A REVIEW OF FOOD OF BALAENOPTERID WHALESAKITO KAWAMURA
Faculty of Fisheries, Hokkaido University, Hakodate, Hokkaido

ABSTRACT

In order to elucidate what species among so many kind of marine organisms are likely to be consumed largely by the balaenopterid whales, the existing evidence on the food habits of baleen whales is reviewed. To meet withthis primary purpose the report was mainly focussed on to describe qualitative
aspects of food species having been known to date from the notable whaling
grounds over the world rather than documenting quantitative subjects.' One
of interesting facts noticed throughout the contribution was that there exists
fairly intense diversity in the assembly of food species composition by regions
such as; northern hemisphere vs. southern hemisphere, Pacific region vs.
Atlantic region, inshore waters vs. offshore waters, embayed waters vs. open
waters, where the former usually shows more div'ersed complexity than thelatter. The fact however suggests that although the composition of food species locally varies over the various whaling grounds, the food organisms astaxonomical groups are very similar one another even in locally isolated whaling grounds when the food organisms and their assemblies are considered by
the family or genus basis. In this connection many evidences given in the
text may suggest that the balaenopterid whales as a whole may substantially
live on quite simply compositioned forage assembly in comparison with tremendous variety of organisms existing in the marine ecosystems. One of important aspects of the baleen whales food must be found in their characteristics of forming dense swarms, schools, and/or aggregations in the shallower
enough layers to be fed by the whales. The present and past status of larger
baleen whales as the mighty monarch through their evolutional pathways
may entirely depend upon the spatial distribution pattern of possible foodorganisms, i.e. the animal aggregations.

CONTENTS

Introduction .................................................. 156
The balaenopterid whales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 7
Baleen plates and feeding . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 159
Feeding types . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163
Kind of food organisms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 166
Southern Seas .............................................. 166
a. Antarctic and subantarctic .............................. 166
b. South African waters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 170
c. Australian and New Zealand waters ...................... 174
d. South Pacific and its environs. . . . . . . . . . . . . . . . . . . . . . . . . . . . 175
e. Brazilian waters ........................................ 175
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Indo-Pacific176
a. Arabian and Malaysian seas ........................... . 176
Northern Seas-North Pacific ............................... . 176
a. North Pacific and Bering Sea ........................... . 176
b. Far Eastern Seas ..................................... . 180
c. Subarctic to subtropics of pelagic Pacific ................. . 182
d. Coastal waters of Japan and East China Sea regions ..... . 184
Northern Seas-North Atlantic ............................... . 186
a. Northeast Atlantic ..................................... . 186
b. Northwest Atlantic ................................... . 189
Summarizing addendum ....................................... . 191
References ................................................... . 193
INTRODUCTION

