A REVIEW OF SPERM WHALE FOOD
TAKEHIKO KAWAKAMI
Tokai Regional Fisheries Research Laboratory, Tokyo

ABSTRACTPapers on the food of sperm whales caught in various parts of the world
hitherto published were reviewed, and the outline of the informations obtained
are as follows:
The composition of the items in the sperm whale food is various according
to region, season, year, etc., depending on the composition of the food animals
inhabiting the area. The most important food items are squids, followed byfishes.
Among squids, numerically important species are histioteuthids, gonatids,
onychoteuthids and octopoteuthids. In addition to these squids, cranchiids arealso very important in some regions. Seeing their importance from the view
point of weight, the important species are octopoteuthids and histioteuthids
nearly all over the world except some regions, and onychoteuthids, architeuthids and ommastrephids are important in some regions.
Though the species of fishes in the food of sperm whales are various, they
are less important than squids. They are rather important in the northeastern part of the North Pacific, New Zealand waters and the northern part of
the North Atlantic.
The important fishes as the food of sperm whales are rock fishes, cod,sharks, rag fish, skate, dory-type fish, ling, lumpsucker, etc., though the degreeof the importance of them are different according to region.
Sperm whales sometimes attack and catch large squids, e.g. Architeuthis sp.
and Moroteuthis robusta of 2-3 m in mantle length, and also large fishes of 13 m in total length such as sharks Cetorhinus maximus, Somniosus sp. etc., Dissostichus mawsoni, etc.
The amount of the stomach contents of sperm whales has been consideredless than 300 kg, and the foods consumed in a day have been considered differently by person as 2-3 tons or 1 ton. And also there is another estimate:
300 kg and 200 kg for average-sized male and female sperm whale respectively. The total weight of squids eaten by sperm whales in a year has been
estimated to be 110-320 million tons.
INTRODUCTION

The sperm whale (Physeter macrocephalus Linnaeus) has 20-28 teeth in each side of thelower mandible, the diameters of which attain 10 cm (Nishiwaki 1965). These
teeth have the function to attract and catch cephalopods, particularly the large
squids, which are the staple food of the sperm whales. Fishes are, as Clarke, R.
(1956) said, subsidiary, but not negligible, items of food.
The foods of sperm whales in various regions have long been studied by many
Sci. Rep. Whales Res. Inst.,

No. 32, 1980, 199-218

KAWAKAMI200
TABLE I.

OCCURRENCES OF SQUIDS AND FISHES IN THE STOMACHS
Number and percent (of the stomachs

Locality

Japan
Japan

Japan
KurilesNumberstomachs
observed

Numberstomachs
with food

555
1365
1379
2744
65
360?

338
809
818
1627
65
360?

Kuriles (Okhotsk)(Pacific)
Kamchatka
Bering Sea(Komandorskiye)
67(Aleutorskiye)
29
14(Northern)
110(Total)Bering Sea and Aleutian coast-180")(( l 80"- l 60"W)
Coast of Alaskan Gulf
( l 60"- l 3G°W)
British Columbia
Central California
New Zealand
IcelandAzoresAntarctic

South Georgia andSouth Africa

Octopus

Squid
No.

%

No.

%

323*
725
788
1513
65
>250*

96
90
96
93
100
>69

10
16
3
19

3
2
0

6

11

!'3-100
4- 97
21
67
26
14
107

66 (50)21 (14)10 ( 5)
97 (69)99 (75)81 (54)71 (36)91 (64)

256
148
612
125

240
129
434
116

94
8
71
93

139

45
35
52
95

32
;;:;70
96
81

50
128
133

;;;;so

57
39

27
28?

55
100?

129

49
28
56
115

74

63

63

100

54
118

scientists. A detailed review of their results was made by Berzin in 1971. Since
then the several researches on this problem have been made, especially on squids(Clarke, M. R. 1972, 1977; Clarke, M. R. et al., 1974, 1976; Kawakami, 1976;
Okutani et al., 1976, 1978). Recently the taxonomic studies of squids by beaks
have been developed by Clarke, M. R. et al. and the studies on the food of sperm
whales have advanced with a lot of beak samples removed from the stomachs ofsperm whales caught in various parts of the world, and their results were summedup (Clarke, M. R., 1977, 1980).
I have again reviewed the published reports on the foods of sperm whales inSci. Rep. Whales Res. Inst.,

No. 32, 1980

201

SPERM WHALE FOOD, REVIEWOF SPERM WHALES TAKEN FROM DIFFERENT PARTS OF THE WORLD
Number andpercent (of thestomachs observed)
empty stomachs

with food) stomachs with
Others

Fish
No.

%

5
68
25
93

8
3
6

llO**

30

4

No.

2
2

%

No.

%

0
0

217
556
561
1117
0
O?

39
41
41
41
0
O?

25 ( I)
46 (19)64 (29)36 ( 9)
7
13
29
7

94
;;;;45
33
93

68
90?
61
79

48
4
7
6

98
14
13
5

7

11

Omura 1950
Mizue 1951

Males
Females
Total*: Only squid

Kawakami 1976
Betesheva andAkimushkin 1955** : Including
squid remainders
Tarasevich 1963

57-10013- 501917 ( I)
12 ( 5)
9 ( 4)
38 (10)
16
19
178
9

SourceRemarks

Tomilin 1957

3

10

3

3

In parentheses, foronly squids
or fishes

Berzin 1959

1960
1963
1960
1963

Okutani and
Nemoto 1964

1963
2

8
O?

14
O?

15

ll

8
ll

14
28

H

ll

Males
Both sexes
Males
Males

ll

15

Both sexes

Pike 1950
Rice 1963
Gaskin andCawthorn 1967
Roe 1969
Clarke, R. 1956
Korabelnikov
1959
Matthews 1938

various regions, and summarised them. But I could not fully refer to Dr Clarke's
latest paper (1980) as I could get it only after the completion of this manuscrpt.
RELATIVE IMPORTANCE OF SPECIES IN THE FOOD OFSPERM WHALES
The frequencies of occurrence of various food items of sperm whales are different
according to the region, ~eason, year, etc., and also it may be varied by sex, age, orbody length, but various species of cephalopods predominate almost always andSci. Rep. Whales Res. Inst.,
No. 32, 1980

