SQUIDS AS THE FOOD OF SPERM WHALESIN THE BERING SEA AND ALASKAN GULF
TAKASHI OKUTANI* AND TAKAHISA NEMOTO

It is well known that sperm whales, Physeter catodon (Linnaeus), feed on squids inthe world seas. There are many reports on the feeding of sperm whale and squids
found in the sotmach of that whale (Mathews, 1938: Mizue, 1950: Clarke,
1955: Betesheva & Akimushikin, 1955: Clarke, 1962). Betesheva & Akimushkin(1955) described the food species of sperm whales in the waters of Kurile Island,and Pike (1950) and Rice (1963) reported the food animals of whales in the west
coast of America.
The pelagic whaling by Japan and U.S.S.R. have been operating in the Aleutian
waters and Bering Sea, and sperm whale is one of the important species of thecatch therefrom. However, the foods of the sperm whales caught in the northern
North Pacific have seldom been examined, although a report on the stones and otheraliens found in the stomachs of the whales was published by Nemoto and Nasu
( 1962).
The stomach contents of the sperm whales from the waters around AleutianIslands, Bering Sea and Alaskan Gulf were collected in 1963. These specimens
were examined taxonomically, and their occurrences were studies in viewpoint of
distributions of the squids and the predators in the north Pacific. Here this reportdeals with the preliminary results of the investigation, and this study will be continued to contribute the knowledge on distribution and taxonomy of squids in the
North Pacific.
MATERIALS
The present samples were collected through the survey carried out in the North
Pacific on board the factory ship. This investigations are mainly carried out in thewaters around Aleutian Is., Alaskan Gulf and the Bering Sea. The field observations
are made on all sperm whales caught, and the contents of stomachs are descriminatedas squids, fish and other items. The observation was made on the first stomach
content which is in fairly better condition than that elsewhere. However, sometimes second stomach was erroneously dissected for examination. The content
found in the first stomach in worse condition might be the second stomach content
surged back into the first. The squids have been preserved in 10 % formalin sea
water, and they are tentatively classified into types A,B,C,D,E and G. A part ofsamples taken out from the whale stomach was brought home for close examination.

* Tokai Regional Fisheries Research Laboratory, Tokyo.
Ill

~: !
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I
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I

112

T. OKUTANI AND T. NEMOTOSYNOPSIS OF THE SPECIES IDENTIFIED
As was mentioned in the preceding section, at the field observation six forms
discriminated were classified into A, B, C, D, E and G. On the basis of the samples
brought home, each group was found to be composed of the species as follows :A: Moroteuthis robustus (Verrill, 1876)
B & D: Gonatopsis borealis Sasaki, 1923, makko, n. subsp.
C: Mastigoteuthis (?) sp.
Stigmateuthis sp.
Galiteuthis armata Joubin, 1898
Taonius pavo (Lesueur, 1821)
E: Gonatus (Berryteuthis) magister Berry, 1913
G: Gonatus fabricii (Lichtenstein, 1818)
Only C-Group was found containing four species. This is because these
species have gelationus integuments which are easily matilated badly when takenby the predators. Sometime fragments of transparent pens of some unidenified
Cranchiids were remained undigested.
Synoptic notes on the results of examination on the specimens are present in
the following lines.
OEGOPSIDA

Onychoteuthidae

Moroteuthis robustus Verrill, 1876
Because of huge size, reddish variegation and rippled skin, this species is easily
discriminated at the field observation on board the factory ship. One of thespecimens brought home was measured about 700 mm in dorsal mantle length.
Both tentacles were removed, but other characteristics are quite agreeable to thedescription by Sasaki (1929). Another specimen of head-foot part (mantle andvisceral parts lacking) is much larger than the other, and the tentacular club iswell preserved as is illustrated in Fig. 1 a. The specimens examined for the present study are generally gigantic in size, but a unique example of 300 mm-individual was found in the course of field observation. (seep. 118.)
Histioteu thidae

Stigmateuthis species
A badly matilated specimen was examined. The mantle length measured 100 mm
and the longest arm, 300 mm. A part of the integument with some photophores
is preserved. But identification to the species cannot be made because of lack oftentacle.
Gonatidae

Gonatus (s.s.) fabricii (Lichtenstein, 1818)
A single specimen brought home presented a very large size: Dorsal mantle
length, 300 mm; width of the mantle, 56 mm; length of the fin, 170 mm; widthof the fin taken together, 150 mm.