This contribution firstly prepared in response to the request by the CRC Press Inc.,
U.S. A. under the title, "Diets for Balaenopteridae" as one of the contents should
be included in the "Handbook Series in Nutrition and Food". In December 1979,
however, the publisher informed me that they were unable to include this contribution in the volume to be published near future due to excess holds of contributions
submitted. In response to this situation, submission of article was switched to the
Scientific Reports of the Whales Research Institute No. 32, 1980 by courtesy of Dr Hideo
Omura, the Director of the Whales Research Institute, Tokyo.
Since the main purpose of this article was subjected to describe the known evidence on the food habits of balaenopterid whales, the viewpoint was mainly focussed
on to collect and give qualitative information in a manner of encyclopedia rather
than documenting quantitative and logical discussion. On the other hand, the evidence around food and feeding habits of larger whales from the ecological viewpoint
has been well documented by Nemoto (1959) and later by Gaskin (1976). The
former treated largely the case found in the North Pacific and its environs, while
the latter covered nearly all kind of whale species occurring over the world oceans,and summerized widely scattered evidences into a very comprehensive knowledge
from the viewpoint of evolutional ecology.
At present, it may be considered that the whaling is at its lowest but somewhat
stationary activities being operated in very limited regions, and that the kind offood items taken by baleen whales does not seem to increase so largely as the past
several decades when both whaling ground and whale species changed drasticallyfrom year to year. Since the study by Gaskin (1976), however, some additional
evidence has been known through the study of Bryde's whales from the tropical seas(e.g. Kawamura, 1977), and these are considered to be added as a recently known
evidence to the general knowledge of food habits of baleen whales.
This contribution aims to present simply what kind of organisms are likely to
be taken selectively by the larger whales occurring over the world oceans. Different from treating materials by the organic carbon basis, qualitative knowledge by
species basis must be characteristic and indispensable factors in elucidating theSci. Rep. Whales Res. Inst.,
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structure and function of marine ecosystems since the amount of organisms annually consumed by the larger whales is undoubtedly so enormous as estimated the casein the antarctic krill, Euphausia superba (e.g. Laws, 1978). As the biological environmental factors local food condition strongly relates to the movements and migration of whales (Kellog, 1928; N emoto, 1959; Kawamura, 197 5 ). One of anotherimportant aspects of the dietary habits is the state and condition of stomach contents actually observed in carcasses, which indicate ecological characteristics of food organisms such as individual density, biology of animal aggregation and patchy
distributions, neither of these are hardly known by the conventional net samplings(Omori et al., 1972; Kawamura, 1974; Brodie et al., 1978). In this connection, thebaleen whales and their feeding characteristics as discussed by Klumov (1962) can
be considered really functional and effective 'biological sampler' for marine zooplankters and micronektonic organisms.
Each evidence given in the text may indicates that considerably diversified
food habits are exhibited by locality, seasons, and by each whale species even in ataxon, Balaenopteridae. Ever known facts on the food and feeding ecology may
indicate guide lines which suggest a possibly existing inter- and intraspecies relationships between larger whales and small planktonic or nektonic organisms oflower trophic ladder.
In order to unify the family name of food organisms under the similar forms,
such expression as euphausiids, for example, in the tables quoted elsewhere in the
text was changed into Euphausiacea without notifying this in each corresponding
table.
THE BALAENOPTERID WHALESThe order Cetacea is comprised of two suborders, Mysticeti and Odontoceti. The
former is the baleen or whalebone whales, which includes three taxonomicalfamilies, i.e., Eschrichtiidae, Balaenopteridae, and Balaenidae, while the Odontoceti, the toothed whales are comprised of five families (Table 1).
As it is suggested by the common name, all member of mysticete whales thatare known as. the baleen whales or whalebone whales, are furnished with 'subtriangular horny baleen plates in the upper mandible as the filtering apparatus
instead of tooth in collecting forages although the shape and structure of baleen
plates vary considerably by each family and species. As relatively large mouth
proportion, say, about I /5 long of their total body length suggests, the mysticete
whales could be considered the greatest 'filter-feeders' or 'strainers' among all
animal ever lived on this earth. It is their mouthful row of baleen plates thatmake mysticete whales to be the quite distinct animals both in biological and ecological aspects, especially in gathering their food more powerfully in the aquatic
environment. Of three mysticete families, the Balaenopteridae is the representative taxon which includes so-called great whales or 'rorquals' i. e., the blue, fin(finback), sei, Bryde's, minke whales and humpback whale.
Apart from two another families, the balaenopterid whales are supposedly best
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adapted animals to aquatic environments, and distributed widely throughout theworld oceans. Their habitat is usually found in the pelagic waters while twoothers, Balaenidae and Eschrichtiidae, are common in the pelagic to neritic waters.
Sometimes these whale group invade deeply into the inlets and lagoons. Becauseof their well adapted structure to the aquatic environments, the balaenopterid
whales could perhaps have been maintained their largest populations than any othermysticete families in both northern and southern oceans, which undoubtedly due totheir powerful, and consequently more advantageous feeding habits as filter-feeders
in fulfilling the daily nutritional requirements. Because of their gracefully developped but mysterious way of life, the great whales have long attracted scientificinterests, and their dietary habits must have been refered to as one of those mysteries. Inversely, it was unfortunate for the animals of distinct zoological characteristics, that they were doomed later to be the target of modern whaling, but it
was also the beginning of intensive harvest of those giants over the world oceans.
TABLE 1.