202

KAWAKAMITABLE 2. RATIO OF OCCURRENCE OF SQUIDS AND FISHES IN THE STOMACHS
OF SPERM WHALES TAKEN FROM DIFFERENT PARTS OF THE WORLD

Locality
Japan
Japan

Japan
Kulile Islands
Bering Sea(Komandorskiye)(Aleutorskiye)(Northern)(Total)Bering Sea and Aleutian Coast-180")( l 80- l 60"W)

Coast of Alascan Gulf
(160-130°W)
British Columbia
Central California
New Zealand
IcelandAzoresSouth Georgia and South Africa
Notes:

Ratio of occurrencein percentage

Sex ofwhalesYearobserved

Squid

Fish

98
91
97
94
100
69

2
9
3
6
0
31

80
64
53
72

20
36
47
28

1957

94
87
71
93

6
13
29
7

1960
1963
1960
1963

32
;:;;446151
36
87;:;;
90

68
56;:;;
39
49
64

1963
1948-49
1959-62
1963-64
1967
1949
1932-35

~13

10

Males
Females
Total

1948-49
1946-48""
1772
1951-52?

Males
Both sexes
Both sexes

1) The ratios were calculated with the numbers of stomachs having squid and fish.
2) Sources are the same as those in Table l.

everywhere.
The composition of the food items in sperm whale depends on the composition of the food animals inhabiting the area where the whales take food (Berzin1971), but it does not always represent the fauna of the area. This may be thinkable from the fact that species composition of the families is different between net
samples and whales' stomach contents, and many of those taken by nets are smaller species and not young specimens of the large species sampled by whales (Clarke,
M. R., 1977).
The most important food items of sperm whales are cephalopods, followed byfishes as are shown in Tables I and 2. In addition to cephalopods and fishes,
crabs, shrimps, krill, lobsters, colonial tunicates, jelly-fishes, sponges, starfishes, sea
cucumbers, vibrissae, gorgonians, etc. were reported from the stomachs of sperm
whales (Mizue 1950, Clarke, R. 1956, Tarasevich 1963), but these are not considered to be the normal food items.
The families and species of cephalopods and fishes. reported as the food of
sperm whales by many scientists are summarised in Appendix Tables I and II.
Sci. Rep. Whales Res. Inst.,
No. 32, 1980

SPERM WHALE FOOD, REVIEW

203

Cephalopods as the food of sperm whales
As are shown in Appendix Table I, the cephalopods consist of 36 genera(about 56 species) belonging to 19 families, of which only 5 genera (about 6-7 species) belonging to 4 families are octopods and vampyromorph, and the other 31
genera (about 50 species) belonging to 15 families are decapods. Thus squids areof far greater irhpotance than octopods almost everywhere. But in the littoral
waters, the significance of octopods for sperm whales is sometimes greater, especially of those which have a pelagic mode of life, e. g. Amphitretus sp. and Alloposus mollis. They are sometimes found in 50% of the total number of stomachs examined(Berzin, 1971).
Most decapods in Appendix Table I are pelagic species except Loligo sp. whichis the littoral species and was found in the stomachs of sperm whales in the northeastern part of the Atlantic (Clarke, R., 1956). This may be a very rare example.
Octopus spp. are also littoral species and were reported from the stomachs of sperm
whales caught in the littoral waters of the North Pacific (Berzin, 1971).
It is noteworthy, as Berzin (1971) stated, that some of the species inhabiting
the surface waters in some regions, e. g. Todarodes pacificus which is the widespread
and abundant species in the western part of the North Pacific and serves as foodfor the most varied pelagic animals (including baleen whales), do not form apart of the sperm whales' diet.
The data on the compositions in numerical and weight percentages of familiesof cephalopod constituents in sperm whale food are available in the waters around
Japan, Kurile Islands, California, New Zealand, Peru and Chile, Iceland, Azores,
Vigo and Madeira as are shown in Table 3.
The numerically important species of cephalopods are as follows: In theJapanese waters, Histioteuthis dojleini is the most important and Octopoteuthis sp.,Moroteuthis robusta and Ommastrephes bartrami are also important (Omura, 1950; Mizue,
1951; Berzin, 1971; Kawakami, 1976; Okutani et al., 1976, 1978). In the Kurile
Islands waters, the species belonging to Gonatidae such as Gonatopsis borealis, Gonatus
magister and Gonatus fabricii, Galiteuthis armata and Histioteuthis sp. are important(Beteshe'va and Akimushkin, 1955; Berzin, 1959, 1971; Tarasevich, 1963). In the
Okhotsk Sea, Gonatus fabricii is important (Tarasevich, 1963; Berzin, 1971 ). In the
Bering Sea and the adjacent waters, Moroteuthis robusta, the species belonging to
Gonatidae such as Gonatopsis borealis, Gonatopsis makko, Gonatus magister and Gonatusfabricii, and Galiteuthis armata are important (Berzin, 1959, 1971; Okutani and Nemoto 1964). Off British Columbia and California, Moroteuthis robusta, Gonatopsis
borealis and Gonatus fabricii are important (Pike, 1950; Rice, 1963; Berzin, 1971 ;Fiscus and Rice, 1974; Clarke, M. R., 1979). In the Tasmanian and New Zealandwaters, Moroteuthis sp. and Onychoteuthis sp. are important (Gaskin and Cawthorn,
1967; Berzin, 1971; Clarke, M. R., 1977). Off Peru and Chile, Histioteuthis sp.,
Chiroteuthis spp., Octopoteuthis sp. and Dosidicus gigas are important (Berzin, 1971;
Clarke, M. R. et al., 1976; Clarke, M. R., 1977). In the eastern North Atlantic,
Histioteuthis bonelli is the most important and followed by Taningia danae (Clarke, R.,
1955 1956 ,· Clarke, M. R., 1962, 1977 ,· Keil 1963 · Roe 1969 · Berzin 1971 ·'Sci. Rep. Whales Res. Inst.,
No. 32, 1980

'

'

'

'

'

'

204

KAWAKAMITABLE 3.