FOOD OF SPERM WHALES

113

Mantle is robust, purplish in color in preserved state, elongated cylindricaland length is about five times diameter. The fins are muscular and rhomboidal
in outline. They seem to present more elongated shape than those of younger
specimens taken from the North Pacific on other occasions. Arms have two seriesof hooks on inner rows and two series of small suckers on outer rows. Armformula is 3, 1, 2, 4.
The distal parts of both tentacles which may be the most important key ofidentification, were already lost when the specimen was taken out from the whale
stomach. However, rows of small fixing pads disposed on both margins of suckerbearing face of the tentacualr carpus were observed. These rows of pads in
G. fabricii reached far proximally than those in the next species. From suchcharacter and shapes of locking cartilages led the present authors to the conclusive
identification of the present specimen.

Gonatus (Berryteuthis) magister Berry, 1913
Specimens examined are mostly adults measured about 200-250 mm in dorsal
mantle length. They are seldom found broken into the body and head-foot parts, as
their locking apparatuses may be fairly tighter than those of other species.
This species is readily identified by muscular and broad fin and moderately
squat body. At a glance, it may be taken as Ommastrephids, if four-series ofbrachial armatures and peculiar tentacular armatures are overlooked.
Nothing could be added to the careful descriptions by Berry (1912 as a partof G.fabricii; 1913, an amendment of the name) and Sasaki (1929).
Gonatopsis borealis Sasaki, 1923 makko*, n. subsp.
Oegopsida of moderate size; integument soft in touch and smooth all over.
Mantle loligiform, soft, subcylindrical, tapering gradually posteriorly to an acutely
attenuated point ; anterior margin truncated, projecting very slightly at the middorsum, broadly emarginated ventrally leaving rounded lateral angles on bothsides of the funnel ; the widest part of the mantle about one-third the dorsal
mantle length. No sculptures and color patterns present externally but dark
violet internally (in preserved state).
Fins muscular but rather soft with thinned margins, one-thirds as long as thedorsal mantle length, subrhomboidal, length equal to the width of both fins taken
together, obtusely angulated at three-tenths posterior from the anterior margin.
Head as broad as the mantle opening. Neck moderately constricted, probably marginated by a circum-nuchal ridge, which is indistinct in the type specimen.
N uchal locking cartilage elongated rectangular with rounded corners, feebly narrowed at the middle, width a quarter long. Funnel excavation demarked bydepressed nuchal folds on both sides. Funnel moderate in size, semilunar valve
presents inside. Dorsal funnel organ V-shaped, the posterior tips of rami reachas far as the anus. Ventral organs small and ovoid. Funnel locking cartilage
lanceolate, slightly curved, tapering anteriorly and widened posteriorly and then

* The subspecific name, makko, means sperm whale in Japanese.