THE MEMBER OF MYSTICETI (MYSTACOCETI: BALEEN WHALES)

Scientific name
Family Balaenidae
Balaena mysticetus
Balaena glacialis glacialis
Balaena glacialis australis
Caperea marginata
Family Eschrichtiidae
Eschrichtius robustus
Family Balaenopteridae
Balaenoptera musculus nBalaenoptera physalus
Balaenoptera borealis
Balaenoptera edeni 2lBalaenoptera acutorostrata3l
Megaptera novaeangliae

Common namebowheadnorthern right whale, black right whale, right whale
southern right whale, black right whale, right whale
pygmy right whale
gray whale
blue whale
fin whale
sei whale
Bryde's whale
minke whale
humpback

1) There exists a subspecies, pygmyblue whale, B. musculus brevicauda, which is an endemic species inthe southern ocean (Omura, et al., 1970).
2) The name B. brydei is still valid relating to the identity of two forms of this animal (Omura, 1977).
3) B. bonaerensis has been used for the southern animals. Between B. acutorostrata and B. bonaerensis
there are morphological and biological differences by a magnitude of hardly separable (Ohsumi etal., 1970; Omura, 1975, but see also Doroshenko, 1978). But, Rice (1977) proposes three subspecies,
B. a. acutorostrata, B. a. davidsoni, B. a. bonaerensis.

Because immediate ancestry animals for the mysticete whale are unknown atpresent, it is very hard to suppose that at the beginning of adaptive radiation whatmade the land-dwelling animals driven so as to invade into the aquatic environment
during the Oligocene to Miocene periods. However, it seems to be one of the best
way for the ancestry animals evolving toward the aquatic animals that feed mainlyon the abundant minute crustaceans which undoubtedly had been widely obtainable over the epicontinental shallow seas since the suturing of the supercontinent
Pangaea in the Permo-Triassic periods onward (Schram, 1977), and subsequentSci. Rep. Whales Res. Inst.,
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rapid evolution might perhaps be accelated by the increased marine production by
upwelling of the sea, although the latter is a hypothesis at the present state (Lipps
and Mitchell, 1976).
Although each food item for the balaenopterids may show geographicai andseasonal variations along with the abundance of suitable food even within the same
whale species, the filter-feeder as an adaptation to aquatic environments mustbe perhaps an great advantageous way oflife in competing for food with many kindof another animals. Among so many diversed species composition with the variety of radiation in the phocid seals, the crabeater seal, Lobodon carcinophagus, for example, is the unique animal. The complexed teeth and its general arrangementsin this animal serve as the sieve in collecting minute crustaceans such as Euphausia
superba in the Antarctic waters (Bertram, 1940). The unquestionably larger population size of L. carcinophagus presumably indicates the successful evolutional pathways of this animal through out the order Carnivora (Kawamura, 1972).
The balaenopterid whales are really cosmopolitic animals that undertake alarge scale seasonal migrations of several thousands kilometers, which connect between feeding and breeding grounds while the balaenid whales, Balaena mysticetus,
Balaena glacialis glacialis, E.g. australis and Caperea marginata are considered somewhat
neritic and/orendemically established species. Although the famous gray whale,
Eschrichtius robustus migrates for a great distance of such as between Lower California
and the Arctic Ocean, this animal must be considered really neritic and bottom orbenthos feeders (Rice and Wolman, 1971), which enable them to feed any time andplace enroute their migrations though the major feeding seems to occur in the higharctic regions. The balaenopterid whales as fully developed planktono- and ichthyophagous animals complete their migration by utilizing deposited lipids as anenergetic sources. As it has been pointed out by many workers, the balaenopterids
usually do not or can not feed while they are far outside from the proper feeding
grounds, but recent observations (Gambell et al., 1974; Kawamura, 1975, 1977)
strongly suggest the feeding activity of baleen whales even in the warmer seas.
This again suggests the balaenopterid whales to be the most fully adapted filterfeeders among many others since they can deposit possibly enough amount oflipids
for the nutritional requirements in addition to that used in metabolism during four
to five month stay in the feeding grounds.
BALEEN PLATES AND FEEDING