COMPARISON IN NUMERICAL AND WEIGHT PERCENTAGE BY
Kurile Ialands

Japan

California

Family(1976)(1977)

Enoploteuthidae

0.1

Octopoteuthidae

11. l

19.9

Onychoteuthidae

11.4

3.9

1.3

4.0

(Okhotsk)(Pacific)

3.4

1.5

3.2

8.4

8.8

42.9

64.5

50.2

59.5

17.9

6.8

3.1

3.8

1.0

10.9

4.4

Cycloteuthidae
Gonatidae
Psychroteuthidae
Lepidoteuthidae

0.1Architeuthidoe

0.2

Histioteuthidae

30.4

38.2

Ommastrephidae

14.5

0.5

0.1

1.3

2.1

5.0

25.2

11.2

Chiroteuthidae
Cranchiidae

3.7

5.6

27.0

26.5

8.7

Pholidoteuthidae
Unidentified squid

1.5

39.3

Vampyroteuthidae
Octopodidae

2.0

0.9

0.5

Alloposidae

1.6

Amphitretidae

1.5

Others0.7

Total
Source**

100.0

100.0

86.2*

114. 3*

100.0

100.0

(1)

(2)

(3)

(3)

(4)

(5)

Notes 1) The values in parentheses are the percentage by weight.
2) *: These values are calculated by the present author in the same way as the original author
did.

Sci. Rep. Whales Res. Inst.,.No. 32, 1980

SPERM WHALE FOOD, REVIEW205

FAMILY OF THE CEPHALOPOD CONSTITUENTS IN SPERM WHALE FOOD
New
Zealand

(-)

(-)
70.7
(89. 7)

(-)
(-)
(-)

Iceland

Peru, Chile

5.5
(2.6)
9.2
(1.9)
0.1
(0. I)

(-)
3.2
(1.4)
0.1
(0.1)

AzoresCompletecollection

Mixedsample

0.2
(0.1)
3.4
(25. 3)
0.8
(5.9)
0.6
(0.4)
0.3
(0.1)
0.1
(0.0)

0.3
6.8

39.1

3.3

1.8

(-)

(-)

(-)

13.3
(1.0)
8.0
(5.6)

50.3
(56. 3)4.1(31. 7)
16. l
(1.0)
0.8
(0.5)

(-)
(-)
(-)

(-)

8.0
(3. 7)

7.6
(l.1)
3.2
(3. 3)

(-)

2.9
(0.3)
14.3
(83.0)

1.0
(0.3)
0.5
(14.2)

(-)

(-)

(-)

(-)

(-)

(-)

12.9
0.2

(-)

0.2
(3.4)
73.9
(46.5)
0.1
(0.0)
0.1
(0.0)
1.9
(0.6)
0.1
(0.0)
15.8
(10.1)

Madeira

0.7

1.0
(-)

Vigo

0.3
60.5

59. l

(-)

(-)

5.7
(2.3)
1.4
(6.6)
67.2
(5.6)

1.3
(4.0)
I. 7
(40.6)
88.3(36.1)

(-)

(-)

0.3

3.3

(-)

(-)

5.7
(0. I)

(-)

(-)

(-)

(-)

(-)

(-)

(-)

0.7
7.0

(-)
2.6

(-)

(-)

(-)

(-)

(-)

(-)

2.8
(7. 5)

2.9
(2.1)

0.8
(1.5)

(-)

(-)

(-)

(-)

(-)

(-)

(3.4)
100.0
(100.0)
(II)

(-)

(-)

(-)

(-)

100.0
(100.0)(6)100.0(100.0)(7)100.0(100.0)
(8)

100.0

100.0

(8)

(9)

100.0
(100.0)(10)3) **: (!) Okukani et al. 1976 (2) Okutani and Satake 1978 (3) Tarasevich 1963 (4) Betesheva
and Akimushkin 1955 (5) Rice 1963 (6) Gaskin and Cawthorn 1967 (7) Clarke, M. R. et al.
1976 (8) Clarke, M. R. and MacLeod 1976 (9) Clarke, R. 1956 ( 10) Clarke, M. R. and
MacLeod 1974 (11) Clarke M. R. 1962, amended by Clarke, M. R. 1974.

Sci. Rep. Whales Res. Inst.,
No. 32, 1980

206

KAWAKAMI

Clarke, M. R. and MacLeod, 1974). In the Indian Ocean, histioteuthids are themost important (Matthews, 1938; Berzin, 1971; Clarke, M. R., 1972, 1977). In
the Antarctic, onychoteuthids and cranchiids are very important (Matthews, 1938;
Korabelnikov, 1959; Berzin, 1971; Clarke, M. R., 1977).
The data on the amount of squids as the food of sperm whales are rather
numerous (Clarke, M. R., 1962; Clarke, M. R. and MacLeod, 1974, 1976; Clarke,
M. R. et al., 1976), but most of them have not been treated quantitatively (Table 3).
According to Clarke, M. R. (1977) who reported the quantitatively important squid families estimated by the beaks removed from the stomachs of sperm
whales, octopoteuthids, including the large Taningia danae, are important in all
regions except the Antarctic and off Peru and Chile, and histioteuthids, including
several species, are important everywhere expect the Antarctic. In addition tothem, onychoteuthids, architeuthids, ommastrephids such as Dosidicus gigas, Todarodes sp., Ancistrocheirus of enoploteuthid, and pholidoteuthids are also important insome regions.

