THESCIENTIFIC REPORTS
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THE WHALES RESEARCH INSTITUTETOKYO· JAPANSEPTEMBER 197 5l Sci. Rep. Whales j
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THE
SCIENTIFIC REPORTS
OF
THE WHALES RESEARCH INSTITUTE
No. 27

THE WHALES RESEARCH INSTITUTETOKYO, JAPANSEPTEMBER, 197 5

THE WHALES RESEARCH INSTITUTE
1-3-1, ETCHUJIMA, KOTO-KU, TOKYO

THE SCIENTIFIC REPORTS OF THE WHALESRESEARCH INSTITUTE
CONTENTS OF NUMBER 27Page

OMURA, H.
Osteological study of the minke whale from the Antarctic
WILLIAMSON, G. R.

37

Minke whales off Brazil
KAWAMURA, A.
A consideration on an available source of energy and its cost for locomotion
in fin whales with special reference to the seasonal migrations . . . . . . .

61

KAsUYA, T. and N1sHIWAKI, M.

81

Recent status of the population of Indus dolphin .
KASUYA, T.
Past occurrence of Globicephala melaena in the western North Pacific . . . . .

95

McCann, C.
A study of the genus Berardius Duvernoy . . . . . . . . . . . . . . . . . . 111
N1smwAKI, M. and KuREHA, K.
Strange organ in the anal region of the finless porpoise. . . . . . . . . . 139

MACHIDA, S.
Distribution of sperm whale catches m the southern Indian Ocean . . . . 141

OSTEOLOGICAL STUDY OF THE MINKE WHALE FROM
THE ANTARCTICHIDEO OMURA
Whales Research Institute, Tokyo

ABSTRACTTwo skulls and postcranial bones of the minke whale from the Antarctic
were studied, comparing with two specimens from the North Pacific. Characters noted in the skull are: breadth of skull is narrow, rostrum is also narrowand long and the flank is rounded and fiat in profile, anterior margin of nasals
is concave, at the vertex of skull triangular region is not formed by frontal andparietal between nasals and supraoccipital or in a more advanced stage oftelescoping, lachrymal is large and rectangular, hamular process of pterygoid
is short and broad. In the vertebrae the 7th cervical lacking parapophysis.
Lateral tubercle of the pelvic bone situated towards mid-length of the bone in
male.
Above are the main differences which separate it from the minke whalein the northern hemisphere, though the whale in the North Atlantic has also
comparatively long rostrum and is separated by this character, together withdifference in form of the white band of flipper, from that in the North Pacific.
In the light of the present stage of knowledge it is not concluded that theminke whale from the Antarctic (Balaenoptera bonaerensis) is a distinct species
from B. acutorostrata.

INTRODUCTION

The problem whether the little piked or minke whale from the Antarctic (Balaenoptera bonaerensis) is a distinct species from the minke in the northern hemisphere(B. acutorostrata) has not been finally solved yet, though Utrecht and Spoel (1962)
and Ohsumi et al. (1970) report the name B. bonaerensis is a synonym of B. acutorostrata. The conclusion of these authors are mainly based on the external morphological characters, and in addition the latter authors have counted number of vertebrae of three small foetuses and found no distinction in this respect. Detailed
osteological study on adult specimen is needed in the light of the present status ofaffairs.
In the 1971-72 season Dr S. Ohsumi had been in the Antarctic on board
Jinyo Mam, a minke whaling expedition and has collected two complete sets ofthe skeleton. These bones are the main material of this paper. In addition tothese, in 1969 I fortunately could collect two sets of skeleton of the minke whalefrom the North Pacific, by courtesy of Mr T. Miyodori, owner of minke whaling
catcher boat operating on the coast of Ja pan. These bones are also studied forcomparison. Dr S. Ohsumi has also collected a number of hyoid bones of theSci. Rep. Whales Res. Inst.,, No. 27, 1975, 1-36.

2

OMURA

minke whale from the Antarctic in the same season. A taxonomic study basedon these hyoid bones has already been published by Satake and Omura (1974).
MATERIAL
The particulars of the specimens treated in this paper are shown in Table 1. The
specimen 71]2793 and 71]2883 were brought from the Antarctic, kept in a cold
storage chamber of Jinyo Maru, and unloaded at Tokyo port from where they
were transported to the laboratory in fresh condition on 30 March 1972. A few
amount of meat and other soft parts were still attached to bones. Baleen plates
were also remained on the beak of the skull. After removal of these, most bones
were hurried in earth in order to extract oils contained in them. Flippers wereenveloped with nylon mosquito nets before burying, in precaution against missingof small phalanges. They were dug out from the earth in December 1973. Fromsmall bones e.g. small caudal vertebrae, hyoid bones, chevron bones, sterna, etc.
oils were extracted by boiling in laboratory and then soaking in water.
TABLE 1.

PARTICULARS OF THE MINKE WHALE SPECIMENS

Specimenno.

Body
length (m)SexDate of catch

Position of catch

Age

71J2793
71J2883
AY69B
AY69A

8.5
9.8
6.6
5.4

M
F
M
M

Feb. 12, 1972
Feb. 16, 1972
Apr. 12, 1969
Apr. 28, 1969

65°-2l'S, 76°-37'E
65°-00'S, 89°-37'E
Coast of Japan
Coast of Japan

Ad. 25 years*
Ad. 39 years*
Juv.
Juv.

* Age determined by Dr S. Ohsumi by means of ear plug.
Bones of the specimens AY69A and A Y69B were buried in sand of beach of
Ayukawa by Mr T. Miyodori. They were dug out from sand after 15 months
and transported to W.R.I. The specimen AY69A was mounted for display andnow being kept at Suginami Kagaku Kyoiku Center, Tokyo, an educational institution for school children in Suginami ward. Other specimens have been kept
at W.R.I.
As shown in Table 1 the age of the specimens is quite different between
samples from the different hemispheres. Two specimens from the Antarctic are
adult and all vertebral epiphyses are completely fused to their centra, whereas two
specimens from the North Pacific are very young and none of the epiphyses is fusedto the body of the vertebra. Age of the specimens from the Antarctic was determined by Dr S. Ohsumi as being 25 and 39 years respectively by means of ear plug,
but for the specimens from the North Pacific no age determination was made.
This difference in age between specimens from different hemispheres makes
it difficult to arrive at correct conclusions from only the direct comparison of thesebones.

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

OS
TEOLOGY O ド M J
NKE W HALE

3

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OSTEOLOGY OF MINKE WHALE

5maxillaries are arranged roughly in line, nearly parallel to the margin of thesupraoccipital bone. Frontal is present as a narrow band between these bones,
but the parietal does not appear in this region of the skull. It covers frontal on theside of the fronto-nasal process of the maxillary. On the other hand in the specimen from the North Pacific the posterior end of nasals is situated more anteriorly,
and premaxillaries end posteriorly than nasals, and maxillaries more posteriorly
than those bones. Thus triangular region is formed by frontal, but frontal itself
is also represented by narrow band and between frontal and supraoccipital there
present parietal, also triangular in shape. In Figs. 1 and 2 only the photographof the specimen AY69B is shown, but in the specimen AY69A too the shape ofnasals and arrangement of bones at the vertex of the skull is quite similar. In conclusion above the minke whale from the Antarctic has attained a more advanced
stage of telescoping than that from the North Pacific.
Omura (1957) reports three skulls of the minke whale from the coast of Japan,
and this time these skulls were reexamined, especially on characters stated above.

2
Fig. 3. Schematic drawing of vertex of >kull of the minke whale.
I. From North Pacific.
2. From Antarctic.
Pm ...... Premaxillary,
N ...... Nasal,
M, ...... Maxillary,F...... Frontal,
Pa ...... Parietal,
So ...... Supraoccipital.

In addition I examined three more skulls preserved at various places in Japan.
Places where these six skulls preserved are: one at the National Science Museumin Tokyo, one at the Tokyo University of Fisheries, two at the Whale Museum in
Ayukawa, one in Shiogama, and one in Matsushima. Length of these skulls areranged from 1,060 mm to 1,520 mm. The largest specimen is the specimen kept
at the National Science Museum, but in this specimen too vertebral epiphyses are
only fused to their centra completely in the first three cervicals and last ten caudals.
Thus all of the specimens are thought juvenile or semi-adult. In all of the specimens the form and position of nasals are quite similar to those shown in Figs. 1
and 2. The parietal is triangular form at the vertex of the skull in general, but theforward projection on the mid-dorsal line is bifurcated in the specimen AY69A, andin a specimen kept in Matsushima most part of this triangular region of the parietal
is covered by supraoccipital (Fig. 4). In the above two specimens, however, theSci. Rep. Whales Res. Inst.,
No. 27, 1975.

6

Or‑t
URA

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OSTEOLOGY OF MINKE WHALE

7frontal posterior to nasals, premaxillaries, and maxillaries is triangular in dorsalview.
The skull length of the Matsushima specimen is 1,220 mm and it is thoughtthat this specimen came also from a juvenile animal.
On the inferior side of the skull differences are noted by visual comparison
between specimens from different oceans, at the posterior margin of palatines andin the form of the hamular process of pterygoid. In the skulls from the Antarctic
the posterior margin of palatines, from mid-ventral point to articulating pointwith pterygoid, is roughly straight, whereas in the specimen from the North Pacificthis line is much concave. The hamular process of pterygoid is short and broadin the specimens from the Antarctic, but it is slender and pointing in the specimen
from the North Pacific (Fig. 5 ). The posterior margin of palatines, however, subject to individual variation, and in the specimen AY69A this line is convex rather
than concave. Among other specimens noted before the lines are concave in fivespecimens and convex in one specimen. But the hamular process of pterygoid isslender and pointing in all specimens, in which this part of the skull remaining
unbroken.
Skull measurements and calculated percentages against skull length are
shown in Table 2. There are rather wide ranges of individual variation, and insome measurements values of the two Antarctic skulls and the two North Pacificskulls are overlap each other. But there are a number of measurements inwhich values are not overlap between the specimens from different oceans. In
the specimens from the Antarctic the length of the rostrum and breadth at its middle are larger than others (Measurement nos. 7, 10, and 11 ), though practicallyno difference is noted in its width at base (M. no. 12). These differences are large,
especially in the specimen 71J2883, the larger whale, but see also Fig. 6. Mandibles are also larger than in the specimens from the North Pacific (M. nos. 3336), and they have a bit higher coronoid process (M. nos. 37 and 38). Thereseems no proportional difference in the breadth of the skull (M. nos. 16, 17 and18), but the breadth of occipital bone is larger in the specimens from the North
Pacific (M. no. 21 ). Size of occipital condyles as well as foramen magnum arealso larger proportionally (M. nos. 22-26). But the proportion of these bones inposterior part of skull subject to age. In the black right whale proportional
length in these bones decreases with age (Omura et al., 1971).
The breadth of the vertex of skull or breadth across the fronto-nasal processesof maxillaries is broader in the North Pacific whale (M. nos. 13 and 14), and lengthof orbit, measured at distal end of frontal, is larger also. But no conclusion could
be reached due to scanty number of material. Anterior breadth of nasal seems
to be greater in the North Pacific specimens (M. no. 9), but this can not be concluded
so, because it is very difficult to measure correctly in the specimens from theAntarctic.
On the inferior side of the skull significant differences are noted. In theskulls from the Antarctic the vomer extends more forward, hence larger (M. no.
28), and palatines and pterygoids are situated more posteriorly than in the speciSci. Rep. Whales Res. Inst.,
No. 27, 1975.

8

OMURA
TABLE 2.

SKULL MEASUREMENTS OF THE MINKE WHALE
Actual length

1.
2.
3.
4.
5.

6.
7.
8.
9.
I 0.
11.
12.
13.
14.
15.
16.
17.
18.
19.
20.
21.
22.

23.
24.
25.
26.
27.
28.
29.
30.
31.
32.
33.
~

35.
36.
37.
38.
39.
40.

Measurements

Antarctic
71]2793
M 8.5mAd.

Antarctic
71]2883
F 9.8mAd.

Codylo-premaxillary length
Length of premaxillary, right, left,, maxillary, superior, rightleft
Tip of premaxillary to vertex,, nasals
Length of nasals, median
Breadth of nasals, anterior
Length of rostrum
Breadth of rostrum at middle
Breadth of rostrum at base
Breadth across maxillaries at vertex
Breadth of frontal across nasals
Breadth between maxillaries at nares
Breadth of skull, squamosal,, , frontal",,,, , maxillaries
Length of orbit, frontal, right,, , left
Breadth of occipital bone
Breadth across occipital condyles
Height of occipital condyle, right, kft
Breadth of foramen magnum
Height of foramen magnum
Length from foramen magnum to vertex
Tip of premax. to anterior end of vomer, median,, palatine, median2, 115
1,490
1,510
1,450
1,390+
1,520
1,364
133
98
1,354
399
630
187
231
237
1,075
1,047
991
188
190
760
203
131
124
70
97
605
258
1,505
I, 782
1,935
210
2,063
2,075
2,247
2,260
303
302
201
204

2,350
1,710
I, 7071,645
1,645
1, 710
1,550
170
137
1,630
534
770
237
261
307
1,256
1,219
1,144
207
207
886
228
145
142
79
127
650
290
1,642
2,055
2, 150
236
2,285
2,265
2,454
2,445
311
315
220
233

"

,, posterior ,,"",,,, pterygoid
Breadth across hamular processes of pterygoid
Length of mandible, straight, right.~, curved, right, leftHeight of mandible at coronoid, right, left,, condyle, right, leftmens from the North Pacific (M. nos. 29, 30 and 31).
Above are the major differences noted in Table 2. Tomilin (1967) presents
a table comparing cranial indices of the Atlantic and Pacific populations in thenorthern hemisphere, and noted that the Atlantic individuals are characterized
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

OSTEOLOGY OF MINKE WHALE

9FROM THE ANTARCTIC AND NORTH PACIFIC% of skull length

in mm

N. Pacific
AY69B
M 6.6m

N. Pacific
AY69A
M 5.4m

Juv.

Juv.

1,382
986
984
939
933+
1,017
872
94
97
869
252
441
152
174
178
708
703
625
156
157
563
157
98
96
83
90
376
219
856
1, 102
1,205
127
1,330
1,330
1,400
1,390
170
179
135
132

1,140
797
798
780
755
834
692
87
71
687
210
367
138
137
115
613
597
535
146
146
473
154
120
100
80
85
297
206
700
910
990
1,005
1,010
1,045
1,050
148
150
115
116

71J279371]2883

AY69B

AY69A

100.0
70.4
71.4
68.6

100.0
72.8
72.6
70.0
70.0
72.8
66.0
7.2
5.8
69.4
22.7
32.8
10.1
11.1
13.1
53.4
51.9
48.7
8.8
8.8
37.7
9.7
6.2
6.0
3.4
5.4
27.7
12.3
69.9
87 .4
91.5
10.0
97.2
96.4
104.4:
104.0
13.2
13.4
9.4
9.9

100.0
71.3
71.2
67 .9

100.0
69.9
70.0
68.4
68.0
73.2
60.7
7.6
6.2
60.3
18.4
32.2
12.1
12.0
10.1
53.8
52.4
46.9
12.8
12.8
41.5
13.5
10.5
8.8
7.0
7.5
26.1
18.1
61.4
79.8
86.8

71.9
64.5
6.3
4.6
64.0
18.9
29.8
8.8
10.9
11.2
50.8
49.5
46.9
8.9
9.0
35.9
9.6
6.2
5.9
3.3
4.6
28.6
12.2
71.2
84.3
91.5
9.9
97.5
98.1
106.2
106.9
14.3
14.3
9.5
9.6

73.6
63.1
6.8
7.0
62.9
18.2
31.9
11.0
12.6
12.9
51.2
50.9
45.2
11.3
11.4
40.7
11.4
7 .1
6.9
6.0
6.5
27.2
15.8
61.9
79.7
87.2
9.2
96.2
96.2
101.3
100.6
12.3
13.0
9.8
9.6

88.2
88.6
91. 7
92.1
13.0
13.2
10.1
10.2

by a relatively longer rostrum.
In Fig. 6 I have compared several cranial indices of the minke whales fromthe North Atlantic, North Pacific, and from the Antarctic. Figures for the North
Atlantic and North Pacific are cited from Tomilin (1967). For the North PacificSci. Rep. Whales Res. Inst.,
No. 27, 1975.

10

OMURA

whales, among four specimens he used, two were cited from True (1904) and two
from Cowan (1939), and my previous data (Omura, 1957) were not included. In
Fig. 6, therefore, all data obtained from the coast of Japan are also shown, but
separately from Tomilin's figures. Further he grouped specimens into two categories of juvenile (skull length less than 133 cm) and adult (skull length over 150 cm)
and calculated mean value for each group, but since samples are limited in number
so in Fig. 6 only ranges of value are shown. From his table only figures for adultwhales are cited, because the specimens from the Antarctic are all adult. But
specimens from the coast of Japan are all juvenile, except one specimen in which
the skull length is 1,520 mm and exceeds his criteria.
Measurement -- - -----~~·-~·-­
No.