An adaptation of the balaenoptrid whales as the filter-feeder and, despite their ingenious structure in harvesting those minute planktonic crustaceans, the gross daily
nutritional and energetic requirements are enormous although the energy cost for
locomotion as an aquatic animals would be very smaller than any other terrestrial
animals (Schmidt-Nielsen, 1972; Kawamura, 1975). For instance, the southern
fin whale is estimated to consume only 0.022 kcal/gram body weight/km (Kawamura, 1975). As it is supposed generally in the food items for the balaenopterid
whales, there seems to be difficult in obtaining suitable amount of food over theSci. Rep. Whales Res. Inst.,
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vast oceans except those spacially very limited places so-called the feeding grounds.
The baleen whales, from the beginning of their ancestory evolution towards aquatic animals, are largely dependent upon the organisms occurring with large biomass
and forming dense swarms in the upper subsurface waters (Hjort, 1933). Although
the planktonic crustaceans, small gregarious fish, fish larvae, and some others likesquids may occur widely in the oceans, there are only very limited groups of possible food organisms being actually utilized by the baleen whales. The productivity of oceans shows quite biassed aspect in its distributions. The rich primary
production which finally links to the rich secondary production is usually found in
the colder seas of north and south latitudes higher than 40° latitudes and in the
upwelling regions. The famous feeding ground ever known for baleen whales,
therefore, may be regarded as the most possible and easiest place in locating aggregations and/or schools of prey organisms (Fig. 1). As mentioned before, theFig. I. World whaling grounds for mainly rorquals. The original figure by Mackintosh (1965, Fig. 7) to which recent and some past principal occupations of whaling grounds for sei and Bryde's whales are additionally demonstrated by mesh.
The whaling grounds in the mid-latitude offshore waters indicate the catch placeof Bryde's whales during exploratory fishing (1976/77-1978/79) under scientificpermit (see Kawamura, 1977, 1980; Ohsumi, 1979).

evidence that the baleen whales under breeding migrations rarely feed is certain tosome extent, but this actually seems to be the result due to absolute scarceness of
prey in the warmer seas where they give birth to a calf and mate. Whenever thewhales may encounter to the place where large but local food stocks are available
they undoubtedly prey upon them voraciously (Kawamura, 1977). This suggeststhat poor feeding activity of baleen whales during the breeding migrations does
not seem to be the result of their ecological and physiological characteristics but due
rather simply to the poorer standing stocks of possible food organisms in the breeding grounds or warmer sea regions.
To meet on to their nutritional requirements the baleen whale developed aSci. Rep. Whales Res. Inst.,
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quite characteristic ·apparatus-the baleen plates or whalebone. The subtriangular baleen plates with fine inner fringes and their curtain like arrangements
in two sets of row on the upper jaw form a fuge filtering or straining apparatus tosieve off some several cubic meters of water containing the aggregation of food organisms. An importance of well developed baleen plates and the row is obvious.
Once the baleen filter was damaged by some reasons such as the infection of parasites (Rice, 1967), the animal may sometime become very poorly nutritioned beingcaused by far little food ingestion due to less effective feeding apparatus.
In the balaenopterid whales the baleen plates usually count 260-400 in number on one of two sides of upper jaw. Each baleen plate is arranged closely one
to the next to form a comb-like row with an intervals of about 0.5-1.3 cm although
TABLE 2.

AVERAGE NUMBER OF BALEEN PLATES IN ONE OF TWO SIDESIN THE NORTH PACIFIC AND ANTARCTIC BALEENWHALES (Nemoto, 1959; Ohsumi et al., 1970)
Blue

Fin

North PacificRange
300-400 300-400
Approximate
360
355
mean
Antarctic and southern hemisphereRange
260-400* 260-480*360
Approximate
320
mean

Bryde's

Littlepiked

Sei

Humpback,

Right

Gray

260-370300

260-300280

320-380340

300-370330

230-260245

130-180160

250-280

261-359289

300-410* 300-370* 220-240*
345* After the data by Discovery research since 1929.
No. of baleen plates
286 +18* /262 cm
14.1 m, Male

45

~

"'c.

12 ~

2"'

c.
"'
c:

"'
"'

E

10 .§.

35

2"'

-a

<;;
..c

8

0

6~

-

c:

<1>

..c

i"' 30

4 'O

::J

z

(ij

>

2~

~
40 60 80 100 120 140 160 180 200 220 240
Distance from the anterior end of baleen row (cm)..E

251.......L-L...L..L-l-L...L..L..l'-:-'-:-'--:C:--:-'-:-7:-'-:~-'-:-:-:-'-:C:-:-'-:-:-::-'-:~'-'