Fishes as the food of sperm whales
Though the species of fishes obtained from the stomachs of sperm whales arevarious as are shown in Appendix Table II, namely 55 genera (about 68 or more
species) belonging to 49 families, the occurrence prortion of fishes in the food is not
so large as squids, and is different by region and year like cephalopods as are shownin Table 1. The ratios of occurrence of squids to fishes in the food of sperm whales
by region are shown in Table 2. In the North Pacific the fish occurrence is between 1%and68% of the stomachs having fishes and/or squids in them (Omura, 1950;
Pike, 1950; Mizue, 1951 ; Betesheva and Akimushkin, 1955; Tomilin, 195 7; Tarasevich, 1963; Rice, 1963; Okutani and Nemoto, 1964). It is only 1-9% in the Japanese waters, 31 % in the Kurile Islands waters, 6-47% in the western part of theBering Sea and Aleutian coast west of 180°, 7-29% in the Bering Sea and Aleutian
coast between 180° and 160°W, 68% off the coast of Alaskan Gulf, not less than 56%
off British Columbia and 39% off California. In the Bering Sea and Aleutian coastwest of 180° it is, as is shown in Table 2, 6-13% according to Okutani and Nemoto(1963), but it is 20%, 36% and 47% in the waters of Komandorskiye, Aleutorskiye
and northern part of the Bering Sea respectively according to Berzin (1959). Thus
fishes as the food of the sperm whales mostly predominate in the eastern waters along the Alsakan Gulf and the east Bering Sea as was stated by Okutani and Nemoto(1964), and in the northern part of the both sides of the North Pacific.
In the New Zealand waters the sperm whales take squids and fishes in nearly
the same ratio, but about one third of them feed both fishes and squids, and theratio of fishes to squids by weight was 1: 1.69 (Gaskin and Cawthorn, 1967).
In the Iceland waters of the Atlantic, fishes occur in 98% of the sperm whale
stomachs with food, in which 54% contain both squids and fishes (Roe, 1969), andthe occurrence ratio of squids to fishes is 36: 64. In the Azores waters fishes occurin 13% or less (Clarke, R. 1956). For the sperm whales in the North Atlantic,
fishes are more important in the northern part of the Ocean.
Sci. Rep. Whales Res. Inst.,
No. 32, 1980

SPERM WHALE FOOD, REVIEWTABLE 4.

FishesLamprey
Shark
Skate
Rag fishSardineSalmon-like fish
Lanternfish
Conger eel
Saury
Dory-types
South kingfish
GroperAnarrhichus minor
Ling
Rock fish
Sable fish
Lingcod
Lumpsucker
Cod
Angler fish
Total
Source207

COMPARISON IN NUMERICAL PERCENTAGE OF THE
FISHES IN SPERM WHALE FOOD
Japan

Japan

BritishColumbia

California

New
Zealand

Iceland

25.0
54.2

6.9

6.3
3.1

4.2

l. 7
13.8

2.2
3.2
20.0
35.6
4.3
6.7

6.5
29.3
12 .1
15.5
3.1
20. 7
80.0

55.9

25.0

35.6
8.3
8.3

20.0

30.1

100.0
Omura
1950

100.0
Mizue
1951

100.0
Pike
1950

100.0
Rice
1963

100.0
GaskinandCawthorn
1967

31.3
18.8
12.5
100.0
Roe
1969

Note: The percentages were calculated with the numbers of stomachs in which the fish were found by
the present author.

In the waters of South Georgia and South Africa, the fishes occur in 11 % of
the sperm whales' stomachs with food (Matthews, 1938), and in the Antarctic, in
5-13% (Clarke, R., 1956; Korabelnikov, 1959) (Table 1).
Thus, generally speaking, the occurrence proportion of fishes is not so large
as squids, and in many cases fishes are found mixed with squids. These conditions
are various according to the region, and the regions where fishes are rather important are the northern and the northeastern parts of the North Pacific, the New Zealand waters and the northern part of the North Atlantic.
The data on the composition in numerical percentage of fish constituents insperm whale food are available in the waters of Japan, British Columbia, California, New Zealand and Iceland as are shown in Table 4. The important fishes ineach region are rock fish and cod for the Japanese waters, rock fish and rag fish for
the British Columbian waters, skate and shark for Californian waters, dory-type fish
and ling for the New Zealand waters, and lumpsucker and rock fish for the Icelandwaters.

Sci. Rep. Whales Res. Inst.,
No. 32, 1980

KAWAKAMI

208

THE FEEDING TIME OF SPERM WHALES

It may be thinkable that sperm whales actively feed during night when squids comeup to the surface, and do not move about so actively in the daytime, judging fromthe catch data of the Antarctic whaling (Matsushita 1955 ), but Okutani and N emoto(1964) and Clarke, M. R. (1980) found no evidence for special feeding time in the
North Pacific and off Durban respectively.
THE SIZE OF FOOD ANIMALS EATEN BY SPERM WHALESSquidsThough the size of cephalopods usually eaten by sperm whales does not appear so large, sperm whales are considered to attack sometimes the gigantic squids.
This is considerable from the large scars up to 20 cm in diameter on the skin of
sperm whales caused by large suckers of squids, and also from the reports of huge
fragments of squids vomitted by dying whales or recovered among the stomach
contents, although Roe (1969), who recorded the sucker scars of 1.7-11.5 cm in diameter, doubted whether the very large scars such as 11.5 cm in diameter had been,
in fact, caused by squids, and considered it was possible that they were old sucker
scars which had stretched as the whale had grown (Matthews, 1938; Roe, 1969;
Berzin, 1971).
TABLE 5. SIZES OF SQUIDS FROM STOMACHS OF SPERM WHALESEXAMINED AT HORTA IN 1949 (Quoted from Clarke, R., 1956)
Sizes of squids
Length group
Small
MediumMedium-large
Large
Total
Average size of squids

Standard
length
0.6 to 0.9 m
0.9 to 1.5
1.5 to 1.8
1.8 to 2.4

Female whales

Male whales

No. of suids

No. of squids

%

41
26
2
1
70
0.95m

59
37
3

28
13
0

100

42
0.92m

%
67
31
0
2
100

Kondakov (1940) considered that the suckers as large as 20 cm in diameter
might belong to Architeuthis of 18 m in total length. The large Architeuthis whose total
length and weight were 10.5 m (mantle length was 2 m), 184 kg, and 12 m, 200 kg
were actually removed from the stomachs of sperm whales from the Azores in 1955
and from the Antarctic in 1964 respectively (Clarke, M. R., 1955; Berzin, 1971).
An architeuthid 2.4 m in mantle length was obtained from the stomach of the
sperm whale in the Azores waters (Keil 1963). In addition to these architeuthids,
Moroteuthis robusta of 2.9 m, 2.4 m and 2.4 m long were reported from the stomachsof the sperm whales caught off Kommandorskiye Islands, off British Columbia andin the Antarctic respectively (Pike, 1950; Clarke, R., 1956; Berzin, 1971 ).
As the large scars and scratches are usually found on the head of male whales,
and females are smaller in body length than males, it is often considered that male
Sci. Rep. Whales Res. Inst.,
No. 32, 1980