I

16

=

10

0

I

ll!!I

---------=~----~-=~-------------t

12i

12Zlm

~~~-D~-m-~------------------·--j

111

:
I

33
34
37
38

x

~

~

29

t

l2ZI

31

I

I

10

20

30

40

50

I

I

I

60

70

80

.3

90
100
110
% against skull length

Fig. 6. Comparative cranial indices of the minke whale populations in the North
Atlantic, Antarctic, and North Pacific. 'White ... North Atlantic,
Hatched... Antarctic, Double hatched ... North Pacific (from Tomilin), Black ...
North Pacific (coa~t of Japan). Each square denotes ranges of value, and crosssingle value. For measurement number see text.

The measurements compared in this fugure are the following.
Measurement no. 16.
Breadth of skull, squamosal.
,,,, 10.
Length of rostrum.
,, 12.
Breadth of rostrum at base.
",, 11.
Breadth of rostrum at middle.
"
33, 34 Length of mandible, curved.
" 37, 38 Height of mandible at coronoid.
"
" 29.
"
Tip of premaxillaries to anterior""
palatine.

end ofSci. Rep. Whales Res. Inst.,
No. 27, 1975.

OS
・J"EOLOGY OF MIKKI
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OSTEOLOGY OF MINKE WHALE

13

ference is noted between specimens from different oceans.
In Fig. 7 are shown the tympanic bullae of the minke whale from the Antarctic, compared with those from the North Pacific. As seen from the photographand measurements in Table 3 no special difference is noted between bullae fromdifferent oceans. Bullae of the specimen 71]2883 are larger than other specimens,
but in the remaining specimens the size is nearly the same. Compared with otherthree specimens reported by Omura (1957) also no special feature is noted in thesize of bulla, notwithstanding differences in the size of whale body.
The malars and lachrymals are shown in Fig. 8. As seen in the photographmalars are quite similar in shape in general. but lachrymals are of some interest.
In the Antarctic specimen lachrymals are comparatively large and roughly rectangular in shape, like in other balaenopterid whales, but those from the North
Pacific are short and one end is pointing and they resemble closely to anotherspecimen reported by Omura (1957). It is suggested, therefore, this character
is of some importance in taxonomic consideration. Measurements of malars andlachrymals are shown in Table 4.
TABLE 4. MEASUREMENTS OF MALARS AND LACHRYMALS OF THE MINKEWHALE FROM THE ANTARCTIC AND NORTH PACIFIC (in mm)
Lachrymal

Malar
Length 1)

Breadth 2)

LengthBreadth~..______,._

~

~

~

Specimen

Antarctic
71J2793
7IJ2883
N. Pacific
AY69B
1) Greatest

Right

Left

Right

Left

Right

Left

Right

Left

244
254

240
253

31
40

30
38

264
247

258
265

81
99

82
104

169

181

18

17

115

113

63

61

2) At mid-length

VERTEBRAE (Plates VI-IX)

The vertebral formulae of the present specimens are as follows:
Antarctic specimen71]2793
C 7, D 12, L 10, Ca 20, Total 49.
71]2883
C 7, D 11, L 12, Ca 20, Total 50.
North Pacific specimen
AY69B
C 7, D 11, L 12, Ca 18, Total 48.
AY69A
C 7, D 11, L 11, Ca 19, Total 48.
Thus among these specimens from different oceans there is a difference in total
number by 1-2. In two specimens from the North Pacific reported by Omura(1957) the total number of vertebrae are 48 and 47. Both have 7 cervicals, 11
dorsals, 12 lumbars, and caudals are 18 and 17 respectively. Tomilin (1967)
describes that " Vertebral formula reduced as compared to other rorquals: C 7,
D 11, L 12, Ca 18, total 48. However, the vertebral count may reach 50 owing
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

14

OMURA

to additional vertebrae in the lumbar (up to 13) and caudal (19-20) regions".
True (1904) lists up the vertebral formulae by various authors then known
to him, and among them the largest number is 50, reported by Turner (1892)
from Granton, Scotland. To this specimen he describes that " The enumerationof Sir Wm Turner is probably the most accurate, having been made under favorable circumstances, and with the intent of correcting previous errors ". This may
be the basis of the above statement of Tomilin. In any case 50 might be rather
exceptional number in the northern hemisphere.
The present specimens from the Antarctic have 49 and 50 vertebrae in total
respectively, suggesting a difference in this respect. But Ohsumi et al. (1970)
counted number of vertebrae of three fetuses of the minke whale from the Antarctic.
Among these two had 48, and one 49 vertebrae. These fetuses are small and theirbody length has ranged from 46.5 to 50.5 cm, and the number of caudal vertebrae
was uniformly 18. This number of 18 caudals is smaller by two, compared withthe present specimens. Further study is needed, therefore, and it may be premature at present to conclude definitely in this character.
The two specimens from the Antarctic have already attained their physicalmaturity, and vertebral epiphyses are ankylosed to their centra completely. On
the other hand the specimens from the North Pacific are both immature, and noneof the epiphyses fused to their centra.
Anterior view of cervicals of the specimens 71]2793, 71]2883, and AY69B are
shown in Plate VI. As seen in these photographs there are some individual variation in the stage of development of neural spines. In the specimen 71]2793 theneural spine is not developed completely in the 3rd and 4th cervicals. In thesevertebrae the right and left spines do not united at their tips, whereas in other two
specimens they fused completely. Development of the diapophyses and parapophyses is also subject to individual variation, though it seems certain that thedevelopment subject to growth of whale body in general. For example theseprocesses do not fused at their tips on the left side of the 3rd cervical of the specimen71]2793, whereas complete ring is formed in the specimen 71]2883. But in the
5th cervical complete ring is formed on both sides in the former specimen, whereas
ring is not formed on both sides in the latter specimen. In the North Pacific specimen complete ring is formed in the 5th cervical on both sides and on the right sideof the 6th. In another two specimens from the North Pacific reported by Omura(1957) none of these rings is formed, even in the axis. A 25 feet (7.5 m) malespecimen had cervicals of lesser develpoment than in a 6.6 m male whale.
Accordingly differences in form of cervicals, especially of the development
stages of processes, are not thought as important from the taxonomic stand point.
However there is one thing to note. In the specimens from the Antarctic the 7th
cervicals are lacking parapophysis and there is no sign of tubercle. But in the
North Pacific specimen this process is present also on the 7th cervical, as shownin Plate VI. In the another two specimens from the North Pacific also parapophysis is present on both sides, though it is reduced to a tubercle (Omura, 1957).
Cowan (1939) states that "in 7 (7th cervical) it (parapophysis) is reduced to a
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

OSTEOLOGYOF MIJ'¥KE WHALE

1
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.2λ1975.

OMURA16

and the last vertebra with spmous process is lOth and 9th caudal respectively.
However these character may subject to individual variation and less importantfrom the taxonomic standpoint.
The specimen 71]2883 from the Antarctic is of some interest from the pathological view. In this specimen 2nd, 3rd, and 4th caudal vertebrae are developed abnormally, possibly from some pathological cause (Fig. 9). The posterior
and inferior part of the centrum or the articulating facet with chevron bone developed extraordinarilly, forming tubercles. The articlating part of the chevronbone also developed into tubercular large face. The posterior margin of the 3rd

mm
240
220
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1--C--l f---D--l f---L---1 f---ca----l

VERTEBRAE
Fig. 10. Comparison of size of vertebrae of minke whales from the Antarctic andNorth Pacific. Upper two lines denote specimens from the Antarctic, and lower
two lines specimens from the North Pacific.

caudal also developed and it covers anterior margin of the 4th partly. Similar
abnormality was also observed in 2nd and 3rd caudals of a ,Ziphius cavirostris (Omura,
1972)
Measurements of vertebrae are shown in Appendix. From these measurements I have calculated the mean length of each centrum using the following
formula reported before (Omura, 1971), in order to compare volume of eachvertebra.
Mean length = (a x bx c ) 1 13
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

OSTEOLOGY OF MINKE WHALE

17

where a, b, and c are the breadth, height, and length of the centrum
respectively.
Results of calculation are shown in Fig. 10. I also calculated percentages
against their first lumbar, but in this case four lines are overlapping in most parts,
except in cervical and 4th-6th caudal regions. The 4th caudal of the specimen
71J2883 shows somewhat higher value compared with neighboring vertebrae, but
this is partly due to extraordinary growth as stated already. Also in the specimen
71J2797 the 5th caudal shows high value. In this vertebra the breadth of centrum
is the largest among the series of vertebrae. On the other hand two specimens fromthe North Pacific draw more smoothed curve in these regions. Such differences
may be attributable to the difference of age partly, but in general the trend offour curves does not differ materially and such differences shown by Omura (1971)
among different species of baleen whales are not noted.
TABLE 5.
Specimen

SKULL AND VERTEBRAL LENGTH OF THE MINKE WHALE FROM
THE ANTARCTIC AND NORTH PACIFIC
Skull71]2883

2,350

71]27932, 115

AY69B

1,382

AY69A

1,140

71]288371]2793
AY69B
AY69A

25.85
26.48
23.40
22.34

CervicalDorsal(Actual figures in mm)
l, 335
315
(11)
(7)
1,389
280
(12)
(7)
935
223
(7)
(11)
862
209
(7)
(11)
3.47
3.50
3.78
4.10

(Percentage)
14.68
17.39
15.83
16.89

LumbarCaudal

Total

2,309
(12)
1, 764
(10)
1,646
(12)
1,302
(11)

2, 782
(20)
2,439
(20)
1, 720
(18)
1,590
(19)

9,091

25.40
22.09
27.87
25.51

30.60
30.54
29.12
31.16

7,987
5,906
5, 103

100.0
100.0
100.0
100.0

Figures in parentheses show number of vertebrae.

In Table 5 are shown the total length of skull and vertebrae of the minke whalefrom the Antarctic and North Pacific. The vertebral length is the total of thelength of each centrum. Accordingly this length does not show actual length insitu. Further the vertebral length is shown dividing into four regions of cervical,
dorsal, lumbar, and caudals. The length of skull and each region of vertebrae
are also shown their percentage figures against total length of skul and vertebrae.
In comparing these figures it will be noted that in the length of cervicals and caudals
there is no significant difference between specimens from the Antarctic and North
Pacific. The skull length is somewhat larger in the specimens from the Antarctic
than those from the North Pacific. The dorsal and lumbar regions show individual
variation, reflected by different number of vertebrae, but when the two regions
are considered together, then the specimens from the North Pacific have somewhat
larger dorsal and lumbar regions than those from the Antarctic. The combined
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

8
1mνfURAehomt
rn7リ2793f
me
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er Sp
. Upp
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heminkewftso. Chじvronbone
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19

OSTEOLOGY OF MINKE WHALE

percentages of these regions are around 40 percent in the specimen from the Antarctic and the corresponding figure from the North Pacific exceed 42 percent.
In the specimen reported by Omura ( 1971) this figure is 44.66 percent of the total
length, and the skull length occupies 22.25 percent of the total.
The proportional length of skull and vertebrae may differ between adult andjuvenile specimens even in the same species or population, and definite conclusion
is very difficult to reach. But the last mentioned specimen is the largest among
I have ever examined of the minke whale skeleton from the North Pacific andsemi-adult as stated already. Its body length is 7.5 m and the skull length is
1,520 mm. It is suggested, therefore, the Antarctic minke whale has more larger
TABLE 6.

MEASUREMENTS OF CHEVRON BONES OF THE MINKE WHALE
FROM THE ANTARCTIC AND NORTH PACIFIC (in mm)71]288371]2793~....__,_

AY69B,,---·

AY69A,,..---•

LengthHeightLengthHeightLengthHeightLengthHeight

*{43
47

*{80
77

*{38
52

*{93
97

*{5?

*{8?

*{5?5

*{9?

*{124
132
202
213
186
149
119
101
80
46

91

266

78

164

45

101

3
4
5
6
7
8
9
10

*{73
79
103
147
176
172
153
146
121
74

162
190
211
207
171
169
138
87

270
246
229
197
154
120
89
55

108
112
110
99
104
86
52

156
144
124
107
80
60
30

82
88
84
74
65
62

134
128
112
96
82
67

11

*{32
29

*{22
22

*{V

*{3?3

2

* Not united. Question mark means missing.

skull compared with that from the North Pacific, and this is a reflection of more
longer rostrum as shown in Fig. 6. In the external body proportions this is alsosuggested in the proportional length from tip of snout to center of eye (Ohsumi
et al., 1970).
The chevron bones of the two specimens from the Antarctic and a specimen
from the North Pacific are shown in Fig. 11. From the minke whale from theAntarctic 11 chevron bones were secured from the both specimens. In the specimen71]2793 the first two and the last one, and in 71]2883 the first and the last are
not united of the right and left laminae. In the specimen from the North Pacific
9 chevrons were obtained and only in the first two laminae separated. Chevronbones are of little taxonomic value, but chevrons of the specimen 71]2883 areof some interest. As seen in Fig. 9 in the 2nd, 3rd, and 4th chevrons there arebony bridges connecting right and left laminae at their proximal ends. In the
2nd it developed abnormally as stated already, but in the 3rd and 4th no suchabnormality is observed. One explanation may be, therefore, that the bony bridge
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

20

OMURA

is formed when the whale has attained very high age. Of course this is not conclusive and further study is needed. In Table 6 measurements of chevron bones
are given.
RIBS AND STERNUM (Plates X and XI)
Ribs of the minke whale from the Antarctic are massive in general compared withthose of the North Pacific specimen, but this may of course be attributable to the
TABLE 7.

No.

2
3
4
5
6
7
8
9
10
II
12

STRAIGHT LENGTH OF RIBS OF THE MINKE WHALE FROM THE
ANTARCTIC AND NORTH PACIFIC (in mm)71]279371]2883

AY69B

AY69A

~

~

r---"----,

~

Right

Left

Right

Left

Right

Left

Right

Left

721
1,060
I, 216
1,279
I, 163
1,210
1,163
1,096
1,030
*910+
*770+

730
I, 103
1, 229
1,263
1,262
1,206
1,156
1,085
1,025
987
*826+
*448+

794
I, 133
1, 335I, 384
1,410
1,355
1,326
1,209
*I, 160+
1,200
990

*570+
1,153
1,333
1,380
1,381
1,389
1,295
1, 219I, 155I, 160
1,202

505
733
880
907
895
865
838
*715+
805
780
*752+

510
740
875
915
910
875
863
830
805
755
*800+

435
627
730
755
753
735
730
682
643
630
427

417
630
742
762
748
718
714
690
642
622
352

* Broken and not estimated.
TABLE 8.

LengthBreadthMEASUREMENTS OF STERNUM OF THE MINKE WHALE FROM
THE ANTARCTIC AND NORTH PACIFIC (in mm)71]279371]2883

AY69B

AY69A

394
348

388
365

275
205

205
167

difference in age. There is no difference in number, having 11 pairs of ribs inspecimens from the Antarctic and also from the North Pacific, but as stated alreadya specimen from the Antarctic (71]2793) had one small accessory rib in addition
to 11 pairs. In one specimen from the Antarctic (71]2793) from 2nd to 9th rib
are doble-headed, but in other specimen (71]2883) clear double-head is observed
only in the 2nd and 3rd. Specimens from the North Pacific are similar to thelatter specimen in this respect. Measurements of ribs are shown in Table 7.
The form of the sternum is cruciate in general in the specimens both fromthe different oceans, but it varies individually and has less taxonomic value. Photographs ofsterna of the specimens 71]2793, 71]2883, and AY69B are shown in Plate
XI, but quite similar forms are already reported from the minke whale from theNorth Atlantic (True, 1904; Tomilin, 1967). Measurements of sterna are shown
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

21

OSTEOLOGY OF MINKE WHALE

m Table 8.
SCAPULA AND FLIPPER BONES (Plates XII-XVI)

Scapullae of the two specimens from the Antarctic and one specimen from theNorth Pacific are shown in Plate XII. They are typical of the balaenopterid
whales in form. Acromion and coracoid are well developed. In the specimen71]2793 the superior margin is somewhat depressed compared with other two
specimens. The upper curvature may subject to individual variation, though
TABLE 9.

MEASUREMENTS OF SCAPULA OF THE MINKE WHALE FROM THE
ANTARCTIC AND NORTH PACIFIC (in mm)

Greatest breadthGreatest height
Length of acromion, inferior
Breadth of acromion, distal end
Length of coracoid, inferior
Breadth of coracoid, distal end
Length of glenoid fossa
Breadth of glenoid fossa
Ratio of breadth against height

71J2793

71J2883

AY69B

AY69A

~

~

~

~

Right

Left

Right

Left

Right

Left

Right

Left

812
449
219
65
87
50
163
121
1.81

805
452
217
69
84
52
163
118
I. 78

874
491
253
72
95
59
182
133
I. 78

864
491
251
79
98
60
192
133
I. 76

535
312
127
39
65
26
126
79
I. 72

544
307
132
42
69
28
129
80
I. 77

394
260
102
25

391
258
92
26

142*
69
1.52

138*
68
1.52

* Including coracoid.

Tomilin (1967) describes that in the juveniles, the contour of this part of the body
is more convex, and in the adults, more or less straightened. In Table 9 measurements of scapullae are shown, together with ratio of breadth against height of thescapula. As seen in this table the specimen AY69A shows somewhat smaller value
in this ratio, whereas in othei: specimens no remarkable difference is noted. But
the relative size of the bone increases with age, growing much more intensively
in width than in height (Tomilin, 1967). No special difference is noted between
scapullae from the different oceans.
Humerus, radius, and ulna (Plates XIII and XIV) also present no noticeable feature between samples from the Antarctic and North Pacific. In theformer specimens epiphyses are united to the body in humerus completely, and inradius and ulna only the proximal epiphyses are united, and distal epiphyses are
not ankylosed in both specimens. In the specimens from the North Pacific noneof the epiphyses is united to the body. Of course this difference is due to the difference in age, and complete ankylosis in these regions of body may occur when
the whale reached very high age, long after the completion of vertebrae. Measurements of humerus, radius, and ulna are shown in Table 10.
Carpals are of no special feature. They are shown in Plates XV and XVI,
together with phalanges. Measurements of phalanges are shown in Table 11.
As seen in Plates and Table no distinction is noted between specimens from
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

OMURA

22

TABLE 10. MEASUREMENTS OF HUMERUS, RADIUS, AND ULNA OF THE
MINKE WHALE FROM THE ANTARCTIC AND NORTH PACIFIC (in mm)71]279371]2883

AY69B

AY69A

~

~

,...-----'~,_----J'--..

LengthBreadthLengthBreadthLengthBreadth

152
151

303
306

164
161

232
229

110
109

191
190

118*
118*

549
550

89
88

573
577

119
117

379
383

67
67

299
301

50
48

501
499

68
67

540
544

85
84

342
345

45
44

301
306

36
35

LengthBreadth284
288

Humerus
Right
Left
Radius
Right
Left
Ulna
Right
Left

Note: Length and breadth measured at middle.
* Greatest.

TABLE 11.

LENGTH OF PHALANGES OF THE MINKE WHALE FROM THE
ANTARCTIC AND NORTH PACIFIC (in mm)
Left

Right
Specimen

Phalanx
II

III

IV

II

III

IV

71.J2793

1
2
3
4
5
6
7
8

75
75
62
46
25

99
85
62
43
29
21
15
10

80
77
62
44
32
23
17

56
53
46
34

75
76
62
46
29

101
88
62
46
31
23
18
11

79
77
62
46
35
24
16
7

56
55
46
31

71]2883

1
2
3
4
5
6
7
8

86
96
77
56

109
112
80
59
40
29
-*
*

89
98
84
63
47
34
21

61
60
63
50

89
96
78
53

109
114
84
60
41
28
20
14

91
100
84
63
47
34
21

61
61
63
50

51
49
39
26

64
57
42
28
19
13
7
-*

53
50
40
28
18
11

45
39
31
17

52
49
39
25

64
56
41
29
19
12
8
6

55
51
41
28
18
11

44
38
31
16

AY69B
2
3
4
5
6
7
8

Note: Metacarpals are included,* Possibly missing.

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

E!NKE WHALTEOLOGY OF恥f
OS

23

s
cimensi
pe
es
s
he
ft
lao
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eat
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etwos
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engthandt
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eor8.
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b
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e
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h
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r
o
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n
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l
a
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ds
e
ound mark
f
f
eo
renc
e
f
f
i
ld
a
xu
e
es
b
i
r
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s
. (1971)alltas. Omurae
e
tus
e
le f
a
e wh
u
and bl
s
e
n
o
b
r
e
g
r
a
l
h
c
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emal
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h
nt
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twhale
h
g
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kr
c
a
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h
nt
si
cbone
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pel
.
emale
nf
ani
h
t
e 200
r
sa
c bonelvi
etpfet and l
igh
fr
th o
eng
he l
pecimen 71J2793 t
hes
nt
I
ng
pondi
s
e
r
r
o
. Thec
y
el
v
ti
c
e
sp
e
e46and42mm rr
rwidthahei
and204m mandt
e
Th
.
y
l
e
v
i
t
c
e
p
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e
r
m
m
9
1
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8
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r
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n
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f
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h Pa
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heNor
pecimensfrom tnthesI
handwidthor
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n
e
hel
g,t
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o
4m l
.
r5
eo
nAY69A,amal
pecimenAY69B, buti
s
t
i
sab
zei
i
hes
y. Thust
l
e
v
i
ct
spe
e
,27m m r
7,30,and84
e8
rcbonesavi
bothpel
cimen
e
p
es
h
ot
st
e
embl
s
e
cimen 7リ2883. Theform r
pe
hes
ft
to
a
h
han t
rt
e
g
ar
l
‑
o
sphot
t
n
e
s
e
r
op
s
. Heale4m)mal5.
t(ee
8fa(1957）什oma 1
dbyOmur
e
t
r
o
ep
r
.
.hut
s
sRe
.Rep.Whale
i
c
S
.
5
7
9
,1
7
.2
o
N

24

O M URA

g
r
aphs
howi
暗 t
hepe
lvi
ebone
sofa25feel(
7.
5m)mal
e(F
i
g.1
3)
. Int
hi
ss
p
e
cime
n
t
he p
e
lvi
c bones arc much el
ongat
ed. Thiswhal
e wasasemiadu
lta
sal
r
eady
s
t
at
e
d,butpossi
blyt
hewhalewas ma
tu
r
es
e
xualy
, beeauseaver
age body l
ength
a
ts
e
xualmaturityi
s22‑23f
e
e
t(
6.
6‑6.
9m)i
nmal
e
s(
OmuraandSaki
ur
a
,1
956).
Whent
hep
cl
vi
ebone
soft
h
iss
peeimenar
ecompar
edwi
t
ht
ho
s
eo
ft
h
es
p
e
cimen
7リ2793 a c
li
f
f
er
enee i
snot
ed i
nt
h
e
i
rf
orm. In t
h
ef
o
rme
rs
pe
c
imen from t
h
e
Nor
t
hPac
i
f
ict
hel
engt
hofpe
l
vi
cbone
sa
r
e1
7
4and 181m mr
e
spe
c
t
i
v
e
l
yandabi
t
s
mal
l
e
rt
han t
hel
at
t
e
rs
peeimen from t
heAn
t
a
r
et
ic
, bu
ts
t
i
l
l qu
it
eeo
mpa
r
a
b
l
e.
暢

F
i
g
.1
3
. Pel
v
i
cbonesoftheminkewhalefromtheNorthPacific. A m
a
l
e
‑of7
.
51
1
1
l
ong(
ci
t
e
df
r
om Omu
r
a
,1
9
5
7
)
.

I
nt
h
iss
pe
cimen t
het
uber
c
le
s
,t
uber
culumJ
a
t
e
r
al
e
,a
r
cl
e
s
seminentands
i
t
u
at
ed
mor
epost
e
ri
or
lyt
hant
hes
pe
c
i
men fromt
h
eAnt
a
r
c
ti
c. Furtherinthespeeimen
f
r
om t
heNor
t
h Pa
ci
f
i
et
he c
r
an
ialp
r
o
c
e
s
s
e
sa
r
el
ong and muchslenderthan inthosefromt
heAnl
ar
e
ti
c. Theper
eentagef
i
gur
e
sofwi
d
thag
a
in
stl
e
ngtho
fr
i
ght
andl
e
仕p
e
lvi
cbone
sa
re1
6and1
4pe
r
c
en
tr
e
spe
c
ti
v
e
l
yi
nt
hes
pe
c
i
men from theNorth Pac
i
日e
,whe
r
e
ast
hecor
re
sponding f
i
g
u
r
e
s oft
hes
peeimen f
r
om t
h
e An
‑
t
a
r
ct
icar
e23and 21per
c
en
tr
e
spe
ct
ivel
y. Thisd
i宵er
enc
ei
nr
at
i
oi
sar
ef
le
ct
ion
o
fl
e
s
sdeveloped l
at
e
r
altub
er
cl
e
si
nt
hes
pe
c
imen from t
heNorth Pa
c
i
自c
. Fo
r
t
hef
emalenoc
omparab
l
ema
t
e
r
i
ali
sa
v
a
il
abl
ef
ort
heNor
t
hPa
ci
f
i
cmi
nkewhale.
DI
SCUSSION
In t
h
ef
o
regoi
ng chaptersthe cha
r
a
c
t
er
is
t
i
c
soft
he minke whale 仕om t
heAn・
t
ar
c
t
ica
r
ed
is
eus
s
ed mai
n
ly c
ompar
i
ng wi
t
ht
he whalef
rom t
heNort
h Pa
ei
f
i
c.
Theminkewhal
ef
rom t
heNor
t
hPa
ci
f
icwasnamedBa
l
a
e
n
o
pt
e
r
ad
a
v
i
d
s
o
n
ibyScam‑mon(1872). True(
1904)compared twoskullsfromt
h
eNor
t
hPa
c
i
自c
,i
nel
udi
n
t
het
ypeofピ
i
a
v
i
d
s
o
n
i
,wi
t
ht
ho
s
eofα
c
u
t
o
r
o
s
t
r
a
t
af
r
om t
heNorthAt
lant
ie. Hiscon‑
c
lus
i
o
n was therecannotbes
aidt
obeacons
t
an
tdi
f
f
e
r
e
n
c
ei
nanyoft
hepro‑
port
ionsbet
weent
hes
pe
cimensfr
om t
het
wooceans". Cowan(
1939)andO mura
(
1
957)s
uppor
t
ed t
h
iseonclusion,but Tom
i
l
i
n(
1
967)f
ound that Cr
a
n
i
a
li
n‑
Sc
i
.R
e
/
1
.Wh
a
l
e
sRe
s
.I
n
s
t
.
,
}
'
I
匂.2
7
, 197
5.

OSTEOLOGY OF MINKE WHALE

25

dices of both populations show that the Atlantic individuals (adult and, particularly,
juvenile) are characterized by a relatively longer rostrum ". And he concluded" We consider this character, which is manifested also by the general proportionsof the body (Table 61 ), as a significant difference between the North Atlantic subspecies Balaenoptera acutorostrata acutorostrata Lecepede, 1803-1804 and the North
Pacific Balaenoptera acutorostrata davidsoni Scammon, 1872 ". In this connection I
like to point out the difference in the form of the transverse white band on the
flipper. This problem was first raised by Scammon (1872), but his description
is not explicit as pointed out by True (1904) and his figure (Scammon, 1874) ofwhite band is not correct.
In the minke whale from the North Pacific the anterior margin of this whiteband runs nearly perpendicular to the flipper and there is a promontory or
projection of white area into the black, towards the mid-line. This is well shownin Plate II of Omura and Sakiura (1956). Almost all minke whales from theNorth Pacific investigated by myself and my colleague exhibit this pattern. The
minke whale taken at Los Angeles Harbor also has similar pattern (Fig. 2 of Fry,
1935). On the other hand in the minke whale from the North Atlantic the marginof the white band draws roughly semi-circle and no forward projection of whitearea is noted. This form is well demonstrated in the upper photograph of Plate
I of Moore and Palmer (1955) and also in Fig. 3 of Sergeant (1963).
In the skull no other difference than that pointed out by Tomilin seems topresent between the specimens from the North Pacific and North Atlantic. Judged
from the photographs shown by True (1904) the anterior margin of the nasals is
flattened as described by Tomilin (1967). The triangular region formed by frontal
and parietal on the vertex of the skull, between nasals and supraoccipital whichshows lesser degree of telescoping, seems also present, although I could not findout any description on this particular point of the skull in other literatures. The
form of the hamular process of the Massachusetts specimen is quite similar to my
specimen from the North Pacific. In the post-cranial bones differences are noted
in two characters between specimens from the Antarctic and North Pacific. One
is the presence of parapophyses on the 7th cervical in the specimens from the North
Pacific, but lacking in those from the Antarctic. The North Atlantic specimens
have also this processes on the 7th cervical, as already stated. Another difference
is in the form of the pelvic bones, but I could not examin this on the North Atlantic
specimen.
The number of vertebrae does not differ between specimens from the North
Atlantic and North Pacific, but further counting is needed in whales from theAntarctic. A distinction is noted in the hyoid bone between specimens from theAntarctic and North Pacific, the former specimen having more longer stylohyals
proportionally (Satake and Omura, 1974), but nothing is known of the specimen
from the North Atlantic.
In the external morphological characters which separate minke whales fromthe Antarctic from those from the northern hemisphere are: 1) no white bandpresent on flipper, and 2) baleen plates are white at the front of the series andSci. Rep. Whales Res. Inst.,
No. 27, 1975.

26

OMURA

1. Skull.
a. Breadth.

Broad (54.6-57.3%) - - - - - - - - - - 1
Narrow (50.8-53.4%) ------------------~ 1
b. Rostrum: I
Dorsal view. Flank straight and pointed -----~~
Rounded or curved ------------------: I
Profile. Curved downwards
Flat '-------------------------------------1 I
I
ILength. Long (60.4-69.4%) --------------------,
1
Short (61.8-64.0) ----------~1
Breadth at base. Broad (32.7-36.6%) - - - - - - - + - i
Narrow (29.8-32.8%) ----------~ I
1
Breadth at middle. Broad (23.1-24.3%)'
Narrow (17.0-22.7%)7c. Lachrymal. Short and one end pointing,-·
1
Large and rectangular-----------------~ I
d. Nasals: I
Anterior margin. Convex or flattened ______ _;._,
Concave----------------------~ I , --, 1
Inferior surface. Grooved - - - - - - - - - - - - - + - ' ,·Not grooved ------------------~ I
e. Apex of skull, between nasals and supraoccipital
I I
Triangle region is formed by fronta I and parietal ~ --lNo triangle region present ----------------------~ I ~ acutorostrata
f. Hamular process of pterygoid
I
Slender and pointing ______ - - - - - - - - t - 1 ,___ davidsoni
Short ·and broad-------------------------------, I
I I
2. Vertebra:--r---- bonaerensis
g. 7th cervical. Parapophysis present _~~11
absent --------------~ I--+_,==---=---=--1--1 ____!

3. Pelvic bone (Male).
h. Tuberculum lateral e.
4. Hyoid bonei. Stylohyal.

l I
I i , ---1

Situated posteriorly - - - - ,11towards mK:ldle -----~

,-

·

J /

---1

1

1

!
ii

Short
Long ----------------------------------: I
5. External characteristics
l I
j. Baleen plates. Uniformly yellowish-white -----~~
Gray and white at back series -----~ I
k. Flipper. White band present: I
Prominence at proximal margin
Lacking prominence:
White band absent ____________________ J---f-J

Fig. 14. Relationship among the minke whale populations from the North Atlantic(acutorostrata), North Pacific (davidsoni), and Antarctic (bonaerensis).

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

OSTEOLOGY OF MINKE WHALE

27

gray and white at the back, whereas uniform yellowish-white in whales in thenorthern hemisphere (Williamson, 1961; Utrecht and Spoel, 1962; Kasuya andIchihara, 1965; Ohsumi et al., 1970).
In Fig. 14 the relationship among the minke whale populations from theNorth Atlantic (acutorostrata), North Pacific (davidsoni), and Antarctic (bonaerensis)
is shown. In the figure the skull indices are the percentages figures against skull
length, and all from the mature specimens, cited from Tomilin (1967. Table 61)
for the North Atlantic and North Pacific specimens. These figures are the rangesof one or two groups, for example the breadth of skull shows ranges of two populations of acutorostrata and davidsoni altogether on one hand (broad) and ranges ofanother population bonaerensis on the other (narrow).
Fig. 14 is a summary of this study and the next problem is the validity of thesecharacters. The most weak point in osteological study of large whales is only
limited number of samples are available, and it is difficult to ascertain whether
the difference is specific or subspecific or only individual or by growth. There arerather wide individual variations in measurements of skulls and other bones. Formof rostrum and other particular bones would also subject to variation. Relation
between form of nasals and flat or bending down rostrum is of some interest. In
the sei whale (Balaenoptera borealis) nasals are convex and rostrum is bending, whereas
in the Bryde's whale (B. edeni) nasals are concave and rostrum is flat. It is thought,
therefore, these are connected with food and feeding or other ecological mattersof whales concerned. But no conclusion cannot be reached at present. Accordingly further study is needed in order to clarify these characters listed in Fig. 14.
But even at this stage it can be said that the Antarctic population bonaerensis is more
distinctly separated from other two populations in the northern hemisphere, i.e.
acutorostrata and davidsoni, though this distinction may not specific level. Further
it has attained a more advanced stage of telescoping than the populations in thenorthern hemisphere.
The North Pacific population davidsoni is separated from the North Atlantic
population acutorostrata in having relatively shorter rostrum and different patternof white band on the flipper. The Antarctic population bonaerensis is separated
from the above two populations in having relatively narrow skull and rostrum,and in other 9 characters, as shown in Fig. 14. The habitat of this population
is not confined within the Antarctic waters and it follows south and north migration, but possibly it is limited in the southern hemisphere.
Williamson (1961) found great similarity of this population (his « Balaena »whales) Balaenoptera bonaerensis Burmeister, 1867, and the likelihood that B. huttoni
Gray, 1874 is also of the same species, and proposed that all three types be regarded
as specimens of B. bonaerensis provisionally. But he added that " It has yet to bediscovered whether this latter is a true and separate species or a subspecies of B.
acutorostrata ". As already stated the number of vertebrae is not differ, at leastat this stage of study, between B. acutorostrata and B. bonaerensis, though muchdifferences are found in other particular points, it cannot be concluded that B.
bonaerensis is a distinct species from B. acutorostrata.
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

28

OMURA

Another question is the occurrence of the minke whale with white band on
flipper in the southern hemisphere (Williamson, 1961). Taylor (1957) reports
observation of rorquals in pools in sea-ice in the Crown Prince Gustav Channel,
Antarctic peninsula, and describes that "some of the rorquals had light patcheson the upper surface of their flipper ". But no information is available at present
to know the relationship between this population of the minke whale in the southernhemisphere and B. acutorostrata, nor between this population and bonaerensis.
ACKNOWLEDGMENTS
I am much indebted to many persons during the course of this study. First of all
my sincere thanks are due to Dr S. Ohsumi, Far Seas Fisheries Research Laboratory,
who kindly secured two sets of skeleton of the minke whale from the Antarctic,
and to Mr T. Miyodori, owner of minke whale catcher boat operating on the
coast ofJapan, who supplied also two sets of skeleton from the North Pacific. Thesefour skeletons form the basis of this study.
During the preparation of bones as complete sample many persons helpedme greatly, including Dr S. Ohsumi and Dr T. Kasuya and his co-workers at
Ocean Research Institute of the University of Tokyo. Dr A. Kawamura, Mr
S. Machida and Miss Yuko Satake of Whales Research Institute have assisted not
only in preparation of bones but also in taking measurements and photographs.
I am very grateful for them all.
I am much indebted to Dr M. Nishiwaki of the Ocean Research Institute
who gave me very valuable advise throughout this study. My thanks are also
due to crew of Jinyo Maru and staff of Taiyo Gyogyo K. K. They have donevery troublesome works on the deck of the factory ship when sampling the bones
and transporting them to W.R. I.
REFERENCES
CowAN, I. McT., 1939. The sharp-headed finner whale of the eastern Pacific. ]. Mamm., 20(2): 215-225.
FRY, ]R., D. H., 1935. Sharp-headed finner whale taken at Los Angeles Harbor. ]. Mamm., 16(3): 205-

207.
HosoKAWA, H., 1951. On the pelvic cartilages of the Balaenoptera-foetuse~, with remarks on the specificaland sexual difference. Sci. Rep. Whales Res. Inst., 5: 5-15.
KASUYA, T., and T. IcBIHARA, 1965. Some informations on minke whales from the Antarctic. Sci. Rep.
Whales Res. Inst., 19: 37-43.
MoORE, J.C., and R. S. PALMER, 1955. More piked whales from southern North Atlantic. ]. Mamm.,

36(3): 429-433.
0HSUMI, S., Y. MASAKI, and A. KAWAMURA, 1970. Stock of the Antarctic minke whale. Sci. Rep. Whales
Res. Inst., 22: 75-125.
OMURA, H., 1957. Osteological study of the little piked whale from the coast of Japan. Sci. Rep. Whales
Res. Inst., 12: 1-21.
OMURA, H., 1971. A comparison of the size of vertebrae among some species of the baleen whales withspecial reference to whale movements. Sci. Rep. Whales Res. Inst., 23: 61-69.
OMURA, H., 1972. An osteological study of the Cuvier's beaked whale, :(iphius cavirostris, in the northwest
Pacific. Sci. Rep. Whales Res. Inst., 24: 1-34.

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

OSTEOLOGY OF MINKE WHALE

29

OMURA, H., and H. SAKIURA, 1956. Studies on the little piked whale from the coast of Japan. Sci. Rep.
Whales Res. Inst., 11: 1-37.
OMURA, H., M. NISHIWAKI, and T. KASUYA, 1971. Further studies on two skeletons of the black right whalein the North Pacific. Sci. Rep. Whales Res. Inst., 23: 71-81.
SATAKE, Y., and H. OMURA, 1974. A taxonomic study of the minke whale in the Antarctic by means ofhyoid bone. Sci. Rep. Whales Res. Inst., 26: 15-24.
SCAMMON, C. M., 1873. On a new species of Balaenoptera. Proc. Cal. Acad. Sci., 4(20): 269-270 (Printed
in advance, 1872).
SCAMMON, C. M., 1874. The marine mammals of the north-western coast of North America, described and illustrated,
together with an account of the American whalejishery. San Francisco. 319 pp.
SERGEANT, D. E., 1963. Minke whales, Balaenoptera acutorostrata Lacepede, of the \'\lestern North Atlantic.
]. Fish. Res. Bd. Canada, 20(6): 1489-1504.
TAYLOR, R.J.F., 1957. An unusual record of three species of whale being restricted to pools in Antarctic
sea-ice. Proc. ,Zool. Soc. Land; 129: 325-331.
ToMILIN, A. G., 1967. Cetacea. Mammals of the U.S.S.R. and adjacent countries. IX. Israel Programfor Scientific Translations. Jerusalem. 717 pp.
TRUE, F. W., 1904. The whalebone whales of the western North Atlantic compared with those occurring in European
waters with some observations on the species of the North Pacific. Smithsonian Contributions to Knowledge.
XXXIII, 332 pp.
TURNER, WM., 1892. The lesser rorqual (Balaenoptera rostrata) in the Scottish seas, with observations on its
anatomy. Proc. R. Soc. Edinburgh, Sess: 36-75.
UTRECHT, W. L. VAN, and VAN DER SPOEL, 1962. Observation on a minke whale (Mammalia, Cetacea)
from the Antarctic. ,Zeit. Siiugetier, 27(4): 217-221.
WILLIAMSON, G. R., 1959. Three unusual rorqual whales from the Antarctic. Proc. ,Zoo{. Soc. (London),133: 135-144.
WILLIAMSON, G. R., 1961. Two kinds of minke whale in the Antarctic. Norsk Hvaifangst-Tid., 50(4): 133141.

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

30

OMURA
APPENDIX.
1.

MEASUREMENTS OF VERTEBRAE OF THE MINKE WHALE FROM
THE ANTARCTIC AND NORTH PACIFIC (in mm.)

SPECIMEN 71]2793 FROM THE ANTARCTIC
Serialno.

Vertebralno.

Greatest
breadthGreatestheightCentrum
Breadth
(a)

c

383

222

207

220

182

2

2

527

3
4
5
6
7

3
4
5
6
7

454
461
454
441
455
479
470
469
527
608
699
758
791
801
804
790
781
736

8
9
10
11
12
13
14
15
16
17
18
19
20
21

D
2
3
4
5
6
7
8
9

10
11
12
L

2

22

3

23
24
25
26
27
28
29
30
31
32

4
5
6

33

34
35
36
37
38
39
40
41
42

7
8
9
10

Ca
2
3
4
5
6
7
8
9
10
11
12

13

176
173
185
211
228
240
273
332
358
381
408
424
438
435
445
455
474
492
493
730
723
519
698
511
678
501
646
525
525
639
618
555
601
550
585
550
521
549
482
518+
445
515
384
403+
326
388
279
330
231
265
T.P. disappear
252
216
189
143
S.P. disappear

157
149
143
141
142
149
166
158
159
162
161
165
161
165
166
168
168
172

176
179
182
186
187
190
194
196
202
209
208
207
212
234
219
216
190
165
155
137
124
109

HeightLength(b)(c)

R. 131(axbxc)113{L. 133
R. 125{L. 126

81

130

29

76

106
112
115
118
118
118
117
117
119
119
124
122
121
122
123
125
130
141
143
145
148
152
157
159
163
165
167
170
180
182
181
193
186
182
177
181
172
133
109
96

30
30
32
35
43
53
68
88
100
111
121
128
135
140
142
149
154
153
161
167
170
173
175
181
190
195
199
203

79
79
81
84
87
98
110
118
124
129
134
137
138
141
143
146
150
155
162
163
166
170

212

215
213
210
204
198
184
170

130
88
74
69

173

176
183
185
189
193
199
201
201
213

203
198
184
172
151
117
100
90
Continued ...

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

31

OSTEOLOGY OF MINKE WHALE
APPENDIX.

Continued.
Centrum

2.

Serialno.

Vertebral
no.

43
44
45
46
47
48
49

14
15
16
17
18
19
20

Greatest
breadthGreatestheight(axbxc)li3

Breadth(a)Height(b)

Length
(c)

94
81
65
51
40
35
18

83
70
54
42
31
21
15

62
54
45
36
30
25
17

67
54
43
33
26
17

R. 146{L. 137
115
121
123
124
132
132
132
132
137
140
140
142
142
142
147
149
151
165
167
167

85

141

39
34
34
36
39
48
56
70

99
92
90
92
96
103
109
119
134
141
144
147
151
155
161
162
165
170
173
178
183
186
189
193
194
197
202
210
218
223
225
225
241
236
233
220

77

SPECIMEN 71]2883 FROM THE ANTARCTIC.

c
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

2
3

4
5
6
7
D

2

3
4
5
6

7
8
9

10
L

11
1
2
3
4
5
6

7
8
9

10
11

12
Ca

1
2
3
4
5
6

7

386

247

227

677
476
559
511
541
546
575
579
538
621
705
798
855
896
918
924
918
786
798
755+
767
760
634+

260
217
206
221+
263
266
258
324

211
187
173

723

705
665+
575+
564+
612
561+
534
512
468
400+
332
266

373

408+
429
445
462
468
480
486
502
516
533
565
575
511+
589
624
622
623
483+
451+
616
553+
598
561
424+
435+
350+
325+

172

170
171
174
181
182
183

178
174
173
178
187
183
185
183
186
193
197
201
202
203
207
212
217
224
234
239
234
234
253
268
266
244

172

173
179
186
179
178
183
188
194
199
205
204
236
208
206
200

96

109
120
129
140
148
151
156
160
164
167
176
180
186
187
191
198
202
209
220
229
234
239
239
235
237
231
219

Continued ...

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

32

OMURA
APPENDIX.

3.

Serialno.

Vertebral·no.

Greatest
breadthGreatestheight

38
39
40
41
42
43
44
45
46
47
48
49
50

8

T.P. disappear

290
254
220
S.P. disappear

9

10
11
12
13
14
15
16
17
18
19
20

Continued.
Centrum
BreadthHeightLength(axbxc)113

(a)

(b)

(c)

219
187
171
153
145
117
100
70
68
57
45
47
24

195
201
199
174
127
112
98
81
66
52
35
22
14

203
186
157
107
88
80
75
66
54
46
38
30
18

205
191
175
142
117
102
90

R. 108{L. 98
76
79
81
83
85
85
85
82
82
83
84
83
81
82
85
85
87
91
94
95
98
101+
106
105
109
110
113
115
118
123

41

90

72

62
51
39
31
18

SPECIMEN AY69B FROM THE NORTH PACIFIC.

c
2

2
3

3
4
5

4
5
6

6

7

7

8

D

9

10
11
12
13
14
15
16
17
18
19
20
21
22
23
24

2

3
4
5
6

7
8
9

10
11
L
2
3
4
5
6
7
8
9

25

26
27
28
29
30
31

10
11
12
Ca

272

176

167

364
319
307
320
314
336
366
343
345
382
420
466
505
538
559
573

188
147
138
156
174
185
193
232
272
288
308
320
330
340
344
360

152
122
119
117
116
117
122
117
121
116
116
116
116
118
129
121
120
124
124
128
129
130
133
132
133
135
141
141
147
148

572

377

558
564
562
582
546
548
533
525
est. 490
est. 462
429
392
est. 352

388
400
414
421
447
445
440
422
419
425
415
392

46
23
29
25
29
30
38
49
69
78
84
91
96
102
106
109
113
117
120
124
128
131
134
137
141
145
153
156
160
161

81
61
65
62
66
67
73
78
88
91
94
96
97
100
105
104
106
110
112
115
119
120
124
124
127
129
135
136
141
143
Continued ...

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

OSTEOLOGY OF MINKE WHALE
APPENDIX.

4.

Serialno.

Vertebralno.

Greatest
breadthGreatestheight

32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48

2
3
4
5

325
W2
255
221
178
146
T.P. disappear

366
TI7
286
252
225
200
175
147
108
S.P. disappear

6

7
8
9

10
11
12
13
14
15
16
17
18

33

Continued.
Centrum
BreadthHeightLength(axbxc)113

(a)

(b)

(c)

145
151
157
157
153
144
126
117
101
87
74
67
54
43
35
25
20

123
126
129
128
126
128
127
121
100
83
68
58
.52
41
30
22
16

162
161
157
154
149
142
129
105
72
62
55
51
45
39
31
27
18

142
145
147
146
142
138
127
114
90

34

83

47
24
24
26
26
28
34
48
63

83
60
60
61
62
63
67
76
83
85
87
88
91
96
95
98
98
101
103
105
108
110
112

77

65
58
50
41
32
25
18

SPECIMEN AY69A FROM THE NORTH PACIFIC.

c
2
3
4
5
6
7
8

260
2
3
4

5
6
7
D

9

2

10
11
12
13
14
15
16
17
18
19
20
21

3

22

23
24
25
26
27

4

5
6
7
8
9

10
11
L

1
2
3
4
5

6
7
8
9

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

305
286
288
285
293
306
312
308
321
368
404
444
466
494
502
496
480
est. 460
460
460
452
449

441
429
414
390

170

159

169
148
149
163
174
175
180+
204
235
248
254
264
273
283
290
297
303
314
320
326
335
292+
353
350
350
349

152
115
111
108
109
110
114
119
119

115
112
110
110
116
113
113
112
114
115

116

117
117
122
121
124
130

R. 107{L. 107
81
80
81
82
83
82
79
78
77

75
75
74
76
80
78
84
82
86
89
92
95
98
99
99

102
103

72

78
85
89
94
96
100
103
105
108
110
113
115

117
119
122
127

113

116
119
Continued ...

OMURA

34

APPENDIX.

Continued.
Centrum

Serial
no.

28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48

Vertebralno.

Greatest
breadthGreatestheight

JO

377
361
318
285
265
230
197
173
144
120

343
333
329
304
275
250
229
206
179
154
132

II
Ca I

2
3
4
5
6
7
8
9
10
II

12
13
14
15
16
17
18
19

T.P. disappear

S.P. disappear

Breadth(a)Height(b)

Length
(c)

131
133
132
129
131
135
138
140
135
119
103
94
82
68
65
59
51
39
23
17
10

108
112
115
115
115
115
117
116
114
114
115
103
82
69
58
50
41
31
23
14
9

131
135
138
140
144
140
136
133
129
120
108
80
61
53
49
45
38
31
21
16
8

(axbxc)l/3

123
126
128
128
129
130
130
129
126
118
109
92
74
63
57
51
43
33
22
16
9

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

OSTEOLOGY OF MINKE WHALE

35

EXPLANATION OF PLATESPLATE I
Dorsal view of skull of the minke whale from the Antarctic and North Pacific.
Fig. I. Specimen 71]2793 from the Antarctic.
Fig. 2.
71]2883
AY69B,, North Pacific.
Fig. 3.
PLATE II
Ventral view of skull of the minke whale from the Antarctic and North Pacific.
Fig. I. Specimen 71]2793 from the Antarctic.
Fig. 2.
71]2883
Fig. 3.
AY69B,, North Pacific.
PLATE III
Lateral view of skull of the minke whale from the Antarctic and North Pacific.
Fig. 1. Specimen 71]2793 from the Antarctic.
Fig. 2.
71]2883
AY69B,, North Pacific.
Fig. 3.
PLATE IV
Posterior view of skull of the minke whale from the Antarctic and North Pacific.
Fig. I. Specimen 71]2793 from the Antarctic.
Fig. 2.
71]2883
AY69B
North Pacific.
Fig. 3.
PLATE V
Dorsal view of mandibles of the minke whale from the Antarctic and North Pacific.
Fig. I. Specimen 71]2793 from the Antarctic.
Fig. 2.
71]2883
AY69B,, North Pacific.
Fig. 3.
PLATE VI
Anterior view of cervical vertebrae of the minke whale from the Antarctic and North
Pacific.
Fig. I. Specimen 71]2793 from the Antarctic.
Fig. 2.
71]2883
Fig. 3.
AY69B,, North Pacific.
PLATE VII
Lateral view of vertebrae of the minke whale from the Antarctic.
Fig. 1. Cervical and dorsal vertebrae.
Fig. 2. Lumbar vertebrae.
Fig. 3. Caudal vertebrae.

Specimen 71]2793.

PLATE VIII
Lateral view of vertebrae of the minke whale from the Antarctic.
Fig. I. Cervical and dorsal vertebrae.
Fig. 2. Lumbar vertebrae.
Fig. 3. Caudal vertebrae.

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

Specimen 71]2883.

36

OMURA
PLATE IX
Lateral view of vertebrae of the minke whale from the North Pacific.
AY69B.
Fig. 1. Cervical and dorsal vertebrae.
Fig. 2. Lumbar vertebrae.
Fig. 3. Caudal vertebrae.

Specimen

PLATE X
Ribs of the minke whale
Fig. 1. Specimen 71J2793.
Fig. 2.
71J2883.

from

the

Antarctic.

PLATE XI
Ribs and sternum of the minke whale.
Fig. 1. Ribs of specimen AY69B from the North Pacific.
Fig. 2. Sternum of specimen 71J2793 from the Antarctic.
Fig. 3. Sternum of specimen 71J2883 from the Antarctic.
Fig. 4. Sternum of specimen AY69B from the North Pacific.
PLATE XII
Scapullae of the minke whale from the Antarctic and North Pacific.
Fig. 1. Specimen 71J2793 from the Antarctic.
Fig. 2.
71J2883,,
Fig. 3.
AY69B,, North Pacific.
PLATE XIII
Humerus, radius, and ulna of the minke whale from the Antarctic and North Pacific.
Right side.
Fig. 1. Specimen 71J2793 from the Antarctic.
Fig. 2.
71J2883
Fig. 3.
AY69B,, North Pacific.
PLATE XIV
Humerus, radius, and ulna of the minke whale from the Antarctic and North Pacific.
Left side.
Fig. 1. Specimen 71J2793 from the Antarctic.
Fig. 2.
71J2883
AY69B,, North Pacific.
Fig. 3.
PLATE XV
Carpals and phalanges of the minke whale from the Antarctic and North Pacific.
Right side.
Fig. 1. Specimen 71J2793 from the Antarctic.
Fig. 2.
71J2883
Fig. 3,
AY69B,, North Pacific.
PLATE XVI
Carpals and phalanges of the minke whale from the Antarctic and North Pacific.
Left side.
Fig. 1. Specimen 71J2793 from the Antarctic.
Fig. 2.
71J2883
Fig. 3.
AY69B,, North Pacific.

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

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No.2λ1975.

MINKE WHALES OFF BRAZILGORDON R. WILLIAMSONHeather Cottage, Kessock, Inverness, Scotland, Britain

ABSTRACT

Minke whales are the main catch off Costinha 7°8 at the NE tip of Brazil.
The whales congregate each September-October-November over the continental slope in water 27°C. About 700 whales are caught per year. All are of
the southern hemisphere subspecies Balaerwptera acutorostrata bonaerensis.
Caught whales are lengths 7.5-10.7 metres (average 8.5 m) which correspond to ages 7-50 (average 15 years old). All the males and about 90%of the females are mature. In the catch females outnumber males 2 ~~: 1 (j'.
No foetuses of visible size occur. Lactating females with calf are seldom seen.
Available data do not permit a firm conclusion regarding the length of
the reproductive cycle. Some data suggest a one-year cycle, some data suggest a two-year cycle. Calves appear to be born in waters of intermediate
temperature and intermediate latitude.
Minke whales are common off Brazil and Durban yet rare off west south
Africa. Perhaps they migrate north mainly up the western side of the South
Atlantic, unlike sei whales which migrate up both sides.
Suggestions for future research are made.

INTRODUCTION

Tropical whaling station winter catches are of special interest to biologists because
they reveal what whales are doing during the reason in which mating and birthoccur.
7°S from the equator is the whaling station of Costinha, at the NE tip ofBrazil, near Joao Pessoa and 140 km north of Recife (Fig. 1). Brazil has been amember of the International Whaling Commission during 1951-66 and since 1973
and biological data on each whale killed have been supplied to the Bureau of International Whaling Statistics at Sandefjord since 1953 Some basic statistics on
Costinha whale catches are given by Paiva (1961) and Paiva and Grangeiro (1965,1970).
This report gives a general account of the Costinha whaling station and describes the catch of minke whales, which is now the main species caught. The
data from Costinha are of exceptional interest because• the season is the tropical winter• catching occurs during six months unbroken period• the effort is constant; one catcher all the time
• the catching area is the same all season long
• the minke whale stock is in virgin condition.
Sci. Rep. Whales Res. Inst.,

No. 27, 1975, 37-59.

G. R. WILLIAMSON

38

Minke whales are caught in great numbers in the Antarctic and about 32,000
have been caught in the southern hemisphere since minke whale hunting began inearnest about ten years ago (Table 1).

75"w

60"w

45"w

30"w

15"w

o·

15"E

30"E

Fig. 1. Atlantic Ocean showing currents (arrows) and October water temperatures.

MATERIAL USEDThis paper is based on data on whaling operations and whale catches off Costinha
supplied through the kindness of the owners of the station, Compania de Pesca