114

T. OKUTANI AND T. NEMOTO

rapidly narrowed to an angulated posterior end; groove very shallow, widened
posteriorly ending an open end. Mantle locking cartilage very weak and ridgelike in shape.
Arms robust, very powerful in appearance; the ventral pair decidedly shorter
than the others and as long as two-thirds the longest arm; the arm formula 2, 3, I, 4;
the longest arm attains two-thirds the dorsal mantle length. All of the arms nearly
the same strength, gradually tapering distally ; the sucker-bearing faces flat meeting with the side faces in a rectangle ; the aboral side roundish. lntegumental keel
poorly developed on the third lateral arms. Web not well developed except at
the proximal portion of the ventral arm. Armatures of arms composed of hooks
and suckers. In the ventral pair, they present only suckers with several teeth
at the distal margin of the horny rings. In other pairs of arms, the proximal
several rows are loosely arranged suckers, there followed about thirty transverserows of two small suckers outer side and two large hooks of inner side each. The
outer suckers are borne on the thick and long pedicels and bear horny rings withfew triangular teeth at the distal margin. The inner hooks are very sharp withvestigial sucker-like structure proxiamally. Tentacle completely absent. Buccal
membrane papillated and dark violet in color (in preserved state) with sevenprojections on the margin, each suspended by long rib.
Gladius slenderly penniform, about one-fifteenth as broad as long.
Radulae not examined.
Types*: Holotype, male, found from the stomach of a sperm whale ; the integument
is somewhat matilated probably by action of digestive fluid of the whale. Dimensions of the type specimen are as follows : dorsal mantle length, 350 mm ;width of the mantle, 55 mm; length of the fin, 100 mm, width of the fin taken together, 100 mm; length of the dorsal arm (right), 200 mm; second lateral arm(right), 210 mm; third lateral arm (right), 225 mm; ventral arm (right), 170 mm.
Another specimen (paratype) with the dorsal mantle length 300 mm long is
also matilated and separated into two parts. The third specimen (catch number
39: 52°26N, 175°11E) was only head-foot part remained and was the largest of all.
The longest arm measured 300 mm in length and the arm formula was 2, 1=3, 4.
Remarks: At a glance, the present new subspecies is very similar to the typical
Gonatopsis borealis Sasaki which is abundant in the northern North Pacific. However,some external morphological characteristics do not coincide with those of G. borealis,
unless G. borealis presents a remarkable individual changes in proportion of various
parts of the body.
An abridged comparison of those two types are as follows :typical borealis
makko, n. subsp.
General shape of the mantle Cylindrical, somewhat
Cylindrical, verysquat; skin thickslender; skin thin
Fin length
2/3 total breadth or
The same as the total
2/5 mantle length
breadth or 2/3 mantle
length* Deposited in the National Science Museum, Tokyo, Reg. Nos: Mo. 13311 (Holotype), Mo. 13312(Paratype).

FOOD OF SPERM WHALES

115

N uchal locking cartilage

Panduliform, with aElongated rectangular
gentle constriction
near the middle
Funnel locking cartilage
Panduliform, 1/3 asCurved lanceolate,broad as long
1/4 as broad as longArm
2/5 the mantle length
2/3 the mantle length3, 1=2=4
2, 3 ( =) 1, 4
The criteria common to both forms are armature of arms which composedof four series of suckers and hooks all the way through and the sagittate outline of
the fins. In contrast to these, another known Gonatopsis, G. octopedaus Sasaki has
8-12 series of small suckers on the distal parts of the arms and a reiniform fins.
Distribution: Probably a bathyelagic zone of the northern North Pacific.
Mastigoteuthidae
Mastigoteuthis ( ?) sp.
A single strange specimens in a half digested state is taken to be a Mastigoteuthid.
The specimen has degenerated tentacles which present only muscular short
appendages between the third lateral and ventral arms. Degeneration oftentacles are hitherto known on Gonatopsis (Gonatidae) and Octopodoteuthis (Octopodoteuthidae) and none on any other families. Therefore, it is still inconclusive whether the present specimen belongs to Mastigoteuthidae.
Cranchiidae
Taoniuspavo (Lesueur, 1821)
Galiteuthis armata (Joubin, 1898)
Two above-mentined Cranchiids were found mixed with the preceding species
and Stigmateuthis sp. As they have very delicate and gelationous integuments,
they are found to be completely destroyed.
DISTRIBUTIONAL IMPLICATIONThe catch of sperm whales have been concentrated to the coastal waters of the
Aleutian Is., the Bering Sea and Alaskan Gulf. This is corroborated by Clarke(1956) who described that sperm whales congregate to the neighborhood of theislands in the mid oceans and continental shelves. The distributions of sperm
whales and their foods are illustrated in Figs. 1 and 2 based on the results in 1960
and 1963, respectively.
The male whales occupy the major part of the catch, and very small numberof females were caught as are so for the Antarctic catch of sperm whales. Matthews (1938) suggests the difference in food between the male and fameal is possiblypresent as the males are usually large in size and more powerful in catching preys
by deep sea diving. But no clear difference between the size of food squids intaken
by male and female has been proved. (Matthews, 1938: Clarke, 1956).

T. OKUTANI AND T. NEMOTO

116

0

2J-39

0

5-19

0

J-4

Fig. I. Distribution of squids and fish in the stomachs of sperm whales from the Bering
Sea, 1960. White-Squids, Black-Fish.
1'10''/,

160"

040i

to
j

0

0

J

~i

:_1~-'

E

180'

160'V;

20-39

0

~0-19

0

5-9

0

J-4
J4C/ ·,v

Fig. 2. Distribution of squids and fish in the stomachs of sperm whales from the Bering
Sea and the Alsakan Gulf, 1963. White-Squids, Black-Fish.