SPERM WHALE FOOD, REVIEW

209

sperm whales take larger squids than females (Matthews, 1938). Clarke, M. R.
(1980) also considered that males favour larger species than females by a x2 test of
the data of the sperm whales off Durban. But there is also another opinion thatmales and females take food of the same size as shown in Table 5 (Clarke, R. 1956).
The size of the food items in the diet may be different according to the ground,and an example of the difference was shown by Clarke, R. (1956) as follows: Theaverage standard length of squids from the sperm whales caught in the Azores was
0.94 m and that of the Antarctic (Moroteuthis ingens) was 1.3 m.

FishesFishes are, as stated above, generally not so important food for sperm whales
as squids, but sometimes large fishes 1-3 m in body length have been removed fromthe stomachs of sperm whales as follows: a 3 meter shark, the species of which was
unidentified, was reported off South Africa (Chabb, 1918; fido Berzin 1971), abasking shark Cetorhinus maximus 2.5 m long, at the Azores (Clarke, R., 1956), agreen shark Somniosus sp. 2.1 m long, another shark Squalus acanthias 1.3 m long,and Alepisaurus aesculopius 1.3 m long, in the Kurile waters (Betesheva, 1961 ; fido
Berzin 1971), fishes belonging to Notothenidae up to 1.5 m long and a ray Rajagriseocandata I.Im long, near Tierra del Fuego (Semskii, 1962, fido Berzin 1971),
Dissostichus mawsoni up to 1.7 m long (as a rule 1.2-1.4 m), near the Balleny Island(Berzin, 1971 ), fan tailed ragfish lcosteus aenigmaticus 1.3 m long, off British Columbia(Pike, 1950).
THE WEIGHT OF DOOD OF SPERM WHALES

The weight offood in a stomach of sperm whales
The weight of food in a stomach of sperm whales may be different according
to the size of whales, and is generally considered to be less than 300 kg. According
to Betesheva and Akimushkin (1955) and Betesheva (1961), it did not exceed 200
kg. Other data showed that stomachs contained 201-500 squid (Gonatus fabricii)(Berzin, 1971 ), the weight of which may be calculated as 80-200 kg, supposing the
average weight of this squid is 400 g. The weight of fresh food of the sperm whalesin the Cook Strait region of New Zealand was reported to have varied from 12. 7 to
105 kg (Gaskin and Cawthorn, 1967).
On the difference of the quantity of food for male and female sperm whales,
there are two different opinions. The quantity of food of sperm whales in theJapanese waters was not different between males and females (Mizue, 1950), but
the uncomplete data from the Azores suggested (not concluded) that females werefeeding rather less heavily than males, and this might be explained as follows:
females are hampered to feed heavily by calves which cannot dive deep (Clarke,R., 1956).

The amount ef sperm whale food consumed in a day and in a year
The food of sperm whales consumed in a day was considered to be at least 2Sci. Rep. Whales Res. Inst.,
No. 32, 1980

210

KAWAKAMI

3 tons of squids for a sperm whale 13-14 m long by Sleptsov (1952), and one ton
a day by Berzin (1971).
Clarke, M. R. (1977) considered the amount of food consumed in a day by
sperm whales would be 2-4% of their body weight and calculated as 300 kg
and 200 kg for males and females respectively, supposing the average body weights
were 15 tons and 5 tons for males and females respectively. As he stated that theweight of sperm whales is between 8 and 50 tons, if the largest whale is considered,
the amount of food consumed in a day will be one ton.
The total weight of cephalopods eaten by sperm whales each year has been
estimated by Clarke, M. R. (1977) to be over 110 million tons on the basis of thedata on estimated number of whales and estimated weight of food consumed bywhales, and also as a muximum figure the amount of cephalopods eaten was estimated to be 320 million tons, accepting a value of 15 tons for the average whale
and a consumption of 3.5% of body weight per day as well as an average calorificvalue of 75% of that for mascular squids.
NUTRIENT DISPERSAL BY SPERM WHALESThis problem has scarecely been studied. Sperm whales seem to eat food actively
near the surface during night as stated above, but it may be true that they certainly eat several midwater species and also eat species in spawning condition on thebottom. Therefore, it could be considerable that sperm whales probably carry nutrient upwards by eating deepsea squid and defaecating near the surface as Clarke,
M. R. (1977) stated.
Many families of cephalopods are said to have large amount of ammonia intheir tissues and coelom, and 53-78% of the number of cephalopods consumed bysperm whales are said to be ammonical squid (Clarke, M. R., 1977), which areprobably not good for human food as they are.
ACKNOWLEDGMENTS

My sincere thanks are due to Dr H. Omura, Director of the Whales ResearchInstitute, who kindly enocouraged me to make this review. I would also like toexpress my very warm thanks to Mrs Y. Takahashi of the Whales Research Institute and Mrs M. Hara of Ocean Research Institute of Tokyo University who kindly helped me in borrowing papers reffered.
REFERENCES
Aim, T., 1978. Keys to the Japanese fishes fully illustrated in colors, Hokuryukan, Tokyo, pp. 358. (in Japanese)
BERZIN, A. A., 1959. On food of sperm whale in the Bering Sea. /zvestia TINRO, 47: 161-165. (in Russian)
BERZlN, A. A., 1971. The sperm whale (Translated into English in 1972). Israel Program for Scientific
Translation, pp. 394.
BETESHEVA, E. I. and I. I. AKIMUSHKIN, 1955. Food of the sperm whale (Physeter catodon L.) in the Kurile
Sci. Rep. Whales Res. Inst.,
No. 32, 1980