Norte do Brasil, Mr. Kiyohisa Kanda president and Mr. Issao Ishigami manager.
The catcher gunner Captain Sato and station foreman Mr. Masahiro Saito supplied much information. Mr. Kanda visited Britain to discuss details with me
and two British scientist friends of mine visited the whaling station briefly.
Each whale landed at Costinha has the following particulars recorded• date killed• position killed• length in feet in straight line from snout tip to tail notch·sex• whether foetus of noticeable size is present• stomach contents (in some seasons).
Blubber. The thickness of the blubber on the back of the body just below
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

39

MINKE WHALES OFF BRAZILTABLE 1. CATCHES OF MINKE WHALES IN THE SOUTHERN HEMISPHERE 1950APRIL 1975 (DATA FROM BUREAU INTERNATIONAL WHALING STATISTICS)
Year

BrazilAntarctic*

1950
51
52
53
54
55
56
57
58
59
1960
61
62
63
64
65
66
67
68
69
1970
71
72
73
74
75
Total

Total

Durban

9

9

3

3

42
46
493
103
205
162
2
21
101
7
10
18
605
59
30
44
3054
5745
8000
7000
25759

42
46
493
105
205
162
2
24
146
76
367
512
1158

2

2
44
67
352
488
456
617
701
900
702
650
765

2
5
6
97
112
171
204
135
173
117

5747

1024

788
902
1148
3891
6568
8882
7000
32530

* Antarctic catches are listed as follows : 1961/62 catch is listed under 1962 etc.

DISTANCE FROM COAST
(")

0

)>

en
--{

35km

I

I

50km

I

70{m

lOOkm

I

SURFACECONTINENTAL SHELF

Fig. 2. Distribution of minke whales off Costinha. Dots represent whales. This
diagram shows relative abundance of whales, not actual numbers. (Data from
whaling company.)
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

40

G. R. WILLIAMSON

the dorsal fin was measured to the nearest 0.5 cm on all whales caught between
August 16th-December 20th, 1974 (see Fig 7).
Testis samples. To determine the degree of spermatogenesis in the testes,samples of testis tissue were preserved from three male whales during each monthJune-December of the 1974 season. The samples were airmailed for histologicalexamination to Dr. R. F. Macadam of Raigmore Hospital Pathology Department,
Inverness, Scotland.
Vagina smear samples. To determine the nature of the "white mucus" inthe vagina of female whales, smear slides were made of the contents of the vaginasof three female whales of mature size during each month June-December of the1974 season. The smears were airmailed to Dr. R. F. Macadam for histologicalexamination.
The Bureau of International Whaling Statistics, Sandefjord, Einar Vangstein
director, has supplied data where needed.
Data on minke whales off west south Africa and off Durban have been supplied respectively by the Sierra Fishing Agency manager Mr. RH Shepperd and
Dr. Peter Best.
HYDROGRAPHY
Costinha whaling station is located 750 km (4 70 miles) south of the equator at latitude 7°S on the NE corner of Brazil. The continental shelf is narrow about 30 km
wide, and drops steeply into the abyss (Figs. 1 and 2).
A warm equatorial current approaches the coast from the east and becomes
divided by the land into north-flowing and south-flowing branches.
During the main whaling season September-October-November, sea surface
temperatures over a wide area off Costinha are about 27°C.
Frech bite wounds of the type made by the 40-cm-long whale-biter shark
Isistius are common on whale off Costinha, suggesting this tiny pelagic shark must
be common in the area. (The discovery of the causal agency of these wounds wasmade by Jones (1971).) Large sharks 1.5-2.5 m long are plentiful over the continental shelf within 30 km of the coast and cause much trouble biting chunks outof the dead buoyed whales, but in oceanic water further out from the coast deadwhales are not attacked by large sharks.
HISTORY AND DAILY OPERATIONS AT COSTINHA
Whaling at Costinha started in 1910. Since 1951 inclusive the station has been
operated by Compania de Pesca Norte do Brasil (COPESBRA) with head officein Recife. This is a subsidiary of Nippon Reizo K.K. (Japan Ice Storage Co.)of Tokyo and the key personnel are Japanese.
One antarctic-type catcher Seiho Maru No. 2 (306 tons) is used. It leaves
port about 0400 hours each day, hunts all day at distance of 30-140 km from thecoast, and returns with its catch to port after dark about 2000 hours-midnight. The
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

MINKE WHALES OFF BRAZIL

41

whales are cut up during the night. The meat is salted and dried and is soldlocally in Paraiba state and the oil is sold for leather tanning in south Brazil. The
whaling season is from June to December.
The virgin relative abundance of whale species off Costinha appears from thecatches (Table 2) to have been:minke, sei, humpback and sperm-common
bryde-occasional
fin and blue-very rare
At Costinha, whalers killed the easy-to-kill and large species first and catches
have been dominated in turn by humpback, sei and now finally minke and sperm
whales (Table 3). Sei and bryde's whales were not recorded separately in the catchrecords prior to 1967, but have been separated since then. During 1967-74 thecatch of sei and bryde's totalled 251 and 25 whales respectively, that is, sei were
90 % of the catch of the combined species. The percentage of sei was presumably
higher prior to 1960 before antarctic hunting reduced the sei stocks.
Paiva (1961) and Paiva and Grangeiro (1965, 1970) give data on the 1960/67
whale catches of all the species, mainly recording numbers caught each month,mean lengths each month, sex ratios, sexual condition and stomach contents.
Off Cabo Frio whaling station further south in 23°S more fin whales occur
than off Costinha but otherwise the relative abundance of whale species is similar
at the two places (Table 4 ).
The once-large stock of humpback whales features little in the catch of either
station in recent years due to the fact that it was greatly reduced by hunting in the
1904-1920 period.
TABLE 2.

NUMBER OF WHALES KILLED OFF COSTINHA, BRAZIL 7°S
DURING 1910-74. (DATA FROM TABLE 3)

English Name

Brazilian Name

No Killed
1910-74

Present Status

Blue
Fin
Sei
Bryde
Minke
Humpback
Spermbaleia azul
baleia fin
baleia espardate
baleia cabeca de suenga
baleia minkebaleia preta
cachalote1
3
about 3,600
about
350
5,747
1,542
517

rare
rare
rare
rare
very common
rarecommonMINKE WHALES

Existing knowledge on minke whales· Minke whales are the commonest remaining species of baleen whale in thesouthern hemisphere and are also common in the northern hemisphere. In thesouthern hemisphere the latest estimate of the population is 300,000 minke whales(Ohsumi and Masaki, 1974a).
The taxonomy of minke whales became complicated by the discovery that
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

42

G. R. WILLIAMSONTABLE 3. WHALE CATCHES OFF COSTINHA 1910-74 (1924-49 DATA FROMPAIVA 1961, OTHER YEARS DATA FROM BUREAU INTERNATIONAL WHALING STATISTICS)

Year

1910
11
12
13
14
1915-23
1924
25
26
27
28
1929-46
1947
48
49
1950
51
52
53
54
55
56
57
58
59
1960
61
62
63
64
65
66
67
68
69
1970

Humpback

Bryde*

Minke

Blue

Fin

Sperm

Total

Catch taken but no record kept

102
342
352
317

102
342
352
317
No catch taken, station closed

62
42
32
47
40

62
42
32
47
40
Catches taken but no records kept

11
21
15
24
28
9

8
18
6
14
5
8
10

11
8
10

71

72
73
74
Total

Sei*

1,542

14
10
18
98
151
153
161
183
198
196
115
118
294
500
504
272
253
256
149
72
49
6
58
6
6
56
3
23
2
18
5
6
2
about 3,600 about 350

2

3
2
4
11

2
44
67
352
488
456
617
701
900
702
650
765+
5,747

5
4
7
4
13
24
20
39
75
76
55
66
75
29
517

3

25
32
34
122
179
163
170
202
205
214
117
128
315
511
521
284
272
304
229
448
563
559
754
803
975
774
732
797
11,768* Sei and Bryde's whales were not distinguished prior to 1957, but Sei probably composed 90% of
the catches : see text.
+ Includes 3 minke killed but lost at sea. Anomalous differences in whale totals in many seasonsare caused by circumstances like this: the total killed exceeds the total measured.

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

MINKE WHALES OFF BRAZIL

43

southern hemisphere whales have a different colour pattern from northern hemisphere whales (Williamson, 1959; Ohsumi et al., 1970). In nearly all other respects
the minke whales of the two hemisphere are identical and it is probably reasonable
to consider the whales of the two hemispheres as being no more than separate subspecies with the following names:
Southern Minke Whale Balaenoptera acutorostrata bonaerensis Burmeister, 1867.
Northern Minke Whale Balaenoptera acutorostrata acutorostrata Lacepede, 1804.
Knowledge of the biology of southern minke whales advanced greatly
with the publication of the reports of Ohsumi et al. (1970) and Ohsumi and Masaki(1974a and 1974b). The following summary is based on these reports augmented
by information from the studies of northern minke whales by Omura and Sakiura(1956) andJonsgard (1951).
Southern minke whales are born about 2.8 m long after a gestation period of
about ten months. Calves are suckled on milk for about four months and areweaned when about 4.5 m long. They become sexually mature at about six yearsof age when males are about 7.2 m long and females 8.0 m long. Males in which
the larger of the two testes weighs more than 0.4 g are usually sexually mature.
Minke whales grow to about fifty years maximum age and to maximum lengths of
9.8 m in males and 10. 7 m in females (Table 5).
In the Antarctic, most of the whales are large. Of whales caught 54 % arefemales. About 55 % of the females and 86 % of the males are mature. Of maTABLE 4. WHALES CAUGHT AT CABO FRIO WHALING STATION 23°S NEARRIO DE JANEIRO 1960-63 BY SOCIEDADE DE PESCA TAIYO LIMITADA.
(DATA FROM BUREAU INTERNATIONAL WHALING STATISTICS)
Year

Hum back

Sei and
Bryde

Minke

1960
61
62
63
Total

3
2
3
2
lOt

250
453
338
93
1134tt

.s-5

Blue

"C)

~

Oi 0.

(.) 8

zo~Oi

Fin

Sperm

Total

21
10
49
4
84

28
97
81
35
241

302
562
472
134
1470

t Humpbacks had already been greatly reduced in numbers.
tt Proportion of Sei and Bryde in the combined species catch are not known.

TABLE 5. AVERAGE LENGTH AT SEXUAL MATURITY AND MAXIMUM LENGTH
OF MINKE WHALES. SEXUAL MATURITY OCCURS AT ABOUT 6 YEARS OF AGE.

Hemisphere

Area

Mean Length
at sexual
maturity (m)Maximumlength (m)Author~3Southern Hemisphere

Antarctic""
Northern
HemisphereBrazilNewfoundland
Norway
Japan

"
"

"

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

7.2

6.9
6.9

~
8.0

3

~

9.8

10.0

7.3
7.3

9.8
9.1
8.5
8.8

10. 7
9.1
9.1
9.1

Ohsumi et al. (1970) and
Ohsumi and Masaki (1974a)
Whaling company
Sergeant (1963)
Jonsgard (1951)
Omura and Sakiura (1956)

44

G. R. WILLIAMSON

ture females about 90 % are pregnant, and resting mature females are seldom foundin the Antarctic. Immature whales are most numerous in the Antarctic in theearly summer (November-December).
Ovulation rate is estimated to be 0.89 per year (Ohsumi and Masaki 1974b).
88 % of mature females ovulate in spring August-Deptember when they arestill lactating (P. Best, pers. comm. based on examination of 16 lactating females
killed off Durban by special permit).
Minke whales are believed to bear one calf per year. A wide range of foetus
sizes amongst pregnant females in the Antarctic indicates a long mating season.
The migrations of minke whales are probably an annual oscillation about acentral latitude of perhaps 40°S, in which the magnitude of oscillation increases
steadily as a whale gets older and larger. Calves, most immature whales, and mature females in resting condition keep to the 20°S-50°S latitudes of mild water temperatures. It is the pregnant females, the larger of the immature females and themature males which go far south. Whales caught south of 60°S are of average age
15 years and average length 8.5 m (age range 7-50 years, length range 6-10 m).
Mature whales probably make an annual migration from close on the equatorto the high Antarctic and back, a distance of about 14,500 km (8,000 nautical
miles) round trip. At the southern end of their migration they feed on krill andconcentrate right at the edge of the pack ice in 60°S-70°S latitude. They are soaboundant and concentrated that Japanese catchers could kill more than 20 minke
whales per day if they wished (Ohsumi, pers. comm.).
Minke whales characteristically divide themvelves into groups dominated bya certain sex or age of whale, more so than in other baleen whale species. This
habit makes it very difficult for scientists to obtain the random samples which arerequired in order to make accurate calculations of the population size etc.

Costinha catches of minke whales
All minke whales caught are of the southern hemisphere type of minke whale
Balaenoptera acutorostrata bonaerensis as described by Burmeister (1867), Williamson(1959) and Ohsumi et al. (1970). See Plate I.
Each year the first minke whales arrive from the south in late June and July.
They appear singly about 100 km off the coast over deep ocean water and moveslowly north. Later groups appear. Groups of whales usually contain 4 or 5
individuals, which are sometimes of one sex, sometimes mixed sexes. Less frequentpairs of whales are usually one male with one female. The whales mostly swim
over 2,500-meter-deep water about 10-30 km beyond the steep edge of the continental shelf, about 40-60 km from the coast (Fig. 2).
Maximum numbers of minke whales are present during September-OctoberNovember (Table 6, Fig. 4) and highest catches and highest catches per day are
obtained in October. Whales of progressively larger sizes enter the area eachsuccessive month and they depart in the same sequence (Fig. 4 and Table 7). In
November numbers of whales decrease, groups are no longer seen, only pairs andsingle whales, and the last individuals leave in December.
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

45

MINKE WHALES OFF BRAZIL

The sex ratio among caught whales is 2 ~ ~ 1 : ~ (67 %~ ). The largest individuals caught were several 9.8 m males and several 10. 7 m females.
(Sei whales off Costinha, by contrast, during the years 1950-65 when big
catches were taken, were caught in greatest numbers during August and September
and 40-50 % were females in typical years).
Water temperatures are about 27°C during the September-October-November season of abundance (Fig. 1).
Some minke whales can be seen as far north as the extreme NE tip of Brazil(5°S latitude, 250 km north of Costinha) but the whaling company state that none
turn left along the north coast of South America.

°"'°"'0

100

U1

90
(f)
UJ 80
-'
<i::
:i:: 70
3::
u. 60
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50
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en 40
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LENGTH (METERS)

Fig. 3. Lengths of minke whales in a typical season's catch at Costinha (650 whalescaught in 1973). The arrows show the mean lengths at which sexual maturity is
attained. The males are probably all mature and the females 90% mature.
(Data from Bureau International Whaling Statistics.)

Minke whale catches by year, month and sex are given in Table 6 and length
frequencies of catches in typical seasons are given in Figs. 3 and 4 and Table 7.
Correlation of the length frequencies with the results of Ohsumi et al. (1970) and
Ohsumi and Masaki (1974b) allows the ages and maturity of the catches to be deduced and the following conclusions reached:• at the beginning of the season, small numbers of immature females anda very few immature males are present. Most of these are not shot because
they are too small to the worth shooting. They leave the area by midspring; mid-September.
•mature whales start to arrive in the area in July, and from mid-September
onwards nearly all minke whales in the area are mature.
·of the total catch offemales about 90 %are mature, of the total catch of males
about 100 % are mature.
·the ages of whales in the catch range 7-50 years old with average age 15
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

G. R. WILLIAMSON

46

~

0,.

0
5

~

JULY

~

CJ)

0

f'TI

><
c

)>

r
r

-<

10

-;

5
0

s:
)>

c

:::0
fTI

I

I
I

SEPTEMBER

15

I

10

I
I

5

I

g
<(

I

3

u...
0

I
I

Cf)

0::
IJ.J

15 ~
10 ~

I

OCTOBER
I

I

5
0

I

I
I
I

.

I
I

15

NOVEMBER

I

10
5
0

10
5

DECEMBER

0

8

9

10
8
LENGTH (FEET AND METERS)

9Fig. 4. Length frequencies of each month's catch of minke whales in a typical
Whales of progressively larger
season at Costinha (762 whales caught in 1974).
sizes enter the area each successive month and depart in the same sequence. (Datafrom Bureau International Whaling Statistics.)
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

47

MINKE WHALES OFF BRAZILTABLE 6.

MINKE WHALES KILLED IN EACH MONTH OFF COSTINHA,
BRAZIL 1969-74. (DATA FROM WHALING COMPANY)
No. minke whales killed

June

July

Aug

Sept

Oct

Nov

Dec

year

%Q

Q
Total

5
6

21
28
49

34
54
88

49
88
137

48
102
150

48
105
153

6
28
34

207
410
617

67%Q

3

2
6
8

47
52
99

49
86
135

44
110
154

44
104
148

32
104
136

6
15
21

224
477
701

68%Q

3

6

Q
Total

'O

38
62
100

92
159
251

114
199
313

43*
110*
153*

*
*
*

325
575
900

64%Q

6

32
45
77

3

3
2
5

10
12
22

33
51
84

57
108
165

49
138
187

61
117
178

18
43
61

231
471
702

67%Q

4
2
6

26
62
88

45
99
144

64
146
210

60
108
168

11
23
34

210
440
650

68%Q

4
3
7

31
38
69

42
89
131

53
137
190

55
127
182

44
84
128

17
38
55

246
516
762

68%Q

16
16
32

145
177
322

222
404
626

340
701
1041

374
816
1190

288
628
916

58
147
205

1443
2889
4332

67%Q

3
1969

1970

Q

Total
1971

1972

Q
Total

3
1973

Q
Total

3
1974

Q
Total

6-year
total

3
Q
Total

* Catch stopped due to Brazilian local quota having been reached,
TABLE 7. MEAN LENGTHS OF MINKE WHALES IN EACH MONTH CAUGHT AT
COSTINHA DURING 1966-67. (FROM PAIVA AND GRANGEIRO 1970)

Month
July
AugustSeptember
OctoberNovemberMalesmean length m (no. measured)

Femalesmean length m (no. measured)

1966

1967

1966

1967

7.8 (13)7.9 (11)
8. 2 (25)
8.6 (33)
8. 7 (25)

7 .9 (14)8.0 (21)
8.4 (27)
8.3 (46)
8.4 (40)

8.1 (12)8.3 (17)
8. 3 (64)9.0 (98)
9 .1 (52)

7 .9 (22)
8.5 (39)8.8 (83)
8.8 (133)
8.9 (67)

years old, the same as typical Antarctic catches.
During September-October-November a whitish mucus is found in the vagina
of all female whales, also occasionally in late August and early December. The
lips of the vagina of these females are a reddish colour. None of the female whales
contain a visible embryo in the uterus.
About twelve mothers with calf are seen each year.
The stomachs of nearly all the whales are empty. A little krill is found in thestomachs of about 3 % of the whales.
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

48

G. R. WILLIAMSON

5,747 minke whales have been killed off Costinha so far plus a further 200
whales have been taken by factory ships in the Antarctic region l5°W-70°W which
presumably are from the same stock. So far there are no signs of depletion of theminke whales off Costinha: average lengths of whales and catch per day's hunting
are not decreasing (Table 8).
In August 1972 the catcher made a single voyage 400 km eastward out into
the open ocean. Although the main concentration of minke whales was locatedclose beyond the continental shelf edge, a few minke whales were seen all the way
out as far as the catcher went.
TABLE 8. CATCH OF MINKE WHALES PER DAY'S HUNTING IN PEAKMONTH OF SEASON (i.e. October) AND AVERAGE LENGTH OF SEASONSCATCH OFF COSTINHA 1966-74. (DATA FROM WHALING COMPANY).
average length mYear

No. caught per
day's hunting
in Octobermales

females

1966
67
68
69
1970
71
72
73
74

4.2
5.1
4.3
4.8
4.8
5.0
6.0
6.8
5.9

8.3
8.3
8.3
8.3
8.3
8.2
8.7
8.3
8.2

8.8
8.7
8.7
8.6
8,8
8,8
9.2
8.9
8.6

Blubber thickness results. The method of measuring the blubber thickness
was the same as used by Ohsumi et al. (1970), in order to obtain comparable
results. Due to the fact that off Costinha whales of progressively larger size occur
in successive months, simple averaging of blubber thickness measurements does notproduce a useful result. Therefore I selected for analysis only whales of a certain
length range, and selected the range 8.4-9.3 m (27.5-30.5 feet), as plenty whales ofthis size occurred in all months. About one half of all the whales caught were inthis length range. The blubber thickness of males and females in this length range
are shown in Figs. 5 and 6.
The results allow the following conclusions to be made:• females have slightly thicker blubber than males. This is the opposite to
Ohsumi et al's (1970) findings in the Antarctic, where males had thicker blubber than females .
• the blubber offemales is thicker than the blubber of males by about 0.5 cm.
among whales killed in August and increases steadily to 1.1 cm thicker
among whales killed in December .
• although male whales after September become thinner in successive months,
among females the blubber is thicker in whales killed in successive months.
The whales are not feeding, therefore cannot be getting fatter due to feeding.
The increase in blubber thickness among females is presumably due toSci. Rep. Whales Res. Inst.,
No. 27, 1975.

MINKE WHALES OFF BRAZIL

49

5.0
4.5
-'4.0

'E
~

(/)
(/)

3.5
3.0

w

z
:.::
S2

SEPT

AUG

DEC

NOV

OCT

:r:

I-

a::

w

5.5

::::>

5.0

co
co

....J

co

4.5
-4.0

0

10

NUMBER OF WHALES

Fig. 5. Blubber thickness of whales m length range 8.4-9.3 m (27.5-30.5 feet)caught off Costinha during August 16-December 20th 1974. 118 males and 215
females were in this length range, which is about half of all whales killed during
this period.

5.0

E'

.o----o~.

If)fi] 4.5
z
:.::
(.)
::c

,,.P'......

females off Brazil ,,,,
a" ...

,..df-

ffi 4.0
CJ
CJ

::::>

...J

CJ

3.5

J

M

A

M

J

J

A

s

0

N

D

MONTH

Fig. 6. Mean blubber thickness of southern minke whales in length range 8.4-9.3 m
in different months of the year. August-December data from off Brazil calculated
from Fig. 5, December data from Antarctic (Ohsumi et al 1970, Fig. 4).
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

50

G. R. WILLIAMSON

greater fatness among the new whales which enter the area each month.
•female minke whales caught during August-December off Brazil appear tobe fatter than those caught in the Antarctic during December (Fig. 6.) andmales are of similar fatness. This contrasts with other rorqual species.
However this may be due to accidented differences between my and Ohsumi's technique for measuring the blubber.
Testis results. Dr. R. F. Macadam reports "Testis specimens" from only
seven mature whales caught off Brazil were well preserved and these included
specimens from August, September, October and Novembers. These specimens,
plus testis specimens from five mature whales caught in the Antarctic during
November-December 1971 (kindly supplied by Dr. S. Ohsumi) were examined.
The specimens were fixed in 10% formol saline at the whaling station, andlater embedded in paraffin wax, sectioned 5 microns thick and stained with Haematoxylin and Eosin and also by the Masson trichrome technique. Plate 4 shows
photographs of testis specimens representative of whales caught in Brazil and in the
Antarctic.
Testes from the seven whales caught off Brazil during August-November all
showed many sperm in the tubules and active spermatogenesis in progress. These
whales must have been capable of fertilizing femals. Plate 5 shows a single tubuleand the numerous sperm in it magnified XlOOO.
Testes from the five whales caught in the Antarctic during November-December were not so well preserved as the Brazil specimens. All five whale,s showed alow degree of spermatogenetic activity and only a few sperm."
Vagina smear results. Dr. R. F. Macadam reports "the smears were wellpreserved and consisted of mucus containing many vaginal epithelial cells, many
micro-organisms and no sperm. In humans, sperm in the female vagina disintegrate 2-3 days after copulation. Therefore the absence of sperm in the vaginas ofthese 18 females only indicates that none of them had mated within three days prior
to being caught. They may nonetheless have mated during other days of themonth."

Fig. 7. Position on body at which blubber thickness was measured. The position
fulfils the requirements of being accessible on a hauled-up carcase, on the line of
one of the main flensing cuts and identifiable with accuracy.

West South Africa and Durban catches of minke whales
Off west south Africa, minke whales are uncommon. Gunners operating out
of Saldanha reported that they never saw minke whales there (Williamson, 1959).
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

51

MINKE WHALES OFF BRAZILTABLE 9. LIST OF ALL MINKE WHALES SEEN IN AREA 0°-30°S 0°-17°E OFF
SW AFRICA DURING 2! YEARS ALMOST CONTINUOUS WHALING BY
FACTORY SHIP "SIERRA" NOV 1972-MAY 1975. (ALL MINKEWHALES SEEN WERE KILLED. DATA FROME SIERRAFISHING AGENCY)
LengthSex Sexual Condition
m

Date

Lat

Long

Temp
Sea Surface

March 17, 74
June 11, 73
Nov
8, 73

13°33'S
5°46'S
7°02'S

11°2l'E
10°36'E
11°23'E

23°C
24°C
26°C

9.4
7.3
8.2

M
F
M

Nov

20, 7414°27'S

11°27'E

21°C

8.8

F

Nov

21, 72l6°lO'S

l0°55'E

22°C

9. 7

F

mature?

maturemature, novisible foetus
mature, novisible foetus

StomachContents
empty
empty
empty
empty
some krill

TABLE 10. COMPARISON OF CONDITION OF MINKE WHALES OFF COSTINHA
7°S, DURBAN 30°S AND ANTARCTIC OCEAN 55°-65°S. (COSTINHA DATA FROMWHALING COMPANY, DURBAN DATA FROM P. BEST PERS. COMM.,
AND ANTARCTIC DATA FROM OHSUMI ET AL 1970,
AND OHSUMI AND MASAKI 1974b).
Durban S Africa
30°S

Antarctic Ocean
55°S-65°S

all months somepresent
20-25°C

Dec-Jan-Feb

70

97

mature and immature present (seetext)
small numbers during August-September
?

0

0

90

about 60%

unknown

67% Q

30% Q

3

?

0.27 m (December)Great variations indifferent areas
54% Q
but great variations indifferent areas
55

Many small groups
and a few single
whales. Most groups
are of 4-5 whales.

Small groups and a
few single whales.
Some lactating mothers with young
calf during AugustSeptember.

Many groups of 2-6
whales, a few single
whales and a few
groups of up to 100
whales. Groups usually
dominated by one sex.

Costinha, Brazil

1°sMonths in which minke
are most abundant
Temperature of sea surface
during season of abundance% of all whales caughtthat are sexually mature

Sept-Oct-Nov% of mature females that3

are accompanied bycalves% of mature females thathave no calf and novisible foetus% of mature females thatare pregnant with visible
foetus and have no calf
Mean length of foetuses% female among whales
at sea% female among killed
whales% stomachs

containing
foodGrouping of whales at sea

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

27°S
95

0°C

0

7

52

G. R. WILLIAMSON

Mr. R.H. Shepperd reports that the factory ship "Sierra" during 2! years almost
continuous whaling Nov 1972-May 1975 in the area 0°S-30°S, 0°-l 7°E off west
south Africa has only sighted five minke whales (Table 9). They were all shot andproved to be mature animals.
During the same period the " Sierra " sighted several thousand sei whales.
Off Durban 30°S (Table 10) Dr. Peter Best reports that minke whales arepresent in moderate numbers all year round in waters which range 20°-25°C annually. They appear to be most numerous during winter and spring (JuneNovember). The annual catch of about 150 minke is a mixture of sexually matureand immature animals. During August-September in sea of temperature about
20°C some lactating mothers with young calves are seen each year.
DISCUSSION

Separation of north and south atlantic stocks.

South Atlantic minke whales migrate as
far north as NE Brazil 5°S and on the African side odd individuals migrate to off
Angola with 5°S the extreme northernmost record to date (Table 9). The nearest
localities from which north atlantic minke whales have been recorded are Florida
and the Mediterranean (Moore and Palmer, 1955; Scattergood, 1949). About
5,000 km separates the nearest records, thus it seems unlikely that members of thenorth and south hemisphere stocks ever meet.
Route of northward migration. By what routes do whales travel from the Antarctic
to NE Brazil? After the feeding season, most whales all round the Antarctic headnorth. In wide areas of the Indian and Pacific Oceans there is no land and thewhales must meet and mate successfully in mid ocean. In the western South
Atlantic most whales, after a northward swim of varying length seem likely to strikethe sloping coast of South America at some point. The slope of the coast "funnels"
the whales together and causes them to become concentrated as they proceed northwards. For those whales which continue furthest into warm waters, the NE
corner of Brazil is the focal point where maximum concentration occurs.
It seems probable that most minke and sei whales which arrive off Costinha
have followed the continental shelf edge from varying distances further south, whilea smaller number have arrived direct from the open ocean. The minke whales off
Costinha concentrate about 20 km beyond the edge of the continental shelf (Fig.
2). Further out to sea and on the shallow continental shelf there are only a few
minke whales.
The low-frequency noises made by rorquals can be heard at distances of atleast 180 km (Cummings and Thompson, 1971, p. 1197) and thus minke whaleson the open ocean, when they get within this range of the continental slope, probably
hear and are attracted to the concentration of whales.
The virtual absence of minke whales off west south Africa, and yet their presence off the eastern southern Africa (Durban) and eastern south America,
suggests an assymmetry in the autumn northward migration from the Antarctic.
It appears that minke whales migrate chiefly up the western side of the southern
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

MINKE WHALES OFF BRAZIL

53

atlantic, heading into the south-flowing Brazil current, but keep clear of the northftowing Benguela current.
If this is the case, the coast of Brazil may be the mating ground of minke
whales from a wide sector of the Antarctic.
Sei whales in contrast migrate up both sides of the South Atlantic.
Length of the reproductive cycle. Studies to date conclude that minke whales bear one
calf each year and have a one-year reproductive cycle; thus reproduce twice as fast
as all the larger rorqual species.
In truth, available data are not conclusive on this matter. Some facts suggest
a one-year cycle, some facts suggest a two-year cycle.
Evidence which suggests a one-year cycle:• about 88 % of mature female minke whales ovulate during the latter partof the lactation period, in August-September (see section ' Existing knowledge on minke whales'). This makes it possible for minke whales to become pregnant during late lactation, thus producing one calf per year.
Overlapping of lactation with pregnancy does not occur in the larger
rorquals, all of which have a resting period of several months between theend of lactation and mating (Fig. 5 ).
Evidence which suggests a two-year cycle:• The 27 cm average length of foetuses in December Antarctic catches (Table10) indicates that mating must occur in about September (compare Omura
and Sakiura, 1956, Fig. 23). A ten-month pregnancy would result incalves being born in July and the four month lactation would last till
November. However, during September-October-November, hundreds ofcalf-less mature females arrive off Costinha. These females cannot haveborn a calf recently, therefore it seems impossible that they are producing
one calf per year.
• The 0.89 per year ovulation rate of minke whales is basically similar to,although somewhat higher than, the ovulation rates of the other rorqual
species (Table 12). (It must be remembered that all these rates are estimates
only). If minke whales really reproduced twice as fast as other rorquals,
their ovulation rate should be double that of other species-about 1.4
ovulations per year. But it is not near this value.
The 0.89 per year ovulation rate of minke whales could not allow all thefemales of the population to bear a calf every year but it is nonetheless compatible with the existence of a basic one-year cycle. An individual female
during a ten year period could have 8 calves at one year intervals and one
calf after a two year interval (assuming all ovulations produce calves).
As pregnancy and lactation total 14 months duration, a one-year cycle
can only exist if new pregnancy occurs two months before the end of lactation, that is; the cycles overlap by two months.
The fact that in the Antarctic 90% of mature female minke whales
are pregnant, compared with 66% among blues, 80% among fine and
69% among sei (Table 11) does not clarify the reproduction rate. It is
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

54

G. R. WILLIAMSON

compatible either with minke whales having a higher reproductive rate
than the other species (though hardly double the rate of the other species)
or with minke whales having a two-year cycle in which resting mature
females have the habit of seldom entering the Antarctic.
By examination of the ovaries of mature females caught off Costinha itshould be possible to obtain the vital missing evidence.
If these calf-less whales are only just ovulating or have not yet ovulated,
they must have passed a true resting stage since parting from their calf andthus must have a two-year reproductive cycle.
If the whales have already ovulated (and in this case they will presumably be in early pregnancy) then it is likely that they ovulated and matedpuring lactation, and this overlap would allow a one-year reproductive cycle.
Further data are needed to clarify the reproductive cycle of minke
whales.
TABLE 11. COMPARATIVE DATA ON SOUTHERN HEMISPHERE RORQUAL
SPECIES DURING THE COOL-WATER SUMMER FEEDING SEASON
Species

greatest numbersof adults
furthest south% femalesin catch% mature in catch % of mature...-----r----..

66

22

females thatare pregnant

timing of migrationof immatures

Blue
December1
47 1
66 1
later than matures2
Finlater than matures 2
January 1
50 1
80 1
Sei
February 3
94 4
92 4
69 3
54 5
866
55 6
905
Minke
?
earlier than matures 5
Sources of data. 1 =Mackintosh 1942, p 270, Table 25 and Fig. 27.
2=Laws 1961, p 341.
3=Gambell 1968 p 39-40 and Table 5.
4=Doi et al 1967, table 14.
5=0hsumi and Masaki 1974b.
6=0hsumi et al 1970 p 107.
TABLE 12.
SpeciesOVULATION RATES PER YEAR OF RORQUAL WHALES

average numberof ovulations
per year

evidence used to calculate th valueauthorAverage of the average number of recent
Laurie in Purves and Mountovulations in several hundred antarctic
ford 1959 p 139
whales of three sexual classes (pregnant,based on Laurie 1937, p 250
resting and recently-ovulated non-pregnant).
Average of ovulation rates from threeFin
0. 72tLaws 1961, p 463
types of 2-year breeding cycles, found bycounting corpora albicantia in several
hundred antarctic whales.
Average rate in 6 marked antarctic
0. 77
Ohsumi 1964, p 66
whales captured 19 or more years later,
found by counting corpora albicantia.
No of oestrous cycles in 2-year breeding
Sei
0.69
Gambell 1968, p 123
cycle x average number of ovulations per
oestrous, found by counting corpora
albicantia in 644 whales caught off
Durban.
Minke
0.87
No of corpora albicantia in 1359 antarcOhsumi and Masaki 1974,
tic whales of known age
p9
t value halved from original 1.43 consequent to Roe's (1967) discovery that ear plugs of southernhemisphere fin whales accumulate only one (not two) growth layers per year-see Gambell(1968) p 124.
Blue

0.69

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

MINKE WHALES OFF BRAZIL

55

FIN
SEI

MJJASONDJFMAMJJASONDJFMAM
MONTH

Fig. 8. Reproductive cycles of rorqual whales: comparison of timing of events.
(Blue and Fin data from Mackintosh and Wheeler 1929 and Laws 1959, Sei datafrom Gambell 1968, Minke data from Omura and Sakiura 1956 and present report).

Location, season and temperature ofgiving birth. The presence of calves with their mothers
in 20°C water off Durban 30°S, but seldom in 27°C water off Costinha 7°S suggeststhat minke whales give birth in relatively cool and southerly waters and do not
usually, bring their calves into the warmest waters. This is interesting because
scientists traditionally believe that all whales bear their calves in the warmest partof their range. In minke whales it is chiefly the mature adults which enter the truly
hot tropical waters. What is the purpose of this?
Newly weaned minke calves offJapan are not found in water colder than about
l5°C (author's data) thus presumably suckling occurs in waters of about 20°-15°C
and females do not enter waters cooler than l5°C until after parting from theircalf.
What catch of minke whales can be sustained indefi,nitely off Costinha? If the southern hemisphere population estimate of300,000 minke whales is correct, and if they are equallydistributed around the longitudes, then about 45,000 are living in the l5°W700W western part of the South Atlantic and about 75,000 in the whole South
Atlantic.
The annual yield that this population could sustain cannot be determined atpresent but it will have to be shared between Costinha and the factory ships whichhunt the same stock in the Antarctic.
Future research
1 At Costinha a biologist should spend a season and:• examine the uteri of all females to check whether tiny embryos are present
or absent• examine ovaries of all females to determine presence or absence of corpora
lutea• take samples of testes in each month to determine state of spermatogenic
activity• take samples from every female of the vagina contents, examine them for
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

56

2

3

G. R. WILLIAMSON

sperm and thus discover whether any of the whales are actually mating.
Whale-searching cruises and tagging should be used to investigate:• is there a belt of minke whales stretching between Brazil and Africa during
the winter season?
• what numbers of minke whales are present in winter in mid-ocean areas,
off the west coast of S. America and off the east and west coasts of Australia?
Do northward migrating minke keep to the west warm-current sides of thesouthern oceans?
• in what temperature and latitudes does mating occur?
• in what temperatures and latiudes are calves born?
A factory-ship should be asked to catch samples of minke whales at all latitudes
on its way to and from the Antarctic.
SUMMARY OF NEW KNOWLEDGE ABOUT MINKE WHALESThe data from Costinha reveal the following new facts• immature whales precede mature whales in the migration to the north(Fig. 4).
• as whales get older and larger they migrate north later in the season (Fig. 4,
Table 7). The main concentration occurs during September-OctoberNovember.
• the 2 ~~: 1 ()' sex ratio among caught whales at Costinha, even making allowance for gunners selecting large and therefore female whales preferentially,suggests that there is a real imbalance of the sex ratio off Costinha and thatmany of the smaller mature males do not migrate so far north• mature whales go much further north to 7°S and into warmer water 27°C
than do immature whales• calves are born and suckled somewhere to the south of and in less warm
temperatures than the waters in which the adults congregate off Costinha• immature whales precede mature whales in the migration to the feeding
grounds (whereas in blue and fin whales the mature animals move south
before the immatures; see Table 11 ).
• the simultaneous presence during November-December of considerable
numbers of minke whales both off Brazil and in the Antarctic 7,000 km to
the south indicates either or both of the following: that there is great variation in timing of migration between different individual adult whales and/or
separate high latitude and low latitude stocks of minke whales exist. The
results of analysis of blubber thickness in different months also support thisconclusion.
ACKNOWLEDGMENTS

The bulk of the new data presented in this paper was supplied by the Companiade Pesca Norte do Brasil, operators of the Costinha Whaling Station: Senor KiyoSci. Rep. Whales Res. Inst.,
No. 27, 1975.

MINKE WHALES OFF BRAZIL

57

hisa Kanda, President, Senor Issao lshigami, Manager, Senor Masahiro Saito, Factory Manager and catcher boat Captain A. Sato. Visits to the whaling station weremade by my friends Dr. Barry Cox, Dr. William Ledingham Dr. Jim Owen and
Mr. Tony Hall, helped by the British Council.
Data on minke whale catches in other areas and valuable criticisms of mypaper were given by Dr. Seiji Ohsumi of Japan (Antarctic catches), Dr. Peter Bestof S. Africa (Durban catches) and Mr. R.H. Shepperd of Sierra Fishing Agency(West South Africa catches).
Mr. Sidney Brown, Mr. Einar Vangstein, Dr. Melquiades Paiva, Dr. Soloncy
de Moura and Dr. David Sergeant helped me with data and criticism.
Dr. R. F. Macadam of Raigmore Hospital Pathology Department, Inverness,
Scotland carried out the histological examinations of the vagina smear and testis
samples.
Larry Foster of General Whale, California made the beautiful drawing of thewhale (Plate I).
REFFERENCES
BURMEISTER, H., 1867. Preliminary description of a new species of Finner whale (Balaenoptera bonaerensis).
Proc. zoo!. Soc. Lond., 1867: 707-713.
CUMMINGS, W. C. and P. 0. THOMPSON, 1971. Underwater sounds from the blue whale Balaenoptera
musculus. J. Acoustical Soc. America, 50 (4, part 2): 1193-1198.
Dm, T., S. 0HSUMI and T. NEMOTO, 1967. Population assessment of Sei whales in the Antarctic. Norsk
Hvalfangst-Tid., 56(2): 25-41.
GAMBELL, R., 1968. Seasonal cycles and reproduction in sei whales of the southern hemisphere. Discovery
Rep., 35: 31-134.
HARRISON, R. G., 1969. Reproduction and reproductive organs in Andersen; H. T. (editor), The biology ef
marine mammals. Academic press, New Tork and London, 511 pp.
]ONES, E. C., 1971. Isistius brasiliensis, a squaloid shark the probable cause of crater wounds on fishes andcetaceans. US Nat. Mar. Fish. Serv. Fish. Bull. 69(4): 791-798.
]ONSGARD, A., 1951. Studies on the little piked whale or minke whale, Balaenoptera acutorostrata Lacepede.
Norsk Hvalfangst-Tid., 40(5): 209-232.
LAURIE, A. H., 1937. The age offemale blue whales and the effect of whaling on the stock. Discovery Rep.
15: 223-284.
LAws, R. M., 1959. The foetal growth rates of whales with special reference to the fin whale Balaenoptera
physalus Linn. Ibid., 29: 281-308.
LAws, R. M., 1961. Reproduction, growth and age of southern fin whales. Ibid., 31: 327-486.
MACKINTOSH, N. A., 1942. The southern stocks of whalebone whales. Ibid., 22: 197-300.
MACKINTOSH, N. A., andJ.F.G. WHEELER, 1929. Southern blue and fin whales. Ibid., 1: 257-540.
MooRE, J.C. and R. S. PALMER, 1955. More piked whales from southern North Atlantic. J. Mamm.,
36(3): 429-433.
0HSUMI, S., 1964. Examination on age determination of the fin whales. Sci. Rep. Whales Res. Inst., 18: 4988.
0HSUMI, S., and Y. MASAKI, 1974a. Status of whale stocks in the Antarctic, 1972-73. 24th Rep. Intern.
Whaling Comm.: 102-113.
0HSUMI, S., and Y. MASAKI, 1974b. Biological parameters of the antarctic minke whale in virginal population level. IWC Small Cetacean Meeting at Montreal, report Ml 7 pp 1-12 (mimeo ).
0HSUMI, S., Y., MASAKI and A. KAWAMURA, 1970. Stock of the antarctic minke whale. Sci. Rep. Whales
Res. Inst., 22: 75-125.
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

58

G. R. WILLIAMSON

Omura, H., and H. SAKIURA, 1956. Studies on the little piked whale from the coast of Japan. Ibid., 11:
1-39.
PAIVA, M. P., 1961. Recursos basicos da pesca maritima no nordeste Brasiliero. Bol. Est. Biol. Mar. Univ.
Cceara, 3: !Opp (in Portuguese).
PAIVA, M. P., and B. F. GRANGEIRO, 1965, Biological investigations on the whaling seasons 1960/63 off northeastern coast of Brazil. Arq. Est. Biol. Mar. Univ. Ceara, 5(1): 29-64.
PAIVA, M. P., and B. F. GRANGEIRO, 1970. Investigations on the whaling seasons 1964/67 off north eastern
coast of Brazil. Arq. Cien. Mar. Univ. Ceara 10(2): 111-126.
PURVES, P. E., and M. D. MOUNTFORD, 1959. Ear plug laminations in relation to the age composition of
a population of fin whales (Balaenoptera physalus). Bull. Br. Mus. Nat. Hist., Zoo/., 5(6): 123-161.
RoE, H.S.J., 1969. Seasonal form~tion of!aminae in the ear plug of the fin whale. Discovery Rep. 35: 1-30.
SCATTERGOOD, L. W., 1949. Notes on the Little Piked Whale. The Murrelet, 30(1): 3-16.
SERGEANT, D. E., 1963. Minke Whales Balaenoptera acutorostrata Lacepede off the western North Atlantic.
J. Fish. Res. Bd. Canada, 20(6): 1489-1504.
TAYLOR, R.H.F., 1957. An unusual record of three species of whale being restricted to pools in the Antarctic sea ice. Proc. :(ool. Soc. Lond., 129: 325-331.
WILLIAMSON, G. R., 1959. Three unusual rorqual whales from the Antarctic. Ibid., 133(1): 135-144.

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

MINKE WHALES OFF BRAZIL

EXPLANATION OF PLATESPLATE I
Southern Minke Whale Balaenoptera acutorostrata bonerensis Burmeister 1867.
Drawn
by Larry Foster of General Whale, California from photographs of alive and deadspecimens.
PLATE II
Baleen plates of a typical Southern Minke Whale caught off Brazil. Note the assymetry of the pigmentation; the pale plates extend further back on the right sidethan on the left side, as also occurs in the fin whale.
PLATE III
Unusual case of minke whales trapped over-winter in pack ice in a coastal fjord of
Graham Land, Antarctica. In normal winters these individuals might be off
Brazil. (From Taylor 1957, photos copyright of British Antarctic Survey.)
Plate IV
Comparison of teJtis of two mature whales caught off Brazil with testis of two mature
whales caught in the Antarctic. In the Brazil whales, active spermatogenesis isoccurring; in the Antarctic whales very little spermatogenes is is occurring. Spermare visible only in the Brazil specimens and are the tiny black dots in the tubles.
Magnification Xl50. Photographs by Dr. R. F. Macadam.
Plate V
Testis tubule showing many sperm (the tiny black dots are the nuclei of the sperm
heads). From 9.1 m whale caught off Brazil, November 7 1974. MagnificationXlOOO. Photograph by Dr. R. F. Macadam.

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

59

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~OSWV!Tl l i\へ ·~ ·oA CONSIDERATION ON AN AVAILABLE SOURCE OF
ENERGY AND ITS COST FOR LOCOMOTION IN FIN
WHALES WITH SPECIAL REFERENCE TO THE
SEASONAL :MIGRATIONSAKITO KAWAMURA
Whales Research Institute, Tokyo

ABSTRACTThe energy cost for locomotion and possible distance of the migrations
to and from the feeding grounds for southern fin whale were estimated being
based on the amount of whale oil by girth measurements. A 18.1 meter fin
whale keeps about 90 m 2 of the blubber area as substantially important for
expending the preserved whale oil, and about 1.8 m 3 of oil is preserved in
2.0 cm thick of the blubber. The energy cost for locomotion by swimmingin material fin whale was estimated as 0.022 Kcal- 1 ·Kg- 1 ·Km- 1 by extrapolating parameters onto the known regressions of body weight-energy cost relationships in aquatic animals, and an integrated distance of migrations whichis covered by following above mentioned parameters would be somewhere
between 20,000 and 30,000 Km. These figures lead to a consideration that
it is not unlike but generally quite possible to do have some food intake
through the breeding migrations of whales since energy budget for locomotion in large rorquals seems to be scarcely well balanced in general patternof their movements. These overall situations regarding the energy for locomotion were discussed along with the seasonal distributions of whales and alsowith the so-called problem of the breeding grounds.

INTRODUCTION
In the consideration on the seasonal changes of nutritive condition in California graywhale, Eschrichtius robustus, Rice and Wolman made a calculation on the expectedamount of energy expenditure throughs wimming locomotion being based on the
body weight difference due to the combustion of tissue before and after theirmigratory journey, and the weight loss was assumed as a function of oxidation of fat(Rice and Wolman, 1971). Being found that the overall energy budget for locomotion seems well balanced, Rice and Wolman (1971) concluded that gray whalemust be able to perform migrations of such large distance as invade far deep into
the Bering Sea and/or the Chukchi Sea without causing any malnutrition, anddescribed, " there is no reason to assume that gray whale must feed while on
the winter grounds ". Since gray whale is one of a well known regular migrators
among large whales, the above mentioned result itself would be appropriate in
the subject of controversy concerning to energy budget of migrating whales. VerySci. Rep. Whales Res. Inst.,
No. 27, 1975, 61-79

62

KAWAMURA

recent aerial observations, however, revealed that gray whale clearly feed on small
fishes on the way of their southbound migrations (Sund, 1975).
On the other hand, however, the baleen whales other than gray whale actuallydistribute with more larger numbers in the world oceans. They are balaenid andbalaenopterid species of which the latter would be historically most important
relating to human activities as an industorial resource. The general ecology aroundbalaenopterid whales especially in so-called rorquals much differs from that ingray whale, that is to say, we are still in need of a verification in rorquals on thismatter whether or not we will find similar conclusion to the results by Rice and
Wolman. Under these circumstances I made a calculation and assessed the energy
cost in relation to the swimming locomotion along with the overall energy budget
through the seasonal migrations in southern fin whale, Balaenoptera physalus, andthen extended a consideration to the seasonal distribution of whales.
NOTES ON THE SEASONAL MIGRATIONS OF WHALESThe conception that large whales are very likely to make seasonal migrations
which extend over those vast oceans seems to have come in general knowledge as aresult of accumulating a great deal of fragmental evidences brought on by navigators,
fishermen and whalemen through many past years, but its substantial features
remains still obscure even today. One of well dealt descriptions on the seasonal
distributions in a north and south direction in the North Atlantic balaenopterids
species was worked out by Scoresby (1820), by which our knowledge on the migrations of whales made much progress. Discussing on the evidences of whale sightingsin nineteenth centuries, Scammon (1874) could describe only a possible north- andsouthbound migratory habits for humpback whale, Megaptera novaeangliae, andright whale, Eubalaena gracialis in northern hemisphere is unlikely to migratecrossing over the equator. There are another evidence that some blue whale,
Balaenoptera musculus remains in the waters off California throughout the year. The
seasonal migrations and the movements of whales are such variable and complicatedby each whale species and place from time to time that the matter cannot be said
as the comprehensively well explained subject.
Although it was not the aim in his discussion, True ( 1904) described that some
North Atlantic humpback whale migrates down to at least l8°N latitude. Kellogg(1929) well reviewed on the migrations of six species of baleen whales based on theavailable evidences by that time. Although his descriptions were mainly confinedto the northern species of whales, he (Kellogg, 1929) also mentioned but briefly on
the migrations in southern species. By his above mentioned review the generalidea concerning to the migrations of whales became much clear but still mysterious
especially in recognizing the movements of whales as a large scaled and successive
event performed through a life span of each whale individual, that is, the migrations of animals in broad sence. The comparative field study by Mackintosh andWheeler (1929) on the biological and morphological characters in southern blueand fin whales of both South Georgia and Saldanha Bay made our knowledge on