20

Fig. 2. Variation of the number of baleen plates in sei whale. Arrow shows the
spot where maximum length of baleen plates was found (Kawamura, 1974).
(* fringes)
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the number of baleen plates and their intervals to the next may vary in details withspecies and the age of animals (Kawamura, 1974). (Table 2, Fig. 2). Usually,
the younger the animal, the more thickly intervaled in baleen plate arrangements.
One of baleen row measurements as found in the southern sei whale is demonstrated
in Fig. 2. The inner fringe of baleen plate is furnished with fine horny bristles(=fringes) of about 3-5 cm long which, by overlapping one to the other, to form
the sieving meshes or 'fibrous mat' as called by Gaskin (1976) that directly concerned with retaining the food organisms on them. The thickness of baleen bristles
Blue (adult)

1--------------1----------4

Blue (young)Fin (adult)
Bryde's (adults, Saldana Bay)
Bryde's (adult, North Pacific)
Bryde's (young, North .Pacific)
Littlepiked (adult)
I Sei (adult)~
Sei (young)Humpback (adult)
Grey
1-W-t Right (young). ()
Fin young1----1_________,
1----1---------t,_1-1----1

1.5

1.0
Diametre (mm)

0·5

0.1

1·B

Fig. 3. Diameters of baleen fringes of baleen whales measured at the center position of baleen plates. Black belts show the ranges of dominant sizes of baleen
fringes. Adult: whales after the sexual maturity, Young: sexually immature andbaleen plates are not chipped (Nemoto, 1959).

6.0
5.0

oEg
• Kawamura(Present study)+Williamson(1970)o Nemoto(1970)

0Bm
N" 4.0

.sro
Q)

~ 3.0
o

Bp

2.0

Bp
Bm
Bp
Be
Bb

Eg

1.0

~
Ba

Mn
Ba

10

15

20

Blue
Fin
Bryde's
Sei
Right
Humpback
Minke

25

Body length(m)Fig. 4. Total baleen filter area in baleen whales (Kawamura, 1974). Both filled
and open triangles show the filter area for the Bryde's whales in the North Pacificand southern oceans respectively (Kawamura, 1978a).