SPERM WHALE FOOD, REVIEW

211

Islands region. Truqy Inst. Okeanol. 18: 86-94. (in Russian)
CLARKE, M. R., 1962. Stomach contents of a sperm whale caught off Madeira in 1959. Norsk HvalfangstTid., 51(5):173-191.
CLARKE, M. R., 1966. A review of the systematics and ecology of ocean squids. Adv. mar. Biol., 4: 99-300.
CLARKE, M. R., 1972. New technique for the study of sperm whale migration. Nature, 238: 405-406.
CLARKE, M. R., 1977. Beaks, nets and numbers. Sym. Zoo!. Soc. London, 38: 89-126.
CLARKE, M. R., 1980. Cephalopoda in the diet of sperm whales of the southern hemisphere and their
bearing on spearm whale biology. Discovery Rep. 37. pp. 324.
CLARKE, M. R. and N. MACLEOD, 1974. Cephalopod remains from a sperm whale caught off Vigo, Spain.
J. mal. biol. Ass. U.K., 54: 959-968.
CLARKE, M. R. and N. MACLEOD, 1976. Cephalopod remains from sperm whales caught off Iceland. J.
mar. biol. Ass. U.K., 56: 733-749.
CLARKE, M. R., N. MACLEOD and 0. PALIZA, 1976. Cephalopod remains from the stomachs of sperm
whales caught off Peru and Chile. :?,ool., Lond, 180: 477-493.
CLARKE, R., 1955. A giant squid swallowed by a sperm whale. Norsk Hvalfangst-Tid., 44 (10): 589-593.
CLARKE, R., 1956. Sperm whales of the Azores. Discovery Rep., 28: 237-298.
FAR SEAS FISHERIES RESEARCH LABORATORY, 1972. Colored illustrations of bottom.fishes collected by Japanese trawlers. Japan Deep Sea Trawlers Association, Tokyo, pp. 145. (in Japanese)
FAR SEAS FISHERIES RESEARCH LABORATORY, 1976. Colored illustrations ef bottomjishes collected by Japanese trawlers. II. Japan Deep Sea Trawlers Association, Tokyo, pp. 188. (in Japanese)
Fiscus, C. H. and D. W. RICE, 1974. Giant squids, Architeuthis sp., from stomachs of sperm whales captured
off California. Calif. Fish and Game, 60 (2): 91-93.
GAMBELL, R., 1972. Sperm whales off Durban. Discovery Rep. 35, 199-358.
GASKIN, D. E. and M. W. CAWTHORN, 1967. Diet and feeding habits of the sperm whale (Pkyseter catodon L.)
in the Cook Strait region of New Zealand. N.:?,. J. mar. Freshwat. Res. 1 (2): 159-179.
lsHIKAWA, C. and Y. WAKIYA, 1914. Note on a gigantic squid obtained from the stomach ofa sperm whale.
J. Coll. Agric. Imp. Univ. Tokyo, 4 (7): 435-443.
lwAI, E., 1956a. Descriptions on unidentified species of dibranchiate cephalopods. I. An oegopsiden squid
belonging to Genus Architeuthis. Sci. Rep. Whales Res. Inst., 11: 140-146.
IWAI, E., 1956b. Descriptions on unidentified species of dibranchiate cephalopods. II. A Cranchiidae
squid of the Genus Taonius. Sci. Rep. Whales Res. Inst., 11: 153-158.
KAWAKAMI, T., 1976. Squids found in the stomach of sperm whales in the northwestern Pacific. Sci. Rep.
Whales Res. Inst., 28: 145-151.
KEIL, A., 1963. Riesentintenfische aus dem Pottwal-Magen. Natur und Museum, 93 (8): 319-323.
KoRABELNIKOV, L. V., 1959. Diet of sperm whale in the Antarctic. Priroda, 3.59: 103-104. (in Russian)
McCANN, C., 1974. Body scarring on Cetacea-Odontosetes. Sci. Rep. Whales Res. Inst., 26: 145-155.
MATSUBARA, K., 1979. Fish, morphology and hierarchy, Ishizaki-Shoten, Tokyo, pp. 1605.
MATSUSHITA, T., 1955. Daily rhythmic activity of sperm whales in the Antarctic Ocean. Bull. Japanese
Soc. Sci. Fish., 20 (9): 770-773. (in Japanese with English abstract)
MATTHEWS, L. H., 1938. The sperm whales, Physeter catodon. Discovery Rep., 17: 93-168. 9 pls.
MrzuE, K., 1950. Food of whales in the adjacent waters of Japan. Sci. Rep. Whales Res. Inst., 5: 81-90.
NrsmwAKI, M., 1965. Whales and Pinnipeds, University of Tokyo Press, Tokyo, pp. 82.
0KUTANI, T., 1973. Guide and keys to squid in Japan. Bull. Tokai Reg. Fish. Res. Lab. 74: 83-111. (inJapanese with English abstract)
0KUTANI, T., 1975. Supplements and amendments for" Guide and keys to squid in Japan". Bull. TakaiReg. Fish. Res. Lab., 83: 41-44. (in Japanese with English abstract)
0KUTANI, T. and T. NEMOTO, 1964. Squids as the food of sperm whale in the Bering Sea and Alaskan Gulf.
Sci. Rep. Whales Res. Inst. 18: 111-121.
0KUTANI, T., Y. SATAKE, S. 0HSUMI and T. KAWAKAMI, 1976. Squids eaten by sperm whales caught off
Jo ban district, Japan, during January-February, 1976. Bull. Tokai Reg. Fish. Res. Lab., 87: 67-113.
0KUTANI, T. and Y. SATAKE, 1978. Squids in the diet of 38 sperm whales caught in the Pacific off northern
Honshu, Japan, February 1977. Bull. Tokai Reg. Fish. Res. Lab., 93: 13-27.
Sci. Pep. Whales Res. Inst.,
No. 32, 1980

212

KAWAKAMIOMURA, H., 1950. Whales in the adjacent waters of Japan. Sci. Rep. Whales Res. Inst., 4: 27-113.
PrKE, G. C., 1950. Stomach contents of whales caught off the coast of British Columbia. Prog. Rep. Pacif.
Cst. Stns, 83: 27-28.
RrcE, D. W., 1963. Progress report on biological studies of the larger cetacea in the waters off California.
Norsk Hvalfangst-Tid., 52 (7): 181-187.
RoE, H.S.J ., 1969. The food and feeding habits of sperm whales (Physeter catodon L.) taken off the west coastof Iceland. j. Cons. int. Explor. Mer., 33 (!): 93-102.
TARASEVICH, M. N., 1963. Data on feeding of sperm whales in the northern area ofKurile waters. Trudy
Inst. Okeanol Akad. nauk SSSR. 71: 195-206. (in Russian with English summary).