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

LOCOMOTION ENERGY AND MIGRATIONS

63

the migrations of whales much concrete. Shortly, it can be said that pattern of
the whale migrations to the extent as have currently been recognized conception
may be basically established by their work. However, movements of whales andwhere they may stay during winter or so-called non-feeding seasons are almost
unknown even today except those traditional expressions of somewhere of " warm
temperate or tropical waters."
According to Mackintosh and Wheeler (1929) connective tissue lies beneath thevery surface skin of whales is the main body for depositing fatty substances, andthe thickness of the blubber itself varies by season in southern species, that is, morethin layered blubber is found just after the winter and get more fat by an approachof autumn as a result of voracious feeding through the past several months in the feeding ground, whereas little feeding takes place in South African waters. With thesefacts along with a possible identity of whale populations which occur in both South
Georgian and South African waters, it was obvious that both blue and fin whales
are likely to make meridional migrations by season through which they do not
feed but can continue swimming around by expending the blubber oil as the energy
source for their locomotion. The general idea of the seasonal migrations of large
baleen whales became clear to such somewhat fixed forms as having been recognized.
At the same time the reason why they have to migrate into those " warm temperate
or tropical waters " was well explained by supposing that cold feeding grounds ofalmost freezing point would be disadvantageous for parturition and for new born
calf to keep well its body temperature. The conception seems fairly well establishedand there have found no additional evidences or explanations regarding this mattereven in recently published articles which in part dealt seasonal distributions, feeding
and breeding migrations for large whale species of the world oceans (e.g. Slijper,
1958; Jonsgard, 1966; Mackintosh, 1965, 1966; Nishiwaki, 1966; Matthews, 1968).
However, Kellogg (1929) considers that there must be some fin whales throughout
winter season in the South Georgian waters, and Mackintosh (1965) thinks thestatus of whale migrations as a possibly very loose and/or somewhat random
movements in general, that is, some individuals may stay in the feeding grounds
throughout a whole year while the other may not undertake those annual back andforth movements so regularly. Reviewing on the geographic ranges and seasonal
distributions for large whales in the North Atlantic, Jonsgard (1966) consideredthat some whales may stay in the boreoarctic part of the North Atlantic, but he
did not deal much about the breeding migrations by reason of less documented
subjects. On the other hand, Mackintosh (1966) described two kinds of interestingand suggestive ideas on the status of migrating whales whether they are apt to concentrate or disperse in the breeding grounds. Matthews (1968) explained the seasonal migrations of whales schematically but he remained within showing a " warm
temperate or tropical seas " as the destination of migrating whales. Recent description by Gaskin (Gaskin, 1972) concerning to whales migrations made a subject
clear in a concrete form that the sea region where the reproduction of fin whaletakes place would be found in the offshore waters within 150 miles or thereabouts.
Where are the large whales during winter or reproductive season is still mysSci. Rep. Whales Res. Inst.,
No. 27, 1975.

64

KAWAMURA

terious subject which could be approached only by accumulating a scattered evidenceson whale sighting. In the North Pacific region, for instance, a possible place forover wintering in fin and sperm whales seems to be still beyond of our scientificknowledge. Although an approximate movements of migrating whales may berecognized after the general principle as mentioned above, it seems still in need offurther investigations and considerations to prove the matter more clearly. For

@®

CD©

®

®

Fig. 1. Points for girth measurements.
Table 1 and Fig. 3.
TABLE 1.

Encircled numerals correspond to those in

GIRTH MEASUREMENTS IN THE SOUTHERN FIN AND

Ser. No.
1656t

167ltt

1673

1676

Sex
Body length (m)Thickness of blubber (cm)

F
21.1
7.0

F
17.5
6.5

M
17.6
6.5

M
17.5
5.0

Date of catch (month/day)

3/11
49-09
77-42

3/12
49-16
79-32

3/12
49-16
79-32

3/13
49-28
80-31
200

810

900

860
850

780

644

560

496

286

238

674
560
448
330
208

630
530
464
300
208

Position of catch ~ ~ ~Highest point of lip
Center of blow hole
Eye
Angle of gape
Anterior base of flipper
Tip of flipper
Posterior end of ventral grooves
Unbilicus
Center of genital apertureAnus
11 Center between dorsal fin and base of tail flukes
12 Base of tail flukes
2
3
4
5
6
7
8
9
10

* Measurement at 140 cm anterior from the base of tail flukes
110
**
150
***
1) Measurement at 238 cm posterior from the base2) Posterior margint) Pregnant: foetus of 281 cm, femalett) Not pregnant

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

65

LOCOMOTION ENERGY AND MIGRATIONS

instance, whether or not non-feeding activity during seasonal migrations explains or
actually fills overall locomotion energy required for migrating into those "warmtemperate or tropical seas" only by expending the blubber oil without causing
malnutrition.
MATERIAL AND METHOD
During the course of 1971/72 Antarctic whaling season when I was on boad of themother ship " Tonan Mam No. 2 " of the Nippon Suisan Co. Ltd., girth of fin andminke whales was measured at three to nine points of the carcasses (Fig. 1). True
(1904) noted several girth measurements on the basis of one side or whole of carcasses, and Captain Roys gave 39 ft. of girth for 95 ft. blue whale in 1800's (Scammon, 1874). However, since very little girth measurements have been done in
the ordinal program of the biological examinations on whale carcasses, I would like
present here the whole row data of the girth measurements in nine fin whales (6male and 3 female) and four male minke whales for the convenience of later use(Table 1). As shown in Table 1 both fin and minke whales were caught in the
mid March of 1972. Two-meter measure was used for actual measurement by
MINKE WHALES CAUGHT IN 1971/72 ANTARCTIC SEASON
Fin

Minke

1678tt
F
19.3
6.5

1679

1692

1702

1705

16-1

M
17.4
5.5

M
20.9
6.0

M
17.7
5.5

M
18.1
7.0

3/13
49-28
80-31

3/13
49-28
80-31

3/14
49-46
80-03
240

3/14
49-46
80-03
180

3/14
49-46
80-03
220
7702'

M
8.4
4.6
(Avg.)
3/16
53-58
85-16

16-2
M
8.2
4.6
(Avg.)
3/16
53-58
85-16

4.6
(Avg.)
3/16
53-58
85-16

26-2
M
8.4
4.6
(Avg.)
3/16
53-58
85-16

316

316

314

354

420
480

426
492

408
448

412
470

490
450
372
296
234
100

500
480
376
290
216
108

440
426
346
262
200
108

452
432
356
280
216
100

23-1
M

310
860

596
558
380*
268

560
480
296**
186

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

720
626
560
354
232

794
854
8041'

790

640
540
470
300*
192

920
600
532
320***
220

66

KAWAMURA

sliding the scale like a looper. The air which had been filled in the ventral cavity
to make the carcasses afloat was discharged when the carcasses were pulled up
onto the £lensing deck except no. 1673 fin whale which kept some amount of air
but not fully. Each girth measurements given in Table 1 are obtained by doubling the actual figure from one side measurement. Measurements at nos. 1 and 3
points show only the upper half, that is, only a part of the skull.
Since no measurement for so-called body proportion analysis on the carcasses
concerned in Table 1 was conducted, the proportion data obtained in the South
Georgian fin whales by Mackintosh and Wheeler (1929) were adopted as the pos-

t'

l.18
1.07)
18.86
(18.94)24.45(24.83)28.51
44.57 (28.71)(44.35) 45.82(45.59)Fig. 2. Average body proportions (in percentage figures) for the southern fin
whales. Figures for female are given in brackets (Calculated from the data by
Mackintosh and Wheeler, 1929).
SQUARE METER

I· 17.97 + 17.39 +

®
I
I

®

®

19.67

+ 14.36 .j:·o~1 • 9.42 •I• 4.05 ·!

0

®@

@

@
I

"II

1+-----------13.57 M - - - - - - - - + t

Fig. 3.

Surface area of body trunk for a southern fin whale (18.l m, male).

sibly most well representing figures to know relative position of the girth measurements (Fig. 2). Of six fin whales 18.1 meters of male individual (no. 1705 specimen
in Table 1) which was most well measured was selected for the calculation of bodysurface area, and consequently, the volume of the blubber. By this treatment thewhole body surface area was calculated section by section as given in Fig. 3. Anterior part from blow hole and tail flukes were excluded from the calculation sincethese two are considered unlikely to relate to the question to be discussed here. DisSci. Rep. Whales Res. Inst.,
No. 27, 1975.

LOCOMOTION ENERGY AND MIGRATIONS

67

tance between notch of flukes and the anterior base of tail flukes was hardly determined clearly, and 4.0 percent was adopted tentatively for its proportion: thenthe actual body length which concerns to a consideration is 13.57 meters, and thewhole body surface area corresponds to 13.57 meter body length is calculated as
89.924 square meters (ea. 90 m 2 ).
The thickness of the blubber with connecting tissue beneath the epidermis anddermis considerably varies with the seasons. In southern fin whales caught in theSouth Georgian waters averaged difference in the thickness of the blubber against
body length in percentage figures was 0.37 for summer and 0.27 for winter respectively (Mackintosh and Wheeler, 1929). The material of 18.1 meter fin whale
showed 7.0 cm thick blubber, that is, 0.39 percent against its body length while it
was only 5.0 thickness (ea. 0.29 percent) in another material of 17.5 meter male fin
whale. Comparing these figures with those obtained in South Georgian waters,
TABLE 2.

DATA AND ASSUMED CONSTANTS CONCERNING TO THE
CALCULATIONS FOR FIN WHALE LOCOMOTION*

Species
Position of catch
Date of catch
Body length
Sex
Thickness of blubber
Area of blubber concerned**
Body weight***

Fin whale : Balaenoptera physalus
49°46'S, 80°03'E
March 14, 1972
18.l mMale
7.0cm
89.9 m 2
34.0 tons

Thickness of blubber expended
Caloric value for blubber oil
Specific gravity of blubber oil
Energy cost for locomotion
Efficiency of muscle in performing
external work
Condition of swimming
Estimated migration distance

2.0cm
9.34-9.45 Kcal/gr.
0.92
0.022 Kcal/gr./Km
25%
10 knots, laminar flow
2.1x10'-2.3X10 4 Km

* for further detail see Schmidt-Nielsen (1970, 1972) and Kawamura (1973a, b)** body trunk proper: anterior part from blow hole and appendages are excluded*** after the formula by Ash (1952)

the 7.0 cm thick blubber in 18.1 meter fin whale would possibly be at its upper
most extremes as the function of depositing whale oil, and similarly, 5.0 cm thickblubber would be at its lower extremes. These facts suggest that the blubber oil
which corresponds to 2.0 cm difference in the thickness of the blubber is likely to
be consumed through the migratory journey. According to Slijper (1958) about
40.0 percent of the blubber is in general consisted of connecting tissues. So if wetake this figure into considerations the total volume of whale oil preserved in the
2.0 cm thick blubber for 18.1 meter fin whale would be calculated as about 1.80
cubic meters. Assuming five months between May and September as for the duration of whale migrations, the blubber oil as the energy source for locomotion would
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

68

KAWAMURA

be about 0.36 m 3 ·month-1, or 1.2 X 10 4 ml·day- 1 • Since generally known caloric
value and specific gravity of fin whale oil is 9.34-9.45 Kilocalories • gram-1 and 0.92
respectively, the unit volume of consumable whale oil could be converted to 1.1 X
104 gram·day-1, or 10.27Xl04 -10.39xl0 4 Kilocalories·day-1 • The grand totalof consumable energy during five month migrations in 18.1 meter fin whale
would be about 1.55X10 7 -1.57 X 10 7 Kilocalories. Data and some constants concerning to above mentioned calculations are given in Table 2.
ENERGY COST FOR LOCOMOTION IN FIN WHALESThe energy cost for swimming locomotion in large whales is still unknown. However, it seems possible to speculate the energy cost for locomotion by the aid of known
data in small toothed whales and in some another kind of aquatic animals (e.g.
Schmidt-Nielsen, 1970, 1972). As I showed previously (Kawamura, 1972), theenergy cost in large whales seems to be very less expensive when it is comparedwith the case found in another species of aquatic animals. Only four percent orthereabouts in daily feeding rate in sei whale (Kawamura, 1974) and possibly
similar figures in another species of baleen whales would support very less energy
expenditure for locomotion in those large whales. According to Schmidt-Nielsen(1972) the energy cost in a dolphin of 100 Kilogram body weight is about 0.1
Kilocalories·Kilogram- 1 °Kilometer-1 when it swims through laminar flow of waterover the body at 10 knots in speed if we assume its conversion efficiency of muscle
in performing external work would be 25.0 percent whereas 22.0 percent was assumed by Gawn (1948). In general the energy cost for locomotionin animals is
closely related to the body weight of animals, and changes by following after the
exponential like curve (Schmidt-Nielsen, 1970, 1972). Then the energy cost or
metabolic rate such as 0 2 ml·gr-1 ·h-1 in relation to the body weight of animals canbe expressed under logarithmic scale by three different regressions each of whichshows difference by the kind of living media; whether the animal flys, runs, and/or
swims. Since these corelations can be followed in aquatic animals and seems to
be not unreasonable for extrapolating treatment, the energy cost for locomotion ( =swimming) in large whales would be known reasonably by extrapolating theirbody weight onto known energy cost-body weight relationships.
Then the energy cost for swimming in 18.1 meter fin whale which weighs about
34 tons is estimated as 0.022 Kilocalories·Kilogram- 1 °Kilometer-1 if the same
parameters are assumed as in the case of dolphins, that is, the animal swims at the
speed of 10 knots under laminar flow of water over the body and 25 percent in theefficiency of muscle in performing external work. (See Schmidt-Nielsen, 1972,
fig. 7) Similarly a blue whale of 100 tons of body weight would expend about
0.014 Kilocalories°Kilogram-1 °Kilometer-1 as its energy cost for swimming. The
data indicate that the large rorquals seem to expend some 1/5 to 1/7 times of energy
cost than the dolphins.
The actual energy cost in 18.1 meter and 34 tons of fin whale may be 7.48 X
102 Kilocalories • Individual- 1 • Kilometer-1, and some 2.10X10 4 -2.36 X 104 times
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of these cost would be required for five months migration since its totaled energy
corresponds to 1.55 X 10 7 -1.57 X 10 7 Kilocalories. These facts indicate that the18.1 meter fin whale can migrate for the distance of some 2.1X10 4 -2.3x10 4
Kilometers by expending its blubber oil preserved in 2.0 cm thickness of the blubber.
Actually, however, slightly more energy expenditure must be took into considerations since the metabolic rate as examined in human body increases for 10-20 times
according to the physical conditions whether he moves vigourously or rests (Yoshikawa, 1959). Jumping, leaping or breaching (Racovitza, 1903) behaviours in large
whales are often observed on high seas, and several times of these kind of violent
movements which may be induced by playing, chasing, and/or escaping from theenemy would possibly be a quite daily event in the wild life. All of these areresponsible for increasing the energy expenditure, since whales were assumed as
migrate constantly under 10 knots of swimming speed throughout their migratory
journey. In gray whale, however, Rice and Wolman (1971) have found well- balanced energy budget being based on the "actively swimming animal", and 10
knots in swimming speed would also be allowed for general considerations as a
totally averaged figure in energy expenditure.
CONSIDERATION ON THE AMOUNT OF WHALEOIL AND THE ENERGY COST
The calculation as has been described in previous section was introduced by following after the several assumptions: 1) Whale oil preserved in the blubber aroundhead and ventral grooves, and that in bones and internal tissues are not utilized
as the energy source for locomotion, 2) Whale oil which corresponds to 2.0 cm
thickness of the blubber which covers over main body trunk is consumed as theenergy source for locomotion, 3) Energy cost for swimming locomotion in large
rorquals was estimated by extrapolating the parameters into the known regression
line in aquatic animals and, 4) As it was assumed in the case of dolphins, the efficiency of muscle in performing external work would be 25 percent, and the animal
swims at 10 knots under laminar flow of water over the body.
Concerning to an absolute amount of whale oil itself, the whales body under
alive would contain much more than that considered in this report. For instance,
Omura et al. (1942) showed that the 18 m meter fin whale caught in the coastal
waters of Ja pan yielded 0. 73., 0.5 and 0.5 tons of whale oil from the blubber, bones
and internal organs respectively. Arai et al. (1950), on the other hand, showedthat 53 percent of the blubber around whales head contained whale oil and 65
percent was in the blubber around the back. Following to these oil rich parts themuscle around lumber to tail flukes, ventral grooves and/or abdomen contains ratherrich (Feltmann et al. 1948) but less than 20 percent or so of whale oil. That is, thepotential in preserving the whale oil as the energy source would be reduced toabout 1/3 when it is compared with oil rich parts, and the amount of whale oilfrom an individual must be more greater than usually expected, whereas theirimportance as the energy source for locomotion would be less than the blubber oil,
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since most of whale oil preserved in bones or some other parts other than the
blubber is considered to be unlike from physiological reasons to be consumed forthe purpose of locomotion.
It is well known fact that whales caught in the beginnings of Antarctic whaling
season show slender body trunk due to not only their thin layered blubber but also
possible consumption of oil and fatty substances preserved in the whole body.
Considering on this fact along with a comparison in metabolic physiology found inanother animals, the whale oil preserved in the tissues other than the blubber
would be consumed to some extent as the energy source for locomotion. And if
we assume some physiological analogy between whales and human body (Yoshikawa, 1959), whales under some peculiar occasions or situations may consume
not only their oil but also do even proteins of body tissues.
Whale bones also contain a considerable amount of whale oil, but it is presumably not consumable as the energy source for locomotion by reason of thefunction of bones themselves. The strength of animal bones is the function of theirsectioned area and it increases only by following after a square number relations withbody length whereas body weight of animals which must be suspended by the bones
increases by cube number relations. Although the whales, a typically well adapted
animals for aquatic environment, do not need such strength as terrestrial animals, andthe bones of whales are not structured well like those animals. Then the function
or physiological process which makes the strength of bones reduce much by consuming the contained whale oil may have to be kept as little as possible. Still
more, the deposit of whale oil among many organs seems to occur firstly in the internal organs, fatty layer and then in the bones by this order (Omura et al. 1942).
This fact also supports the assumption that whale oil preserved in the bones seems
to be hardly consumed as the energy source for locomotion.
From these discussions it would be reasonable to consider that there are somestill more amount of consumable whale oil in living animals while some are notavailable as the energy source for locomotion. As it was described previously thewhale oil only from the blubber can make the whales swim for 20,000 km or thereabouts. Then if some additional energy source for locomotion is expected, thedistance could be moved potentially without reducing any physiological functions
such as a malnutrition in large rorquals would be somewhere between 20,000 Km
and 30,000 Km. On the other hand, Mackintosh (1966) supposed the migrating
distances covered by large rorquals must be somewhere between 1,000 and 3,000
miles. These figures, however, correspond to only about 17 to 50 degrees of latitudes, and are considered rather under estimations.
There are no ways to prove how much extent the energy cost obtained by an
extrapolation for large rorquals is appropriate or not. The estimated energy costof about 1/5 of dolphins, however, can be discussed little more in the point of
metabolism of animals. The basal metabolism of animals closely relates to the bodysurface area. Bottlenosed dolphin, Tursiops truncatus which weighs 100 Kg. has
1.85 m 2 of body surface area (Ridgway, 1972). If it is assumed that the surface
around its head occupies about 20 percent of the whole, the surface around body
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trunk would be 1.5 m 2 • On the other hand, 18.1 meter fin whale has about 90 m 2
of body surface area which actually corresponds to 13.5 meter long of body trunk
only, and weighs 34 tons. Comparing with these figures of both bottlenosed dolphin and fin whale, the ratio in body weight shows 340 times larger in fin whale
whereas only 60 times in the ratio of body surface area. The well averaged numberof respiration in dolphins and phocids is 2 to 3 respirations •minute-1 while it is 1 to
2 respiration(s)·minute-1 in fin whales (Scholander, 1940). The large whales
consume comparatively little amount of oxygen than the dolphins, and are able toswim under very well controlled energy coefficient. In this respect a slight change
in body length or body surface area affects well on the whole metabolism of animals
by following after the Rubner's surface law. Actually, since the body surface areaof fin whale would reduce to some extent while they are enroute through migrations, the energy cost for locomotion may also reduce. This may be one of thereasons why large rorquals perform seasonal migrations by less energy expenditurethan the dolphins and also possibly than the figures estimated by extrapolation. It
is possible, therefore, that the energy cost and body weight relationships in large
whales may not expressed by a simply extended regression lines but by a slightly
curved or the folded lines with different coefficient.
However, these relations which have been known in another animals are well
expressed by setting up a simple regression even for between mouce and horse. Concerning to this extrapolating problem, Schmidt-Nielsen (1972) described as "Although we have no inherent right to extrapolate from the salmon to the dolphin, we
can safely assume that both animals are hightly adapted and effective swimmers,
and extrapolating from a I-kilogram salmon to a 90-kilogram dolphin is not unreasonable ". Similarly it was considered that large whales also could be extrapolated onto the regression line. In concluding this section the followings may
give a standards: the energy cost for swimming locomotion in average sized fin
whales is 0.02 Kcal.·Kg- 1 °Km-i, and overall distance which could be moved
without feeding both on breeding grounds and on the way would be somewhere
between 20,000 km and 30,000 km.
MIGRATIONS OF WHALES AND ESTIMATEDDISTANCE TO BE PERFORMED
Generally recognized destination for migrating rorquals is " warm temperate or
tropical seas" if not exclusively. Therefore, it would be worth to discuss whether
or not the distance which was estimated by the available amount of energy forlocomotion and its cost for swimming in fin whales is an accessible distance alongwith the known evidences regarding the migrations of whales to the warmer waters.
If we assume that the whales leave the Antractic feeding grounds at 60°S andstart their northbound migrations heading for the equator and again return to thestart line along the same meridional pathways, the distance covered by this round trip
is 13,320 km. That is, the whales are able to move or play around more for only
7,000 km or corresponding energy expenditure. Since little is known on the pattern
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of daily life in large whales except several but fragmental observations (e.g. Schevill
and Watkins, 1966; Cousteau and Diole, 1972), it seems still in some difficulty toestimate even an integrated distance performed by a whale in a day. It is, however, reasonable to consider that whales are unlike to swim straight but trace aquite complicated pathways which gives far more doubled distance than is consideredby an actual situation, since the daily behavior of animals may possibly have notnecessarily any objects from one end to the other.
Following fact is very suggestive concerning to the daily movements of whales.
In the whaling ground which is one of the terminal places in whales migrations, where
the whales stay for some duration within a certain range to feed by moving backand forth, it is often experienced that the whales having been spotted in the previous evening are hardly found again in the next day. In this case it is well supposedthat the whales had moved for a considerable distance during the previous night.
To make the matter clear it would be convenient to assume the case that thosewhales were sighted again after 3 hour scouting through surrounding waters at the
speed of 15 knots. Then about 2 hours out of 3 hour scouting would be the component giving away from the starting point, that is, the whales moved at least for 2
hours as to give the distance between catcher boat and themselves. J.-Y. Cousteau
attached a specially designed marks on the back of a fin whale to make him easy forchasing it through the night, and he observed that the whale usually continued tomove at the speed of 6 to 7 knots though the whale stopped several times during theobservations (Cousteau and Diole, 1972, p. 72). Their observations give us aknowledge that the whale may considerably move around even through a night.
From these facts it would be quite natural to consider that the whales are apt tomove at least for several tenth miles within a night. If the whale migrates from
60°S to the equator at 10 knots in swimming speed, 15 days would give them enoughtime to get to their destination; the terminal sea regions of so-called "warm temperate or tropical seas " where the whales stay for the rest 4 to 5 months. Thereis no reason to doubt that the daily behaviour of whales differs much both in feeding
and breeding grounds. The estimated distance for nocturnal movements in large
whales, 30 miles per 10 hours agrees well with the swimming speed found under
the natural or undisturbed conditions, and that distance would be a minimum or
close to the lower extremes, since it was estimated being based upon a possible
zigzag movements. The whales seem to move at least about 72 miles ( = 130 km)
in a day, and 15,000 to 16,000 km would be the integrated distance covered during
4 month stay in the terminal sea regions.
Accordingly the grand totaled distance which is performed through 5 monthround trip migrations would be at least some 28,000 to 30,000 km. As it was shown
previously the 18.1 meter fin whale is able to move only for 20,000 km or thereabouts by expending its blubber oil of 2.0 cm thickness, and may have to face ashortage of locomotion energy unless the whale oil or fat of another source wasconsumed supplementally. These estimations suggest that even well adapted
whales for aquatic environment are hardly possible to perform migratory journeyto those " warm temperate or tropical seas ".
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However, if the supply of whale oil from the muscle and connecting tissues in
addition to that from the blubber could be considered, and/or if slightly more than
2.0 cm thick of the blubber oil was consumed, the budget of energy expenditure
required for those migrations would be well balanced. Actually the large rorquals
distribute all over the world oceans and most of them sternly keep 4 to 5 cm thickof the blubber even after a very long journey for 5 to 6 months. On the other
hand, there are actually the whales which keep very thin blubber even in late March,
the time they have to leave the feeding grounds for breeding. These very slenderbodied whales seem unlike to consume their blubber oil any more as the energy source
for locomotion, since it may cause some physiological difficulties to reduce their
blubber oil deposit, and the statistics on the thickness of the blubber also suggest
its unlikelihood. Then slender bodied whales have to change or modify theirmigratory pattern and fashion of life, which may somewhat different substantiallyfrom that of the majority of normally fatted whales. For instance, poorly fatted
whales cannot but remain within some migratory range and/or have to migrate intosome productive and organism rich coastal waters where they can feed to a certain
extent. The whales may have to swim so economical basis as to reduce theenergy expenditure as little as possible since the energy for locomotion in any kindof whales does not seem to have been provided sufficiently. It is well known fact in
the Antarctic feeding grounds that the whales move longitudinally for rather widely
ranged zone extending for two or three of six sectors, whereas they are apt to follow
the cource along the continent or islands in breeding migrations, and little of themmigrate into the real mid oceans. These general tendencies are well observed bywhale sighting at high seas and by the recovery of whale marks (e.g. Dawbin, 1959;
Ivashin, 1971; Mozer, 1971). Whales which have distributed rather randomly
within the feeding grounds are apt to concentrate along their migratory pathways but
it is far sparce and loosely packed gatherings like a stream and is hardly comparable
to so-called school or herd of animals. Humpback whale apparently shows somegatherings as a unit of local populations enroute their breeding migration (Daw bin,
1964; 1966), but these kind of whale movements if not clear like the humpback
whale may also be found in rorquals. These general tendency in whale movementsor migratory habits may closely related to the productive seas or the occurrence ofrich standing stocks of organisms, and most of which is found in the marginal waters
along the continent or islands. A fertile zone such as an equatorial divergenceregions too attracts whales even if it is found on high seas. Considering on thefood and feeding of baleen whales, Nemoto (1959, p. 281) concluded that the movement of whales in the North Pacific and Bering Sea regions is closely related to thegeographical characters such as the shape of the sea, and fin whale occurring in
those waters can be considered as " ocean and marginal denizen ".
The energy for locomotion in rorquals migrations seems to be balanced on thewhole. It is, however still desirable to minimize the energy expenditure as little
as possible since an amount of whale oil preserved in a body is far less for a luxuariant
consumption. Therefore, it would be better and/or appropriate for most of migrating whales to take linear like pathways both in north- and south-bound migrations.
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For instance, little number of whales in the southern waters of Australia would
migrate to the tropical African coast by crossing the Indian Ocean diagonally.
Sperm whales evidently swim at least 600 miles within 8 days between the Antarctic
and South African coast (Clarke, 1972). This fact strongly suggests that the whaleson the whole, once they begin the migrations, can travel very rapidly even though
they feed on squids on the way.
The migrating whales are considered to be apt to invade into the almost sameor very close region of terminal seas every season. In the coastal waters off Abashiri,
Hokkaido, there once was a fin whale which was easily identified by its characteristically well damaged dorsal fin, and this animal used to be spotted if not entirely inthe same place at least in the beginning of season (Omura and Kawamura, 1968).
Similar evidence or possibility is also documented in the Antarctic fin whale ofArea II based on the recovery of whale marks (Brown, 1962, p. 14), and, thesesituation was expressed as "something like a migration route" (Mackintosh, 1966)
which is somewhat similar to "corridors" called in the migration of birds (Orr,
1970). Since it would be hardly possible to modulate the season and place of
migrations with considerable accuracy, the migration of whales as a everlasting
phenomenon could be recognized only when the migrations of whales were assumed
as follows; in order to perform the migration with those above mentioned kinds ofaccuracy, the whales may have to migrate at full or near at full speed without
wasting time on the way by following well known migratory pathways along withthe aid of well experienced navigation.
GENERAL CONSIDERATIONS IN CONNECTION WITH
THE DISTRIBUTION OF WHALESAs it has been discussed in the previous section, the budget of energy expenditure
for locomotion in large rorquals seems rather complicated, and cannot but consider the subject analogically by the aid of some assumptions, since there are so
little suggestive evidences which were obtained in the field observations. For instance, it is still beyond of our knowledge where and when the perturition andmating of rorquals will actually take place in those vast " warm temperate or
tropical seas ". No other evidences except those have been refered to seem to beleft for the further discussion, and following discussions are a kind of conclusive
view points in which the movements of whales is considered along with the generalfeatures of the distribution of whales.
In rorquals the pattern, habits and the behaviours in the way of life are quite
naturally and generally recognized as a regular migrators which move between coldpolar seas and warm temperate or tropical seas, and these are considered appropriate from the overall budget of their energy expenditure for locomotion, sincemajority of whales are likely to be well balanced in this instance. However, anamount of whale oil or fatty substances preserved in the whales body considerably varies with the physiological status or biological characters endowed with eachwhale species. Still more, the wild animals would presumably have to keep afford
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in the energetic budget to make their behaviour more fruitsfull under the wild
environment whereas the budget in energy expenditure actually seems to be very
tight. In this instance, it would be reasonable and natural to suppose that thewhales usually must feed to a certain extent even if they are on the way of migrations. California gray whale which are supposedly well balanced in the energy
expenditure can perform seasonal migration without taking any foodstuff throughtheir round trip migrations (Rice and Wolman, 1971 ). Even in gray whale, however, feeding activity seems to be not unusual at least in their southbound migration close to the regions of their breeding grounds (Sund, 1975). The idea that
more rorquals do not seem to feed or be under almost starved condition during
their migrations (e.g. Jonsgard, 1966; Mackintosh, 1966) would be hard to believe if not exclusive. In general the metabolic rates increase high in the warmer
environment than in the colder one, and this may become serious in small younger
animals since they swim under relatively high energy cost for locomotion than thewell aged animals. That is why large rorquals do not concentrate much in mid
oceans where a monotonous but well stabilized environment prevails, but morecommonly found in some coastal or offshore waters of structural complexity, thatis, the region of high production or rich in standing stocks of organisms whichis undoubtedly advantageous for occasional feeding. Two ways of the possible
status for migrating rorquals whether they concentrate or disperse in the breeding
grounds (Mackintosh, 1966) are quite suggestive on the whole. In the latterinstance the evidence that many whales are spotted in offshore waters within acertain ranges in lower latitudes strongly suggests a possible feeding activity of
whales as the case explained by Clarke (1962). Even planktivorous species ofbaleen whales apt to feed on shoaling small fishes and their larvae according to thetime and place. A typically similar situation is found in the comparison for food
habits between the North Pacific and the Antarctic (Nemoto, 1959; Kawamura,
1973, 1974). That is, there reason to believe that the baleen whales in breeding
grounds may feed on some organisms such as the larvae of skipjacks, tunas and ofmany other warm water fishes. By taking these general situations along with
presumed physiological conditions and feeding habits in large rorquals into consideration, it seems reasonable to believe the food intake during breeding migrations as the comments by Dr. Sergeant. (Mackintosh, 1966, pp. 143-44).
One of explanations why rorquals migrate to those " warm temperate or
tropical seas " is an advantage for breeding especially for the safety of newborn
calf of very thin blubber and make them easy to keep body temperature from chilly
environment. Antarctic minke whale, Balaenoptera bonaerensis, likely to show rather
irregular breeding cycle (Ohsumi, personal communication) than the other rorquals
which show an approximately one-year cycle. In this instance some individuals of
the Antarctic minke whale may have perturition during their stay within thecolder region, and how is the case for the calving in bowhead whales, Balaena mysticetus, explained. In this respect the Kellogg's pointing out that "it is not the sea
temperature but the food conditions that induce whale migrations " would be quite
suggestive itself (Kellogg, 1929, p. 45). Kellogg's idea also make us consider that
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the migrations of whales is desirable to be treated by each whale species.
Results of whale sighting in the Atlantic and Indian Oceans show the year
round distribution oflarge whales in the temperate and tropical regions (e.g. Town: end, 1933; Brown, 1957, 1958; Zenkovich, 1962). In those waters populationof whales may be consisted of the several different origins: some may come fromnewly migrated population and the other may somewhat from endemic or locallyseparated population such as the fin whales in the East China Sea (Fujino, 1954;
Ichihara, 1957) and sei/bryde's whales in the northern waters off North Island of
New Zealand (Dawbin, 1959, p. 15). Those locally separated whales seem to staythere throughout the year, and consequently, may have to feed on regardless theseasons. Then it is quite unlikely that the only newly migrated whales do not feedon among those taking food in those local waters. In the waters off California blue
whales are usually seen throughout the year (Kellogg, 1929), and they can feedon sardine and prawn in San Quentin region, lower California (Scammon, 1874).
Apart from the seasons, great number of large whales are apt to concentratein the zones between 30° and 40° of both south and north latitudes of the Atlantic
and Indian Oceans (Brown, 1957; 1958), and humpback whale, Megaptera novaeangliae stay in the offshore waters of lower latitudes throughout the year along the west
coast of South America. In the Arctic regions of the North Atlantic some humpback whales stay throughout the winter (Kellogg, 1929). One of another instances
is Pacific black right whale, Eubalaena gracialis which are found close to the ice-packin the Okhotsk Sea and Bering Sea regions, and then fin whale in Queen Charlot
Sound, north of Vancouver Island of approximately 50°N during February (Scammon, 1874). These facts indicate that the actual pattern of distributions in large
baleen whales, if not entirely, do not always coincide with those ideas generally
known as migration cycles. The " tropical seas " is considered unlikely for mostwhales as to be a destination being invaded after the painstaking long journey, andthis make us even doubt whether or not those habits are inevitable for carrying out
an appropriate life of whales. In the general fashion of the distribution of whales,
however, age, sex, and physiological status of each whales would relate to theirmigrating movements though the details on this subject have been known little
except the case in sperm whales, Physeter catodon, which once had caught in all over
the world oceans and are known as very highly socialized animals in the mode ofline among large whales (e.g. Mackintosh, 1965; Gaskin, 1970; Berzin, 1972).
Then the structure of population in baleen whales which occur in " warm temperate or tropical seas" may be a subject to be known in future. The baleen
whales may have to feed to a certain extent in food rich waters regardless seasons,kind and origin of whales stocks. Some blue whales are believed to feed actively
in the waters south and south east of Madagascar, which are located far northfrom the Antarctic feeding ground proper (Gambell et al, 1974). The ideas ofboth non-feeding activity and seasonal migrations in large whales are not primarily
the phenemon of two sides of a thing but independent each other, if not, the actualfeatures in the distribution of whales would hardly be explained.
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CONCLUSION
Some whales keep rather sufficient energy source to perform the migration
between cold feeding ground and warm breeding ground while others are hard toaccomplish their migrations unless they take food to some extent on the way. Migrating population of whales is composed of individuals which greatly vary in energy
potentials. On the whole the budget of energy for locomotion does not always
balance well or clearly unbalanced. Under these circumstances some whales may
migrate to the tropical seas and others may have to take different pathways of less far
or, have to modulate the ways of life although some of them will be fatty in anotheryear and can migrate much distance as the standards. It would be convenient tosuppose that whales feed primarily on the way of their breeding migration to getan additional energy for locomotion although the extent of feeding may actually
vary with each individuals. Some whales which had not preserved much whale
oil cannot but follow the ways along more productive and food rich waters, and/or
have to remain within the waters of rather higher latitudes throughout their breeding
migration. The migrations of whales, therefore, seem to be a stream like movements of animals as called by Mackintosh (1965), and may be performed withmuch flexibility from variable fashions of migratory movements. The " warm
temperate or tropical seas " which have been believed as the destination of breeding migration still remains as an ambiguous conception. As it has been discussed,
however, whales keep potential for making a round trip which corresponds to 120
degrees or more of latitudes, and consequently,. the whales in the archipelago of
Micronesia, for instance, cannot always be considered as the stray from local population but also possible to consider as the migrants from the polar regions. In thisinstance, mingling of the populations between northern and southern hemispheresat one of their terminal sea regions would be quite possible but considered the caseto be very little. Kellogg (1929), however, suggested much the possibility ofmingling in the case of blue whale populations.
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laHIHARA, T., 1957. An application of linear discriminant function to external measurements of whale.
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IVASHIN, M. V., 1971. Some results of whale marking carried out from Soviet ships in the south hemisphere. <;oologicheskii .(;hrnal, L: 1063-1078. (in Russian with English summary).
JoNSGARD, A., 1966. The distribution ofbalaenopteridae in the North Atlantic Ocean. In: Whales,
Dolphins, and Porpoises. K.S. Norris (ed), Univ. Calif. Press, Berkeley and Los Angeles: 114-124.
KAWAMURA, A., 1972. (Marine mammals in the food chains). The Heredity (Iden), XXVI, 11: 21-31.
(in Japanese)
KAWAMURA, A., 1973a. (Some materials of the girth measurements for the southern fin and minke whales).
Geiken-Tsushin, 266: 185-187. (in Japanese)
KAWAMURA, A., 1973b. (Some materials of the girth measurements for the southern fin and minke whales
II. A tentative considerations on the energy cost for locomotion and migrations in large whales).
Ibid.,267: 193-197. (inJapanese)
KAWAMURA, A., 1973c. Food and feeding ofsei whale caught in the waters south of 40°N in the North Pacific.
Sci. Rep. Whales Res. Inst., 25: 219-236.
KAWAMURA, A., 1974. Food and feeding ecology in the southern sei whale. Ibid., 26: 25-144.
KELLOGG, R., 1929. What is known of the migrations of some of the whalebone whales. Ann. Rep.
Smithsonian Instn. 1928. : 467-494.
MACKINTOSH, N. A., 1942. The southern stocks of whalebone whales. Discovery Rep., XXII: 197-300.
MACKINTOSH, N. A., 1965. The stocks of whales. Fishing News (Books) Ltd., London, 232 pp.
MACKINTOSH, N. A., 1966. The distribution of southern blue and fin whales. In: Whales, Dolphins, andPorpoises. K. S. Norris (ed), Univ. Calif. Press, Berkeley and Los Angeles: 125-144.
MACKINTOSH, N. A., and J.F.G. WHEELER, 1929. Southern blue and fin whales. Discovery Rep., I: 257540.
MATTHEWS, L. H., 1968. The whale. George Allen & Unwin, London, 287 pp.
Mi:iRZER BRUYNS, W.F.J., 1971. Field guide of whales and dolphins. uitgeverij tor/n.v. uitgeverij v.h.c.a. mees
ziesenniskade, Amsterdam, 258 pp.
NEMOTO, T., 1959. Food of baleen whales with reference to whale movements. Sci. Rep. Whales Res. Inst.,
14: 149-290.
NISHIWAKI, M., 1966. Distribution and migration of the larger cetaceans in the North Pacific as shown byJapanese whaling results. In: Whales, Dolphins, and Porpoises. K. S. Norris (ed) Univ. Calif. Press,
Berkeley and Los Angeles: 171-204.
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OMURA, H., Y. MATSUURA, and I. MIYAZAKI, 1942. (The whales: its science and current status of whaling).
Suisan-sha, Tokyo, 319 pp. (in Japanese).
OMURA, H., and A. KAWAMURA, 1968. (On the biological investigations of the black right whale in the
Okhotsk Sea). Geiken-Tsushin, 205: 1-22. (in Japanese).
ORR, R. T., 1970. Animals in migration. The Macmillan Co. (Japan. transl. T. Shibuya, Hakuyosha,
Tokyo, 404 pp. 1975)
RACOVITZA, E., 1903. A summary of general observations on the spouting and movements of whales. True
from the Zoology of the Voyage of the Belgica, Cetology, pp. 5-19.
RmE, D. W., and A. A. WoLMAN, 1971. The life history and ecology of the gray whale (Eschrichtius robustus).
Spee. Puhl., 3, Amer. Soc. Mammalogists. 142 pp.
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Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

RECENT STATUS OF THE POPULATIONOF INDUS DOLPHINTOSHIO KASUYA AND MASAHARU NISHIWAKI
Ocean Research Institute, University of Tokyo, Tokyo

ABSTRACTA survey of the population of Indus dolphin Platanista gangetica indi
(Blyth, 1859) was conducted in December 1974. The range of the significant
abundance of this subspecies is restricted to the section between Sukkur andGuddu Barrages, and to the section between Guddu Barrage and Taunsa andPanjnad Barrages. The size of the population in the former section was
estimated to be between 350 and 400 animals. Even if the populations inother sections are included, the total population seems to be only 450 to 600
animals. The calculated mortality rate, reproductive rate and environmental
factors suggest that the population is still decreasing. For the survival of thispopulation, the stop of the killing of this dolphin and the control of water athigher level in winter are suggested.

INTRODUCTION
The dolphin of the genus Platanista is found in the lndus River and in the GangesBrahmaputra Rivers. There is found a difference of the shape of nasal crest between the dolphins in the two separate rivers (Pilleri and Gihr, 1971). Though
Kasuya (1972) reported the difference of the proportion of tail length, as he did
not measure the lndus dolphin but referred the data prepared by Pilleri (1970)
his conclusion is still doubtful.
It seems to be generally accepted that these two separate populations, started
from a single origin, have been isolated long enough to attain some morphologicaldifferenciation. However, there are two contrary opinions on their classification.
One is to consider them distinct species (Pilleri and Gihr, 1971), and the other is toclassify them into two subspecies of a single species (Kasuya, 1972). At the present
status of knowledge on these animals, we consider it reasonable to take the latteridea until more distinct morphological or ecological difference is found between thetwo, or until it is suggested that they do not freely interbreed even when the geographical barrier is not present.
The lndus dolphin was known to have been distributed in the wider range ofthe tributaries (Anderson, 1878). However, since the start of the construction of
permanent barrages for irrigation, the dolphin population have been split up into
separate smaller populations (Pilleri, 1970). And in some sections of the river,the population has been decreased or extinct (Pilleri, 1972; Roberts, 1972; Taberet al, 1967). The alarming situation of this dolphin was already noticed by Pilleri(1972) based on the observation made in 1969 and 1972 at the lower part of theSci. Rep. Whales Res. Inst.,

No. 27, 1975, 81-94.

82

KASUYA AND NISHIWAKI

section blocked by Sukkur and Guddu Barrages. The total population of thisdolphin has been considered not exceeding 700-1,000 (Anon., 1973).
The present study was conducted in December 1974 to know the present rangeof distribution of the dolphin and to estimate the population.
RANGE OF DISTRIBUTION
The presence of the dolphin was checked at various bridges or barrages. However,as the dolphin is usually scarce near the barrage even if it is situated in the section
where the presence of many dolphins is confirmed by other method (e.g. Sukkur or
Kashmore Barrage), it was tried to collect more informations from the fishermenor people living by the river. Furthermore, if possible the length of about 1 to
3 km along the stream was observed at each spot.
Section 1, up of Jinnah Barrage
The Jinnah Barrage is situated near Kalabagh Town (71°34'N, 35°59'E).
This section was observed on Dec. 15. The wide stream was present only within
about 5 km above the barrage, and above the range the river enters into a gorge.
Though it was reported by Anderson (1878) that the dolphin was present in the water
above Kalabagh, the dolphin was not observed by us. The people in the town
denied the presence of the dolphin. The dolphin seems to be absent in this partof the river in recent years.
Section 2, between Chasma and Jinnah Barrages
