additional data could be serve to bring these values higher precision.
8) After estimating relative sizes of populations I and II which actually
migrate to areas A and C, it became to be clarified that population II
has larger stock number than the other. Higher precision in estimatingthese actual figures should be expected by additional data hereafter.
9) Shape of baleen plates of finback whales from area of Okhotsk
and Pacific sides of the north eastern Japan is different from that from
East China Sea and reveals same type as that from areas around
Aleutians, though the data obtained are very scanty so far. Whether
whales from these areas belong to population I or other will be discussed
after obtaining informations available in future.
10) In spite of fairly abundant annual catch, no recapture of whales,
which was marked in the areas around Aleutians has been reported
from areas off British Columbia and off California. This fact mightsuggest existence of another different population in these areas, so thatit can be concluded that at least four different local populations offinback whales distribute in the north Pacific Ocean.

ACKNOWLEDGEMENTS
The author wishes to express his sincere thanks to Dr. Hideo Omura,
Director of the Whales Research Institute, and Prof. Yasuo Suyehiro,
University of Tokyo for guiding this study and reading original manuscript.
Assistance in collecting samples and discussing on investigations ofstaffs of the Whales Research Institute including Dr. Masaharu Nishiwaki,
Messrs. Takahisa Nemoto, Tadayoshi Ichihara, Seiji Ohsumi, and Keiji
Nasu are also gratefully acknowledged.
Sincere thanks are due to Mr. Takehiko Kawakami and inspectors of
the Japanese Fisheries Agency who offered informations on marking
investigation, to the whaling companies who helped the author in giving
facilities for experiments at landstations and on board factory ships, andto Mr. Teijiro Tajima, Meat Inspection Office, Tokyo, who offered serumsof domestic animals.
Mr. Yukio Nose, University of Tokyo kindly gave to the author valuable

SUBPOPULATIONS OF THE NORTH PACIFIC WHALES

139

suggestions on calculation of the results of marking investigations.
Finally the author would like to record their appreciations of Prof.
John E. Cushing, University of California, Dr. George J. Ridgway and
Dr. Lucian M. Sprague, Fish and Wildlife Service, Department of Interior,
U.S.A. who gave to the author various suggestions and discussions on
mutual problems in serology and genetics.

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K.FUJINO142

APPENDIX. FREQUENCY DISTRIBUTION OF NUMBER OF CORPORA
ALBICANS IN PREGNANT FEMALES TAKEN FROM THE NORTHPACIFIC FINBACK WHALES
Off Kamchatka (A)

0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
@
41
42
Sum

1
6
5
2

1
2
1

13 9
12 8
16 9
12 14
11
9
10 8
7 13
10 5

2
2 11
1 5

5
8

4
1

4
2

3
5

1
1

2
1

3

1
2

1

1

1

4
2

9
7

7
4

4

5

6

South of east Aleut. (B)

8
6

4
3

5 12

2

2 16
4 12
5 10
5 12
6 10
4 18
9
3 12

2

8

3

7

4

6
5
6
3

7
3
5
4

3

17

7

6

3

5

1
2
3

1

9

7

4

3
2

1
1

7
4
5

1

2
1
1

1

1

2
1
2
1

3
2

1

2

2 5 10
2 11
4

6
2

2
4

4

1

1
1
1

2

2

1

1

1

2

6
8
4
2
1
5
3
2

3
1
1

2
1

12 8
7 15
16 6
16 4
2 7 4
12 7
9 10
12 6
2

8
7

3
5

1

7

2

3

5

2

2
1
1
2
1
2

1
1
1
1

1
1
1

1

North of east Aleut. (C)

31
23
16
18
9
12
7
6
3

25
40
21
33
21
24
16
14
6

10 10
19 9
20 14
14 7
9 2
10 4
8
1
4
2
5

13
3 10
2 7

4
2

5

1

2

2

6

4

3

2

1

5
2
2

4
1
1

2
2
2

2

5

8

26

6 14
6 15
5 10
4

14

3

7

1

1

1

2

2

1

1

1
1
2

2
1

2

1

1

1

1

1

1

1

1

1
1
1

1

1

1
1
1

1
1

1

1
1
1

1

1
1
1
30 122 101

33 102 76 53 167

458

10 136 80 13

136 219 117 60 44 140