Sci. Rep. Whales Res. Inst.
No. 32, 1980

SPERM WHALES FOOD, REVIEW213

APPENDIX TABLE I. THE CEPHALOPODS FOUND IN THE
STOMACHS OF SPERM WHALES BY AREA*
FamilyGenus Species>

Loliginidae
Loligo forbesi
Enoploteuthidae
Ancistrocheirus lesueuri +Octopoteuthidae
Octopoteuthis longiptera +0. spp.
+Taningia danae+~Cucioteuthis unguiculata Onychoteuthidae+
Onychoteuthis banksii+
0. spp.
+Moroteuthis robusta+
M. liinbergii+
lvl. ingens
lvl. robsoni
M. knipovitchi
M. sp.
Tetronychoteuthis
dussumierii
T. sp.
Kondakovia longimana
K. sp.
Cycloteuthidae
Cycloteuthis akimushkini C. sp.
Gonatidae+
Gonatus fabricii+
G. berryi+
G. antarcticus
G. sp.
+
Ber~yteuthis magister+~ Gonatus magister+
Gonatopsis borealis+
G. makko+
G. octopedatus+
Mesonychoteuthis
hamiltoni
Psychoteuthidae
Psychoteuthis sp.
Lepidoteuthidae
Lepidoteuthis grimaldii +L. sp.
Architeuthidae
Architeuthis japonica
+

N2

>

NE 3

IndianAtlantic

Pacific
NW 1> sw•>

SE 5

>

NE 3

>

> SE5l

so>

+

+

SW 4

Antarctic
Pac

Atl

Ind Ant7l

Off S.
Afr.Bl

+

+
+

+
+

+
+

+
+

+

+

+

+
+
+

+
+

+

+

+

+

+

+

+
+
+

+
+

+

+
+

+
+

+

+

+

+

+

+

+

+

+

+

+
+

+

+
+
+

+
+

+

+

+
+
+

+
+

+
+

+

+
+
+

+
+
+
+

+
+
+
+
+
+
+

+
+
+
+

+
+

+

+

+
Continued .•.

Sci. Rep. Whales Res. Inst.,
No. 32, 1980

214

KAWAKAMIAPPENDIX TABLE I.
FamilyGenus Species

Continued.

Atlantic

Pacific

IndianNWll N 2 > NE 3 > sw•> SE 5> NE 3 >sw•> SE5)

A. physeteris=Dubioteuthis physeteris A. sp.
Histioteuthidae+
Histioteuthis dojleini+=Stigmateuthis dojleini
H. sp.
=S. sp.
=Calliteuthis sp.
H. meleagroteuthi s=Meleagroteuthis+
separata
H. bonelliana
H. bonellii
H. cookiana
Ommastrephidae
Todarodes sagittatus
T. sp.
Nototodarus sloani
Ommastrephes bartrami +=Stenoteuthis bartrami +Dosidicus gigas=Ommastrephes gigas
Symplectoteuthis sp.
Chiroteuthidae
Chiroteuthis ca{yx+C. veranyi+
C. spp.
Mastigoteuthidae
Mastigoteuthis sp.
Cranchiidae
Cranchia scabra+
Taonius pavo+
T. megalops
T. spp.
+Galiteuthis armata+
G. sp.
+
Phasmatopsis li)!moctypus P. sp.
Mesonychoteuthis
hamiltonyPholidoteuthidae
Pholidoteuthis sp.
P. ? boschmai
Vampyroteuthidae
Vampyroteuthis
injernalis

+

+
+

+

+
+

+

+
+
+
+
+

+

+

S6>

+
+

Antarctic
Pac

Atl

+

Ind Ant7 >Off S.
Afr. 8 >

+

+
+

+

+

+

+
+

+

+
+

+
+

+
+

+

+

+
+

+
+
+

+

+

+
+
+
+
+
+

+
+

+
+

+

+

+

+

+

+
+

+

+

+
+

+

+

+

+

+
+
+
+
+

+
+

+
+
+

+

+
+

+
+
Continued .••Sci. Rep. Whales Res. Inst.,
No. 32, 1980

SPERM WHALE FOOD, REVIEWAPPENDIX TABLE I.

Continued.

Atlantic

Pacific

FamilyGenus Species

Indian _ _ _Antarctic_ _ _ _ _ Off S.

NW 1> N 2> NE 3 > SW 4> SE 5> NE 3> SW 4> SE5>

Vampyroteuthis sp.
Octopodidae
Octopus vulgaris
0. gilbertianus
0. sp.
Paroctopus gilbertianus
Alloposidae
Alloposus mollis
A. sp.
Amphitretidae
Amphitretus sp.

+
+
+
+
+

+

+

+
+

215

S 6>

Pac

At!

Ind Antn Afr.B·>

+

+

+
+
+

+
+
+

+

+

Notes:* The list has been drawn up on the basis of data from the works listed in the chapter " REFERENCE." The bar in the table only shows that the shecimen was not reported by this name.
1) Northwestern part of the Ocean.
5) Southeastern part of the Ocean.
2) Northern part of the Ocean.
6) Southern part of the Ocean.
3) Northeastern part of the Ocean.
7) The part of the Ocean was not reported.
4) Southwestern part of the Ocean.
8) Off South Africa.

APPENDIX TABLE II. THE FISHES FOUND IN THE STOMACHS
OF SPERM WHALES BY AREA*
At!. Ind.