This part was not observed by us. But considering the small distance between
two barrages and the scarcity of water below Jinnah Barrage, the dolphin will not
be distributed in this section.
Section 3, between Taunsa and Chasma Barrages
This section was observed at Dera Ismail Khan and Taunsa Barrage on Dec.
14.
The water of the Ind us by Dera Ismail Khan was about 500 m in width andseemed to be fairly deep suggesting the possibility of the presence of the dolphin.
However we could not find it. A man living on the river bed told that he does not
see the dolphin in winter season but he had once seen an animal in summer season.
At the north side of Taunsa Barrage the water was abaut 1,200 m wide. But
there was found no dolphin in the area of about 2 km along the river.
From the above informations we consider that the dolphin might be absent inthis section or if present the number must be almost negligible.
Section 4, from Guddu Barrage to Taunsa Barrage (R. Indus) and to Panjnad Barrage (R.
Panjnad)This section was studied at 5 spots on 7 occasions as mentioned below.
Though no animal was observed at the down stream of Taunsa Barrage (Dec.
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

83

POPULATION OF INDUS DOLPHIN

14) and Panjnad Barrage (Dec. 13 and 17), it does not necessarily indicate theabsence of the dolphin in this section because the water was scarce below the barrages and the dolphin will not migrate there.

32°

+
II)

"

"O

.£

TAB "'-DGK•

+

+

0

50

100

150 200km+
Fig. 1. Map of the Indus. Black bar indicates the barrage, numeral the numberof the section in text, and the arrow the check point of Platanista. BEG, Begari.
CHA, Chachran. CHB, Chasma Barrage. DGK, Dera Ghazi Khan. DIK,
Dera Ismail Khan. EMB, Emerson Barrage. GUB, Guddu Barrage. HYD,
Hyderabad. ISB, Islam Barrage. JHM, Jhang Maghiana. JIB, Jinnah
Barrage. KAR, Karachi. KHU, Khushab. KHW, Khanki Weir. KOB,
Kotri Barrage. MIT, Mithankot. MUL, Multan. MUZ, Muzaffargarh.
PAB, Panjnad Barrage. RAB, Rasul Barrage. ROH, Rohri. SEH, Sehwan.
SHA, Shahpur. SUB, Sukkur Barrage. TAB, Taunsa Barrage.

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

84

KASUYA AND NISHIWAKI

By Dera Ghazi Khan, the observation was made on Dec. 14 at the boat bridgeconnecting Dera Ghazi Khan and Muzaffargarh. The water there was only about
100 m wide and probably less than few meters in depth. No dolphin was foundin the ranges of about 1 km at the both sides of the bridge.
Other observations were made along the stream of the length of about 3 km
at the boat bridge between Mithankot and Chachran on Dec. 13 and 17. On thisspot the water was more plenty than that at Dera Ghazi Khan situating about 150
km north. The width was about 500 m and the current was fast (2 to 3 km/hr ).
At the first visit one small dolphin (about 120 cm in body length) was observed
swimming leisurely at about 100 m north of the bridge, and another slightly larger
animal was seen at about 500 m south of the bridge. In case of the second visit,
two small individuals were sighted near the latter spot.
At the water just above the Guddu Barrage, there was found no dolphin(Dec. 12).
From the above informations it is safe to conclude that some individuals of
the Indus dolphin are distributed in this section of the Ind us River. The range ofthe distribution in winter season will be from Guddu Barrage to the confluence of
the Indus and the Panjnad. The length of this part is about 100 km. As this partof the section receives the water from the two rivers, the water level will not be too
low to support Platanista in winter season.

Section 5, between Sukkur and Guddu Barrages
The presence of the dolphin in this section was confirmed from boats, and theestimation of the population is made in the next section of this report.
Section 6, between Kotri and Sukkur Barrages
The river of this section is devided into two by Ghulam Mohammed Barragesituated about 12 km north of Kotri Barrage. During our trip we only confirmedthat the water is very scarce at just below Sukkur Barrage and at Sehwan Town,
which is situated slightly south of the midpoint between Sukkur and Kotri Barrages.
This indicates the probable absence of the dolphin in the upper part of the section.
However, as reported by Pilleri (1972), Mr. Roberts sighted one animal near thenorth of Ghulam Mohammed Barrage and suggested that the population of thedolphin in the section will be quite small. He also states that Platanista is not expected down stream of Kotri Barrage. These informations are considered to becorrect.
Section 7, from Emerson Barrage to Rasul Barrage (R. Jhelum) and to Khanki Weir (R.
Chenab)The section was studied on Dec. 16 at the bridge on theJhelum River connecting Khushab and Shahpur. This bridge is situated nearly at the midpoint between
the two barrages. At this point the stream was only from 100 to 150 m in widthand shallow. No dolphin was observed.
The other observation was made by the bridge crossing the Chenab River at
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

POPULATION OF INDUS DOLPHIN

85

the north of Jhang Maghiana This bridge is situated about 32 km upper streamof Emerson Barrage. In spite of the close distance from the lower barrage, thewater was only bout 70 m wide and was very shallow.
It is reasonable to expect no dolphin in this section, as already suggested by
Taber et al (1967) through the observation of the Ravi and Chenab Rivers.

Section 8, between Panjnad and Emerson Barrages
The Ravi River, a tributary of the Chenab River, was almost dry in this season(Dec. 16) leaving small ponds on the bed.
At the bridge connecting Multan and Muzaffargarh the stream was about
150 m in width and shallow, and no dolphin was observed on two occasions (Dec.
13 and 16). At Panjnad Barrage an information was collected on Dec. 17 froma fishermen telling that the dolphin is absent in this section.
No dolphin seems to be distributed in this section.
POPULATION IN THE SECTION BETWEEN SUKKUR
AND GUDDU BARRAGESThe field observation of the Indus River of this section was conducted in the period
from Dec. 7 to Dec. 12, using one (from Guddu Barrage to Begari) or two (fromBegari to Rohri) local boats. The size of the boats was about 2.8 m wide and6.5 m long. The boats cruised only in daytime with a stop for lunch, and usually
2 or 3 biologists watched the front direction of the boat while it was slowly rowed
down the stream.
On Dec. 7 1974 our boats left the camp on the east bank of the Indus situating
about 1 km south of Begari and arrived at Begari after 10 minutes. Then we tooka jeep at Begari for Guddu Barrage. At Guddu Barrage a boat was arranged andwe left that place at 1526 of the same day. After spending 2 nights on the boat, wearrived at the camp near Begari at 1900 of Dec. 9. On Dec. 9, as the observation
was stopped at sunset (1735), we could not observe the river of about 3 km. On
the next day (Dec. 10), we changed the boat and left the camp for Rohri, and arrived there at 0920 of Dec. 12 after spending 2 nights on the boats.
Though it was dry season, the width of the water was more than 200 m (estimated by eye) at any part of the section. However the width was small enoughand the weather was calm to keep all the width in the range of observation, except
when the water was split into two streams by sand bank or by island. On such cases
the observation was always made on the larger stream. The speeds of the current
was 2. 7 km/hr (Dec. 8), 1.5 km/hr (Dec. 9), or 5.0 km/hr (Dec. 11) at the shore on
the faster side of the stream.
The stream probably corresponds to the arm of the Indus running the eastside of Tappu Island reported by Pilleri (1970, 1972) was narrow and formed asort of canal, but was distinguished by fairly wide dry river bed. When Pilleri
visited Tappu Island in 1969 the east arm was much larger than the west, but it
reduced the size by 1972 to be as wide as the west arm (Pilleri, 1972). Possibly theSci. Rep. Whales Res. Inst.,
No. 27, 1975.

86

KASUYA AND NISHIWAKI

east arm attained the present condition by continuing the reduction of the size
after Pilleri visited there in 1972.
When the dolphin was found, the number of surfacings was counted for eachindividuals untill the animal passes the level of the boat. The distinction of theindividual was made considering the position and interval of surfacings, the size ofthe animal, and sometimes the swimming direction. Though most of the animals...
2:··--------------10 Dec.·- --- - - -- - - - - - - ··:

0

...

• I

__ _..12Dec .

....
.... --"'\.i ..
..:. •"'~ ~!

Miyassaghot

~

:.~

'.

\

~

'

11 Dec.-----GUB

..

.:i
',

Ferry(Kire)-·-~.._-.a..i&.-~Baiya

2.7km/hr...
8 Dec.

(

Keti Shah Bela)
Forest9 Dec.

060

120min-...... ',' ~.
• 0;~

BEG 7 Dec.

Fig. 2. Schematic figure of the lndus between Guddu Barrage and Begari (bottom)
and between Begari and Rohri (top). Direction of the stream and cruising hours areindicated instead of direction and goegraphical distance. Circle at the left sideof the river indicate the dolphin observed in cruising hours, and that at the rightthe animals observed while anchored. Numbers in parenthesis indicate theroughly estimated number of dolphins and number of surfacings in high density
area. BEG, Entrance of Begari Canal. GUB, Guddu Barrage. ROH, Rohri.

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

87

POPULATION OF INDUS DOLPHIN

seemed to be solitally, the area with high density was usually restricted to some deep
places, and the dolphin was scarce in the shallow and rapid stream. In some cases
many dolphins were found in a small area (e.g. 100 to 500 m in diameter), and thedistinction of the individuals was difficult. Even in these cases it was not sure if
the animals were forming a school or only assembling randomly at one spot. These
cases are shown with parenthesis in Table 1. In this table the lines are arranged in
the order of from the upper stream to the lower stream.

Interval of respiration
One juvenile dolphin (about 120 cm in body length), which was released after

5
50900, 6 Dec.

10
5
L..

0630, 6Dec.

.01930, 5Dec.

QJ

E 5
::J

z

1730, 5Dec.

10
510~'--~~~~;:;Jrn~~fil-_i_-1..._,_-L_l~53-0~.-5~D-ec~.~
5om·~~~~mi::.;~~L...'--'-----'--'--'-_L__L-l._J

0

10

20

30

40

50

60

70

80

Interval in seconds
Fig. 3.

Frequency of respiration interval of a juvenile P. g. indi.

the observation, was driven into a small inlet. And the interval of surfacing for
respiration was measured for 5 to 20 minutes at various hours of a day in order toget some clue on distinguishing the individual dolphins in the river. This resultis shown in Figs. 3 and 4.
The respiration interval showed wide variation from 4" to 81". Judging
from the inconspicuous peaks of the frequency, there seems to be three modes of
respiration interval. The one is at 20" or less, the second at about 40", and the thirdat 80". The frequency of the shorter interval had decreased in the second day.
This may be related to the decrease of excitement caused by the driving.
The observations of freely swimming animals give slightly different feature.
The mean interval of 9 respirations of a median size animal was 64". The 4 intervals of another median size animal ranged from 10" to 80" with a mean of 54". On
the other hand the mean interval of 12 respirations of one small animal, which was
swimming slowly at one spot, was 87". These informations suggest that the ratio
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

88

KASUYA AND NISHIWAKIof the longer interval may increase in the freely swimming animals, but that there
is no significant difference of the mean length of the interval between small andlarger animals. These features are also confirmed by the data reported by Pilleri(1970), in which the highest frequency was at 60", the next peak at 10", and theaverage was 42".
The respiration in night time was checked by the respiration sound. Thisshows that the respiration in the night is strongly biased to the shorter side.

Estimation of the population
The numbers of dolphins and of surfacings observed in the Indus between
1130
11 I I
I I I
11 I I I I 11
11 111II
0900
111 I I I I I 111 I 111 I I I 11 I 111
11 11
I I
0630
11 11 I I 1111 I 11 11 11 1111 11 11 I I I I I I
11
1930
111111111111 11111111 11 I
1730
II II 1111111 1111111 111111 Ill! I 1111 1111111
1530
11 I 11 1111 111 I I I II I 1111 11 I 111 Ill 11 111 I 11

0

100

200

300

400

500

600

700

800

II

11

900 1000 1100

Seconds
Fig. 4.

Respiration pattern of P. g. indi based on same data in Fig. 3.

Guddu Barrage and Rohri are shown in Table 1. The rough estimation of numberof dolphins we sighted is made, as shown bellow, from the total number of surfacingsand number of surfacings with the distinction of each individuals.
( 420+ 117) !~~ =256
However this does not give the total number of dolphins in the observed water,
because the overlooked animals are neglected. Accordingly the following method
was used in this study.
Excluding the count of the dolphins made at the places where the dolphinswere too concentrated to get reliable distinction of each individual, if the numberof dolphins are plotted against the number of surfacings of each individual, there
is found a tendency decreasing toward the left (Fig. 5). The frequency of animals
sighted twice is higher than that expected from the linear decrease. This may be
because, if the first surfacing is sighted, a larger attention is apt to be paid for thesecond surfacing. If the least square is used for the points between 1 to 5 surfacings,
the relationship y= -15.90x+82.l, where y is the number of dolphins and x is
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

POPULATION OF INDUS DOLPHIN

89

the number of surfacings, is obtained. However the frequency of the animals
with 6 or 7 surfacings does not come on the extended straight line. Possibly thesepoints will include the cases where two or more animals were erroneously considered
as a single animal. If each case with 6 (7 cases) or 7 (3 cases) surfacings is considered to be represented by 2 animals with 5 or less surfacings, the real number ofanimals discussed here should be 192, and above formula should be parallelly slided
up for (7+3) x2+5=4.0 and can be shown as follows.

80

(/)

''

c

''

E

'

_9-60

0
'O

0

'

,.

''

'

40

''

L.

<!>

•

.Q

E
20
:::J

''

'

''

'

•'''

z

00

2

3

4

'

•

•

5

6

7

Number of surfacings
Fig. 5. Frequency of the number of observed surfacings of each individual of
Platanista in the section between Guddu and Sukkur Barrages. Solid line indicates
y=-15.90x+82.l, and dotted line y=-15.90x+86.1.
TABLE 1. NUMBER OF DOLPHINS AND OF SURFACINGS OBSERVED
BETWEEN GUDDU BARRAGE AND ROHRI IN DEC. 1974
Date
7
7
8
9
9
7
10
10
10
10
11
11
12
12

Hours ofobservation
2°01 1
8130"
8°241
201
9°041
101
151
4°24'
19'
2°341
131
9°00 1

While at stopsurfacings

dolphins

20

10

61

28

124
(29)
(40)
66
(48)
29

53
(10)
(13)
24
(15)
9

90

48

30
420
(117)

12
182
(38)

Sci. Rep. Whales Res. Inst.,

surfacings

dolphins

8

51

1°421

Total

No. 27, 1975.

While cruising

6

3

3

2

90

KASUYA AND NISHIWAKI

y=-15.90x+86.l .................................... (1)The estimation of the number of overlooked dolphins is obtained from formula (1)as the value of y=86.l corresponding to x=o. The real number of dolphinsrepresented by one surfacing is obtained as follows.
182+7+3+86.l
420

0.661 ................................. (2)From this figure and total number of surfacings counted, the total number of dolphins should have been present in the observed area is obtained as follows.
( 420+ 117) x 0.661 =355.0 ............................... (3)This is the minimum estimate of dolphins in the Indus River between Guddu Barrageand Rohri. In order to estimate the total number of dolphins in the section, thenumber of dolphins in the unobserved area must be added.
There are three occasions where the smaller branch of the stream was not observed in spite of the possible presence of the dolphin. The length of time of thosecases were 20' (Dec. 8), 19' (Dec. 9), and 8' (Dec. 9). And the number ofspoutings
counted in the corresponding main streams were 0, 16, and 0 respectively. Assuming that the same number of dolphins are distributed both in the smaller branch
and larger main stream, the following number is calculated as the number of dolphinsmight have been present in the smaller stream.
(O+ 16+0) x 0.661=10.6 ................................. ( 4)However this can be a overestimate, because it is more reasonable to expect thesmaller number of dolphins in the smaller branch.
While the cruise of about 1 hour at the upper part of Begari, the observation
was not made because of the night time. The number of dolphins in this area canbe estimated from the number of surfacings counted in each one hour period atboth sides of this area. In the upper area there were observed 47 surfacings in one
hour, and in the lower area 64 surfacings. The mean is 50.9 surfacings. This shallrepresent
50.9x0.661=33.6 .................................... (5)However this estimation is also considered to be higher, because the water of theunobserved area was so shallow to have difficulty in cruising at night and the waterat both sides of this area was deeper.
Another estimation must be made for the area between Rohri and Sukkur
Barrage (about 2 km), where we could not cruise. As the number of surfacings
counted during the cruise of the same distance just above Rohri was 14, the numberof the dolphins in the areas between Rohri and Sukkur Barrage is obtained as follows.
14x0.661=9.3 ....................................... (6)Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

POPULATION OF INDUS DOLPHIN

91

By totalling the figures (3), (4), (5), and (6), 409 is obtained as the numberof dolphins between Guddu and Sukkur Barrages. As this figure is a maximum
estimation, more reasonable estimation can be between this figure and that of (3),or about 380 animals.ii

Size composition
Table 2 shows the number of dolphins and of the surfacings classified by thesize of the dolphin in the section between Guddu and Sukkur Barrages. The
juvenile animals smaller than about 140 cm in body length are classified into
" Small ", the large individuals presumably exceeding 180 cm and having theelongated rostrum were classified into "Large'', and the remaining dolphins into
" Intermediate ". Though the frequency of the three sizes are calculated by two
methods, one by the number of observed surfacings and the other by the number ofidentified individuals, there is observed no significant difference between the ratios
obtained by two methods. And the approximate ratio of small, intermediate, andlarge animals is 70 %, 20 %, and 10 % respectively. According to Kasuya (1972)
TABLE 2. RELATIVE ABUNDANCE OF THE THREE SIZES OF PLATANISTA
IN THE SECTION BETWEEN GUDDU AND SUKKUR BARRAGES
No. of surfacings

Small
Intermediate
Large
Total

No. of dolphins

no.

%

no.

%

299
89
40
428

69.9
20.8
9.3
100.0

97
32
17
146

66.4
21.9
11. 7
100.0

the female Platanista in the Brahmaputra River attains the sexual maturity at thelength between 170 and 200 cm and grows about 50 cm larger than the male, andthe rostrum is especially long in mature females. So it is considered that the individuals classified into large animal in Table 2 represent the mature females, andthe intermediate premature animals of both sexes and mature males. This meansthat the number of mature females is only 10 %of the total population. Even under
the unreasonable assumption that all the intermediate animals are sexually mature,
the percentage of the mature femals could be 15 % of the total population (an evensex ratio is assumed). In other words, the number of mature females is considered
to be surely less than 30 % and probably about 20 % of all the females in the section
between Guddu and Sukkur Barrages.
DISCUSSION

It is indicated that the dolphin is distributed also in the lower part of the section at
the north of Guddu Barrage. Though its length is almost same with that of thel) The fitness of Poisson distribution to the points in Fig. 5 is worse.
28 instead of 86 and the population between 269 and 309.
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

It gives overlooked animal

92

KASUY A AND NISHIWAKI

entire section between Guddu and Sukkur Barrages, the water in the former area
is more scarce than in the latter. So the total number of the dolphin in that part
must be smaller than that in the section between Guddu and Sukkur Barrages.
Possibly it will not exceed the half of the population in the latter section or about
200 animals. If the density of 2 animals in 3 km is applied for the area, the population can be about 100 animals. As the number of dolphins in the section between Kotri and Sukkur Barrages and that between Taunsa and Chasma Barrages
is almost negligible, the total number of Platanista in the lndus River is estimated tobe between 450 and 600. This is the lowest estimation has ever been made on thispopulation.
The ratio of sexually mature females to the total females was obtained in theformer section. The corresponding figures of delphinids are 69.1 % (calculated
from the data in Sergeant, 1962) in Globicephala melaena, 68.3 % (Kasuya et al, 1974)
in Stenella attenuata, and 58.5 % (Kasuya, 1972) in S. coeruleoalba. However these may
sometimes have a bias derived from the segregation of the animals by different growthstage (Kasuya et al, 1974). The real maturity ratio of female S. attenuata calculated
from the life table in Kasuya et al (1974, Table 24) is 59.0 %· These values are extremely high compared with that of Platanista in the Indus. A possible explanationof this phenomenon is the high mortality rate. If the mean age at sexual maturity is
known, the mortality is roughly estimated. Kasuya (1972) suggested on Platanista
in the Brahmaputra River that the age at the attainment of sexusal maturity infamales might be 10 years of slightly less as in the case of other toothed whales. If
9 years is applied for the age of the female Platanista in the lndus River at the attainment of sexual maturity, there can be the following equations.
H"e-µx dx
He-µx dx
M=I-e-µNo. of mature females
No. of immature females

Where µ is the annual mortality coefficient, M the annual mortality rate, x the ageof the animal in years. As the number of mature females is 20 %of the total females,
the annual mortality rate of 0.164 is obtained from the above equations. Even for
the assumption of improbably high ratio of mature females 30 %, the annual
mortality rate is 0.125 Though the mortality can change by the age of the animal,
the above calculation will give some idea on the mean mortality rate of Platanista inthe past. There is another problem in this estimation. The mean mortality thus
calculated is largely influenced by the change of the annual recruit, and in the strict
sence this value can indicate the real mean mortality only when the population isstable. In a decreasing population the mortality will be underestimated, and in a
increasing population overestimated. However, as the environment of the dolphin
has been changing worse as mensioned below, it is unreasonable to suspect theincreasing population. Accordingly the mortality estimated here seems to becorrect or an underestimation.
The next problem in analysing the population of the dolphin is the recruitment rate of Platanista. Kasuya (1972) showed that the parturition and mating
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

POPULATION OF INDUS DOLPHIN

93

seasons of Platanista in the Brahmaputra River are in winter season and that thecalves born in the early winter are weaned by the beginning of the next dry (winter)season. If this reproductive cycle is correct for Platanista in the Indus River, themean calving interval will be at least 2 years. Then the gross annual reproductive
rate is calculated as follows, assuming the even sex ratio and the ratio of mature
females 20 % obtained above.

Even in case of the unreasonably high ratio of mature female~ 30 %, the gross annual
reproductive rate is only 0.075.
The net reproductive rate is obtained as the difference of gross reproductive
rate and the mortality rate. It is 0.05-0.164=-0.114 and surely less than 0.0750.125= -0.050. Though this calculation is rough, it will be correct to conclude
that the population of Platanista in the Indus River has been decreasing at the rateof about 10% per annum. As the environment does not seem changing better, thedecrease of the population will be continuing.
One possible reason of the high mortality of this dolphin population can be
the unknown amount of the catch by local fishermen. Another and more importantfactor seems to be the change of the environment caused by the development of theirrigation system as indicated by various authors (Anon., 1973; Pilleri, 1972;
Roberts, 1972; Taber et al, 1967). The cause of the disappearance of the dolphin
in most of the part of the Indus seems to be related directly to the construction of
the barrages for irrigation system. The dolphin population, before the construction of the barrages, will have expanded the range of distribution in summer season
and retreated to main stream in winter season as in the case of Platanista in theGanges-Brahmaputra Rivers (Kasuya and Haque, 1972). However the construction of barrages blocked the seasonal movement and devided the population into
smaller units. Then, as most of the water is used for irrigation in each section, only
the subpopulations in few sections where the water level is not too low in winterseason have survived at the lower population level. It is told that the water decreases even at Begari so low in February not to allow the use of the local fishingboat. Begari situates at the middle of the section between Guddu and Sukkur
Barrages where most of the Platanista population is left. This condition will be badfor the survival of the dolphin. Furthermore, if the utilization of water for irrigation increases in future, the survival of this dolphin population will be impossible.
As the conclusion following two acts are suggested for the survival of Platanista
gangetica indi (Blyth, 1859). One is to stop the poaching of this dolphin. For thispurpose the educational propaganda will be needed to deny the erroneous conception on the medical efficiency of the dolphin oil. The other is to keep the water ofsome section above the certain level even in the winter season. This seems to bepossible only through the accomplishment of the economical utilization of the water
resources.
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

94

KASUYA AND NISHIWAKIACKNOWLEDGMENTS
Greatest thanks are due to Mr. T. A. Khan and Dr. R. L. Brownell Jr., who gave
us valuable suggestions in planning our research trip to Pakistan and kindly checked
the draft. Dr. F. Yamasaki, Dr. T. Kamiya, and Dr. A. Takemura are also acknowledged for their cooperations offered for our study.
REFERENCES
ANDERSON, J., 1878. Anatomical and zoological researches comprising an account of the zoological results of the two
expeditions to western Yunnan in 1868 and 1875 and a monograph of the two cetacean genera, Platanista andOrcella, vol. I. Bernard Quaritch, London, 550 pp.
ANONYMOUS, 1973. Indus dolphin or susu, Platanista indi (Blyth, 1859). in Red Data Book vol. 1 (Mammalia):
2. Inst. Union Conserv. Nat., Morges.
BLYTH, E., 1859. On the great rorqual of the Indian Ocean with notice of other cetals, and of the sirenia
or marine pachyderms. J. Asiat. Soc. Bengal, 28: 481-498.
KASUYA, T., 1972. Growth and reproduction of Stenella caeruleoalba based on the age determination bymeans of dentinal growth layers. Sci. Rep. Whales Res. Inst., 24: 57-79.
KASUYA, T., 1972. Some informations on the growth of the Ganges dolphin with a comment on the Indus
dolphin. Ibid., 87-108.
KASUYA, T., and A.K.M. AMINUL HAQUE, 1972. Some informations on distribution and seasonal movementof the Ganges dolphin. Ibid., 24: 109-115.
KASUYA, T., N. MIYAZAKI, and W. H. DAWBIN, 1974. Growth and reproduction of Stenella attenuata in the
Pacific coast of Japan. Ibid., 26: 157-226.
PILLER!, G., 1970. Observations on the behaviour of Platanista gangetica in the Indus and Brahmaputra
Rivers. Investigations on Cetacea, ed. G. Pilleri, 2; 27-69, Berne.
PILLER!, G., 1972. Field observations carried out on the Indus Dolphin Platanista indi in the winter of 1972.
Ibid., 4: 23-29, Berne.
PILLER!, G., and M. GIHR, 1971. Differences observed in the skull of Platanistagangetica (Roxburgh, 1801)
and indi (Blyth, 1859). Ibid., 3(1): 13-21, Berne.
ROBERTS, T.J., 1972. A brief examination of ecological changes in the province of Sind and their consequences on the wildlife resources of the region. Pakistan J. Forestry, 22: 89-96.
SERGEANT, D. E., 1962. The biology of the pilot or pothead whale Globicephala melaena (Traill) in Newfoundland waters. Bull. Fish. Res. Board Canada, 132: 1-84.
TABER, R. D., A. N. SHERI, and M. S. AHMAD, 1967. Mammals of the Lyallpur region, West Pakistan.
J. Mamm., 48(3): 393-407.

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

PAST OCCURRENCE OF GLOBICEPHALA MELAENAIN THE WESTERN NORTH PACIFICTOSHIO KASUYA
Ocean Research Institute, University of Tokyo, Tokyo

ABSTRACTSix skulls of G. melaena excavated at the northern Sea of Japan and at
the central part of the Pacific coast of Japan indicate that the species waspresent in the western North Pacific at least untill about lOth century.
However, the absence of any indication of the presence of this species in thecollection from recent fauna and in the modern Japanese whaling statistics suggests that the species might have been extinct from the North Pacific or that asmall population is surviving in some area of western North Pacific which havenot been studied by biologist.

INTRODUCTION
The genus Globicephala Lesson, 1828, is constituted by two species G. melaena (Traill,
1809) and G. macrorlrynchus Gray, 1846, (Fraser, 1950; Van Bree, 1971). They havebeen known, except in the North Pacific, to show antitropical distribution, namely
the former species in colder waters and the latter in the warmer waters (Davies,
1960, and 1963).
In the North Pacific, G. scammonii Cope, 1869, and G. sieboldii Gray, 1846,
were reported from Mexican waters and from Nagasaki in the southern Japan
respectively, which were later concluded by Van Bree (1971) to be the synonyms of
G. macrorhynchus. Though it was denied recently (Nishiwaki et al, 1967), there were
published several reports telling the distribution of G. melaena in the Japanese coastal
waters, and produced a long history of confusion on the zoogeography of globicephalid species. So, it will be worth to have, in the below, a review on this problem
for the better understanding of the biological meaning of the present study.
Andrews (1914) is the first scientist who used G. scammoni for the pilot whale in
the Pacific coast of Japan. This was based on an animal taken at Ayukawa (38°18'N, 141°30'E), but he gave no biological reason distinguishing it from G. sieboldii
reported by Gray (1846). Then Nagasawa (1916) used in his list the name G.
scammoni for the pilot whale in the Japanese waters, considering that G. sieboldii
should not be trusted because it was based on a drawings of a young pilot whale
reported by Temminck and Schlegel (1844). This was followed by Takashima(1931 ). However, Ogawa (1937) listed G. melas in his list ofodontoceti in the coastal
waters of Japan, and showed a photograph of so-called G. melas (236 cmQ") taken at
Shiogama (38°19'N, 141°02'E) on 28, May, 1935. In this list he tentatively considered G. melas as the synonym of G. sieboldii. Furthermore in the same
Sci. Rep. Whales Res. Inst.,

No. 27, 1975, 95-110.

96

KASUYA

list, he listed G. scammoni, based on two specimens collected at Ayukawa. And heconsidered the two Japanese names "Naisagoto" and " Magondo" correspond to
G. melas, and " Shiogoto " and " Tappanaga " to G. scammoni. Though he did notgive in his report, the reason of the identification, it is written in his another book
(Ogawa, 1950). As this is the start of the erroneous story telling the presence ofG. melaena in the Japanese coastal waters, some important part is translated here.
[p. 65] I decided to spend the August of this year [i.e. 1935] for the 2nd trip
to collect the specimens ... at Shirahama in Chiba, Taiji in Kii, and Nagasaki in Kyushu .... [p. 76] At Taiji I got 2 Globicephala (380 cm~, 470 cmd'),measured their bodies, counted the number of bones, observed the pigmentation, and sent only the skull(s) back.... They were commonest at Taiji andcolled " Magondo " .... They seemed to have no difference from G. melas(Traill) ... and strengthened my belief that "Magondo (or Naisagoto)"
indicates this species. One of the reasons for it is ... [p. 77] the presence ofanchor shaped pale area in the chest region. I had been interested in thisfact, since I observed this at Shiogama on 2 or 3 Globicephala . ... In the foreigncountries the white or pale area of the shape is reported only on G. melas(Traill). Though Cuvier and True say that it extends to anus along the
midline of the body, that of my specimen ends at the middle of the ventral
region and shows slight difference. There are recorded three different namesor" Naisa '', "Shio '', and "Onan" for" Goto (or Gondo)" [Pilot Whale]in the Japanese oldest book on cetaceans (Yamase, 1760).... Among which thenames "Naisa" and "Magondo" now used at Taiji are considered to besynonyms, and "Onan", which is equal to "Okigondo ", ... is supposed to indicate Pseudorca crassidens (Owen) .... [p. 78] The still unknown is " Shio ".
I was told at Taiji that this species is large in body size, has long flipper, has
white area at the posterior of the dorsal fin, and also called by the name of"Tappanaga" [=long flipper]. And it was also told that the number ofcatch of this species is few, and caught only in the season(s) of equinox....
Before this information I had guessed that " Tappanaga " may correspond to
G. scammoni Cope (Pacific black fish in U.S.) .... When I arrived at Ayukawa
after receiving the news of the catch of" Tappanaga '',the animal had been already processed. So I identified the species only with a part of skeleton. However, Mr. Hasegawa had observed, by my request, the condition of the anchor
shaped pale area on the chest and reported that it was absent. He gave mea photo of the dorsal view of other individual (Fig. 22) .... There was clearlyobserved a white fleck behind the dorsal fin of which I heard at Taiji ....
[p. 79] The photo, shown in the journal of the museum, of a G. scammoni Cope... in the collection made by Andrews in Japan ... has no pale area on thechest. From these fact I considered that "Shiogoto (Tappanaga)" corresponds to G. scammoni Cope.
Above citation shows that Ogawa's misidentification started in the erroneous
idea that some of different common names in different localities must indicate different species. Then he overlooked the fact that all of his samples were collected in
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

G. MELAENA IN THE NORTH PACIFIC

97

the area and seasons influenced by the warm Kuroshio current, and the importanceof the osteological characteristics and· of the external proportions by which the possibility of G. melaena might have been eliminated from his samples, but he put theweight on less reliable pigmentation. And he reached the odd conclusion that G.
melaena is distributed in the south and G. scammonii in the north. This conclusion(Ogawa, 1937) was followed without further confirmation by Kuroda (1938, 1953),
Noguchi (1946), and Nishiwaki (1957). Then, Nishiwaki et al (1967) concluded,
based on the specimens collected by themselves in the Pacific coast of Japan in thesummer season and on the two of Ogawa's specimens, that there will be distributed only G. macrorlrynchus in the area in which the distribution of G. melaena was
expected by former reports. Though Nishiwaki et al (1967; and see Nishiwaki,
1967; and Rice and Scheffer, 1968) found no significant difference between G.
macrorhynchus in the Japanese coastal waters and G. scammonii, they retained the conclusion on the taxonomy of these two species. And now it has become a generally accepted idea that G. melaena is not distributed in the North Pacific (Riceand Scheffer, 1968; Van Bree, 1971).
In 1974, however, I had a chance to examine the specimens excavated by Dr.
Oba of Hokkaido University and his colleagues from two archaeological sites ofOkhotsk culture on Re bun Island in the northern Sea of Japan. Though they arecomposed of many fragmental bones of odontoceti and mysticeti, the most striking
is the presence of skulls of G. melaena and the absence of specimens identifiable to G.
macrorhynchus. This led me to have a discussion on the possibility of existence ofG. melaena in the western North Pacific.
MATERIALS AND METHOD
Four of 6 skulls of G. melaena referred in this study were excavated from the Kabukai
A-site on the east coast of Re bun Island (45°20'N, 141°E ). Though, 14 C datingof charcoal gives 1530±70 B.P. (TK 157) for the lower strata, the date of this site
is suspected from known archaeological data to be between 8th century (lowest
stratum) and 12th century (upper most stratum) (Oba and Ohyi eds., in press).
As shown in Table 1, this site contained the remainings of large whales and ofvarious delphinid species. However, most of them were found in the condition
almost completely broken into small pieces. Other than the remains shown in
Table 1, there were found 174 vertebrae of Phocoe noides, 92 of Lagenorlrychus, 7 of
Phocoena, 29 of Globicephala, and several vertebrae of large cetaceans. As the excavation was made in a restricted area compared with the expected range of thesite, the accurate ratio of the cetacean species consumed by the people is notknown at present. But it is safe to say that the delphinid species in the site are,with the exception of Pseudorca, composed of the boreal species. Table 1 shows thatthe number of globicephalid animals found in the upper strata is higher than thatin the lower strata. This may suggest the improvement of the fishing technology.
Though the fragments of Globicephala skeletons were found in various strata of
Kabukai A-site, most of them were unable to be identified the species. Only 4 skulls,
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

98

KASUYARKA 3658, 3983, 3987, and 6054, were identifiable, among which RKA 3983 wasyoung and badly destroyed to measure. The stratum containing the present specimens situates approximately at the middle of upper strata, and had been depositedin a pit representing a ruin of a house of that date. They were found in a stone
mound together with 6 skulls of Globicephala sp. of various stages of growth andconditions of preservation, 1 Lagenorhynchus skull, 1 left tympanic bulla of humpback whale, and a radius of black right whale ( ?). Further description and interpretation of this site are reported in Oba and Ohyi (eds., in press).
Another skull of G. melaena (no. 675) was excavated by Dr. Oba and his colleagues at an archaeological site at Motochi on the west coast of the same island.
Though the archaeological study of the site is not finished yet, it is considered tobelong to the age between l lth and 13th century (Ohyi, 1972).
The 6th skull of G. melaena was found by Mr. M. Yoshimura and offered mefor taxomic study. This specimen was excavated from an alluvial silt deposit by
TABLE 1.

MINIMUM NUMBER OF CETACEANS REPRESENTED BY
BONE FRAGMENTS IN KABUKAI A-SITE 4>

StrataUpperstrata1>

Middlestrata2>Lower
strata 3>

Total

Globicephala
Lagerwrhynchus obliquidens
Phocoena phocoena
Phocoenoides dalli
Pseudorca crassidens
Physeter catodon
Ziphiidae sp.
Megaptera novaeangliae
Balaenoptera borealis
B. acutorostrata
Eubalaena glacialis

13
3
1

4
2
1
3

2
4

19
9
3
5
3
1
1
6

+
3
1
3

2

2

1

+

2

1) Brom surface to fish bone stratum II/III,
2) From fish bone stratum III/IV, 3)
From fish bone stratum IV to VI,
4) Calculated in each smaller strata. Vertebrae
other than cervical, and bones in flipper are not included, but shown by " + ".

the Hekuri River near Tateyama (34°59'N, 139°5l'E) at the entrance of Tokyo
Bay. This strata belongs to the N uma deposit, and the date of the skull is considered
to be equal with the date of fossil coral and of fossil oyster collected at the same
stratum of same location (Mr. Yoshimura, pers. comm.). The 14 C dating of thecoral and of the oyster gives 6,340±140 B.P. and 6,430±130 B.P. respectively(Araki, 1973).
For the osteological comparison, the measurements of 25 skulls of G. macrorhynchus collected on the Pacific coast of Ja pan were used. They are 7 skulls in thecollection of National Science Museum (Tokyo), 5 skulls of Ogawa's specimen in
the collection of the same museum, 1 skull collected at Ayukawa (no date), 3 skulls
collected at Choshi (35°42'N, 135°56'E, Dec. 1966), and 9 skulls collected at Ararion the Pacific Coast of Japan (39°49'N, 137°46'E, June 1965 and 1967). The lastSci. Rep. Whales Res. Inst.,
No. 27, 1975.

G. MELAENA IN THE NORTH PACIFIC

99

13 skulls are kept in the Ocean Research Institute, University of Tokyo. The skull
measurements of 4 G. scammonii (=G. macrorhynchus) specimens in the collection of
Los Angeles County Museum (EDM F 298, WAW 46, WAW 47, WAW 48) and one
skull kept in the Smithsonian Inst. (USNM 3076), and the measuremnts of 13
skulls from the North Atlantic in the collection of Smithsonian Inst. (USNM nos.
21572, 257412, 257413, 279366, 257414, 37264, 241182, 37261, 22592, 37263,
395712, 22571, 22570) are used here. The measurements of the type specimen of
G. scammonii (USNM 9074) were provided by Dr. R. L. Brownell. Measurementsof 9 skulls of G. macrorhynchus cited from Fraser (1950) and Van Bree (1971, =G.
sieboldii) were used in Figs. 1 and 2.
The measurements of G. melaena were obtained on 21 skulls in the collection ofSmithsonian Inst. (USNM nos. 395373, 395365, 395372, 395364, 395363, 395361,
395362, 395360, 395357, 16298, 20981, 21118, 14418, 14360, 20958, 303018, 12098,
12097, 12099, 12100, 20957, and EDM 1000) and on 4 skulls in the collection of Los
Angeles County Mus. (four EDM specimens).
The catch statistics of Globicephala in the Japanese coastal waters were compiled from Geiryo Geppo [the monthly report of whaling operation] of the period
from 1949 to 1952, when most intensive fishing for Globicephala was operated because of the social demand for the food. This report was originally presented to thegovernment by the owners of the small whaling boats, but now is kept in the
Whales Research Institute in Tokyo. The position of the catch is represented inthis study by the position of the whaling station, and the number of the catch isshown in each areas obtained by combining several nearby stations.
OSTEOLOGY
The difference between G. melaena and G. macrorhynchus is found, on the skull, in the
shape of premaxillae on the dorsal surface of the rostrum (Fraser, 1950). In thelatter species the lateral margins of the premaxillae expand widely at the anterior
region of the rostrum to cover, entirely on adult individuals, the maxillae of thatpart. However, on _G. melaena this expansion is very weak and the maxillae arewidely exposed, even on the adult animal, on the dorsal surface of the anterior partof rostrum. Though Van Bree ( 1971) says that rostrum is elongated in G. melaena
than in G. macrorhynchus, there was found no difference in the length of rostrum between the two species (Fig. 2.). The number of teeth is fewer and the size of
tooth larger in G. macrorhynchus.
The five Globicephala skulls from Rebun Island and Tateyama are, except one
juvenile, those of adult individuals, and the lateral margins of the maxillae and ofpremaxillae of the 5 skulls are almost intact to allow the accurate comparison. And
their feature coincides with that of G. melaena in the above characteristics (PLATES).
The number and size of alveoli are also rather close to that of G. melaena (Table
2). As shown in Fig. 1, the measurements of rostral region show the difference between the two species, and G. macrorhynchus from the Pacific coast of Japanshows the perfect coincidence with those of the same species in other areas. HowSci. Rep. Whales Res. Inst.,
No. 27, 1975.

KASUYA

100

ever the measurements of 5 measureable skulls from Rebun Island and Tateyama
come in the range of G. melaena in the North Atlantic. From these reasons the present Globicephala skulls are classified into G. melaena, and it is concluded that G. me-

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Length of skull in cm
Fig. 1. Scatter diagram of the measurements of rostral region of Globicephala. A:
Rostrum basal width. B: Rostrum width at middle. C: Premaxillae width at
middle ofrostrum. Open circle: G. macrorhynchus (Without bar indicates animalfrom] apanese coast, bar at right that from Ca1ifornia coast, and bar at left that fromother seas). Closed circle: G. melaena (Bar at right indicates animal from Chili,
bar at left that from North Atlantic, and no bar no history). Cross mark: G.
melaena from Japanese coast.

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

101

G. MELAENA IN THE NORTH PACIFIC

0

•

0

5 35
c

§ 30
L.
.......

(/)

0
L.

0

25

..c
.......
0

g120

0

0

_.CIJ

0

"

0

40

45

50

55

60

70

65

Length of skull in cm
Fig. 2. Scatter diagram showing the relationship between length of skull and lengthof rostrum. For marks see Fig. I.

TABLE 2.

SKULL MEASUREMENTS OF G. MELAENA FROM JAPANESE COAST (MM)Specimen no. and Locality
Measurements
1. Condylo-basal length
2. Rostrum length
3. Rostrum basal width
4. Rostrum width at middle
5. Premaxillae width at same point
6. Maximum width of premaxillae
distally
7. Maximum width of premaxillae proximally
8. Width of rostrum at the level of no. 6
9. Width across preorbital angles ofsupraorbitals
10. Width across postorbital angles ofsupraorbitals
11. Width of skull at orbits
12. Zygomatic breadth
13. Width of braincase across parietals
14. End of rostrum to blowhole
15. Length of upper tooth row (to end ofpmx.)
16. Number of upper teeth* Last alveolus rudimental.
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

675
KRA3658
Kabukai Motochi

RKA6054
Kabukai

Awa Mus.
Tateyama

RKA3987
Kabukai

617
315
224
167
138

619
310
223
166
146

623
331
119x2
87x2
76x2

633
329
235
171
151

681
352
257
200
170

140

148

146

151

172

148

156

158

174

168

170

90x2

171

205

352

178x2

375

399

384

415

440

349
375
285
394
Ll63,
Rl54

371
416
273

405

175x2

359
288
407
Ll68,
Rl66
LIO, RIO

401
L201

L175

L9

Lll*

299
466
Ll90,
R205

?:

.

~ ~~

<c ~

......

.,.'1 ~.""::tl"'

~~

;s

~

::tl

sample size
range
mean
standard error
sample size
range
mean
standard error
sample size
range
mean
standard error
sample size
range
mean
standard error

3. Rostrum basal width (o/o)

4. Rostrum width at middle (o/o)

5. Premaxillae width at same point

11. Orbital width (o/o)1) Skulls more than 540 mm are selected.

16. Number of upper teeth

1022.1-28.5
24.9
0.65

5
26.8-29.4
27.6
0.45
5
22.4-25.0
23.9
0.44

61.1-65. 7
62.9
0.76
14
9-11
9.9
0.20

10

10
27 .4-31.9
29.2
0.49

5
36.0-38.2
37 .1
0.42

4
56.4-59.5
58.2
0.65
4
9-11
10.0
0.41

10
37.3-40.l
38.8
0.28

5
50.1-53. l
51.6
0.50

10

580-712633. l13.76
1049.8-54.4
51. 7
0.46

5
617-681
634.6
11.92

2) Both sides are dealt separately.

sam pie size2>
range
mean
standard error

sample size
range
mean
standard error

2. Rostrum length (o/o)(o/o)

sample size
range
mean
standard error

G. melaena
Chili

G. melaena
Japan

14
21.5-28.2
25.2
0.48
10
59.2-66.7
63.1
0.73
16
9-12
10.3
0.24

14
31.2-40.2
36.9
0.71
14
27.3-33.0
29.8
0.45

14
590-695
619.9
11.80
14
47 .2-51.2
50.2
0.31

G. melaena
N. Atlantic

13
29.6-40.4
33.8
0.92
12
62.5-68.6
65.8
0.63
26
7-10
8.0
0.20

13
30.5-40.4
34.8
0.86

13
37 .6-44.6
40.4
0.53

13
544-680
611.8
12.79
13
48.0-52.2
50.4
0.35

G. ma.crorhynchus
N. Atlantic

24
26.7-47.2
32.3
1.02
21
62.2-78.5
68.7
1.02
48
6-9
7.8
0.14

24
540-748
621.6
11.35
24
47 .2-51.0
49.2
0.21
24
35.6-48.6
41.3
0.76
24
28.3-49.0
34.1
l.03

G. macrorhynchus
N. Pacific

COMPARISON OF SOME IMPORTANT SKULL MEASUREMENTS OF TWO SPECIES OF GLOBICEPHALA

1. Condylo-basal length (mm)1>

fABLE 3.

><

>

~
rJ:J
c::

0
t.o

G. MELAENA IN THE NORTH PACIFIC

103

laena was present at the approximate lOth century in the northern part of the Sea ofJapan.
SEASONAL FLUCTUATION OF THE RECENT CATCH
Fig. 2 shows the seasonal fluctuation of the catch of Globicephala in the coastal watersof Japan, compiled from the monthly report of the catch in the years from 1949 to
1952. In these years many small whaling boats operated in the Japanese coastal
waters for minke whale, ziphioid whales, killer whale, and pilot whale. The fish~

-5
I

I

I

I

'o

-5
~o
I

/

I

LLJ_J

lo

z

-150

-..:

~

-100

\
='

-200

-so

\

0

0

0

("')

50
LJ,,~L..L--'---L..L--'---L..L_L_jQ

r---11

I

1

I

I

I

I

I

L.i___i_j

-5
I

_JQ

JFMAMJJASOND

130°

~30
JFMAMJJASONDO

140°

Fig. 3. Seasonal fluctuation of the catch of Globicephala in the years from 1949 to
1952. The number of catch is shown by squares, and the months of whaling
operation by holizontal line. Closed circle indicates whaling station, and star
the position of the occurrence of G. melaena. Whaling area is shown by Roman
numerals.

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

104

KASUYA

ing ground was approximately within the range of 50 nautidcal miles (90 km) fromthe coast (Omura and Sakiura, 1956). Though they prefered minke whale, theycaught other species also, because the demand of whale meat for human consumption was high. Accordingly the seasonal and geographical fluctuation of the catchof Globicephala in these years is considered to reflects the abundance of this species.
The total number of whaling boats of this type operated was 74 (1949), 66 (1950),
68 (1951) and 69 (1952). And the annual number of Globicephala catch was 760
1949), 667 (1950), 591 (1951), and 307 (1952). It is not analyzed yet whether thedecrease of the annual catch is the reflection of the depletion of the population or
the reflection of economical factors. Probably both of the factors will have influenced on it.
As shown in Fig. 3, the catch in the central and northwest regions (except
areas VII to IX) shows, in general, the two peaks one in the autumn and theother in early summer. And the interval between the two peaks is shorter in thenorthern areas. Though the peak of early summer is inconspicuous in the areas VI
and VII, this seems to be the influence of the catch of minke whale. According toOmura and Sakiura (1956), these area is a good whaling ground of minke whale
and the season is from January to July with a peak in May. This will have aeffect to lower the peak of Globicephala in early summer.
However, in the areas VIII and IX, the catch of Globicephala shows only one
peak. In these areas the season of minke whale is between June and October witha peak in July (Omura and Sakiura, 1956). Accordingly the season of the peak ofcatch of Globicephala in these areas will not be biased, even if the total number ofcatch is smaller compared with the number of migrating animals.
This fact suggests that the area where Globicephala densely distributes moves
from the south to the north in the early summer, and after arriving at the latitude of 43°N in July and August, it again moves to the south to arrive at Kii and
Shikoku area (32°N to 34°N) in November and December. Possibly this seasonal
movement of the whaling ground reflects the migration of one species of Globicephala in the Pacific coast of Japan. But as suggested by Fig. 3, some Globicephalaindividuals seem to stay in all seasons in the Pacific coast of the south west regionof Japan (areas I to IV).