FOOD OF SPERM WHALES117

It is evident from the Figs. 1 and 2, squids dominantly occur in the westernpart of the coastal waters around the Aleutian Is. Fish as the food of the whaleare, on the contrary, mostly predominate in the eastern waters along the Alaskan
Gulf and the east Bering Sea. This tendency was also noticed by Ohsumi (personalcommunication, 1956) for the Bering Sea. In 1960, the dominant occurrences ofsquids were observed in the waters around 176 °E longitude and fish are mostly
found in the northern area of the Bering Sea.
The 1963 catch of sperm whales was taken from the Alaskan Gulf where thesperm whales feed on fish in larger quantity than squids. For the foods of sperm
whales in the other parts of the North Pacific Ocean, there are some reports available:
Mizue (1950) described squids and fish as the foods of sperm whales in theadjacent waters to Japan without classifiying them. Betesheve & Akimushkin(1955) described foods of sperm whales from the Kurile waters. The foods speciesof sperm whales in the east Pacific along the American coast were described by Rice
(1963).
TABLE I.

OCCURRENCES OF SQUIDS AND FISH IN THE STOMACHS
OF SPERM WHALES FROM THE NORTHERN PARTOF THE NORTH PACIFIC, 1960 AND 1963
Beiing Sea &Aleutian coast-180°

Bering Sea &Aleutian coast
180°-160°W

1960{ Squids
Fish

240
16

434
178

1963

{Squids
Fish

129
19

116
9

Coast of
Alaskan Gulf
160°W-130°W

45

94

According to Betesheva & Akimushkin (1955) and Sleptozov (1955), the mostimportant species of suqids as the food of sperm whales are Gonatus magister and
Gonatus sp. followed by Gonatus fabricii, Galiteuthis armata, Taonius pavo, Chiroteuthis
veranyi and Meleagrotheuthis seperata. They presented few occurrences of giantsqids, Moroteuthis robustus and Architeuthis Japonica, from the Kurile waters. On theother hand, Moroteuthis robustus occupied 41 % of the total food of sperm whalescaught off America (Rice, 1962), although the materials are rather poor. Gonatopsis
borealis stood the second rank of the foods reported by him.
From the materials treated here, the. following patterns of distributions byspecies are observed:
Type A: Moroteuthis robustus
This was the most important squid in the waters around Aleutian Is. and the occurrences of this species are illustrated in Fig. 3. Moroteuthis robustus was found in
the coastal areas of Alaskan Gulf as far as the Canadian coast. This squid wasreported to be found from the stomachs of sperm whales taken off the west coast of
British Columbia (Pike, 1950: Robbins, Oldham & Geiling, 1937). These obserbation suggest that M. robustus is also common in the California Current area.

118

T. OKUTANI AND T. NEMOTO

There was found no diurnal change in occurrences as is shown in Fig. 5.
Matsushita (1955) stated that the sperm whales in the Antarctic feed on squids inthe early or the late hours of the day when those squids come up to the surface of thesea. Thus, whales need not to dive so deep in the sea to feed on large squids. At
feeding, sperm whales in the Bering Sea may reach at the sea bottom sometimes, assome stones and other sedimentary aliens are found in the stomachs of sperm whalescaught there (Nemoto, & Nasu, 1962). But, the ocean depths are generally not solarge there as in the Antarctic waters.-The coastal waters of the Aleutian Is.
are shallow enough for sperm whales to reach at the bottom, so that Moroteuthis
robustus may be fed by them even in the mid-day at or near the bottom.

0
0
0
0

Type A, l l -,,,,Type E,

5-10
l-4
l-4
j

180'W

re:/~',

l40'W

Fig. 3. Distribution of squids found in the stmaches of sperm whales from the Bering Sea
and the Alaskan Gulf: Type A (Moroteuthis robustus) and Type E (Gonatus magister).