Pacific
English nameLamprey
Pacific lamprey
Brown cat shark
Basking shark
Blue shark
Shark
Spined dog fish

Green shark
Greenland shark
Black shark
Shark

FamilyGenus Species
Petromyzonidae
Entophenus tridentatus
Scyliorhinidae
Apristurus brunneus
Cetorhinidae
Cetorhinus maximus
C. sp.
Carchariidae
Carcharias glaucus
Squalidae
Squalus acanthias=Koinga kirki
Dalatiidae
Somniosus sp.
S. microcephalus
Dalatias licha
D. sp.
Echinorhinidae
Echinorhinus cooki
E. sp.

z ~ z ~rn
+ +
+

~

~

~

rn

'p..°u"' ~zAntarctic
u

U:i

Ol

p..

~

"O
i::

...
i::

<i:

+
+

+
+ +

+
+
+

+

+

+
+

+
+

+
+
Continued ...

Sci. Rep. Whales Res. Inst.,
No. 32, 1980

KAWAKAMI

216

Continued.

APPENDIX TABLE II.

English nameShark, Pacificeangel shark§: " &iz z zRound-herring

SardineSalmonSalmon like fish
Lantern fish

Large eel
Conger eel
Eel
Saury
Dory-like fish
John Dory
Porcupine fish
Mackrel3:rn ~rn 0..."" ~z~Rajidae

Myctophidae
Myctophum humboldti
Tarletonbeania crenularis
Alepisauridae=Plagiodontidae
Alepisaurus aesculapius=Plagyodus aesculapius
A. spp.
Anguillidae
Anguilla sp.
Congridae
Scombresocidae
Cololabis saira
Trachichthidae
Hoplostethus sp.
ZeidaeZeus Jaber
Diodontidae
Diodon sp.
Scombridae
Scomber japonicus
S. tapeinocephalus=S. japonicus
tapeinocephalus

c;,

""

0...

~

"O

~

c

<t;

+
+

Raja smirnovi
R. rhina
R. griseocaudat,aR. sp.
Chimaeridae
Dussumieridae
Etrumeus micropus
Clupeidae
Sardinops melanosticta
Engraulidae
Engraulis japonicus
Salmonidae
Onchorhynchus gorbusha

(.)

(.)

Squatinidae+

Squatina californica
Shark
Ray, Long nose
skate

Antarctic

At!. Ind.

Pacific
FamilyGenus Species

+

+ +

+

+

+ +

+
+

+

+

+
+
+
+

+
+
+ +
+

+
+

+

+
+
+

+

+ +
+
+
+
+
+
+
Continued ...

Sci. Rep. Whales Res. Inst.,
No. 32, 1980

217

SPERM WHALE FOOD, REVIEWAPPENDIX TABLE II.

Continued.
Atl. Ind.

Pacific
English name

FamilyGenus Species

~

z

Tuna
Yellow fin tuna
BarracudaSouthern king fish

GroperLing
Rock fish
Red fishRock fish

Sable fish, Blackcod

ThunidaeThunus albacares
Sphyraenidae
Sphyraena sp.
Acinaceidae
Rexea solandri=Jordanidia solandriHistiopteridae
Pseudopentaceros richardsoni
Serranidae
Po[vprion oxygeneios
Anarrhichadidae
Anarrhichas minor
Ophidiidae
Genypterus blacodes
Scorpaenidae
Sebastes jlammeus
S. iracundus
s. alutus
S. mentella
S. marinus
S. ruberrimus
S. rober=Sebastodes rober
S. sp.
Helicolenus papilosus
Anoplopomidae
Anoplopoma fimbriaz z 3:
~

rJJ

~

rJJ

"'utllii..

~

Antarctic
u

z

U:i

tll

ii..

~

-0

>::
......

i::

~

+
+

+

+
+
+
+
+
+ + + +
+
+
+
+

+
+ +

Hexagramrnidae
Pleurogrammus monopterigius P. sp.
+
Ophiodon elongatus
Greeling, Ling cod
CottidaeLarge gobies+
Myoxocephalus jaok+
M. verrucosus+
M. sp
Agonidae
Percis japonica
Cyclopteridae
Cyclopterus lumpus
Lurnpsucker
Aptocyclus ventricosus+
MoridaePodonema longipes+
GadidaeCod (fish)+
Gadus macrocephalus
+

+
+

+
+

+
+

+

+
+
+
+
+

+

+
+

Continued ...

Sci. Rep. Whales Res. Inst.,
No. 32, 1980

KAWAKAMI218

APPENDIX TABLE II.

Continued.
Pacific

English name

FamilyGenus Species;::z=G. morhua macrocephalus +G. morhua
G. aeglefinus=Melanogrammus
aeglefinus
Pollachius t•irens
Theragra chalcogramma
Walleye pollack+
Eleginus gracilis+
E. gracilis navaga+
Southern poutassou Gadidae
Micromesistius australis
Notothenidae
Dissostichus eleginoides
D. mawsoni
Coryphaenoididae
Macrurid=Macruridae
Coryphaenoides pectoralis+=Laemonema longipes
C. sp.
=Macrurns sp.
Hemimacrurus acrolepis+
Icosteidae
Icosteus aenigmaticus
Fan-tailed rag-fish
Acrotus willoughbyi
Brown ragfish
Angler fish
Lophiidae
Lophius piscatorius
Large (bathy
Ceratiidae
pelagic)
Ceratias holboelli
angler fish
Himantolophidae
Himantolophus groenlandicus
Oneirodidae
Angler fish
Oneirodes sp.
+
Pacific cod

At!. Ind.

Antarctic

~

z z
+

+
+
+
+

+

+

+

+

+
+
+
+
+

+
+

+

+ +

+

Notes:* The list has been drawn up on the basis of data from the works listed in the chapter " REFERENCE." The bar in the table only shows that the specimen was not reported by this name.
1) Northeastern part of the Ocean.
5) Southeastern part of the Ocean.
2) Northern part of the Ocean.
6) The part of the Ocean was not reported.
3) Northeastern part of the Ocean.
7) Southern part of the Ocean.
4) Southwestern part of the Ocean.

Sci. Rep. Whales Res. Inst.,
No. 32, 1980