The catch of Globicephala in the sea of Japan and in the northern coast of K yushu is too scarce to discuss the seasonal fluctuation. But it is safe to say that thecatch is restricted to the summer season in spite of the longer operation of whaling
in this region. As the number of the catch of Globicephala is so few, there may arise
a question if the Globicephala reported by the fishermen from the Sea ofJapan represent real Globicephala. However, there is no doubt in the existence of the genus
in the Sea of Japan, because a school of Globicephala was sighted by myself on 28,
August, 1971 at the center of the Sea of Japan (39°04'N, 134°36'E). The surface
water temperature at the spot was 24.5°C.
In the Okhotsk Sea, the Japanese whaling boats for small whales operated in
the seasons from March to November. But there are no catch of Globicephala recorded in the statistics I analyzed. Omura and Sakiura (1956) showed that theSci. Rep. Whales Res. Inst.,
No. 27, 1975

G. MELAENA IN THE NORTH PACIFIC

105

season of minke whale in this region lasts from March to July with a peak in May,and the same result was obtained even from my data. After the season of minke
whale, they caught Berardius and Orcinus in the Okhotsk Sea. As the capture of
Globicephala is not considered to be more difficult than that of Berardius or Orcinus
even if the size is slightly smaller than the latter, they must have captured Globicephala
if it had been distributed in the sea in that season. According to the personal communication of Mr. S. Miyoshi who has been operating the whaling for more than
20 years mainly in the Okhotsk Sea, he have sighted no Glogicephala in the Okhotsk
Sea.
These informations strongly suggest that Globicephala does not migrate, in theseasons from March to November, to the southern part of Okhotsk Sea where theJapanese whaling boats for small whales operate.
DISCUSSION

On the species of Globicephala constituting the catch in the Pacific coast of Japan,
the specimens of known locality give the information. All the known records ofGlobicephala in this area, i.e. 6 specimens collected at Arari in June 1967 (Nishiwaki
et al, 1967), external measurements of 16 animals and 13 skulls collected at Taiji
33°38'N, 135°56'£) in July and August 1969, 1 Ogawa's specimen caught at Shirahama (34°54'N, 137°53'£), 3 animals in a school stranded at Choshi (35°42'N,
140°52'E) in December 1966, 2 Ogawa's specimens presumably collected at Shiogama in May 1932 and 1935 (Ogawa, 1937; Ogawa, 1932 cited in Kuroda 1938),
2 Ogawa's specimens presumably collected at Ayukawa (Ogawa, 1937), and 1 skull
and skeleton in National Science Museum in Tokyo killed at 39°28'N, 142°19'£
in October 1953, are represented by G. macrorhynchus. These animals were collectedfrom the animals representing both the early summer and the autumn-winter peaks.
The comparison of fishing season and oceanographical condition gives anothersupport on the identity of the species. The boundary of the front of warm Kuroshio Current is indicated by the 17°C surface isotherm (Uda, 1954), and in August
the boundary go up to the latitude of 43°N (Uda and Nasu, 1956; Miyazaki et al,1974) or to the northern limit of the whaling ground of Globicephala analysed here,
and retreats in March to 35°N (Miyazaki et al, 1974) or to the approximate northern
limit of Globicephala ground in winter season. From these informations it is concluded that all the catch of Globicephala in the Pacific coast ofJapan is constituted by
G. macrorhynchus, and that the species lives in the Kuroshio watermass and moves
north and south in accordance with the seasonal movement of the boundary of
Kuroshio front. Accordingly the range of normal distribution of G. macrorhynchus
is expected not to extend into the cold watermass.
There is obtained no data directly indicating the species of Globicephala caught
by recent whaling in the Sea of Japan. However, the existence of the skull of G.
sieboldii [=G. macrorl!)'nchus] caught at Nagasaki in western Kyushu, the fact that
Globicephala was caught in the same season with that of Pseudorca (areas XI, XII
and XVI), one sighting record mentioned in the former section, and the presence
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

106

KASUYA

of warm Tsushima Current in the Sea of Japan (Uda, 1954) strongly suggest that
there is distributed G. macrorlrynchus in the summer season. So I consider that all
the Globicephala appeared in the statistics is represented by G. macrorhynchus.
As the northern extension of Tsushima current is faint in the north western andnorthern coast of Hokkaido, there will be expected no migration of G. macrorhynchus.
Though the water in the eastern part of northern coast of Hokkaido is high in thesummer season (Uda and Nasu, 1956), it is restricted to a small locality and will
not result in the migration of G. macrorlrynchus.
From the above discussions it is supposed that G. melaena found on the Rebun
Island must have been caught in the seasons from late autumn to winter.
Though at present there is no data to have a discussion on the present statusof population or on the migration of G. melaena in the North Pacific. There can be
two possible hypotheses on that problem. The first is to consider that G. melaena
is extinct in the Okhotsk and in the Sea of Japan. Judging from the carvings excavated at Sakhalin and at Nemuro in eastern Hokkaido (Tsuboi; 1908 and 1909;
Yahata, 1943), it is sure that the natives of the Okhotsk coast including the people
on the Rebun Island of that date had the techniques to catch even some mysticeti.
However, as they seems to have used only hand harpoon and small canoe for thewhaling, it is difficult to consider that the G. melaena population was exterminatedonly by the whaling of that date. If the population have been exterminated, weshould expect the interspecific competition between G. melaena and other organisms.
One of the possible competitative mammal species other than man seems to be
Phocoenoides dalli (True, 1885). On this hypothesis, a possibly small population of
G. melaena which might have immigrated to the North Pacific is considered tohave failed in establishing its niche obstacled by the dominant population of Phocoenoides. On the date and the route of the immigration of G. melaena, there can be
two possibilities. The first is, as suggested by Davies (1962), through the easternequatorial Pacific in the Wurm glacier stage. The second is from the North
Atlantic through the Arctic Ocean in more recent age. The vegetation detectedby the pollen analysis and the Alpine timberline show that the climate of all
the northern hemisphere was warmer in the age between 8,000 B.P. and 2,000
B.P. (Atlantic and Sub-boreal periods) than the present (Deevey and Flint, 1957),
and the air temperature at Oslo was suggested to have been higher by 3°-4°C(Tsukada, 1974). The similar result was obtained by Emilliani (1964) by theoxigen isotopic analysis of the Caribbean Sea deposit. Though the exact changeof water temperature in the Arctic Ocean of that date is not known, above climaticchange seems to have provided the passages for G. melaena to North Pacific.
The other hypothesis is to consider that G. melaena is surviving at least in
the western North Pacific and its adjacent seas, and that it lives in the areasand seasons when the Japanese whaling operation or scientific study was notconducted. For this hypothesis the presence of the peak of density of globicephalid species in November in the southern coast of Hokkaido (Kasuya, 1971)
will give some suggestion. As Orcinus 'is 'recorded separately in his aerial sighting records and the water is too cold to expect Pseudorca, the possible species
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

G. MELAENA IN THE NORTH PACIFIC

107

recorded there can be Globicephala especially G. melaena. According to thepersonal communication of Mr. K. Sasaki, who is one of the fishermen operating
the Phocoenoides hunting in January to March in the coast of Iwate and Miyagi prefectures (Pacific coast, 38°N to 40°N in latitude), they sometimes find Globicephalawhile they are operating Phocoenoides hunting. The surface water temperature atthose positions is told to be 7°C to 8°C. As Phocoenoides is mostly found in the watercolder than l6°C and above this temperature all of my sightings of Globicephala andother temperate or tropical species (e.g. Pseudorca, Tursiops, and Stenella) have occurred (Kasuya, umpublished, Miyazaki et al, 1974), the sighting of Globicephalamensioned in the above can represent G. melaena. But this problem is left to be confirmed. Though the information on the globicephalid founa on the Russian coastis desirable, there seems to be no reliable study (Dr. Yablokov, pers. comm.).
Though it is not the purpose of this study to have a discussion on the Japanesecommon names indicating Globicephala, some comments are added here. At first
we must remind that Japanese fishermen can use different names for the animals
of various growth stages as usually found in some species of fish, and that the namecan be modified even by the fishing season. Ui (1942) reported a case that the pilot
whale caught at Taiji in the calm season of November and December is called"Nagi [ =calm]-gondo ". It is, of course, expected that the fishermen at different places use different common names. However, it will be easily concluded
that " Magondo [=common pilot whale] " used by the whalers at Taiji for Globicephala caught at Taiji indicate G. macrorhynchus which were formerly misidentified
as G. melaena. The whalers at Taiji have another name " Tappanaga " indicating
so-called another Globicephala species. According to a former whaler Mr. Mizutani(pers. comm.), " Tappanaga " is rare at Taiji but commonly caught in the summer
season in the northern part of the Pacific coast of Japan. He tells that " Tappanaga " has, as the name indicates, longer flipper than that of " Magondo ", and thesize of the body of" Tappanaga" is usually larger and the content of oil is fewer.
The distinction of the two is told to be difficult. However, as the relative length offlipper of G. macrorl?Jnchus can be expected to increase accompanied with the growthof the animal (Sergeant, 1962), the above distinction is not reliable. Furthermore,
the fishing season of " Tappanaga " in northern Japan corresponds to the seasonof G. macrorhynchus mentioned above. Accordingly it is concluded that the name" Tappanaga " does not indicate G. melaena but may indicate old individual of
G. macrorl?Jnchus. I have no new information on "Naisa" or "Naisagoto" andon " Shio " or " Shiogoto " which had been considered to be the synonyms of"Magondo" and" Tappanaga" respectively (Ogawa, 1937). But I suppose thatthey also indicate G. macrorhynchus because they were reported to be found near
Taiji and the body length and the swelling of the head is larger in " Shio " than"Naisa" (Yamase, 1760), which is considered to be male secondary sexual
character ..

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

108

KASUYAACKNOWLEDGMENTS

Greatest thanks are due to Pro£ H. Ohyi and Mr. T. Nishimoto of Hokkaido University and to Dr. M. Yoshimura of Awa Museum at Tateyama City, who kindly offeredme the privilege to study the specimens of G. melaena and various informations on
the specimens. Dr. L. G. Barnes of Los Angeles County Museum, Dr. R. L. Brownell and Dr. J. G. Mead of Smithsonian Institution, and Dr. A. V. Yablokov ofInst. Developmental Biol. are deeply acknowledged for their cooperation andvaluable suggestions.
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Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

KASUYA

110

EXPLANATION OF PLATESPLATE I
Fig. J. G. melaena, ventral view of skull found in the alluvial deposit by the Hekuri
River at Tateyama (34°59'N, 139°5l'E).
Fig. 2. G. melaena, lateral view of the same specimen.
Fig. 3. G. melaena, dorsal view of the same specimen.
Fig. 4. G. melaena, dorsal view of skull (no. 675) found at Motochi archaeologicalon the west coast ofRebun Island (45°20'N, 141°00'E).
PLATE II
Fig. I. G. melaena, dorsal view of skull (RKA 6054) found at Kabukai archaeological A-site on the east coast of Rebun Island.
Fig. 2. G. melaena, dorsal view of skull (RKA 3658) found at Kabukai A-site.
Fig. 3. G. melaena, dorsal view of skull (RKA 3987) found at Kabukai A-site.
Fig. 4. G. macrorhynchus, dorsal view of skull, caught at Taiji (33°35'N, 135°43'E) in
May 1967.
PLATE III
Fig. I.
Fig. 2.
Fig. 3.
Fig. 4.
Fig. 5.

G. macrorhynchus, dorsal view of skull, California coast, USNM 9076 (?).
G. macrorhynchus, ventral view of the same specimen.
G. melaena, dorsal view of skull, Virginia coast, USNM 303018.
G. melaena, ventral view of the same specimen.
G. melaena, dorsal view of skull, Chili, USNM 395365.

Sci. Rep. Whales Res. Ins1
No. 27, 197

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A STUDY OF THE GENUS BERARD/US DUVERNOY*CHARLES McCANN
8 Kiwi Street, Heretaunga, Upper Hutt, New Zealand

ABSTRACTThe taxonomy and distribution of the two known species of Berardius,
B. arnouxi Duvernoy and B. bairdi Stejneger are reviewed, and some anatomicaland biological aspects of the two species are considered.

INTRODUCTION

The members of the genus Berardius or Porpoise Whales, in spite of their large size,
are little known. Although one of its two closely allied species, B. bairdi Stejneger,
was known to and fished for by Japanese whalers very early in historic times, it was
not scientifically recognised until 1883.
The genus Berardius was founded by Duvernoy in 1851 on a skull from Akaroa,
Banks Peninsula, New Zealand. The specimen was taken to France by M. Arnoux,
who was at the time surgeon to the French corvette Rhin commanded by Captain
Berard. These two gentlemen are commemorated in the generic and specificnames of the type species, Berardius arnouxi Duvernoy.
Although more than a hundred years have elapsed since the original description of B. arnouxi, barely forty specimens have been reported in literature. Bysome mischance, the ' o ' was omitted from Arnoux in the original description.
Since then, some cetologists have retained the original spelling, arnuxi, while others
have amended it to arnouxi. As the original spelling was almost certainly a 'lapsus
calami ', the amended spelling should be adopted.
Flower (1872) gave the first detailed description of the general anatomy ofBerardius from a specimen then in the Museum of the Royal College of Surgeons.
This specimen which had stranded at New Brighton, Canterbury, New Zealand,on 16th December, 1868 had been sent to London by Sir Julius von Haast. Little
has since been added.
The two known species are geographically isolated: B. arnouxi Duvernoy occurs only in the Southern Hemisphere, in the Southern Pacific and Southern
Atlantic Oceans; whilest, B. bairdi Stjneger is confined to the Northern Pacific
Ocean, concentrating seasonally near Japan while strays are occasionally found* This paper was compiled some years ago while the author was still employed as the Vertebrate Zoologist
at the DOMINION MUSEUM, Wellington (now the NATIONAL MUSEUM) under the Directorship of Dr. R. A. Falla (now Sir Robert A. Falla), since retired. Dr. R. K. Dell, is the present Directorof the National Museum.
The specimen registration numbers quoted in the paper are referred to by the originals with theprefix D. M. or Dom. Mus.
Sci. Rep. Whales Res. Inst.,

No. 27, 1975, 111-137.

C. McCANN

112

along the western shores of North America.
Recent papers by Omura et al. (1955) and others have discussed the biology
and whaling of B. bairdi, but no such information is available for B. arnouxi. McCann (1962), in a revision of the genus Mesoplodon discovered that the calves of B.
arnouxi had until then been identified as Mesoplodon hectori (Gray). Hector (1870)
had originally identified the type of M. hectori (Gray) correctly as the young ofBerardius arnouxi Duvernoy, but Gray (1871) declared it a new species of Berardius.
Turner (1872) cautiously transferred B. hectori Gray to Mesoplodon where it hadremained ever since. McCann's discovery not only places the genus Mesoplodon ina more satisfactory systematic position, but indicates that the breeding grounds ofBerardius arnouxi are in the vicinity of the New Zealand Archipelago.
GENUS BERARDIUS DUVERNOY (PLATES I-II)
1851. Berardius

1872.

Mesoplodon

1922.

PaikeaDuvernoy, Ann. Sci. Nat. (3) 15: 52. Type Berardius
arnouxi Duvernoy. Type locality: Akaroa, New Zealand.
Type in Paris Museum.
Turner, Trans. Roy. Soc. Edin. 26: 778. (Berardius
hectori (Gray))
Oliver, Proc. Zool. Soc. Lond. p. 575. Type Mesoplodon
hectori (Gray), Type Brit. Mus. 1677b. 76. 2. 16. 3.
DESCRIPTION

Large whales attaining 7.8 to 12.7 metres (26 to 42 ft) in length. Head large, buffer-like, without nuchal depression but a pronounced rostrum or beak; upper jaw
subacute anteriorly; lower jaw longer than the upper, extremity rounded; twopairs of strongly compressed, functional teeth near the extremity, the anterior larger
than the posterior; teeth erupted in both sexes on reaching maturity; occasionally
only a single (anterior) tooth present in each ramus; normally no teeth in maxillae.
Eye of moderate size. Blowhole median, crescentic. A pair of diverging gulargrooves forming an inverted V-shape, the anterior arms not in contact; occasionallyan additional pair of shorter grooves may appear; no thoracic nor abdominal
pleats. Body fusiform, laterally compressed posteriorly only. Flippers moderately
broad and rounded distally; flukes large, shallowly falcate, without median notch
although occasionally a shallow indentation may appear mesially. Dorsal fin nearer
the caudal, triangular, slightly falcate or represented by a short obtuse elevation.*
SIZE

B. bairdi is larger than B. arnouxi; the former reaches 12.6 metres in length (42 ft),
whereas the latter has not been known to exceed 9.9 metres (33 ft).

Curiously,* Perhaps the result of injury or attack by shark.
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

STUDY OF BERARD/US DUVERNOY

113

although B. bairdi is the larger and bulkier of the two species, it has a relatively
smaller head than B. arnouxi (True, 1910: 67).
SEXUAL DIFFERENCES

Females of B. bairdi are larger than males, but in B. arnouxi both sexes appear to beof very similar size. As the mandibular teeth erupt in both sexes at maturity, theexternal sexual differences are not as well marked as other genera of the family inwhich the teeth erupt in the male alone.
GESTATION

The few breeding records available, suggest that the gestation period is approximately ten months and calving takes place during the late spring or early summer.
ANATOMICAL OBSERVATIONSThe skull (Pls. III-VI)
The skull of Berardius differs from those of other ziphioid genera in its greater
symmetry and simplicity. Its elements are less firmly interdigitated than in othergenera. The mesorostral groove remains unossified throughout life, i.e. the mesethmoid
cartilage does not ossify partially nor wholly as in ,(iphius and Mesoplodon. The
cranium is relatively more depressed in adults than in the rest of the family, being
broader than deep (Pl. VI).
Flower (1872) drew attention to the imperfect closure of the suture between
the basisphenoid and presphenoid bones; the suture being, at times, open 12.5 mm(1 in) or more. This is quite normal, not only in Berardius but also in other members of the Ziphiidae. In Berardius, the suture between the supraoccipital and thefrontal bones is also imperfectly closed, but in the rest of the family there is a more
complete fusion of these elements. The basioccipital and the basisphenoid elements
fuse completely, so that, in all the genera, the line of fusion is not clearly discernible.
In dorsal view (Pl. III, fig. 2), the posterior cranial region is semilunate, theantorbital tubercles forming the ' horns ' of the moon. The rostrum is somewhat
lanceolate, arising from within the antorbital notches and is considerably shorter
than the mandibles. In side view (Pl. III, fig. 3 ), the dorsal profile is almost straight;
the lower profile rising gently from the pterygoids towards the end of the rostrum.
The maxillae are much shorter than the premaxillae, the latter alone forming theextremity. In ventral view (Pl. III, fig. 1) a small portion of the vomer is visible
between the pterygoids and palatines. For a short distance anteriorly, the approximation of the margins of the maxillae obscures the vomer, which reappears once
more as a linear-lanceolate bony wedge between the maxillae and premaxillae(Pl. III, Vo), but ends considerably short of the rostrum extremity. The premaxillae approximate (but do not fuse) in front of the vomer. An open canalSci. Rep. Whales Res. Inst.,
No. 27, 1975.

114

C. McCANN

forms a ridge on the ventral aspect of each premaxilla. From the anterior 'segment ' of the vomer, there is a marked difference in the transverse level between the
maxillae and premaxillae, the latter being nearer the midline, resulting in a certain
amount of flatness in this region.
No basirostral grooves are present, but there is a blind ' pit ' beneath the maxillae just in front of the malar bones (Pl. VII, MP). The position of this pit corresponds approximately to the commencement of the basirostral groove in Mesoplodon
grayi Haast, the species in which the basirostral grooves are best developed in thefamily. There is no maxillary notch or tubercle as in Mesoplodon, but a maxillarycrest is present (PL. 7, MXC).
In Berardous the premaxillae are subequal posteriorly, the right being only
very slightly wider than the left in the narial region. Their extremities are nodular
and are not turned outward as in Mesoplodon nor do they overhang the narial opening in the adult. The large bulbous nasal bones overtop the ends of the premaxillae slightly and over-hang the narial opening, supported below by a wide
'bracket' formed by the posterior wall of the vomer, (Pl. V, fig. 3, Vo). In the
newly-born animal the nasals do not attain the full size and are consequently below
the level of the extremities of the premaxillae. Two large ' blind pits ' (BP) on thewall of the vomer, just below the nasals (one on either side of the septum) arecharacteristic of Berardius and readily distinguish it from any other genus of Ziphiidae. These pits are clearly visible in both old and young (Pl. V, fig. 3, BP).
The mesethmoid is completely fused with the presphenoid posteriorly. Within the narial cavity the mesethmoid is sandwiched and fused between the recurved
walls of the vomer; together they form the narial septum. Anteriorly the mesethmoid emerges from between the vomerine walls and enters the mesorostral groove
as a free projection for a short distance, becoming thicker and roughened along its
dorsal surface. It is free from the vomer.
The walls of the narial cavity are composed posteriorly and mesially (septum)
largely of the vomer; the lateral faces largely of the pterygoid, the premaxillae andsmall wedges of the maxillae and palatines. A large foranien, situated on the internal lateral face of the nares leads by a canal, laterally, to a large foramen in theorbital region.

Foramina (Pl. III): In dorsal view, the skull shows several large formina, some ofwhich are useful accessory diagnostic features. Anteriorly, there are two premaxillary foramina, one in each premaxilla (PMXF); two large maxillary foramina, one (sometimes dual or multiple) in each maxilla (MXF). Both pairs offoramina are in front of the narial cavity; in Berardius they are approximately on
the same transverse level although slight variations in level may occur. At theposterior edge of the maxillary crest or occasionally slightly further back, there isa large foramen connected by a canal with a foramen in the orbital region. This
post-crest for amen (PCF) (as I propose to call it) is peculiar to Berardius and absent inother ziphioids, even in Hyperoodon ampullatus which has a greatly exaggerated maxillary crest. Still further back, on the maxillary plate, there is another large foramen
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

STUDY OF BERARD/US DUVERNOY

115(PMF) (and occasionally an additional one or two); a canal connects this foramen
to another in the orbital region.

Antorbitalforamina (Pls. VI, VIII): In ventro-lateral view, the skull of Berardius has
three large foramina in front of the large infundibulliform optic foramen (OPF).
These three large foramina are not readily seen in any of the other ziphioid genera
as they are concealed within the apparently single 'antorbital foramen' behinda ridge formed by the palatine bone. The anteriormost foramen (Pl. VII, fig. I)

Fig. 1. Arrangement of antorbital foramina (diagramatic).
figures. del. C. McCann.

Lettering as in otherleads by a wide canal to the maxillary foramen (MXF) on the dorsal surface. Occasionally, immediately above this foramen (1), a small foramen perforates thelachrymal bone (L) and passes under it. A short distance behind is another large
foramen (Pl. VII, fig. 2) which perforates the lachrymal bone posteriorly; a large
vessel leads from this foramen to the dorsal surface of the skull where it appears as
the post-maxillary foramen (PMF) (or foramina). Lastly, somewhat below and alittle behind the last mentioned foramen and partially below the optic foramen(OPF) is another large foramen (Pl. VII, fig. 3). From this foramen, a large canalpasses to a large foramen on the lateral wall of the narial cavity (NF) and anotherbackwards through the foramen lacerum (Pl. X, fig. 1) into the brain cavity. Thisis the passage of the 5th cranial nerve.
From within the maxillary foramen (MXF) several secondary canals diverge(text-fig. I): one large division from the maxillary contributes to the formation of
the premaxillary foramen (PMXF); anteriorly a canal passes through the matrixof the maxilla towards the extremity of the rostrum (I term this the median rostral
canal (MRC)); posteriorly, three similar canals pass backwards under the premaxilla towards the vertex, emerging laterally under the posterior extremity of thepremaxilla. Other smaller canals from within the maxillary foramen arise to appear dorsally on or near the base of the maxillary crest. Finally, a large canalSci. Rep. Whales Res. Inst.,
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116

C. McCANN

from within the maxillary foramen emerges as the maxillary crest foramen (MCF)
towards the rear of the crest itself.
From within the premaxillary foramen, a canal is given off anteriorly leading
to the extremity of the rostrum. This canal, which I propose to call the upper rostral
canal (URC), is sandwiched between the premaxilla and the maxilla, particularly
proximally.
Attention has already been drawn to the canals leading from within the large
posterior foramen to the narial cavity and the brain case. Anteriorly, a small canalpasses through the maxtrix of the maxill towards the end of the rostrum; I propose
to call this the lower rostral canal (LRC). These three rostral canals are readily seen
in a cross section through the rostrum.
(In .?,iphius, the lower rostral canal, instead of passing through the matrix ofthe maxilla, passes to the exterior at the anterior edge of the palatine bone andcontinues in a shallow groove to the end of the rostrum underside).
The optic foramen is large and canals lead from it into the cranial cavity, tothe internal optic foramen, and through the foramen lacerum.
Behind the optic foramen (OPF), the external foramen ovale (FO) is situatedbeneath the falciform process (FP) of the squamosal surrounding the tympanic
bulla. A little further to the rear, and at a slightly lower level, is the external auditory foramen. The exterior exit of the carotid artery is somewhat irregular inposition and size; it is situated at the suture between the pterygoid and the alisphenoid.

Brain case (Pl. X, fig. 1): The brain cavity is amply described by Flower (1872).
Relatively, it appears to be larger than in other members of the family. A
well-marked ridge, formed by the orbito-sphenoid and continued upwards by thefrontal, is present in Berardius. This ridge is developed to a lesser degree in Hyperoodon, but absent in both ,Ziphius and Mesoplodon.
The base of the brain case appears to be flatter in Berardius than in .<:,iphius.
The sella tursica and tuberculum sella are both better developed in ,Ziphius (female)than in Berardius, and the walls of the cerebellum fossa appear to be more rugose.
In sagittal section the internal cranial foramina are readily visible. Fromanterior to posterior, they are:
1) A small foramen high up on the wall of the presphenoid close to the midline, entering the nares by a canal. Flower (1872) suggests this may be the rudiments of the olfactory foramen.
2) The large optic foramen perforating the presphenoid.
3) The large sphenoidal fissure (Pl. VII, fig. 1) and the foramen rotundum
slightly behind the optic foramen (Pl. VI, OPF), but obscured by the orbito-sphenoidal ridge. These transport the nerves to the orbit and the middle division of
the 5th cranial nerve.
4) A small foramen for the carotid artery (Pl. IX, ea). On the floor of thebasisphenoid, on a transverse level with the foramen ovale. The canal to the exterior appears as a foramen near the posterior border of the pterygoid.
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

STUDY OF BERARDIUS DUVERNOY

1175) The large foramen ovale further to the rear of the sphenoidal fissure andpartially obscured by a ridge of bone. In Berardius the sulcus leading from the foramen ovale to the sphenoidal fissure is broad and shallow, but in :(,iphius andMesoplodon it is more canalised.
6) A foramen for the seventh nerve to the rear of the foramen ovale. This
enters the funnel of the auditory meatus and exits through the auditory foramen.
7) The condylar foramen, a small foramen a short distance behind the auditory foramen. This perforates the cerebellar fossa, but has its external opening
in the fissure formed by the basioccipital crest and the exoccipital.
8) The auditory foramen, clearly visible to the rear.

Mandibles (Pl. III, fig. 3)
The mandibles in Berardius are much longer than the rostrum and permanently
expose the large anterior teeth. The distal half of each ramus is comparatively
narrow and the upper and lower profiles in this region are almost parallel; theproximal half expands rapidly to form the posterior half. There is no defined' gonus ' in Berardius as in some other ziphioids. The symphysial union is relatively short and the two halves do not anchylose even in old age as in the rest of thefamily. The mental foramen may be single or multiple and is somewhat erratically situated.
Mandibular teeth (Pl. XII)
Unlike other genera of Ziphiidae, Berardius erupts two pairs of functional teeth
in both sexes, the anterior pair being usually larger than the posterior. They arestrongly compressed laterally. In this, both Berardius and Mesoplodon agree; anddiffer from the rest of the family in which the teeth are circular or oval in crosssection.
Viewed laterally, both pairs are roughly triangular, the base being the shortest
side, while the anterior edge is frequently slightly longer than the posterior. The
lateral faces may be flat or variously undulate with numerous furrows and striae
arising along the base and converging towards the apex. Along the antero-posterioredge, the axis is bent outwards slightly. The tip is an acute, enamelled point surrounded by a ' collar ' of dentine. In old animals the teeth are usually so severely
abraded, that their tops are flat, almost level with the gum.
Occasionally, only a single tooth (anterior) is present in each ramus.
Vertebrae:
The formula of the vertebrae in the two species is as follows:
Berardius arnouxi: C 7 (3+4); Th 10; L 12 (13); Ca. 19=48 (49).
Berardius bairdi: C. 7. (3+4); Th. 11; L. 12. ea. 16+=46+.
True (1910) observed that the skeletons of both species of Berardius are very
similar in most details, but there are some specific differences particularly in thenumber of thoracic vertebrae and of ribs.** Hale (1962) records a specimen of B. arnouxi with eleven ribs on one side.
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

C. MaCANN

118

Hector (1878) recorded thirteen lumbar vertebrae in a specimen of B. arnouxi,
which is still in the collection of the National Museum, Wellington. Wellington(Slet Mus. 183) and has unmistakably thirteen lumbar vertebrae. However, twelve
lumbar vertebrae appears to be the normal number in B. arnouxi as shown by otherspecimens in the National Museum, Wellington. True (1910) remarks that the" discrepancy here shown cannot be accounted for at present, but at all events, noneof the formulae of B. arnouxi correspond to that of B. bairdi."
In the number of thoracic vertebrae, the two species differ specifically. B.

Fig. 2. Berardius bairdi Stejneger. Right scapular (714x 525 mm.) (after True, 1910,
pl. 33, fig. 2).

arnouxi has ten and B. bairdi, eleven.
bers of vertebrae respectively.

The numbers of ribs correspond to the num-

Cervical vertebrae: The three anterior cervical vertebrae are completely anchylosed
in the adult to form a single mass in both species. A newly born specimen of B.
arnouxi (Dom. Mus. 614) has only the first two vertebrae completely fused.
Caudal vertebrae: The number of vertebrae in the caudal region appears to varyslightly (eighteen to nineteen appears to be normal). The small terminal elements may easily be lost or dissolved during maeeration.
Chevron bones: There are normally nine chevron bones, the first usually of two
irregular pieces, not united mesially. The fourth and fifth are the largest and bestdeveloped.
Sternum:
The sternum is like that of .Z,iphius and Mesoplodon, but naturally larger. It is
normally composed of five elements, the fifth frequently anchylosed to the fourthand bifid posteriorly.
Scapulae: (Pl. XIII)
Sci. Rep. Whales Res. Inst.,
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STUDY OF BERARD/US DUVERN OY

119Apart from size, the shape of the scapulae is remarkably different in the two
species. The acromion and coracoid processes are contrasted; there is a greater
space between the two processes in B. arnouxi than in B. bairdi, and in bairdi theacromion almost completely overlaps the dorsal margin of the coracoid (text-fig. 2).
SCHOOLING
Omura and his colleagues (1955) state that Berardius bairdi forms schools of ten or
more individuals, males appearing in greater numbers than females.
There is some evidence that the breeding school could be dominated by asingle master bull. Linear weals and other wounds occur on the bodies of males,
particularly old bulls, indicating that there is considerable rivalry among the males-the females seldom exhibit such scarring. Adult females of Berardius probably
add their quota of scarring on the bulls for unwanted attentions.
MALE RIVALRY
Pike (1953) referring to scars in Berardius bairdi offers another explanation of scarring"Numerous white scratch marks appear on the back and flanks of both specimens(Pl. II, Ill). The scratch marks when found on males of other Ziphiidae andDelphinidae, have been attributed to the teeth of rival males. This explanation
seems unlikely to be applicable in this species whose teeth are either buried in thegums [young animals] or are so blunt and protrude so slightly. A more plausible
explanation for all species is that the scratches are caused by the sharp ' beaks ' of
the squids on which they feed. Similar scratches are found on the heads of sperm
whales."This explanation overlooks the fact that the teeth are extremely sharp when
first erupted and only become blunt with age, and that scarring by cephalopods
would not be restricted to males, nor would it occur indiscriminately all over the
body. The present author believes the scars are the result of male rivalry, a viewalso held by Omura, Fujino and Kimura (1955). Over several years, I have examined numerous carcases of Ziphiidae and other whales for scarring. I find thatin male Z,iphius linear scars, which are numerous on back and flanks, less common
on the body, appear as single weals or exactly parallel weals, ranging from a few
centimetres to one metre or more in length. They are produced by one or a pairof teeth ripping along the body, the distance between the parallel scars corresponding to the distance between the mandibular teeth at the extremity of the mandible.
Females rarely exhibit linear scarring.
There are also round, oval or elliptical white scars*, particularly by the vent.

* Recently, my suspicions that the elliptical scars, so commonly met with on the bodies of ziphioids,
particularly in the region of the vent, were probably caused by some epizoic crustation were confirmed,
when I asked Dr. J.C. Yaldwyn, the Assistant Director of the National Museum to produce the largest' copepod ' he was aware of in the museum ' collection. This he did and I immediately recognised itas the epizoan responsible for the scars. It proved to be the female of Livoneca ravnaudi (M. Edwards):CYMOTHOIDAE. A paper on the subject will appear shortly under our joint names.

Sci. Rep. Whales Res. Inst.,
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120

C. McCANN

In ]l,fesoplodon, linear scars are very common on old males, but they are neverexactly parallel as in Ziphius. As the mandibular teeth are nearer the posterior
union of the symphysis in Mesoplodon (except in M. mirus True) both teeth cannot
be brought into play at the same time. (M. mirus could produce scars similar to
those in -?,,iphius). Circular, oval or elliptical scars also appear on the body as in
Ziphius.
In Berardius there are three types of linear weals: (a) a single rip probably
made by a single anterior tooth; (b) parallel line or scars (Pl. II, fig. 5) made by
the two anterior teeth; the distance between the weals being approximately thatbetween these teeth; and, (c) parallel linear scars, one larger and deeper than theother, but the width narrower than to the distance between the two anterior teeth.
These could be produced by both teeth of one ramus making contact at the sametime. All three forms of linear scars have been seen on one old male (Pl. II).
There were also the usual circular, oval and elliptical scars (Pl. II, fig. 5).
The circular, oval or elliptical scars referred to probably result from the attack or
attachment of suctorial animals. Pike (1951) and Nemoto (1955) discuss andillustrate scars made by cyclostomes, particularly the Sea Lamprey (Entosphenus
tridentatus), a species confined to the Northern Pacific Ocean. In the whales I
have examined the scars showed what appeared to be the impressions left by thetransverse lamellae of the discs of sucker-fish. Prolonged attachment could wellbring about permanent discolouration of the skin. Concentrations of scars areinvariably around the vent, which suggests that the fish congregate in the area
to feed on excrement. Squid have been known to scar sperm whales in much thesame manner.
Parasitic crustaceans could also be responsible for some of the scarring, somescars suggesting the erstwhile presence of barnacles, e.g. Coronula. However, whilethese dermal barnacles are commonly epizoic on baleen whles, they have not, as
far as I am aware, been observed on ziphioids. Circular scars could also result
from 'jab ' wounds caused by the tip of an opponent's rostrum.
SYNONYMYBERARDIUS ARNOUX! Duvernoy: The Southern Porpoise Whale
1851.
Berardius Arnuxii
Duvernoy, Ann. Sc. Nat. (3) 15: 51. (Typelocality: Akaroa, Banks Peninsula, New
Zealand. Type in Paris Mus.).
Gray, Cat. Seals & Whales Brit. Mus. p. 348.
Berardius Arnuxi
1866.
Owen, Monogr, Palaeontogr. Soc. p. 31, fig. 11.
.:{,iphius (Berardius)
1870.
Arnouxii
Hector, in Knox, Trans. N. Zeal. Inst. 2: 27.
1870.
Berardius Arnouxi
Hector, Ann. Mag. Nat. Hist. (4) 5: 222. Trans.
1870b. Berardius Arnouxi
N. Zeal. Inst.
Haast, Trans. N. Zeal Inst 2: 190.
1870a. Berardius Arnouxi
Haast, Ann. Mag. Nat. Hist. (4) 6: 348. (Repeat
1870b. Berardius Arnouxiof previous article)
Sci. Rep. Whales Res. Inst.,

No. 27, 1975.

STUDY OF BERARDIUS DUVERNOY

1871.

' :(,iphidae '1871.

Berardius Hectori

1872.
1872.

Mesoplodon Hectori
Berardius Arnouxi

1873.

Mesoplodon J;noxi

1873.
1874.
1876.
1878.
1878.

Mesoplodon Hectori
Mesoplodon Hectori
Berardius arnouxi
Mesoplodon hectori
Mesoplodon hectori

1880.

Berardius arnouxi

1893.
1900.
1900.
1910.
1920.

Mesoplodon hectori
Mesoplodon hectori
Berardius arnouxi
Berardius arnouxi
Berardius arnuxi

1922.
1922.

Berardius arnouxi
Paikea hectori

1924.
1939.
1950.
1957.

Mesoplodon hectori
Berardius arnuxi
Mesoplodon hectori
Berardius arnouxi

1962a.
1962b.
1962.

Berardius arnouxi
Berardius arnouxi
Berardius arnuxi

1974.

Berardius arnouxi

121

Knox & Hector, Trans. N. Zeal. Inst. 3: 125, pls.
13-17.
Gray, Ann. Mag. Nat. Hist, (4) 8: 115-117.
(Type locality: Titahi Bay, N. Zeal., Type Brit.
Mus. 1677b. 76. 2. 16. 3.)
Turner, Trans. Roy. Soc. Edin. 26: 778.
Flower, Trans. Zool. Soc. Lond. 8: 203-234, pls.
27-29.
(part.) Hector in Knox, Trans. N. Zeal. Inst. 5:167 (Part type, Titahi Bay specimen, Brit. Mus.
1677b. 76. 2. 16. 3)
Hector, Trans. N. Zeal. Inst. 5: 170.
Gray, Trans. N. Zeal. Inst. 6: 96-97.
Buller, Trans. N. Zeal. Inst. 8: 407.
Hector, Trans. N. Zeal. Inst. 10: 338, p. 16.
Flower, Trans. Zool. Soc. Lond. 10: 416, pls. 71
and 72 fig. 4.
Van Beneden & Gervais, Osteogr. Cet. pp. 377,
515, Atlas pls. 21-23.
Forbes, Proc. Zool. Soc. Lond. p. 277.
Beddard, Book of Whales, Lond. p. 220.
Beddard, Book of Whales, Lond. p. 228-233.
True, Bull. U.S.N.M. 73: 77.
Marelli, An. Mus. Nac. Secc. Zool. Buenos
Aires. 30: 411-444, pls. 1-5.
Oliver, N. Zeal. Journ. Sci. Tech. 5: 135.
Oliver, Proc. Zool. Soc. Lond. p. 575. (Type,Brit. Mus. 1677b. 76. 2. 16. 3).
Harmer, Proc. Zool. Soc. Lond. p. 542.
Hale, Ree. S. Austr. Mus. 19 (4); 5.
Fraser, Proc. Linn. Soc. Lond. 162: 50, pls. 3, 4.
Taylor, Proc. Zool. Soc. Lond. 129 (3): 325-332,
pl. 3, figs. 1-4.
McCann, Tuatara 10 (1): 13-18, pls. 1, 2.
McCann, Ree. Dom. Mus. 4 (9): 83-94 pls. 1-5.
Hale, Ree. S. Austr. Mus. 14 (2): 231-243, pls.
5, 6, text fig. 1.
McCann, Sci. Rep. Whales Res. Inst., Japan,
No. 26, 145-155, Figs. 1-23, pls. 1-8.
DESCRIPTION

General external characters as in the genus
Teeth (Pl. XII): The anterior mandibular teeth are normally larger than theposterior. Viewed laterally both are roughly triangular in shape, the base being
the shortest side, and the anterior frequently longer than the posterior. The
lateral faces may be flat or undulate with numerous furrows or. striae arising along
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C. McCANN

the base and converging towards the tip. Viewed antero-posteriorly, the vertricalaxis is inclined slightly outwards in its upper portion. The apex is formed of anenamelled point of about 10 to 12 mm surrounded by a 'collar' of dentine. On
the inner face of the enamelled cusp there may be one or two vertical furrows. In
old animals the tooth is worn down to a flat-top level with the gum.
Size: Berardius arnouxi may attain a length of 9.6 m (32 ft). The difference insize between the sexes does not appear as pronounced as in B. bairdi.
Measurements in the flesh, ad. male. Dom. Mus. 1,433 (Col. C. McCann)
Total length 8.50 m; snout to anterior edge of dorsal 6.150 m; Snout togenital 5.60 m; genital to anal 0.30 m; length of penis 0.90 m; circum. of penis
at base 0.23 m; snout to eye 1.50 m; snout to flipper 1.77 m; gape 0.59 m; snout
to blowhole 1.10 m; blowhole 0.13 m; Beak 0.551 m; throat groove 0.55 m; anal
opening 0.23 m; Fluke across 2.444 m.
Neonatals so far recorded do not exceed 3.0 m (11 ft) but the records could
be of animals prematurely born when the true neonatal length be slightly greater.
Colour: " The colour of the whole animal was of a deep velvety black, with the
exception of the lower portion of the belly, which had a greyish colour." (Haast,1870).
" The colour was black with a purple hue, except a narrow band along the
belly which was grey. The muzzle, flippers and tail lobes were intensely black."(Hector, 1878: 338).
An old male from Pukerua Bay (Pls. I-11), although decomposing, clearly
showed a dirty white dorsal surface. This colour extended from the ' crest ' of thehead to shortly behind the dorsal fin. Parts of the dorsal fin and some surrounding areas were even whiter. The rest of the body was brownish black (Pl.
I and 11, fig. 1). As most observers describe the dorsaum is black, it is possiblethat in this individual the dorsum may have bleached during exposure, after deathand before stranding.

DISTRIBUTION
Berardius arnouxi is apparently confined to the South Pacific and South Atlantic
Oceans, between 30°S and Antarctica. Odd specimens have stranded along theeastern shores of New Zealand as far north as 37°S, but these strandings have beenlater than the normal period when this whale visits the New Zealand area.
Specimens: (See text-fig. 3)
New Zealand Waters: Otago Peninsula, 1840 (Otago Mus. Al4.37 mandibles);Stewart Island (Otago Mus. A24.69, skull), 2 teeth (Dom. Mus. 523); Mason Bay,
Stewart Island, February, 1929 (Dom. Mus. 523); Mason Bay, Stewart Island,
February, 1929 (Dom. Mus. 416, skeleton); Centre Island, Foveaux Strait (Dom.
Mus. 528, worn tooth); Chatham Islands (Otago Mus. A339); Coast near WanSci. Rep. Whales Res. Inst.,
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STUDY OF BERARDIUS DUVERNOY

123

ganui, before 1920 (Wang. Mus. A339); Coast near Wanganui, before 1920 (Wang.
Mus. skeleton); Worser Bay, Wellington, January 1870 (Dom. Mus. 7, skull), 12th
January, 1877 (Dom. Mus. 183, skull); 30th January, 1876 (Dom. Mus. 239, 2
na

178"

17"

17?

Three Kings Is.

NEW ZEALAND-SHOWING-

50

25

O

50

100

1'jJ M.tes

13:1(J)EJ3:S:WO!:E~==:;;10J=~~2CD KILOMETRES

75 Miles to an inch.

g

1870~WELLINGTON'/.

ll!~~~+-~~~~t--~~--t~t\-'-:.--~/'.r--__,,&~~~1-~~~~-f-~~---1~

CHATHAMS

x

17U

Fig. 3.

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

17?

174"

17"

Strandings of Berardius arnouxi Duv. in New Zealand.

178'

C. McCANN

124

teeth); Saltwater Creek, north of Banks Peninsula, 27th December, 1873 (Dom.
Mus. 1402, skeleton); New Brighton Beach, Christchurch, 16th December, 1868
(Roy. Coll. Surg., skeleton); Akaroa, Banks Peninsula, 1846 (Type, Paris Mus.),(Brit. Mus. 96.11.20.1, male); South Tapu, Thames Coast, 13th March, 1944
(Auck. Mus.); Te Horo Beach Wellington District, 13th January, 1931, male,
8,400 mm (28 ft); Otaki Beach, Wellington Dist, 4th January, 1934, female, 8,
100 mm (28 ft.), (Dom. Mus. 528, 4 teeth);* Pukerua Bay, Wellington District, 18
th February 1960, old male, 8,850 mm (29-ft), (Dom. Mus. 1433, 2 teeth); **Pencarrow, Wellington Harbour, male 9,000 mm (30 ft, male), female 6,000 (20 ft),ft), (Dom. Mus. 526, 2 teeth); Lyall Bay, Wellington, 1874 (Buller); East Cape,(Dom. Mus. 527, 1 tooth); Titahi Bay, Wellington District, January, 1866, male
2,725 mm (9-ft), (Britt Mus. 1677b 76. 2. 16. 3, Type of B. hectori Gray); Plimmerton, Wellington District, 25th January, 1937, 3,200 mm (10 ft, 8 in.), (Dom. Mus.
614, skeleton); 'New Zealand', (Brit. Mus.), (U.S.N.M. 21511).
Australian waters: Port Lorne, S. Australia, 27th December 1935, 8, 700 mm(29 ft), (S. Austr. Mus. M. 5012)
Antarctic waters: Taylor, April 1955 (Photographs).
Argentine waters: Arroyo del Pescado, Los Talas, 29. 5. 1918, female, 7,800 mm(26 ft), (Mus. Nac. Buenos Aires).
South Atlantic waters: South Shetland Islands, female (Brit. Mus. 1934. 6. 15.
l); Falkland Islands (Brit. Mus. 1949. 8. 19. l); La Plata, 29th May, 1918, female,(Mus. Nac., Buenos Aires).
Of thirty specimens recorded from New Zealand waters, the majority stranded
during the late spring or early summer (December to February-March). Two
neonatals which may have been premature births are recorded as stranding inJanuary 1866 and 1937 (as Mesoplodon hectori (Gray)). The presence of adults andneonatals during the same period of the year suggests that Berardius arnouxi visits the
New Zealand area to breed. On the other hand, photographs of Berardius arnouxi
locked in the ice in April 1955 (Taylor, 1957) show that they return to the Antarctic
region to feed. These observations point to a seasonal north and south migration.
The single recorded Australian specimen, a gravid female was found in thesame period in which Berardius visits New Zealand waters.
The number of Berardius arnouxi visiting New Zealand waters annually is unknown, particularly as their visits do not coincide with the whaling season (Mayto August), and they are not at present of commercial value. Any information we
have is derived from chance strandings.
In the South Atlantic, there appears to be a similar pattern of movement presents itself, but the records are too few for definite conclusions. A single neonatal
has been recorded from one of the smaller islands of the Falklands group (Fraser,1950), but its exact time of stranding is unknown.
As with Mesoplodon, species of Berardius appear to calve near the larger archi-* Porirua Harbour, Wellington District,January 1862, male, 8100 (27 ft), (Fide Knox, 1871: 125)** The sex of these two specimens is in doubt as they were reported and the teeth of the adult collected bya layman.

They may have been a female and calf.

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pelagos of the world in both hemispheres. B. arnouxi in the New Zealand and South
Atlantic archipelagos and B- bairdi around the Japanese Archipelago.
FOOD

Haast (1870) notes that the stomach contents of one animal contained about halfa bushel of the horny beaks of cephalopods (squids). Haast mentions ' Octopus ',but this is hardly likely, as the octopus is an inshore animal, whereas squids are
pelagic and often occur in vast shoals.
ANATOMICAL OBSERVATIONSCranium (Pls. III, IV): The cranium of the adult has been well described and figured by Duvernoy (1851) and Flower (1872).
The skull of Berardius exhibits almost perfect symmetry (Flower 1872). The
same symmetry appears in the newly-born young. The large bulbous nasals area prominent feature of the vertex in the adult, but the nasals are not so conspicuous in the newly born (Pl. IV, fig. 1). The extremities of the premaxillae rise above
the nasals and are slightly curved forwards, not so recumbent as in the adult, nor dothey overhang the narial cavity as in some species of Mesoplodon. Further, they do
not twist outwards as asymmetrically as in Mesoplodon; they are almost equal insize, the right premaxilla being very slightly larger than the left.
The recurved walls of the vomer, with the mesethmoid bone sandwiched between them contribute to the formation of the narial septum, the mesethmoid bone
eventually emerging anteriorly into the mesorostral groove as a free projection
between the margins of the premaxillae. The ossified projection of the mesethmoid
terminates as an irregular, rough or nodular projection, extending as far forwards
as a transverse line drawn through the middle of the maxillary crest. In the flesh,
the mesethmoid bone is continuous with the mesethmoid cartilage and fills themesorostral groove. In Berardius the mesorostral groove is permanently open andis never partially nor wholly filled by bone as in some species of Mesoplodon. Posteriorly, the vomer forms a broad ' bracket ' beneath the large nasals with a large
blind pit (BP) on either side of the septum. The shape of the ' bracket ' and thepresence of the blind pits are distinctive of Berardius.
Anteriorly, the vomer extends about two-thirds the length of the rostrum
forming a trough for the mesorostral cartilage. Ventrally, much of the anterior