The specimens of M. robustus reported by Betesheva & Akimushkin ( 1955)
were mostly ranging in size from 100 cin to 140 cm in total length including thefoot. The largest one reported by Pike (1950) and Robbins, Oldham and Geiling(1937) measured 250 cm in length. In the present study, the large six specimensof Moroteuthis were found in the stomach of a single sperm whale, and thirty specimens of 100 cm or smaller were found from another sperm whale.
Type B & D:

Gonatopsis borealis makko

The position for the largest specimen (Type B) found together with Moroteuthis
was the west Aleutian waters. Other small specimens could not be identified on theFOOD OF SPERM WHALES

119

occasion of field observations. They were described as type D and were found in
the central part of the Bering Sea and the adjacent waters to the Aleutian Is. This
species was also found in the comparatively high latitudes of the Bering Sea. A
single occurrence was also observed in the coast of Alaskan Gul£'t~' :.

•

0Type B,

1

Type D, JO-

0 ,,
0

,,

1 -4

6

Type C, 1-4

xType G,.IG'',\

Fig. 4. Distribution of squids by type found in the stomachs of sperm whales from theBering Sea and the Alaskan Gulf: Type B (Gonatopsis borealis makko), Type D (thesame), Type C (Mastigoteuthis ( ?), Stigmateuthis, Galiteuthis and Taonius), and Type G.
(Gonatus fabricii).

Type C: Mastigoteuthis? sp., Stigmateuthis sp., Galiteuthis armata, and Taonius pavo
Above four species are described as type C on the field note although the shapeand other taxonomical criteria are different. These species were mainly found in
the central part of the Bering Sea as shown in Fig. 4. From the occurrences offragmental integument in the stomachs of sperm whales, these squids were suspected
to be distributed mostly the central part of the Bering Sea.
Type E : Gonatus magister
This species was observed in the coastal waters of Aleutian Is. and Alaskan Gulf
as in Fig. 3. This species is considered to be very common in the north-eastern
Pacific.
Type G: Gonatus fabricii
Gonatus fabricii was recorded as the main food of the toothed whale (bottlenose) in the Atlantic (Hjort & Ruud, 1929). Betesheva & Akimushkin (1955) des-T. OKUTANI AND T. NEMOTO

120

cribed considerable numbers of occurrences of this species from the adjacent waters
to Kurile Is. Pike (1950) also found this species from sperm whales taken off the
Birtish Columbia coast. In the present survey, only a single large specimen was
found from the Alaskan Gulf.
Type D55Type A

5

10

5

6

8

10

12

14

18

Time of the Day

Fig. 5. Occurrences of Type A (Moroteuthis robustus) and Type D (Gonatopsis borealis
makko) in the stomachs of sperm whales from the northern part of the North Pacific byhours of a day.

It is interesting to note that none of Myopsida, mostly coastal and benthonicforms, was found in the stomachs of sperm whales. Myopsida is generally tropic
and temperate water forms and they are naturaly not distributed in the waters ofhigh latitudes.
SUMMARY

The squids specimens found in the stomach of sperm whales taken from the northernpart of the North Pacific (Bering Sea and Alaskan Gulf) were examined.
1. Moroteuthis robustus, Mastigoteuthis? sp., Stigmateuthis sp., Galiteuthis armaa, Taonius
pavo, Gonatus magister, Gonatus fabricii, and a new subspecies, Gonatopsis borealis makko,
were found in the stomachs of sperm whales as their foods.
2. The distributions and occurrences of those squids are discussed.
Moroteuthis robustus, which is considered to be the most important squid for the foodof sperm whales, is common in the adjacent waters to Aleutian Is. Gonatopsis
borealis makko is also common not only in the central part of the Bering Sea, but also
in the northern part. Gonatus magister is found along Aleutian Is. and Alaskan Gulf,
while G. fabricii is found in the Alaskan Gulf. Cranchiids are discovered from hecentral part of the Bering Sea.
3. The distribution of those food squids is considered to be concentrated to thecentral part of the Bering Sea along the chain of Aleutian Is. This coincides withthe major distributional area of sperm whales. On the other hand, fish as the foodof sperm whales, are mainly found in the waters of the northern part of the Bering
Sea and Alaskan Gulf.