end of the vomer is visible, wedged between the premaxillae. This condition is
also found in the newly-born young (Pl. IV, fig. 2, Vo).
In dorsal view, the rostrum is somewhat spearshaped. No distinct maxillaryturbercle is present but there is a maxillary crest. In side view, its dorsal profileis straight; the lower gradually descends from its anterior extremity towards thepterygoids. In ventral view, the broad maxillary prolongations are seen embracing the premaxillary extremities with the vomer wedged between. In transverse
section, the rostrum is considerably broader than deep near the middle.
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C. McCANN

The adult skull is broader than high, but in the newly-born the height is greaterthan the breadth.

Foramina (Pl. IX, fig. 1): The premaxillary and maxillary foramina are on approximately the same transverse level. The premaxillary is very much smaller
than the maxillary which frequently has secondary foramina around it. The postcrest foramen and the secondary foramina are not pronounced in the neonatalskull.
Antorbital region (Pl. IX, fig. 2): The antorbital tubercle is exaggerated by thepresence of a deep antorbital notch. In dorsal view, the tubercle is seen to beformed by the maxilla and a small portion of the malar bone, with the anterior
extremity of the frontal, a short distance behind forming the orbit. In side view,
the broad anterior end of the frontal is shortly behind the end of the maxilla andthe anterior projection of the malar bone. At the lower corner of the frontal, anarrow portion of the lachrymal bone is visible but not on a level with the lateral
margin of the frontal. Ventrally, the malar bone alone appears to form the tubercle; the lachrymal bone falls short of the lateral margin of the frontal. Thereis a triangular wedge between the frontal and the malar.
The antorbital notch is margined by the maxilla alone.
In the newly-born, the arrangement is somewhat different. In dorsal view,the malar borders the antorbital notch; the lachrymal is just visible in front of theanterior extremity of the frontal. The orbital portion of the frontal is not laterally
expanded as in the adult, but forms a narrow margin to the maxilla behind thetubercle. In side view, the large ' cuboidal ' end of the lachrymal abuts the anterior end of the frontal and a very small portion of the malar is also visible. Ventrally, the lachrymal and malar bones predominate in the formation of the tubercle
and malar bones predominate in the formation of the tubercle to the exclusion of
the anterior end of the frontal and maxilla.
Mandibles: The extremity of the mandibles protrudes considerably beyond therostrum, so much so that the anterior teeth are outside the mouth when it is closed.
The distal half of the mandibles is comparatively narrow and the upper and lower
profiles are almost parallel. The proximal half expands rapidly to form the posterior
portion of the jaw. The symphysial union is comparatively short and the two
halves appear never to fuse together as in some other genera. The mental foramen
may be single or multiple and somewhat irregularly situated on either side, even in
the same individual.
Eyes: The relative size of the eye ' sockets,' in relation to the length of the skull,is smaller than in either Ziphius or Mesoplodon. The jugal process is accordingly
comparatively short.
Ear Bones (Pl. XI):

The bulla and the periotic of an adult measure 73 by 48 mm
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and 80 by 45 mm respectively (Dom. Mus. 1406). In the newly-born the same bones
in the same order measure 54 by - - mm, periotic not stated (Brit. Mus. 1677b. 76.
2. 16. 3); 51 by - - mm, periotic not stated (Brit. Mus. 1949. 8. 19. l); 51.5 by
36 mm and 56 by 32 mm (Dom. Mus. 614).
Scapula (Pl. XIII): In B. arnouxi the vertebral border is more or less straight, notconves as in B. bairdi; both the anterior and posterior borders are more verticalthan in B. bairdi. The anterior border is almost straight, whereas in B. bairdi it isconvex dorsally, and slightly concave ventrally; the posterior border is less concave than in B. bairdi. The anterior spinous fossa is relatively narrower than in
B. bairdi.
The acromion is almost level with a vertical dropped from the anterior angle;
the upper and lower profiles are, at first, almost parallel but gradually diverge towards the extremity; in some specimens the extremity is decidedly wider. In B.
bairdi, the acromion falls short of a vertical dropped from the anterior angle andthe lower profile turns abruptly upwards producing an almost semicircular anterior
margin, which gives the entire process a somewhat circular or spiral border.
The coracoid and acromion processes are subequal in length whereas in B.
bairdi the former exceeds the latter. In dorsal view the two processes are divergent,but in B. bairdi there is very little space between the two. The coracoid is narrow
with both upper and lower profiles slightly concave; in B. bairdi the process is broadand slightly curved upwards with the upper profile concave, the lower convex.

SYNONYMYBERARDIUS BAIRD/ Stejneger: Japanese Porpoise Whale
1883. Berardius bairdi
Stejneger, Proc. U.S.N.M. 6: 75-77. (Type locality:
Bering Island; type U.S.N.M. 20992).
Malm, Bihang K. Svenska Akad. Handl. 8 (4): 109.
1883. Berardius vegae
Stejneger, Journ. Amer. Geogr. Soc. N.Y. 18: 317-328.
1886. Berardius bairdi
Beddard, Book of Whales, Lond. p. 233.
1900. Berardius bairdi
True, Bull. U.S.N.M. 73: 60, pls. 26-33. 34, fig. 7, 35,
1910. Berardius bairdi
fig. 7; 36, fig. 7; 39; 42.
Kellogg, Journ. Mamm. 12: 73-77.
1931. Berardius bairdi
Matsuura, Zool. Mag. Zool. Inst. Fae. Sci. Jap. Univ.
1942. Berardius bairdi
54: 466-473.
Matsuura Marine Mamm. (Kaiju). 298 pp.
1943. Berardius bairdi
Scheffer & Slipp, Amer. Midl. Nat. 39 (2): 266.
1948. Berardius bairdi
Scheffer, Pacific Sci. 3: 353, fig. 1.
1949. Berardius bairdi
Pike, Journ. Mamm. 34: 98-104, pl. 1.
1953. Berardius bairdi
Slipp & Wilke, Journ. Mamm. 34: 105-113, pls. 1, 2.
1953. Berardius bairdi
Omura, Fujino & Kimura, Sci. Rep. Whales Res. Inst.
1953. Berardius bairdi
J ap. 10: 89-132 (per errorum baildi)
Omura, Sci. Rep. Whales Res. Inst. Jap. 13: 213-214,
1958. Berardius bairdi
figs. 1, 2.
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C. McCANN

128

DESCRIPTION (PL. XIV FIG. 1)
General external characters as in the genus. There is no marked difference in thesize and shape of the mandibular teeth in either sex; (they are erupted in both sexes
alike on reaching maturity (Omura, 1955). Pike (1953) examined an 8.7 m (29 ft)
females which he regarded as both physically and sexually immature, and found theteeth were " completely buried." His observations support those of Omura (1955 ).
The largest teeth on record are: (anterior) vertical height 89; antero-posterior
width 103 mm; lateral diameter--; (posterior) vertical height 60 mm; anteroposterior width 37 mm; lateral width ? for a male of 9.9 m (33 ft); (interior) vertical height 93 mm; antero-posterior width 72 mm; lateral diameter?; (posterior)vertical height 55 mm; antero-posterior width 33; lateral width ? ; for a 8. 7 m(29 ft) female (Pike, 1953).

Size:

B. bairdi is up to 3.0 m (10 ft) longer than B. arnouxi.

female measured 12. 6 m (42 ft).
than females.

The largest known
Males are about 2.0 m (approx. 7 ft) smaller

Colour:Pike ( 1953) describes a male and female as follows: "In these specimens
the body is black on the head, back, flukes and flippers. The undersurface is of aslightly lighter shade, with white patches at the umbilicus and some white and grey
markings. The male has a conspicuous d~amond-shaped white patch at the umbilicus and some white around the genital opening. The female is lighter incolour and the skin was covered by abrasions caused while the whale was being
towed into the station. Numerous white scratch marks appear on the back andflanks of both specimens (Pls. I, II, V)."
Omura et al. (1955) state: "According to our observations, however, color ofwhole body is pure black or somewhat lighter, even on the under surface of the
body is pure black or somewhat lighter, even on the under surface of the body. It
is not certain, but most whalers say that there are two groups of Berardius, one being
black and another slatish. As there are many white scratch marks (Pls. I-II) on
the skin of Berardius, especially in old bull, the colour looks more lighter than it isin such whales. There are white patches in three regions of the ventral side of thebody, i.e. on the throat, between the flippers and at the umbilicus (Pls. II-IV,
Pl. III, fig. 2). Every Berardius seems to have the white patch on the throat or
between the flippers, there are considerably individual differences."
DISTRIBUTION

Berardius bairdi is restricted to the Northern Pacific Ocean; concentrating in Japanese waters during the breeding season in spring and summer. Occasional strayindividuals or small schools are found off the North American coast.
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Specimens: Bering Island (Type, U.S.N.M. 20992); St. George Island, ad. female(U.S.N.M. 49726); Centerville, California, ad. male (?), (U.S.N.M. 49725); St.
George Island, young male, ( U.S.N.M. 49727); Bering Island ( ?), (Grebnitzki ?)( U.S.N.M.); about 10 miles off Kains Island, Quatsino Sound, 5th July, 1950, ad.
male, 9.9 m (33 ft), (Pike, 1953); 20 miles E.S.E. of Cape St. James, 9th August,
1951, imm. female, 8.7 m (29 ft), (Pike, 1953); one mile south of Ocean City,
Washington, July 1950, 16.5 m (35 ft), (Slipp and Wilke, 1953); Japan, Nat. Sci.
Mus., 10.8 m (36 ft); Santa Cruz, California, June, 1925 (Mus. Calif. Acad. Sci.
skull);COMMERCIAL WHALING
In summer and autumn, B. bairdi concentrates in Japanese waters north of the 30°N
parallel. The Japanese whaling season is from April to November (Omura et al,
1955: 93 ), reaching its peak in the summer months of July and August. Omura
and colleagues found the main concentration in the southern part of the range off
the east coast of Japan proper, the whales moving in northwards, according to thewhalers, as the season advances. After breeding, they apparently leave for feeding
grounds in more northern latitudes.
Whaling commences in the northernmost area in the Okhotsk Sea earlier thanelsewhere and has a second peak in autumn but it is supposed that there is a different
population or community in the Okhotsk Sea from elsewhere.
The small numbers taken along the western shores of Japan may be due tofew schools passing west through the Soya and Tsugaru Straits.
The American season is from April to September or October, but as Berardius
is a sporadic visitor it is less important than in Japan (Scheffer and Slipp, 1948:
256).
BREEDING
Omura et al (1955, fig. 28) extrapolate length records to arrive at the following" the pairing of Berardius takes place in February and the parturition in December
in most individuals, length of gestation being about 10 months." Omura (1958)
however, records an anomalous 12 mm long embryo taken in August, and True(1910) records a newly-born calf discovered in June.
The neonatal length, in most cetaceans, is approximately one-third the lengthof the parent. Accordingly, in Berardius bairdi which can grow to about 12.6 m(42 ft), the newly-born calf should approximate 4.2 m (14 ft). Omura et al (1955)
estimated the neonatal length as 4.5 m (15 ft). The largest foetus they record wasin September and measured 4.2 m (14 ft). Stejneger (True, 1910: 64) records
a newly-born calfof 4.8 m (16 ft) washed up on Bering Island on June 5, 1883, withthe remains of the umbilicus still adhering.

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C. McCANN

FOOD

Pike ( 1953) records the stomach contents of a young female as: " about two gallons
of small rockfish and squid remains."
PARASITES

Omura et al. (1955) record Conchoderma sp. attached to the teeth (Pl. XIV, 2). Thisbarnacle has been recorded on the teeth of Hyperoodon ampullatus and Physeter catodon(Wolff, 1960), and I have found it on a tooth of Mesoplodon grayi. A species of Conchoderma is also found commonly on Megaptera nodosa, never on the animal itself but
attached to the barnacle Coronula, so commonly found on the Humpback. Unlike
Coronula, Conchoderma must have a hard substrate to attach itself to.
Pike (1953) recorded nematodes in the stomach of a specimen of B. bairdi.
ACKNOWLEDGMENTS

The author wishes to thank Dr. R. A. Falla, Director, Dominion Museum, Wellington, for the opportunity of undertaking the work; Dr. R.R. Forster, Director,
Otago Museum, Dunedin, for the loan of material; Dr. R. K. Dell and Mr. B. Hamlin for valuable comments and suggestions on the manuscript; Mr. P. M. Hedgland,
Photographer, Dominion Museum, for the excellent anatomical photographs; Dr.
H. Omura, Whales Research Institute, Japan, for permission to reproduce thephotographs of Berardius bairdi; and Miss M. Hall, Librarian, Dominion Museumat the time, for help in obtaining literature.
REFERENCESBEDDARD, F. E., 1900. A book of whales. London, 320pp. 40 illustrations: 228-233.
BENEDEN, P. J. VAN, and F.L.P. GERVAIS, 1868-1880. Osteogr. cet. osteographie des cetaces vivants et fossiles.
Paris, pp. 377, 515, Atlas, pls. XXI-XXIII.
BULLER, W. L., 1876. Presidential address 7th August, 1875; Trans. N. Zeal. Inst., 8: 407.
CARCELLES, A., 1948. Los cetaceos en las aquas argentinas. Mus. Arg. cien. Nat. (Pub. Extra), 238: 6.
DuvERNOY, G. L., 1851. Memoire sur les caracteres osteologiques des genus nouveaux ou des especes nouvelles de oetaces vivant ou fossiles. Ann. Sci. Nat., 3(15): 1-71.
FLOWER, W. H., 1872. On the recent ziphioid whales, with a description of the skeleton of Berardius arnouxi.
Trans. Z,ool. Soc. Lond., 8(3): 203-234, pls. XXVII-XXIX.
FLOWER, W. H., 1878. A further contribution to the knowledge of the existing ziphioid whales: genus
Mesoplodon. Ibid., 10(11): 416, pls. LXXI-LXXII fig. 4 (M. hectori).
FORBES, H. 0., 1893. Observations on the development of the rostrum in the cetacean genus Mesoplodon,
with remarks on some of the species. Proc. ,(,ool. Soc. Lond., 216-236, pls. XII-XV. (M. hectori).
FRASER, F. C., 1950. Note on a skull ofHector's beaked whale, Mesoplodon hectori (Gray) from the Falkland
Islands. Proc. Linn. Soc. Lond., 162: 50-52, pls. III-IV, figs. 1-5.
GRAY, J.E., 1866. Cat. Seals, Whales and dolphins in the British Museum, 384 pp.
GRAY, J.E., 1871. Notes on the Berardius of New Zealand. Ann. Mag. Nat. Hist., (4)8: 115-117.
GRAY,]. E., 1874. List of seals, whales and dolphins of New Zealand. Trans. N. ,(,ea[. Inst., 6: 96-97.
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STUDY OF BERARD/US DUVERNOY131

HAAST, J. voN, 1870. Preliminary notice of a ziphid whale, probably Berardius arnuxi, stranded on the 16th
December, 1868 on the beach near New Brighton, Canterbury. Ibid., 2: 190-192.
HAAsT,j. VON, 1870. Ditto. Ann. Mag. Nat. Hist., (4)6: 348 (Repeat of previous article).
HALE, H. M., 1939. Rare whales in South Australia. S. Austr. Nat., 19(4): 5-6, photo.
HALE, H. M., 1962. Occurrence of the whale Berardius arnuxi in Southern Australia. Ree. S. Austr. Mus.,14(2): 231-243, pls. V-VI, fig. 1.
HARMER, S. F., 1924. On Mesoplodon and other beaked whales. Proc. Zool. Soc. Lond., 541-587.
HECTOR,j., 1870. List of the bones of Seals and whales in the Colonial Museum, Wellington, New Zealand.
Ann. Mag. Nat. Hist., (4)5: 220-224.
HECTOR, J., 1872. On the whales and dolphins of New Zealand seas. Trans. N. Zeal. Inst., 5: 155-170,
pls. I-VI.
HECTOR,j., 1878. Notes on the whales of New Zealand Seas. Ibid., 10: 331-343, pls. XVI-XVII.
KELLOGG, R., 1931. Whaling statistics for the Pacific Coast of North America. Journ, Mamm., 12: 73-77.
KNOX, F.J., and J. HECTOR, 1871. Observations on the Ziphiidae, a family of cetacea. Trans. N. Zeal.
Inst., 3: 125-129.
MALM, A. W., 1883. Skelettdelar af hval, insamlade under expeditionear med Vega 1878-1880. Bihang.
K. Svenska Akad. Hand!., 8(4): 109.
MATSUURA, Y., 1942. On the northern Porpoise whale, Berardius bairdi Stejneger, in waters near Boshu.
Zoo[. Mag., Zoo!. Inst., Fae. Sci. Tokyo Imp. Univ., 54: 466-473.
MATSUURA, Y., 1943. Berardius bairdi Stejneger ranges the Sea of Okhotsk. Ibid, 55: 237.
MATSUURA, Y., 1943. Marine mammals (Kaiju). Tokyo, 298 pp.
MARELLI, C. A., 1920. Revision osteologica de Berardius arnouxii Duv. An. Mus. Univ. Nac. Buenos Aires.,30: 411-444, pls. I-V.
McCANN, C., 1962. The taxonomic status of the beaked whale, Mesoplodon hectori (Gray)-Cetacea. Ree.
Dom. Mus., 4(0).
NEMOTO, T., 1955. White scars on Whales. (I) Lamprey Marks. Sci. Rep. Whales Res. Inst., 10: 69-77,
figs. 1-10.
NISHIMURA, S., and M. NISHIWAKI, 1964. (Berardius bairdi Stejn.). Pub!. Seto Mar. Biol. Lab., Art. 25, pl.
XXI, fig. 1 (only).
NISHIWAKI, M., and N. 0GURO, 1971. Baird's beaked whales caught on the coast of Japan in recent 10
years. Sci. Rep. Whales Res. Inst., 23: 111-122. pls. 1-11.
0HSUMI, S., 1964. Comparison of maturity and accumulation rate of corpora albicantia between left andright ovaries in cetacea. Ibid., 18: 123-147, pl. I.
OLIVER, W.R.B., 1922a. The whales and dolphins of New Zealand. N. Zeal. Journ. of Sci. & Tech., 5(3):129-141.
OLIVER, W.R.B., 1922b. A review of the cetacea of New Zealand seas. Proc. Z,ool. Soc. Land., 557-585, pls.
I-IV.
OMURA, H.K. FUJINO, and S., KIMURA, 1955. Beaked whale Berardius bairdi of Japan, with notes on Z,iphiuscavirostris. Sci. Rep. Whales Res. Inst. Japan., 10: 89-132, pls. 1-X, figs. 1-30.
OMURA, H., 1958. Note on embryo of Baird's beaked whale. Ibid., 13: 213-214, figs. 1-2.
OWEN, R., 1870. Monograph on the British fossil cetacea from the Red Crag. Monogr. Palaeontogr. Soc., 140, pl. 1-V.
PIKE, G. C., 1953. Two records of Berardius bairdi from the coast of British Columbia. Journ. Mamm., 34:
98-104, pl. I.
SLIPP, J. W., and F. WILKE, 1953. The Beaked Whale Berardius on the Washington Coast. Ibid., 34: 105113, pls. 1-11.
SCHEFFER, V. B., andJ. W. SLIPP, 1948. The whales and dolphins of Washington State with a key to thecetaceans of the West Coast of North America. Amer. Mid!. Nat., 39(2): 257-337.
SCHEFFER, V. B., 1949. Notes on three whales from the Aleutian Islands. Pacific Sci., 3: 353, fig. 1.
STEJNEGER, L., 1883. Notes on the natural history of the Commander Islands including descriptions of
new cetaceans. Proc. U.S.N.M., 6: 75-77.
STEJNEGER, L., 1886. On the extermination of the great Northern Sea-cow (Rytina) a Reply to Professor
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

132

C. McCANN

A. E. Nordenskioeld. Journ, Amer. Geog. Soc. N.Y., 18: 317-328.
An unusual record of three species of whales being restricted to pools in Antarctic
sea-ice. Proc. :(,ool. Soc. Lond., 129(3): 325-332, pls. I-VII.
TRUE, F. W., 1910.
An account of the beaked whales of the family ziphiidae in the collection of the UnitedStates National Museum, with remarks on some specimens in other American Museums. Bull. U.S.
N.M., 73: 1-89. pls. I-ILi.
TURNER, W., 1872. On the Occurrence of :(,iphius cavirostris in the South Shetland Seas, and a cpmparison ofits skull with that ofSowerby's whale (Mesoplodon Sowerbyi). Trans. Roy. Soc. Edin., 26: 759-778, Append.
779-780.
WOLFF, T., year.
Rankefodderne Conchoderma og Coronula pa hvaler. Saertryk af 'Flora og Fauna', 66(1):1--8, 6 photos.
TAYLOR, R.H.F., 1957.

Sci. Rep. Whales Res. Inst.,

No. 27, 1975.

133

STUDY OF BERARDIUS DUVERNOYAPPENDIX I.
A

2
3
4
5
6

B

1392
894
1080
494 !
684
748

7

8

150

c

D

E

F

G

H

I

J

1420
870+
1040
560
720
700
445
169

1240
820+
970+
555
623
675
380
166
108
205
199
87
235
350
78
215
156

1193
780
918
455
615
610
405
158
90
201
195
75
210
370
65
194
137
1065
262
202

1262
800+
1000+
570
710
710
430

1220
780+
900+
518
660
640
410
172
92
192
183
90
200
340
65
186
133
1055
250
213

1200
785
910
475
620
620
395
160
95
180
195
90
202
350
82
192
128
1050
280
193

1270
865
1040
530
625
660

1174
800

9

10
11 21012 10213
14
15
16 21317
18 1236
19 29420 222

211
219
92
240
390
71 ·

1245
350
230

MEASUREMENTS OF SKULLS

218
168
1240
327
210

184

95
208
221
82
215
415
73

194
166
1158
287
212

APPENDIX II.
A

1378
2 880
3
4

5

666

6
7

8

188

B

c

1524
960
1185
563
766
750

1423
925
1130
544
682
675

207

197

215
235
110

195
217
98

165

85
261
193
1334
292

82
228
171
1289
295
230

D

1062(?)
578%
720+
530
520

E
1474
1025
1187
575
716

740
223

155
80

493
577

584
378
149

189

91
180
380
63

80

72
186

1011
245

MEASUREMENTS OF

G

H

I

1440
908
1146
569
693
732
447
208

1343
870
865
525
657
660
397
194

1438
923
1122
582
757
748
445
206

1421

204
238
98

203
239

215
246
102

194
230
98

86
246
180
1334

72
216
173
1190

78
221
170

241

212

222

F

1170
571

686

201

9

10 181
11 238
12
96
13
14
84
15
16 24017 16818 1282
19 145+20 223271

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

187

197
239

83

JOO

83
195
142
883+
145+
175

235
178
1360
310
245

71

94

238
163
1300
276
234

134

C. McCANN

OF BERARDIUS ARNOUX! (MILLIMETERS)
K

L

M

N

1350
903
1078
525
662

1372
919
1097
533
625
671

587E
343
443E
235
228
258

150

152

37

203
69

208
74

49

62
188

61
191

90

1225
303
204

1245+
310
211

481
155
83

505+
280+
383+
252
207
238
140
46
37
82
116
49
84
197
38
93
59
440
125
92

0
601
358
469
240
228
242

p
1260
765
648(?)

35

700
435
168

50

200
218
98

91
498
161
91

220
154
1155
290
230

Type Paris Mus.
Otago Mus. Al4.31
c. Otago Mus. A24.69
D. Otago Mus.
E. Dom. Mus. 7
F. Eom. Mus. 416
G. Dom. Mus. 143
H. Dom. Mus. 1402
Mus. Nac. Bs. AiresI.
J. S.R.N.M. 21511
K. Flower 1868 R.C. 3
L. True
M. Brit. Mus. 1677G, 76.2.16.3
N. Dom. Mus. 614
o. Brit. Mus. 1949-8.A.l
P. S. Aust. Mus. M.5012
A.
B.

BERARDIUS BAIRD! (MILLIMETERS)

A.
B.
C.
D.
E.
F.
G.
H.
I.

Type U.S.N.M. 30992
U.S.N.M. 49736? U.S.N.M. 49725
U.S.N.M. 49727
Bering Is!. U.S.N.M.
33 ft. (Pike, 1953)
29 ft. (Pike, 1953)? 34 ft. 5 in. (Slipp & Wilke, 1953)
36 ft. (Omura, Fujino, & Kimura, 1955)

Key to the numbers, 1 to 20, in the appendix tablesof measurements.
1. Total length of skull; 2. Length. of rostrum;
3. Tip of rostrum to posteror median margin ofpterygoids ; 4. Height of vertex to pterygoids, vertical ; 5. Breadth between centre of orbits ; 6.
Breadth between zygomatic processes ; 7. Breadth at
maxillary notches ; 8. Breadth of rostrum at middle ;JO. Greatest breadth of premaxillaries, proximally ;11. Greatest breadth of premaxillaries in front ofanterior nares; 12. Greatest breadth of anteriornares; 13. Length of temporal fossa; ]4. Breadth
between temporal fossae; 15. Breadth of foramen
magnum; 16. Width of occipital condyles; 17.
Height of occipital condyle; 18. Length of mandible; 19. Length of symphysis; 20. Greatest depthof mandible.

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

STUDY OF BERARD/US DUVERNOY135

EXPLANATION OF PLATESPLATE I
Berardins arnouxi Duvernoy stranded in Pukerua Bay, New Zealand.

Photos E. Pain.
PLATE II
Old male of Berardius arnouxi Duvernoy.
Fig. 1. Stranded animal; Fig. 2. Head showing frontal bas, beak and gular grooves;
Fig. 3 Blowhole and scarring; Fig. 4. genital and anal area; Fig. 5. Dorsal fin, newand old scars and wounds; note the parallel scars and possible shark teeth marks(left lower corner).
Photos McCann.
PLATE III
Skull of adult Berardius arnouxi Duvernoy.
(See index to lettering, page 137).
Photos P. M. Hedgland.

Courtesy National Museum.

PLATE IV
Skull of neonatal Berardius arnouxi Duvernoy.
(See index to lettering, page 137).
Photos P. M. Hedgland.

Courtesy National Museum

PLATE V
Sagittal section of adult Berardius arnouxi Duvernoy.
Figs. 1. and 2. general aspect. Fig. 3. Narial region; note' blind pit' (BP) in vomer.
(See index to lettering, page 137)
Courtesy National Museum
Photos P. M. Hedgland.
PLATE VI
Posterior aspect of Skulls of Berardius arnouxi Duvernoy.
Fig. I. neonatal; Fig. 2. adult.
(See index to lettering, page 137)
Courtesy National Museum
Photos C. Hale Fig. I. & P. M. Hedgland Fig. 2.
PLATE VII
Antorbital region of adult Berardius arnouxi Duvernoy.
(See index to lettering, page 137)
Photos P. M. Hedgland.

Courtesy National Museum

PLATE VIII
Antorbital and aural region of neonatal Berardius arnouxi Duvernoy.
Fig. I. Antorbital foramina and optic foramen; note only foramen No. 2 is visibleat this stage. Fig. 2. aural area.
(See index to lettering, page 137).
Courtesy National Museum
Photos P. M. Hedgland.
PLATE IX
Formation of Antorbital tubercle and associated bones in Berardius arnouxi
Duvernoy (adult).
Fig. I. Dorsal aspect; Fig. 2. ventral aspect; Fig. 3. orbital region.
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

136

C. McCANN(See index to lettering, page 122).
Photos P. M. Hedgland.

Courtesy National Museum·

PLATE X
Adult Berardius arnouxi Duvernoy.
Fig. I. Brain cavity; Fig. 2. Aural region.
(See index to lettering, page 122).
Photos P. M. HedglandCourtesy National Museum

PLATE XI
Adult and neonatal tympanic bones contrasted.
A-adult; B-neonatal.
(See index to lettering, page 122).
Photos P. M. HedglandCourtesy National Museum

PLATE XII
Teeth of Berardius arnouxi Duvernoy.
Upper set, inner faces; centre, outer faces of same set; lower, antero-posterior faces.
Courtesy National Museum
Photos P. M. Hedgland
PLATE XIII
Scapulae of Berardius arnouxi Duvernoy.
Fig. 1. Dorsal aspect of adult and neonatal.
Fig. 2. Ventral aspect of same pair.
Photos P. M. Hedgland.

Courtesy National Museum

PLATE XIV
Berardius bairdi Stejneger.

Fig. I. Adult animal; Fig. 2. Extremity of rostrum; note Conchoderena attached toteeth; Fig. 3. Teeth of adult and of a 2,700 mm foetus; top row, anterior, bottomrow, posterior.
PhotosCourtesy Dr. H. Omura.
PLATE XV
Adult skull of Berardius bairdi Stejneger.
(See index to lettering, page 122).
PhotosCourtesy Dr. H. Omura.

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

STUDY OF BERARD/US DUVERNOY137

KEY TO LETTERING AND FIGURES ON PLATES
Ac-Acromion.
ALS(AS)-Alisphenoid bone.
AN-Antorbital notch.
AOF-Antorbital foramen.
AT-Antorbital tubercle.
AU(auf)-Auditory foramen.
B(ty)-Tympanic bulla.
BOC-Basioccipital crest.
BP-' Blind pit' in posterior wall ofvomer (nares).
BS-Basisphenoid bone.
Ca-Carotid foramen.
CE-Condylar foramen.
Cor-Coracoid.
EO-Exoccipital bone.
ET(e)-Eustacian tube.
F-Frontal bone.
FC-Falx cerebri.
FM-Foramen magnum.
FO(fo)-Foramen ovale.
FP-Falciform process.
FS(fl)-Sphenoidal fissure.
J-Jugal process.
L-Lachrymal bone.
LRC-Lower rostral canal.
M-Malar bone (jugal).
MAS-Mastoid process.
MCF-Post maxillary crest foramen.
MD-Mandible.
ME(MES)-Mesethmoid bone.
MF-Mandibular foramen.
MP-Maxillary pit.
MRC-Median Rostral canal.
MRG-Mesorostral groove.
MX-Maxilla.
MXC-Maxillary crest.
MXF-Maxillary foramen.
N-Nasal bone.
NF-Narial foramen.

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

OBS-Orbito-sphenoid bone.
QC-Occipital condyle.
OF(OPF)-Optic foramen.
OS-Orbitosphenoid.
Pa-Parietal.
Pal.-Rid.-Palatine ridge.
PCF-Post crest foramen.
PE(per)-Periotic bone.
PL(Pl)-Palatine bone.
PMF-Post maxillary foramen.
PMX-Premaxillary.
PMXF-Premaxillary foramen.
PS-Presphenoid.
PT-Pterygoid bone.
S-Nasal septum.
SO-Supraoccipital bone.
SQ-Squamosal bone.
Ty(B)-Tympanic bulla.
Vo.-Vomer.
URC-Upper rostral canal.
Ear bones:
a-Posterior articular surface between bones.
a'-Anterior articular surface between bones.
af-Aquaduct of Fallopius.
al-Anterior lobe of periotic bone.
e-Eustacean tube.
earn-External auditory meatus.
epl-Exterior posterior lobe of tympanic.
fo-Fenestra ovalis.
fr.-Fenestra rotunda.
ipl-Internal posterior lobe of tympanic.
m-Mastoid process.
per-periotic bone.
pi-Posterior lobe of periotic.
ty-Tympanic bulla.
Numbers: Nos. I, 2 and 3 refer to foramina in the
antorbital region.

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STRANGE ORGAN IN THE ANAL REGION OF THE
FINLESS PORPOISE
MASAHARU NISHIWAKI
University of the Rycckyus, Okinawa

AND

KAZUO KUREHA
Himeji City Aquarium, Himeji

The Himeji City Aquarium received a telephone call from Mr. Tsuda of Aioi Yacht
Harbor on the morning of October 13, 1974 asking to take over a finless porpoise
which had been caught by a member of the yacht club. The porpoise was foundfloating up and down the surface by a sailing crew near by Karani Is., one of thescattered islands about 20 km to the west of Himeji City 4 km distant from thebeach. Then the crew hauled it with handy net.
When Kureha arrived at the harbor, the porpoise had already been dead.
So, he carried the carcass back for examination. This 114 cm male specimen had
a small hole in the genital aperture just inbetween the anus and the penis opening.
Investigating with a surgical tool, the examiner felt the cavity was about 2 cm
deep and had closed end. Then Nishiwaki was invited to join the examination. The opening hole was on the center line of ventral surface and looked likea small vagina. At dissection, it was found that the hole was the opening of a blindsac 3 mm in diameter and 24 mm in length inside. The sac narrows immediately
at the close end and connected to the hind part of the urinary bladder with thinconnective tissue. There was no organ such as uterus or ovaries besides it nor any
gland found around the sac by naked eyes.
There has been a description on this opening. Dr. K. Mizue of the Nagasaki
University studied on smaller cetaceans in the western waters of Kyushu about ten
years ago. He reported on the finless porpoises caught in Tachibana Bay in his
research report-XII that "naturally, males of this species has no nipple slits along
the genital opening, but there is a blind sac just inbetween the genital opening andthe anus." He had a interest on it and would study separately in future. Asked
for further information, Dr. Mizue answered that he had seen 12 males at that
research and seen this organ in every specimen. So, he thought it might be quite
common, not a matter of drawing special attention nor he have kept any specimenof this portion. When he thought back, he was not sure whether all the twelve
porpoises had this hole. Although he thought it was a degenerated nipple, now
he thinks it was not.
Because there is only one specimen of this portion of finless porpoise, it might
be better to leave histological study undone until more specimen will have found.
Here the authors intend to report a finding of the strange sac and want to draw
Sci. Rep. Whales Res. Inst.,

No. 27, 1975, 139-140

140

NISHIWAKI AND KUREHA

attention of scientific collegues to get further information and to know whether thisis an aromatic or civet secretion gland or a harmaphroditic vagina.
REFERENCE~hzuE, K., K. YosHIDA, and Y. MASAKI, 1965.

Neomeris phocaenoides, so-called Japanese" SUNAMERI '',caught in the coast of Tachibana Bay, Nagasaki Pref. Studies on the Little Toothed Whales in theWest Sea Area of Kyusyu-XII. Bull. Fae. Fish., Nagasaki University, 18: 7-29.

EXPLANATION OF PLATE
Fig. I. The 114 cm male specimen of the finless porpoise (Neophocoena phocaenoides).
Fig. 2. Ventral surface of the specimen. N: navel, P: penis opening, H: strange
hole, A: anus.
Fig. 3. Dissected genital aparture of the specimen. T: testis, U: urinary bladder,
R: rectum, P: penis, N: navel, A: anus, H: strange hole.

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

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DISTRIBUTION OF SPERM WHALE CATCHES IN THE
SOUTHERN INDIAN OCEANSABURO MACHIDA*
Whales Research Institute, Tokyo

ABSTRACTOver 6,000 sperm whales were caught by Japanese antarctic whaling expeditions in the southern Indian Ocean bounded by latitude 30° -50°S andlongitude 20°-l45°E on the way to and from the antarctic whaling grounds
in the seasons of 1957/58-1972/73. Most catches were made intensively in
November and December before the baleen whale seasons.
The distribution of the species in the area concerned is discussed in relation to the surface temperature field. The relationship between the heavy
catches of the animals and the Subtropical Convergence was found. Spermwhale concentrations are restricted in the convergence region and in temperate waters north of it. They are distributed up to about 41°S in the southwestern, southcentral, and southeastern Indian Ocean and about 47°S south ofAustralia. Virtually no females occur in colder waters south of the convergence region.
INTRODUCTION
Sperm whales have been caught by Japanese antarctic whaling expeditions in middle latitudes in the southern Indian Ocean on the way to and from the antarctic region in the Antarctic Ocean since the 1957/58 season.
There are very few reports about the distribution of sperm whales in pelagic
waters in the Indian Ocean. Especially in the southern part of the ocean only verysmall catches of the species were made by the American whaleships mainly in nineteenth century (Townsend, 1937) and only a few animals were observed by research
and merchant vesseles (Brown, 1957; Slijper et al., 1964). On the other hand we
can see the distribution of the species in the area concerned in the charts by Kasuya(1964), Ohsumi and Nasu (1968), and Berzin (1972), which were mapped basedon the materials from Japanese and Soviet antarctic whaling expeditions respectively. However, the practical study of the sperm whale distribution and thegrounds for the species in relation to the oceanographic conditions is rather scarce.
In any programme of whale research aimed at the reasonable exploitation of
a population a knowledge of the distribution of the animals concerned is invaluable.
In this present report the distribution of the sperm whale catches in the southern
Indian Ocean bounded by latitude 30°-50°S and longitude 20°-145°E has been

* Present address: Japan Marine Fishery Resource Research Center, Tokyo.
Sci. Rep. Whales Res. Inst.,

No. 27, 1975, 141-157.

142

MACHIDA

described in relation to the surface temperature field, using the materials fromJapanese antarctic whaling expeditions in the seasons of 1957/58-1972/73.
RESULTSJapanese antarctic whaling expeditions had caught sperm whales heavily in middle
latitudes in the southern Indian Ocean from 1957/58 to 1972/73 (Table 1). Though
the sperm whale catches were made during the warmer months of November-April,most animals were actually caught in November before the baleen whale seasons.
A few catches of the animals were also made at the beginning of December and fromthe end of March to April again after carrying out catching for baleen whales.
During the mid summer (January-February), at the height of catching for baleen
whales, only very few sperm whales were caught.
TABLE 1. THE SEASONAL AND MONTHLY CATCHES OF SPERM WHALES BY THE
JAPANESE ANTARCTIC WHALING EXPEDITIONS IN THE SOUTHERN INDIAN
OCEAN FROM 1957/58 TO 1972/73. RIGHT FIGURES: FEMALES
SeasonNov.

1957/58
58/59
59/60
60/61
61/62
62/63
63/64
64/65
65/66
66/67
67/68
68/69
69/70
70/71
71/72
72/73

25

Total

Dec.

Jan.

Feb.

Apr.

Mar.

Total
25

62
391
1,750
1,194
254
309
20

7
187
960
684
7
47

302
458
45

137
305
9

4,810

2,343

46

18

8
22

5

-

1
3 27 -

13211
74
394 206
19
71
738

304

77

-

10 -

14 -

18

-

42

-

29
1
64
60
48
5
159
83

449

29
76 105 36
2

2

210

36

29
20
203
630
1,798
1, 199
414
417
69

7
223
962
684
7
52

315
687
439
71

137
379
215
19

6,326

2,685

Fig. 1 shows the relation between the number of sperm whales and surface
temperatures. It can be readily seen from the figure that the catches of the animals
in November, when most catches were made, have relations with temperature. A
large quantity of catches were made in the relatively limited range from 12° to l6°C
with a peak at l 4°C though the whales occur in the wide range of -1° to 23°C.
On the other hand, in other months, when few sperm whales were caught, therelationship between the catch and temperature can not be seen as well as that inNovember.
Fig. 2 shows the distribution of sperm whale catches in the southern Indian
Ocean from November to April in the seasons of 1957/58-1972/73. Most of theSci. Rep. Whales Res. Inst.,
No. 27, 1975.

DISTRIBUTION OF SPERM WHALE CATCHES

143

catches were made in the central part of the area in November. The southern halfof the favorite grounds is located in shallow waters around Newamsterdam Island(about 37°50'S and 77°30'E) and Saint Paul Island (about 38°40'S and 77°35'E).
The second large catches of the species, which were almost carried out at the beginning of December, are made off southwestern Australia. In the western part of
the area, the distribution of the catches are widely scattered and the extent of thefavorite haunts were hardly encountered though the total number of sperm whale
catches in the area concerned are rather large. In general a large quantity ofcatches of sperm whales was distributed in the warmer waters north of 40° to 41°S.
It is well known that the sperm whale has some social schools. We can not,of course, find out the accurate composition of the schools caught from the catchrecords but the catches of females, with which some large-scale social schools arecomposed, are available. As females have been also caught by Japanese antarctic
whaling fleets in middle latitudes in the southern Indian Ocean the catch location
are presented here.
The relationship between the nature of female sperm whales and oceanographic condition is shown in Fig. 3, which gives the one between the monthly
catch of females and the surface temperatures. Though females were frequently
encountered in l0°-23°C, the catches increased abruptly (from 19 to 277 animals)
in temperate waters from 11° to l2°C. A large number of the female catches weremade in November, and they occurred in waters between 12° and 19°C with apeak at l4°C. During December, probably at the beginning of the month, several
females were caught was made in warmer waters (14° to 20°C) than during November. In addition only a few females were caught in warmer temperatures than
17°C in March-April.
Fig. 4 shows the distribution of the female catches, most of which were also
captured in November to the beginning of December and a slight catch again in
March to April. Furthermore most females were caught in the southcentral Indian Ocean and some animals also caught off southwestern Australia. No females
were caught south of 42°S in the southern Indian Ocean except in the eastern extremity south of southeastern Australia, where several females were encountered as
far south as about 4 7°S.
DISCUSSION

Sperm whales are caught heavily in lower latitudes north of about 41°S (Fig. 2),
especially in November, though the catches are carried out over the area concerned. The southern extremity of the heavy catch area seems to be corresponding
with a certain oceanographic condition. In Deacon's (1937) chart the Subtropical
Convergence, which is the boundary between the Subtropical and Subantarctic
Surface Waters, was drawn between about 39° and 43°S in the southern Indian
Ocean and between about 45° and 47°S south of Tasmania Island in the easternmost area of the ocean. According to Orren (1966) the convergence was foundto lie at 42°S ± 1° in the south western part of the ocean. In the southeastern IndianSci. Rep. Whales Res. Inst.,
No. 27, 1975.

144

MACHIDA

Ocean the convergence region was shown in latitudes from about 39° to 42°S in
Burling's (1961) chart. Rochford (1962) illustrated the South Transitional Zone
schematically, that is the transitional zone between the Subtropical and Subantarctic
Zones, which locates between about 35° and 40°S in the southeastern Indian Ocean.
In addition the Japanese whaling fleet also crossed the convergence region in nearlysame latitudes in the southeastern Indian Ocean (Machida, 1974, a, b, unpublisheddata). On the other hand, in the southcentral part of the ocean, according to
Deacon (1937) the surface temperature observations by the rv Valdivia and Gauss
show that the convergence lies between 40° and 41°S in about 76°E. The rv
Anton Brunn crossed the Subtropical Convergence at about 40°S, 75°E in April(Uda and Nakamura, 1973). Deacon's (1963) examination of mean monthly
temperatures of I-degree squares by Doneke (1938), though little is reported about
the seasonal variation of the convergence, suggests that its variations of the position
are rather small, of the same order as these of the Antarctic Convergence. Taking
account into these knowledge of the Subtropical Convergence it is evident that
good catches of sperm whales are related to the convergence region.
According to Fig. 1 the relationship between the good catches of sperm whales
and the Subtropical Convergence will be confirmed. A large quantity of the catches
were made in the range from 12° to 16°C with a peak at l4°C during November,when is outside the baleen whale seasons. Deacon (1963) described that the Subtropical Convergence is generally given in a close distribution of the surface isotherms of 10°-l4°C in winter and l4°-18°C in summer. Though he divided ayear into two seasons, winter and summer, November should be rather grouped
into the transitional season from winter to summer. There, however, are very few
reports on the Subtropical Convergence except the summer season. In the southeastern Indian Ocean (around 40.5°S and 108°E) a Japanese antarctic whaling fleet
crossed the convergence region from 13.5° to 10.4°C in the middle of November(Machida, 1974b). The range of temperature may be classified into the wintercategory according to Deacon's (1963) criterion. November, however, should begrouped into the warmer month or rather into the transitional season from winterto summer, i.e. spring. Simply taking account of the Deacon's (1963) generalcriterion about the Subtropital Convergence in the transitional season from winterto summer it probably lie in the range between 12° and l6°C, which are themeadian of the range corresponding with the convergence in winter and in summer
respectively. In November a large number of sperm whales, as mentioned above,were caught in these temperatures.
On the other hand a slight catch at the beginning of December, of which thegreat majority of the catches were made off southwestern Australia, appears to becorresponding to warmer waters than 14°C. The warmer temperatures than l4°C
are included into the summer category of Deacon's (1963) criterion of the Subtropical Convergence and coincident with the range which corresponds to the convergenceregion in December (Machida, unpublished data).
Though in the southern Indian Ocean only very small catches of sperm whales
were made mainly in the nineteenth century (Townsend, 1937) and only a few animals
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

DISTRIBUTION OF SPERM WHALE CATCHES

145

were sighted by research and merchant vesseles (Brown, 1957: Slijper et al., 1964),
Kasuya (1964) and Ohsumi and Nasu (1968) illustrated the sperm whale distributionin pelagic waters of the southern Indian Ocean from catch records of Japaneseantarctic whaling expeditions. In addition Berzin (1972) charted the distribution of the species in the area concerned based on the recent materials from Soviet
pelagic whaling expeditions with previous works. In these charts it is evident
that the heavy catches of sperm whales are located around the Subtropical Convergence in the southern Indian Ocean during October-March. He describedfurthermore that large concentrations were noted north of Newamsterdam andSaint Paul Islands. A heavy catch of the animals by Japanese fleets was also
made from in shallow waters around both two islands to north. The large catches
was also illustrated off Southwestern Australia as the favourable grounds in thecharts by Townsend (1937), Gilmore (1959), Mackintosh (1965), Berzin (1972), andothers.
The southern extremity of the good catches of sperm whales seems to reflect
the concentrations of the large-scale schools of the species. According to Gaskin(1970), Ohsumi (1971), and others the sperm whale has the distinct social structureof various types of school, i.e. the nursely schools, the harem schools, the immature
mixed sex schools (the juveniel schools), the bachelor schools, the bull school (malepairs) and the lone bull (solitary bull). Of which the nursely, harem and juveniel
schools are composed of a large number of females. In the short period before thebaleen whale seasons catching for sperm whales will be occupied in waters inhabitingthese schools. We can not, of course, find out the accurate category of the social
structure of the species from the catch recrods but the females are recorded in thecatch logs. It is quite possible to suppose the location of the large-scale schools
from that of females, therefore. The distribution of the female catches shownin Fig. 4 may suggest the distribution pattern of the major schools of the species
which are restricted in the Subtropical Convergence.
The females and the youngs are generally accepted being limited to the regions
between about 40°N and 40°S (Slijper, 1962). Gilmore's (1959) charts show thatfemales are limited by the isotherm of 20°C, which seems to be more or less the range
as the convergence. Kasuya ( 1964), who charted the seasonal distribution of maleand female sperm whales in the rectangles by latitude 1° and longitude 5°, using thepart of the same materials as that in this report, assumed that the southern limit of
the female distribution in the southern Indian Ocean is 40°-45°S in latitude andthat the range is roughly corresponding with the Subtropical Convergence. On
the other hand Nasu (1969) pointed out that the southern range of females wascorresponding with the Australasian Subantarctic Front, which is located in theregion between the Subtropical and Antarctic Convergences (Burling, 1961) basedon the distribution of female sperm whales by Ohsumi and Nasu (1968). They
charted the distribution of the female catches in ten degree squares by using the partof the same materials as that in Kasuya (1964) and in this report. Clarke (1972)
studied the stomach contents of sperm whales caught off Durban (South Africa) andreported that few antarctic cephalopod beaks were found from females and small-sized
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

146

MACHIDAmales though larger males having antarctic cephalopod beaks in large number.
From this fact he suggests that females and small-sized males do not go far south
through the Subtropical Convergence. I may take that the southern range offemale sperm whales in the southern Indian Ocean is corresponding with the Subtropical Convergence as far as I discuss the distribution of the female catches andthe surface temperatures in Fig. 3 and 4.
On the other hand there is really several records that the female and young
sperm whales occurred in higher latitudes beyond the Subtropical and Antarctic
Convergences. Salvenson (1915) illustrated one female caught off South Gergia(about 55°S). In the southern Indian Ocean several schools of small-sized sperm
whales, which seemed to be composed of females and youngs, were observed at theend of January at 59°S, 65°E and at 58°30'S, 71°E, respectively (Ivanov, 1972).
Fourteen females were caught at the end of January south of Herd Island (about
56°S) (lvashin and Budylenko, 1970). Though these observations and records
suggests that some females and youngs are tolerable to cold waters south of the
Subtropical Convergence and/or furthermore through the Antarctic Convergence,these are rather scarce and I may regarde them as strugglers.

REFERENCESBERZIN, A. A., 1971. The sperm wha/,e (Eng. transl. by A. V. Yablokov, IPST, Jerusalem, 1972, 394 pp.).
BROWN, S. G., 1957. Whales observed in the Indian Ocean. Mar. Obsr., 27: 157-165.
BURLING, R. W., 1961. Hydrology of circumpolar waters south of New Zealand. N.
Dep. sci. indust.
Res. Bull., 143 (N. Z· oceano. Inst. Mem., 10), 66 pp.
CALDWELL, D. K., CALDWELL, M. C., and D. W. RrcE, 1966. Behaviour of the sperm whale, Physetercatodon L. Whales, Dolphins, and Porpoises (ed. K. S. Norris), 677-717.
CLARKE, M. R., 1972. New technique for the study of sperm whale migration. Nature, 238: 405-406.
DEACON, G. E. R., 1937. Hydrology of the Southern Ocean. Discovery Rep., 15: 1-124.
DEACON, G.E.R., 1963. The Southern Ocean. The Sea 2 (ed. M. N. Hill), 281-296.
DEACON, G.E.R., 1966. Subtropical Convergence. The Encyclopedia of Oceanography (ed. R. W. Fairbridge),
884-885.
GASKIN, D. E., 1970. Composition of schools of sperm whales Physeter catodon Linn. east of New Zealand.
N. Z· Jl. Mar. Freshwat. Res., 4(4): 456-471.
IVANOV, A. P., 1972. Small-sized sperm whales in the Anatrctic. (Jap. transl. by H. Sakiura, Geiken Tsushin,
282: 14)lvAsHIN, M. V., and BuDYLENKO, G. A., 1970. Female sperm whales in the Antarctic? (cited from A. P.
Ivanov, 1972)
KASUYA, T., 1964. Report of biological investigations of whales caught by Japanese antarctic whaling expeditions in
the 1963/64 season. Whales Res. Inst., Tokyo, 79 pp. (in Japanese)
MACHIDA, S., 1974a. Geographical positions of the major oceanographic fronts observed by whale sighting
boat in the Australasian Antarctic in the 1972/73 season. Geiken Tsushin, 269: 1-5. (in Japanese)
MACHIDA, S., 1974b. Temperature observations by the whaling factory 'Chiyo Maru' and whale catcher'Katsu Maru No. 3' on the way to and from the Antarctic in the 1973/74 season. Geiken Tsushin, 275:
51-54. (in Japanese)
MACHIDA, S., (unpublished data) Subtropical Convergence, Subantarctic Front, and Antarctic Convergence observed by Japanese Antarctic Icebreaker' Fuji'.
NAsu, K., 1969. Oceanography in the southern limit of habitat of southern bluefin tuna. Bull. Jap. Soci.
Fish. Oceano., 15: 19-26. (in Japanese)z.

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

DISTRIBUTION OF SPERM WHALE CATCHES

147

0HSUMI, S., and NAsu, K., 1968. Range of habitat of the female sperm whale with reference to oceanographic structure. (cited from K. Nasu, 1968)
0HSUMI, S., 1971. Some investigations on the school structure of sperm whale. Sci. Rep. Whales Res. Inst.,

23: 1-25.
ORREN, M. R., 1966.

Hydrology of the south west Indian Ocean. Invest. Rep. Div. Sea. Fish. S. Afr., 55: 136.
ROCHFORD, M. R., 1962. Hydrology of the Indian Ocean II. Aust. J. mar. Freshw. Res., 13: 226-251.
SALVESON, T. E., 1915. The whale fisheries of the Falkland Islands and dependencies. (cited from D. K.
Caldwell et al., 1966)
SLIJPER, E.J., VAN UTRECHT, W. L., and C. NAAKTGEBOEN, 1964. Remarks on the distribution and migration of whales observations from Netherlands ships. Bijdr. Dierkunde, 34, 93 pp.
TOWNSEND, C. H., 1935. The distribution of certain whales as shown by logbook records of American whaleships. .<:,oologica, 19: 1-50.
UDA, M., and NAKAMURA, Y., 1973. Hydrography in relation to tuna fisheries in the Indian Ocean. Spl.
Puhl. Mar. Biol. Ass. India, 1973, 276-292.

Sci. Rep. Whales Res. Inst.,

No. 27, 1975.

148

MACHIDA2x10

Nov.-Apr.

12
10

~8
:::J

Iii

u

..... 6

Q)

.0

E

:i. 4

0 2 4 6 8 10 12 14 16 18 20 22
Surface Temperature (°C)
2
x 10

12

Nov.

10
8

6

.....

Q)

~ 0 L....c:::i.........r::==--...L--=~-------='-~~
1~~--------"'--:--Dec.

1..--tL[--~-=='-='---'==J'""J...=-'~----J-a_n_.
Feb.

Mar.

[

~~[_ _ ___,,==-~oJ.....Ln~·~·n=-~n..L...J.~_._._,,,=---<=-=-~A~~...__r.
0
Fig. I.

2 4 6 8 10 12 14 16 18 20 22
Surface Temperature (~C)

Catches of sperm whales and surface temperatures in November-April(1957/58-1972/73)
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

149

DISTRIBUTION OF SPERM WHALE CATCHES
Fig. 2(a). Distribution of sperm
whale catches in NovemberApril (1957/58-1972/73). The
fine line represents the contourof 1,500 fathoms.

••

• •
o·•

1 -15

•

.• . 0•• 16-30• 31 -45

•

• 46-

•

·~·
.....• • :··• • •. ......
..
.

.•. •

.

.·•

• • • •••••

2~0____.__·_·~·-·-·__,_·____4~0---'-----''----~0----6'8°

• •

..•••
.

. • . •..
.

••

••

•

.

•••••
.

•

..
•

••

60

100

80

• • •

•

.......

...._~--~~

100
Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

......

~~.......

~~~~--~~..__~__.---~

120~~"'!'lo

140

150

MACHIDA• •••Fig. 2(b) Distribution of sperm
whale catches in November(1~57/88-1972/73).

1 -15• 16 - 30

••
•

•

•

.•

• 0

• 31 -45
• 46-

0

40

20

• •

•

·•••...••
••

40

•

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

151

DISTRIBUTION OF SPERM WHALE CATCHES

Fig. 2(c). Distribution of sperm
whale catches in December( 1957/58-1972/73).

1 -15• 16 - 30• 31 -45
• 46-

40

0

20

40

•
40

60

eo

•

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

152

MACHIDA30
70E 45s

50 40
135E 40

0

s

\)~
20

•

45
50

40

60

Fig. 2(d). Distribution of sperm whale catches in January (1957/58-1972/73).

1 - 15• 16 -30• 31 -45
• 46-

0

40

0

\> ~~..
.

020
Fig. 2(e).

40Distribution of sperm whale catches in February (1957/58-1972/73).

1 - 15• 16 -30• 31 -45• 46Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

153

DISTRIBUTION OF SPERM WHALE CATCHES

Fig. 2(f). Distribution of sperm
whale catches in March (1957/
58-1972/73).

1 -15• 16 -30• 31 -45
• 46-

•

•

.'1

••

0

·•
80

60

10

Fig. 2(g). Distribution of sperm
whale catches in April (1957/
58-1972/73).

1 -15• 16 -30• 31 -45
• 46-

•
0

•

••
• •

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

40•

MACHIDA

154

2

Nov.-Apr.

x10...... 6
.c
Ol
:::J

~ 4
s...

<l1

..a

E
2
:::J

z

8 10 12 14 16 18 20 22
Surface Temperature (°C)
2

x10

Nov.

6

......

4

.c

Ol

~ 2

u

I...

~ 0 L-----l=------L.1_...J__~ 1L~r~----c=-...J.....1-.L-1---'--o-e_c_.
s:iDr-ifMar.

~~[~~~-c:~=--~~..__A~::~·
8 10 12 14 16 18 20 22
Surface Temperature (°C)
Fig. 3. Monthly catches of female sperm whales and the surface temperatures (1957/
58-1972/73).

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

155

DISTRIBUTION OF SPERM WHALE CATCHES

Fig. 4(a). Distribution of female
sperm whale catches in November-April (1957/58-1972/
73).

1 -15
• 16 -30• 31 -45
• 46-

•

•

•• ••••••••••

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

40

MACHIDA

156.--~..,.....~..........-=,,.,..,.--..----i....-~-r--...--.--~--r~--,30

•

•

o·

Fig. 4(b ). Distribution of female
sperm whale catches in November (1957/58-1972/73) .

1 -15• 16 -30• 31 -45
• 46-

•••
• •
•

• ••• •.
••• • -~.
••.I. • •

••

•••

•

40

Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

DISTRIBUTION OF SPERM WHALE CATCHES

Fig. 4(c).
73).

157

Distribution of female sperm whale catches in December (1957/58-1972/

1 -15• 16 -30• 31 -45
• 46-

A A.
A. .
A.
A.

~

0

0

Fig. 4(d). Distribution offemale sperm whale catches in March and April (1957 /581972/73). M: March A: April

1 -15• 16 -30• 31 -45• 46Sci. Rep. Whales Res. Inst.,
No. 27, 1975.

THE SCIENTIFIC REPORTS OF THE WHALES RESEARCHINSTITUTE, TOKYO, JAPANNUMBER I, JUNE 1948
Akiya, S. and Tejima, S. Studies on digestive enzymf" in whale. 3-7
Akiya, S., Ishikawa, Y., Tejima, S. and Tanzawa, T. Studies on tryptas!" from a whale (Balaenoptera
borealis L. ). 8-10
Akiya, S., Tejima, S. and Ishikawa, Y. Studit>s on the utilization of whale m<'at by the use of pancreatic tryptase of whales. 11-14
Akiya, S. and Kobo, F. The test culture of some microorganisms with whale meat peptone. 15-16
Nakai, T. Chemical studies on the freshness of whale meat. I. Evaluation of freshness and changes inquantity of several kinds of nitrogen in whale meat following deterioration of freshness. 17-26
Nakai, T. Chemical studies on the freshness of whale meat. II. On comparison between whale meatand beef on deterioration of freshnf"SS and autolysis. 27-30
Tawara, T. On the simultaneous extraction of vitamin A-D and vitamin B2 complex from the liver ofa fin whale (Nagasu-Kujira, Balaenoptera physalus L.). 31-37
Tawara, T. Studies on whale blood. I. On the separation of histidine from whal<' blood. 38-40
Nakai, J. and Shida, T. Sinus-hairs of the sei-whale (Balaenoptera borealis). 41-47

NUMBER 2, DECEMBER 1948
Ogawa, T. and Arifuku, S. On the acoustic system in the cetacean brains. 1-20
Yamada, M. Auditory organ of the whalebone whales. (Preliminary rt>port). 21-30
Nakai, T. Chemical studies on the freshness of whale meat. III. Effect of hydrogen-ion concentration
on decrease in freshness and titration curve of whale meat with HCl and Na 2 C08 • 31-34
Ishikawa, S., Omote, Y. and Soma, Y. Analytical distillation of vitamin A in the whale liver oil. 35-41
Ishikawa, S., Omote, Y. and Kanno, H. Molecular distillation of sperm whale blubber oil. 42-45
Kaneko, A. Molecular distillation of fin whale liver oil. 46-50
Akiya, S. and Takahashi, K. Determination of tryptophane in whale meat. 51-54
Ishikawa, Y. and Tejima, S. Protein digestive power of sperm whale pancreatic enzyme. 55-60
Tsukamoto, S. Experiment on digestion of whale meat by koji-mould. 61-66

NUMBER 3, FEBRUARY 1950
Ogawa, T. and Shida, T. On the sensory tubercles of lips and of oral cavity in the sei and the fin
whale. 1-16
Ohe, T. Distribution of the red marrow in bones of the fin whale. 17-22
Hosokawa, H. On the cetact>an larynx, with special remarks on the laryngeal sack of the sei whale andthe aryteno-epiglottideal tube of the sperm whale. 23-62
Akiba, T., Tsuchiya, T., Umehara, M. and Natsume, Y. Bacteriological studies on freshness of whale
meat. (Report No. 1). 63-70
Ishikawa, Y. Protein digestive power of sperm whale pancreatic enzyme. II. 71-78
Mori, T. and Saiki, M. Properties of fats and oils contained in various parts of a sperm whale body.
79-84
Tawara, T. and Fukazawa, R. Studies on kitol. I. Pr<'paration of kitol from whale liver oil. 85-88
Tawara, T. and Fukazawa, R. Studies on kitol. II. Influence of kitol fraction on the determinationof the International Unit of Vitamin A. 89-91
Tawawa, T. and Fukazawa, R. Studies on kitol. III. The effect of sunlight, air and heat on tht> vitamin A and kitol fractions. 92-95
Tawara, T. On the respiratory pigments of whale (Studies on whale blood II.). 96-101

ii
Yoshida, M. Research on methionine in whale. 102-105
Mizue, K. Factory ship whaling around Bonin Islands in 1948. 106-118
Mizue, K. and Jimbo, H. Statistic study of foetuses of whales. 119-131
Nishiwaki, M. and Hayashi, K. Biological survey of fin and blue whales taken in the Antarctic season
1947-48 by the Japanese fleet. 132-190

NUMBER 4, AUGUST 1950
Omura, H. On the body weight of sperm and sei whales located in the adjacent waters of Japan. 1-13
Omura, H. Diatom infection on blue and fin whales in the Antarctic whaling area V (the Ross Sea
area). 14-26
Omura, H. Whales in the adjacent waters of Japan. 27-113
Nishiwaki, M. Determination of the age of Antarctic blue and fin whales by the colour changes incrystalline lens. 115-161
Nishiwaki, M. Age characteristics in baleen plates. 162-183
Nishiwaki, M. On the body weight of whales. 184-209

NUMBER 5, JUNE 1951
Akiba, T., Umehara, M. and Natsume, Y. Bacteriological studies on freshness of whale meat. (Report
No. II.). 1-4
Hosokawa, H. On the pelvic cartilages of the Balaenoptera-foetuses, with remarks on the specifical andsexual difference. 5-15
Ohe, T. Iconography on the abdominal cavity and viscera of the Balaenoptera, with special remarks
upon the peritoneal coverings. 17-39
Akiya, S. and Hoshino, O. Isolation of histidine from whale blood using 3, 4-dichlorobenzene sulfonicacid. 41-47
Tawara, T. and Fukazawa, R. Studies on kitol. IV. Purification of kitol by chromatographie. 49-51
Ishikawa, S., Omote, Y. and Okuda, H. Substances related to vitamin A in the whale liver oil. 53-59
Ishikawa, S., Omote, Y., Kijima, M. and Okuda, H. Thermal decomposition of kitol. 61-69
Mizue, K. Grey whales in the east sea area of Korea. 71-79
Mizue, K. Food of whales (In the adjacent waters of Japan). 81-90
Nishiwaki, M. and Ohe, T. Biological investigation on blue whales (Balaenoptera musculus) and fin whales(Balaenoptera physalus) caught by the Japanese Antarctic whaling fleets. 91-167

NUMBER 6, DECEMBER 1951
Hosokawa, H. On the extrinsic t>ye muscles of the whale, with special remarks upon the innervationand function of the musculus retractor bulbi. 1-33
Murata, T. Histological studies on the respiratory portions of the lungs of cetacea. 35-47
Kojima, T. On the brain of the sperm whale (Physeter catodon L.). 49-72
Mizue, K. and Murata, T. Biological investigation on the whales caught by the Japanese Antarctic
whaling fleets season 1949-50. 73-131
Nishiwaki, M. On the periodic mark on the baleen plates as the sign of annual growth. 133-152
Nishiwaki, M. and Hibiya, T. On the sexual maturity of the sperm whales (Physeter catodon) found in
the adjacent waters of Japan (I). 153-165
Nakai, T. Chemical studies on freshness of whale meat. IV. Some informations of Archomobacter ubiquitum
isolated from whale carcass. 167-176
Nakai, T. and Ono, H. The effects of electric shock and fatigue on post-mortem changes in muscle.
177-185

iii
Omote, Y. Complete recovery of vitamin A from molecular distillation residue of whale-liver oil. 187191
Omote, Y. Chemical structure of kitol (I). Double bonds and hydroxyl groups. 193-198
Hirata, M. Experimental investigation on flattened head harpoon. An attempt for restraining ricochet. 199-207

NUMBER 7, JULY 1952
Ogawa, T. On the cardiac nerves of some cetacea, with special reference to those of Berardius bairdii
Stejneger. 1-22
Akiya, S., Hoshino, 0. and Motohashi, N. On an attempt to preserve whale meat freshness with 5nitro-furfuriden aminoguanidine from decay. 23-30
Akiya, S. and Sawamura, R. Colorimetric determination of 5-nitro-2-furfuridene aminoguanidine. 31-36
Tomiyama, S. and Takao, M. Studies on utilization of higher fatty alcohol from sperm whale oil. 3746
Omote, Y. A rapid method for the separate determination of vitamin A and kitol in the whale-liveroil. 47-50
Arai, Y. and Sakai, S. Whale meat in nutrition. 51-67
Yamaguchi, K. and Fujino, K. On the serological constitution of striped dolphin (Prodelphinus caeruleoalbus (Meyen)) (I). 67-77
Nishimoto, S., Tozawa, M. and Kawakami, T. Food of sei whales (Balaenoptera borealis) caught in the
Bonin Island waters. 79-85
Nishiwaki, M. On the age-determination of Mystacoceti, chiefly blue and fin whales. 87-119
Nishiwaki, M. and Hibiya, T. On the sexual maturity of the sperm whales (Physeter catodon) found in
the adjacent waters of Japan (II). 121-124
Ohno, M. and Fujino, K. Biological investigation on the whales caught by the Japanese Antarctic
whaling fleets, season 1950/51. 125-188

NUMBER 8, JUNE 1953
Yamada, M. Contribution to the anatomy of the organ of hearing of whales. 1-79
Omura, H. Biological study on humpback whales in the Antarctic whaling areas IV and V. 81-102
Fujino, K. On the serological constitutions of the sei-, fin-, blue- and humpback-whales (I). 103-125
Ogawa, T. On the presence and disappearance of the hind limb in the cetacean embryos. 127-132
Nishiwaki, M. and Yagi, T. On the age and the growth of teeth in a dolphin (Prodelphinus caeruleoalbus). (I). 133-146
Kakuwa, Z., Kawakami, T. and Iguchi, K. Biological investigation on the whales caught by the Japanese Antarctic whaling fleets in the 1951-52 season. 147-213
Nishiwaki, M. Hermaphroditism in a dolphin (Prodelphinus caeruleo-albus). 215-218

NUMBER 9, JUNE 1954
Akiya, S., Hoshino, 0. and Motohashi, N. Attempt to preserv" freshness of whale meat with germicides. II. 1-10
Ogawa, T. On the musculature of the sinus venosus and its continuation with the so-called conducting
system of the whale's heart. 11-35
Yamada, M. Some remarks on the pygmy sperm whale, Kogia. 37-58
Yamada, M. An account of a rare porpoise, Feresa Gray from Japan. 59-88
Omura, H. and Fujino, K. Sei whales in the adjacent waters of Japan. II. Further studies on theexternal characters. 89-103
Fujino, K. On the serological constitution of the sperm- and haired beaked-whales (I) Blood groups of

iv
the sperm- and haired beaked-whales. 105-120
Fujino, K. On the body proportions of the fin whales (Balaenoptera physalus (L)) caught in the northern
Pacific Ocean (I) (Preliminary report). 121-163
Nishiwaki, M., Hibiya, T. and Kimura, S. On the sexual maturity of the sei whale of the Boninwaters. 165-1 77
Uda, M. Studies of the relation between the whaling grounds and the hydrographical conditions (I).
179-187

NUMBER 10, JUNE 1955
Hosokawa, H. Cross-sections of a 12-mm. dolphin embryo. 1-68
Nemoto, T. White scars on whales (I) Lamprey marks. 67-77
Omura, H. and Nemoto, T. Sei whales in the adjacent waters of Japan. III. Relation between movement and water temperature of the sea. 79-87
Omura, H., Fujino, K. and Kimura, S. Beaked whale Berardius bairdi of Japan, with notes of ZJphiuscavirostris. 89-132
Fujino, K. On the body weight of the sei whales located in the adjacent waters of Japan (II). 133141
Nishiwaki, M. On the sexual maturity of the Antarctic male sperm whale (Physeter catodon L.). 143149
Ohta, K., Watarai, T., Oishi, T., Ueshiba, Y., Hirose, S., Yoshizawa, T., Akikusa, Y., Sato, M. andOkano, K. Composition of fin whale milk. 151-167

NUMBER 11, JUNE 1956
Omura, H. and Sakiura, H. Studies on the little piked whale from the coast of Japan. 1-3'7
Nishiwaki, M., Hibiya, T. and Kimura, S. On the sexual maturity of the sperm whale (Physeter catodon)
found in the North Pacific. 39-46
Fujino, K. On the body proportions of the sperm whales (Physeter catoaon). 47-83
Fujino, K. On the serological constitution of the fin whales II. Furth~r studies on blood groups. 8598
Nemoto, T. On the diatoms of the skin film of whales in the northern Pacific. 99-132
Hoshina, T. and Sugiura, Y. On a skin disease and a nematode parasite of a dolphin, Tursiops truncatus(Montagu, 1821). 133-138
Iwai, E. Descriptions on unidentified species of dibranchiate cephalopods. I. An oegopsiden squid belonging to the genus Architeuthis. 139-151
Iwai, E. Descriptions on unidentified species of dibranchiate cephalopods. II. A cranchiidae squid of thegenus Taonius. 153-161
Uda, M. and Nasu, K. Studies of the whaling grounds in the northern sea-region of the Pacific Ocean
in relation to the meteorological and oceanographic conditions. (Part I). 163-179
Kimura, S. and Nemoto, T. Note on a minke whale kept alive in aquarium. 181-189
Ishikawa, Y. A characteristic property of whale oils concerning the absorption of gases. I. On theabsorption of carbon dioxide by whale oils. 191-213

NUMBER 12, JUNE 1957
Omura, H. Osteological study of the little piked whale from the coast of Japan. 1-21
Nishiwaki, M. Age characteristics of ear plugs of whales. 23-32
Nemoto, T. Foods of baleen whales in the northern Pacific. 33-89
Nasu, K. Oceanographic conditions of the whaling grounds in the waters adjacent to Aleutian Islands
and the Bering Sea in summer of 1955. 91-101
Kimura, S. The twinning in southern fin whales. 103-125

v
Ichihara, T. An application of lin<"ar discriminant function to external measurements of fin whale.
127-189
Nishiwaki, M. Very small embryo of cetacea. 191-192
Nishiwaki, M. One-eyed monster of fin whale. 193-195
Ogawa, T. and Kamiya, T. A case of the cachalot with protruded rudimentary hind limbs. 197-208
Uda, M. and Dairokuno, A. Studies of the relation between the whaling grounds and the hydrographic
conditions. II. A study of the relation between the whaling grounds off Kinkazan and the boundary of water masses. 209-224
Abe, T. Notes on fishes from the stomachs of whales taken in the Antarctic. I. Xenocyttus nemotoi, a
new genus and neow species of zeomorph fish of the subfamily Oreosonimae Goode and Bean, 1895.
225-233
Tsuyuki, H. On the oils contained in various blubbers of northern elephant seal, Mirounga angustirostris.
235-240NUMBER 13, SEPTEMBER 1958
Omura, H. North Pacific right whale. 1-52
Nishiwaki, M. and Kamiya, T. A beaked whale Mesoplodon stranded at Oiso Beach, Japan. 53-83
Nishiwaki, M. and Randa, C. Killer whales caught in the coastal waters off Japan for recent 10 years.
85-96
Ohsumi, S. (Kimura), Nishiwaki, M. and Hibiya, T. Growth of fin whale in the northern Pacific.
97-133
Nishiwaki, M., Hibiya, T. and Ohsumi, S. (Kimura). Age study of sperm whale based on reading of
tooth laminations. 135-153
Nishiwaki, M., Ichihara, T. and Ohsumi, S. (Kimura). Age studies of fin whale based on ear plug.
155-169
Fujino, K. On the serological constitution of fin whale. III. Human B blood group substances in erythrocytes and some notes on anti-fin Ju specific antibodies. 171-184
Nemoto, T. Cocconeis diatoms infected on whales in the Antarctic. 185-191
Nemoto, T. and Nasu, K. Thysanoessa macrura as a food of baleen whales in the Antarctic. 193-199
Ichihara, T. Gray whale observed in the Bering Sea. 201-205
Ohsumi, S. (Kimura). A descendant of Moby Dick or a white sperm whale. 207-209
Nasu, K. Deformed lower jaw of sperm whale. 211-212.
Omura, H. Note on embryo of Baird's beaked whale. 231-214
Uda, M. and Suzuki, N. Studies of the relation between the whaling grounds and the hydrographic
conditions. III. The averaged conditions of the whaling grounds and their trends of variation during
1946-55. 215-229
Seki, Y. Observations on the spinal cord of the right whale. 231-251
Kamiya, T. How to count the renculi of the cetacean kidneys, with special regard to the kidney ofthe right whale. 253-267
Hosokawa, H. and Sekino, T. Comparison of the size of cells and some histological formations between
whales and man. 269-301
Ogawa, T., Tsunoda, T. and Osawa, M. Amino acid composition of whale meat. 303-307
Ishikawa, Y. A characteristic property of whale oils concerning the absorption of gases. II. On theabsorption of nitrogen by whale oils. 309-321 Tsuyuki, H. Component fatty acids of northern elephant seal oil. 323-332

NUMBER 14, SEPTEMBER 1959
Omura, H. Bryde's whale from the coast of Japan. 1-33
Nishiwaki, M. and Kamiya, T. Mesoplodon stejnegeri from the coast of Japan.
Nishiwaki, M. Humpback whales in Ryukyuan waters. 49-87

35-48

vi
Cushing, John E., Fujino, K. and Takahashi, K. Glycerol-freezing technique as an aid in blood typingof whales. 89-100·
Fujino, K. and Cushing, John E. Blood typing of dried whale erythrocytes with 131 1 labelled antibody.
101-106
Ichihara, T. Formation mechanism of ear plug in baleen whales in relation to glove-finger. 107-135
Nasu, K. Surface water condition in the Antarctic whaling Pacific area in 1956-57. 137-143
Ohsumi, S. (Kimura). A deformed fin whale foetus. 145-147
Nemoto, T. Food of baleen whales with reference to whale mevements. 149-290
Yamada, M. and Yoshizaki, F. Osseous labyrinth of cetacea. 291-304
Nakai, T. Distribution of amino acid in proteins from various parts of whale body. 305-326

NUMBER 15, NOVEMBER 1960
Nishiwaki, M. Ryukyuan humpback whaling in 1960. 1-16
Ohsumi, S. Relative growth of the fin whale, Balaenoptera physalus (Linn.). 17-84
Fujino, K. Immunogenetic and marking approaches to identifying subpopulations of the North Pacificwhales. 85-142
Nasu, K. Oceanographic investigation in the Chukchi Sea during the summer of 1958. 143-158
Tokita, K. and ECG Research Group. Electrocardiographical studies on bottlenosed dolphin (Tursiops
truncatus). 159-165

NUMBER 16, MARCH 1962
Omura, H. Bryde's whale occurs on the coast of Brazil. 1-5
Omura, H. Further information on Bryde's whale from the coast of Japan. 7-18
Nishiwaki, M. Ryukyuan whaling in 1961. 19-28
Nemoto, T. A secondary sexual character of fin whales. 29-34
Omura, H., Nishiwaki, M., lchihara, T. and Kasuya, T. Osteological note of a sperm whale. 35-45
lchihara, T. Prenatal dead foetus of baleen whales. 47-60
Nishiwaki, M. Mesoplodon bowdoini stranded at Akita Beach, Sea of Japan. 61-77
Nishiwaki, M. Observation on two mandibles of Mesoplodon. 79-82
Sinclair, John. An early dolphin embryo (Stenella caeruleoalbus) in serial sections. 83-87
Nemoto, T. Food of baleen whales collected in recent Japanese Antarctic whaling expeditions. 89-103
Uda, M. Subarctic oceanography in relation to whaling and salmon fisheries. 105-119

NUMBER 17, FEBRUARY 1963
Nishiwaki, M., Ohsumi, S. and Maeda, T. Change of form in the sperm whale accompanied withgrowth. 1-14
Ohsumi, S., Kasuya, T. and Nishiwaki, M. The accumulation rate of dentinal growth layers in the
maxillary tooth of the sperm whale. 15-35
Ichihara, T. Photometric method for counting laminae in ear plug of baleen whale. 37-48
Yoshikawa, T. and Suzuki, T. The lamination of the masseter of the humpback whale. 49-52
Fujino, K. Intra-uterine selection due to maternal-foetal in compatibility of blood type in the whales.
53-65
Cushing, John E., Fujino, K. and Calaprice, N. The Ju blood typing system of the sperm whale andspecific soluble substances. 67-77
Nr>moto, T. New records of sperm whales with protruded rudimentary hind limbs. 79-81
Nemoto, T. and Nasu, K. Stones and other aliens in the stomachs of sperm whales in the Bering Sea.
83-91
Nishiwaki, M. Taxonomical consideration on genera of Delphinidae. 93-103

vii
Nasu, K. Oceanography and whaling ground in the subarctic region of the Pacific Ocean. 105-155
Nemoto, T. Some aspects of the distribution of Calanus cristatus and C. plumchrus in the Bering and its
neighbouring waters, with reference to the feeding of baleen whales. 157-170
Tsuyuki, H. and Naruse, U. Studies on the oil of black right whale in the northern Pacific Ocean.
171-190
Yagi, T., Nishiwaki, M. and Nakajima, M. A preliminary study on the method of time marking withleadsalt and tetracycline on the teeth of northern fur seal. 191-195

NUMBER 18, MARCH 1964
Fujino, K. Fin whale subpopulations in the Antarctic whaling areas II, III and IV. 1-27
Ichihara, T. Prenatal development of ear plug in baleen whales. 29-48
Ohsumi, S. Examination on age determination of the whale. 49-88
Nemoto, T. School of baleen whales in the feeding areas. 89-110
Okutani, T. and Nemoto, T. Squids as the food of sperm whales in the Bering Sea and Alaskan Gulf.
111-122
Ohsumi, S. Comparison of maturity and accumulation rate of corpora albicantia between the left andright ovaries in cetacea. 123-148
Omura, H. A systematic study of the hyoid bones in the baleen whales. 149-170
Nishiwaki, M. Revision of the article "Taxonomical consideration on genera of Delphinidae" in No. 17.
171-172
Tsuyuki, H. and Naruse, U. Studies on the lipids in brain of black right whale in the northern Pacific
Ocean. 173-180

NUMBER 19, APRIL 1965
Ohsumi, S. Reproduction of the sperm whale in the north-west Pacific. 1-35
Kasuya, T. and Ichihara, T. Some informations on minke whales from the Antarctic. 37-43
Nemoto, T. and Kasuya, T. Foods of baleen whales in the Gulf of Alaska of the North Pacific. 45-51
Nishiwaki, M., Nakajima, M., and Kamiya, T. A rare species of dolphin (Stenella attenuata) from Arari,Japan. 53-64
Nishiwaki, M., Kasuya, T., Tobayama, T., Kamiya, T. and Nakajima, M. Feresa attenuata captured at
the Pacific Coast of Japan in 1963. 65-90
Nakajima, M. and Nishiwaki, M. The first occurrence of a porpoise (Electra electra) in Japan. 91-104
Hosokawa, H. and Kamiya, T. Sections of the dolphin's head (Stenella caeruleoalba). 105-133
Ohsumi, S. A dolphin (Stenella caeruleoalba) with protruded rudimentary hind limbs. 135-136

NUMBER 20, SEPTEMBER 1966
Ohsumi, S. Sexual segregation of the sperm whale in the North Pacific. 1-16
Ichihara, T. Criterion for determining age of fin whale with reference to ear plug and baleen plate.
17-82
Kasuya, T. Caryotype of a sei whale. 83-88
Kasuya, T. and Ohsumi, S. A secondary sexual character of the sperm whale. 89-94
Nishiwaki, M. and Norris, K. S. A new genus, Peponocephala, for the odontoceti cetacean species Electra
electra. 95-100
Nishiwaki, M., Nakajima, M. and Tobayama, T. Preliminary experiments for dolphin marking. 101107
Nemoto, T. Thysanoessa euphausiids, comparative morphology, allomorphosis and ecology. 109-155
Nasu, K. Fishery oceanographic study on the baleen whaling grounds. 157-210
Ichihara, T. and Nishiwaki, M. External measurements and weight of a southern elephant seal. 211212

viii
Tsuyuki, H. and Itoh, S.
221

Studies on the oils contained in blubber of a southern elephant seal.

213-

NUMBER 21, JUNE 1969
Omura, H., Ohsumi, S., Nemoto, T., Nasu, K. and Kasuya, T. Black right whales in the North Pacific.
1-78
Nishiwaki, M. and Hasegawa, Y. The discovery of the right whale skull in the Kisagata shell bed.
79-84
Ohsumi, S. Occurrence and rupture of varginal band in the fin, sei, and blue whales. 85-94
Hosokawa, H., Igarashi, S., Kamiya, T. and Hirosawa, K. Morphological characteristics and myelinization of acoustic system in the dolphins (Stenella caeruleoalba). 95-123
Mo!, T. and van Bree, P.J.H. A short contribution to the history of whaling in Japan during the 17th
century. 125-129
Tsuyuki, H. and Itoh, S. Fatty acid composition of finless porpoise oil. 131-135
Tsuyuki, H. and Itoh, S. Fatty acid composition of many toothed pilot whale oil. 137-141

NUMBER 22, JUNE 1970
Omura, H., lchihara, T. and Kasuya, T. Osteology of pygmy blue whale with additional information
on external and other characteristics. 1-27
Nishiwaki, M. and Kasuya, T. Recent record of gray whale in the adjacent waters of Japan and aconsideration on its migration. 29-37
Kasuya, T. and Rice, D. W. Note on baleen plates and on arrangement of parasitic barnacles of graywhale. 39-43
Nishiwaki, M. and Kasuya. T. A Greenland right whale caught at Osaka Bay. 45-62
Nasu, K. and Masaki, Y. Some biological parameters for stock assessment of the Antarctic sei whale.
63-74
Ohsumi, S., Masaki, Y. and Kawamura, A. Stock of the Antarctic minke whale. 75-125
Kawamura, A. Food of sei whale taken by Japanese whaling expeditions in the Antarctic season 1967/
68. 127-152
Nemoto, T. and Yoo, K. I. An amphipod, Parathemisto gaudichaudii as a food of the Antarctic sei whale.
153-158
Tobayama, T., Uchida, S. and Nishiwaki, M. Twin foetuses from a blue white dolphin. 159-162
Machida, S. A sword-fish sword found from a North Pacific sei whale. 16:3-164
Tsuyuki, H. and Itoh, S. Fatty acid components of black right whale oil by gas chromatography. 165170

NUMBER 23, SEPTEMBER 1971
Ohsumi, S. Some investigations on the school structure of sperm whale. 1-25
Kawamura, A. Influence of chasing time to stomach contents of baleen and sperm whales. 27-36
Kasuya, T. Consideration of distribution and migration of toothed whales off the Pacific coast of Japan
based upon aerial sighting record. 37-60
Omura, H. A comparison of the size of vertebrae among some species of the baleen whales with specialreference to whale movements. 61-69
Omura, H., Nishiwaki, M. and Kasuya, T. Further studies on two skeletons of the black right whalein the North Pacific. 71-81
Nishiwaki, M. and Kasuya, T. Osteological note of an Antarctic sei whale. 83-89
Hosokawa, H. and Kamiya, T. Some observations on the cetacean stomachs, with special considerations
on the feeding habits of whales. 91-101

ix
Lugassy, A. A., Korostoff, E. and Rabinowits, J. L. Influence of incremental lines upon the compressive
strength of sperm whale dentin. 103-l lO
Nishiwaki, M. and Oguro, N. Baird's beaked whales caught on the coast of Japan in recent lO years.
111-122
Mizue, K., Nishiwaki, M. and Takemura, A. The underwater sound of Ganges river dolphins (Platanista
gangetica ). l 23-128
Kasuya, T. and Nishiwaki, M. First record of Mesoplodon densirostris from Formosa. 129-137
Kawamura, A. and Kashita, K. A rare double monster of dolphin, Stenella caeruleoalba. 139-140
Tsuyuki, H. and Itoh, S. Fatty acid components of Ganges river dolphin oil. 141-147

NUMBER 24, NOVEMBER 1972
Omura, H. An osteological study of the Cuvier's beaked whale, :(,iphius cavirostris, in the northwest
Pacific. 1-34
Nishiwaki, M. and Oguro, N. Catch of the Cuvier's beaked whales off Japan in recent years. 35-41
Nishiwaki, M., Kasuya, T., Kureha, K. and Oguro, N. Further comments of Mesoplodon gingkodens.
43-56
Kasuya, T. Growth and reproduction of Stenella caeruleoalba based on the age determination by meansof dentinal growth layers. 57-79
Kasuya, T. and Oguro, N. A new tagging method of dolphins. 81-85
Kasuya, T. Some informations on the growth of the Ganges dolphin with a comment on the Indus
dolphin. 87-108
Kasuya, T. and Aninul Haque, A.K.M. Some informations on the distribution and seasonal movementof the Ganges dolphin. 109-115
Tsuyuki, H. and Itoh, S. Fatty acid component of various blubber oil of Ganges River dolphin. l 17-125
Naito, Y. and Nishiwaki, M. The growth of two species of the harbour seal in the adjacent waters of
Hokkaido. 127-144.
Ichihara, T. and Yoshida, K. Diving drpth of northern fur seals in the feeding time. 145-148

NUMBER 25, SEPTEMBER 1973
Kasuya, T. Systematic consideration of recent toothed whales based on the morphology of tympanoperiotic bone. 1-103
Omura, H. A review of pelagic whaling operations in the Antarctic based on the effort and catch datain 10° squares of latitude and longitude. 105-203
Nasu, K. Results of whale sighting by Chiyoda Maru No. 5 in the Pacific sector of the Antarctic andTasman Sea in the 1966/67 season. 205-217
Kawamura, A. Food and feeding of sei whale caught in the waters south of 40°N in the North Pacific.
219-236.
Ohsumi, S. Find of marlin spear from the Antarctic minke whales. 237-239
Seki, Y. An anatomical study on the lower extension of the dorsal vagal nucleus to the upper cervical
cord in the sperm whale. 241-249
Tobayama, T., Nishiwaki, M. and Yang, H. C. Records of the Fraser's Sarawak dolphin (Lagenodelphis
hosei) in the western North Pacific. 251-263
Miyazaki, N., Kusaka, T. and Nishiwaki, M. Food of Stenella caeruleoalba. 265-275
Nishiwaki, M. Possible vestigial teats of killer whale. 277-278
Williamson, G. R. Counting and measuring baleen and ventral grooves of whales. 279-292
Tsuyuki, H. and Itoh, S. Fatty acid component of blubber oil of Amazon River dolphin. 293-299
Naito, Y. Comparison in colour pattern of two species of harbour seal in adjacent waters of Hokkaido.
301-310x

NUMBER 26, NOVEMBER 1974
Omura, H. Possible migration route of the gray whale on the coast of Japan. 1-14
Satake, Y. and Omura, H. A taxonomic study of the minke whale in the Antarctic by means of hyoidbone. 15-24
Kawamura, A. Food and feeding ecology in the southern sei whale. 25-144
McCann, C. Body scarring on cetacea-odontocetes. 145-155
Kasuya, T., Miyazaki, N. and Dawbin, W. H. Growth and reproduction of Stenella attenuata in the
Pacific coast of Japan. 157-226
Miyazaki, N., Kasuya, T. and Nishiwaki, M. Distribution and migration of two species of Stenella in
the Pacific coast of Japan. 227-243
Kamiya, T. and Pirlot, P. Brain morphogenesis in Stenella coeruleoalba. 245-253
Castello, H. P., Torno, A. P. and Panizza, J. S. First Antarctic record of a killer whale stranding. 255258
Kamiya, T. and Miyazaki, N. A malformed embryo of Stenella coeruleoalba. 259-263
Kamiya, T. and Yamasaki, F. Organ weights of Pontoporia blainvillei and Platanista gangetica (Platanistidae). 265-270
Machida, S. Surface temperature field in the Crozet and Kerguelen whaling grounds. 271-287
Machida, S. The voyage of the Konan Maru No. 16 to the Antarctic whaling grounds. 289-302
Itoh, S. and Tsuyuki, H. Fatty acid component of different blubber oil of finless porpoise. 303-306
Itoh, S. and Tsuyuki, H. Fatty acid component of Senegal manatee fats. 309-311.
Naito, Y. The hyoid bones of two kinds of harbour seals in the adjacent waters of Hokkaido. 313-320

NUMBER 27, SEPTEMBER 1975
Omura, H. Osteological study of the minke whale from the Antarctic. 1-36
Williamson, G. R. Minke whales off Brazil. 37-59
Kawamura, A. A Consideration on an available source of energy and its cost for locomotion in fin
whales with special reference to the seasonal migrations. 61-79
Kasuya, T. and Nishiwaki, M. Recent status of the population of lndus dolphin. 81-94
Kasuya, T. Past occurrence of Globicephala melaena in the western North Pacific 95-108
McCann, C. A study of the genus Berardius Duvernoy. 111-137
Nishiwaki, M. and Kureha, K. Strange organ in the anal region of the finless porpoise. 139-140
Machida, S. Distribution of sperm whale catches in the southern Indian Ocean. 141-159

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