FOOD OF SPERM WHALES121

REFERENCESAKIMUSHKIN, I.I. (1955). The feeding of the Cachalot. C.R. Acad. Sci., U.S.S.R. IOI (6): 1139-40.
BERRY, S.S. (1912). A review of the cephalopods of western North America. Bull. Bur. Fisher. 30,
Doc. 761: 267-336. 25pls.
- - (1913). Notes on some west American cephalopods. Proc. Acad. Nat. Sci. Phil. 65: 72-77.
BETESHEVA, E.I. & AKIUMSHKNI, I.I. (1955). Food of the sperm whale (P. catodon) in the Kurile Islands.
Trans. Oceanogr. Inst., Moscow 18: 86-94.
CLARKE, R. (1955). A giant squid swallowed by a sperm whale. Norsk Hvalfangst-Tid. 44(10): 589-93.
- - (1956). Sperm whales of the Azores. Discovery Rep., 28: 237-98. Ip!.
CLARKE, M.R. (1962). Stomach contents of sperm whales caught off Madeira. Norsk Hvalfangst-Tid.,
51(5): 173-91.
HEEZEN, B.C. (1957). Whales entangled in deep sea cable. Deep Sea Res., 4: 105-15.
HJORT, J. & RUND, J.T. (1929). Whaling and fishing in the North Atlantic. Rapp. Proc. Vverb. Re. 56:
5-123.
MATTHEWS, L. H. (1938). The sperm whale (Physeter catodon). Discovery Rep. 17: 93-168. 9 pls.
MATSUSHITA, T. (1955). Daily rhythmic activity of the sperm whales in Antarctic Ocean. Bull. Japan. Soc.
Sci. Fish., 20(9) : 770-3.
MIZUE, K. (1950). Food of whales (in the Adjacent waters of Japan.) Sci. Rep. Whales Res. Inst., 5:81-90.
NEMOTO, T. & NAsu, K. (1962). Stones and other aliens in the stomachs of sperm whales in the Bering Sea.
Sci. Rep. Whales Res. Inst., 17: 83-91.
RICE, D.W. (1963). Progress report on biological studies of the larger cetacea in the waters off California.
Norsk Hvalfangst-Tid., 52(7): 181-7.
PIKE, G.C. (1950). Stomach contents of whales canght off the coast of British Columbia. Prag. Rep. Paci.fie
Cosat St. 83: 27-8.
ROBBINS, L.L., F.K. OLDHAM, & E.M.K. GEILING (1937). The stomach contents of sperm whales canght
off the west coast of British Columbia. Rep. British Columbia Museum 1937: 10-20, Ip!.
SASAKI, M. (1923). On a new eight-armed squid from Hokkaido, Gonatopsis borealis n. sp. Ann. :(,ool. Jap.
10: 203-207- - (1929). A monograph of the dibranchate cephalopods of the Japanese and adjacent waters. J. Coll.
Agr., Hokkaido Imp. Univ., 20: 1-357. 30pls.
SLEPTOzov, M.M. (1955). On the biology of cephalopod molluscs of the far-eastern seas and the northwestern Pacific. Trans. Ocenolog. Inst., Moscow, 18: 69-77.

122

T. OKUTANI AND T. NEMOTO

EXPLANATION OF THE PLATESPLATE I
Fig. 1. Moroteuthis robustus (Verrill), dorsal mantle length 700 mm.
la. Tentacular club of another specimen.
PLATE II
Fig. 2. Gonatus (s.s.) fabricii (Lichtenstein), dorsal mantle length 300 mm.
Fig. 3. Stigmateuthis sp. (separata Sasaki?), dorsal mantle length 100 mm.
PLATE III
Fig. 4. Gonatus (Ben:_yteuthis) magister Berry, dorsal mantle length 250 mm.
Fig. 4a. Tentacular club of the same specimen.
PLATE IV
Fig. 5. Gonatopsis borealis makko, n. subsp., dorsal mantle length 350 mm. (Holotype)Fig. 5a. Left lateral arms of the same specimen.
Fig. 5b. A funnel locking cartilage.
Fig. 5c. Nuchal locking cartilage.
PLATE V
Fig. 6. Taonius pavo (Lesueur), anterior part of the mantle torn out, dorsal mantle length 250+x mm.
Fig. 7. An unidentified squid, probably Mastigoteuthis sp.

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