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in terms of their diameter also varies with whale species; then the coarsest filter
among the balaenopterids may be :round in blue whale whilst the finest one in sei
whale (Fig. 3). Further details on the baleen filter can be consulted to Nemoto(1959, 1970), and Kawamura (1974).
As the figures and tables clearly show, the general morphological character offiltering apparatus in the mysticete whales varies from family to family, and from
species to species, although a greater variation may be found in the family Balaenopteridae. The whole structure and the function of filtering apparatus is closely related to those above shown individual characteristics. Among all above mentioned the average along with maximum length of the baleen plates may decides thetotal filter area of animals, which may finally characterize the food and feeding
habits of each whale species (Fig. 4).
Putting together these figures and tables, it is noticed that the balaenopterid
whales as a taxonomical group may be recognized from both eschrichtiids and balaenids by distinguishing relative shorter but finer filtering apparatus. Although
there are several another morphological characteristics relating to the dietary habits of whales, the author believes that the structure of filter itself and its total but
actually functional filter area is among all the most important agents that determine
the feeding habits of each whales species. Further but more general details on thissubject may be consulted to Scoresby (1820), lngebrigtsen (1929), Mackintosh(1965), Slijper (1962), Nemoto (1970), Kawamura (1974) and Gaskin (1976).
FEEDING TYPESThe morphological characteristics of filtering apparatus in baleen whales may
extremely well related to the feeding types and feeding behavior, which largely
determine the kind or the group of principal prey organisms although the latter
again varies with the geographic regions.
Two different feeding types employed in baleen whales have been proposed,(Nemoto, 1959, 1970): i.e., swallowing and skimming types, or swallowers and skimmer (Mitchell, 1974), both of which were perhaps derived from the field observations by lngebrigtsen (1929), although Hjort (1933) noted swallowers as to "play"against the so~called skimmers. His description is so suggestive as to imagine thewhales feeding, and considered worth to be quoted; he (Ingebrigtsen, 1929) describes,"During whaling operations in 1905 Ihad the opportunity of seeing that the humpback is far more intelligent than other species of whale. It employed two methodsof capturing "krill" when the latter was on the surface of the water. One wasto lie on its side on the surface and swim round in a circle at great speed, while itlashed the sea into a foam with flukes and tail and so formed a ring of foam. The
frightened " krill " gathered together in the circle. This done the humpback dived
under the foam-ring and a moment later came up in the centre to fill its open mouthwith "krill" and water, after which it lay on its side, closed its mouth, and thecatch was completed."."The other method'', Ingebrigtsen (1929) continues, "was to go a short disSci. Rep. Whales Res. Inst.,
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tance below the surface of the water, swimming a ring while at the same time it
blew off. The air rose to the surface like a thick wall of air bubbles and theseformed the " net". The "krill" saw this wall of air bubbles, were frightened into the centre, and then the manoeuver of the first method was repeated."" When the whale lay on its side and shut its mouth the rifled belly formed anenormous distended bag before it was contracted and the water pressed out between the whaleboneplates.""When one saw that enormous bag, which was many times larger than thereal mouth from the gullet to the nose one could understand that the humpback,fin-back, sei-whale and blue-whale have the rifled belly from the point of the chinto the middle of the body, solely in order to be able to take in an enormous quantity of water containing food. With the straight jaws and rather straight palatalregion possessed by these 4 whales as compared with the right whales, which have
curved jaws and palatal region, there would be little room for water in the mouth,
if there was taut, smooth blubber under the chin and belly, as its the case with theright whales, the Greenland whale and the North Cape whale." He also describes, " The blue-whale, fin-back and humpback turn over, often with part of the head
above water, when feeding." Very recently Jurasz and Jurasz (1979; observed the
humpbacks in Alaskan waters use three different feeding methods, i.e., lunge feeding, bubblenet feeding, and flick feeding depending on the school mode of available feed. In the lunge feeding there also noticed that the method includes
variations of 'lateral', 'vertical', and 'inverted' lunge feeding.
" The sei-whale, on the contrary, ' skims ' the food. It swims at great speed
through the swarms of copepods, with half open mouth, its head above water tojust behind the nostrils. The copepods rush in with the water and are filtered fromthe water by the whale-bone plates. When a suitable mouthful of copepods has
been taken the whale di~es, shuts its mouth and swallows the food. It is especiallyin the evening and early in the morning, when the copepods are most at the surface,that' skimming' takes place." (Ingebrigtsen, 1929).
The balaenopterids are usually known as to perform a swallowing type feeding.
The whale engulps a mouthful of food containing water by the aid of ventral grooves, and sieve off through the slit of baleen row by giving a pressure on the waterwith its tongue. Then the food organisms retained on the baleen filter are swallowed through throat and oesophagus of several inches across. Recently Gaskin(1976) showed a series of very clear figures offeeding in balaenopterid whales. As
it is noticed in Figs 3 and 4, the sei whale shows rather closer feeding habits to thebalaenid whales with relatively finer baleen bristles and larger filter area, and is considered to demonstrate both skimming and swallowing types of feeding (See also
Table 3). Although the humpbacks as quoted above usually perform the swallowing type feeding, there are clear indication of occasional bottom feeding that is suggested by a significant occurrence of demersal fish and crustaceans along with large
amounts of' fine pebbles' in the stomach (Zenkovich, 1936 cited from Gaskin, 1976).
The minke whale as Norwegian call them "Vaaghval" (=Bay whale) sometimes
enters deeply into the river, inlets and the bay such as the Thames, St. Lawrence,
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FEEDING TYPES IN THE MYSTICETI WHALES(Nemoto, 1970; Mitchell, 1974)

Swallowing type or swallowers
blue whale and pygmy blue whale
fin whale
Bryde's whale
humpback whale
minke whale
Antarctic minke whale
Skimming type or skimmers
right whale (northern and southern right)
Greenland whale (bowhead)pygmy right whale
Skimming and swallowing typesei whale
gray whale
TABLE 4. THE ORDER OF SELECTION OF THE FOOD IN BALEEN WHALES*(Nemoto, 1970) (=shows equivalence, and >shows the dominance to the left)
Blue whale
Fin whale
Bryde's whale
Sei whale
Humpback whale
Minke whale
Right whale

Euphausiacea
Euphausiacea = Copepoda (large)= Gregarious fish >Copepoda(small) >Cephalopoda (squids)
Euphausiacea=Gregarious fish>Copepoda (small)
Copepoda ;;;;;Amphipoda;;;;; Euphausiacea =Swarming
fish=Cephalopoda (squids)
Euphausiacea =Gregarious fish> Demersal fish and crustacean
Swarming fish= Euphausiacea > Copepoda
Copepoda > Euphausiacea >Plank tonic Pteropoda* slightly changed from the original.

freshwater tributaries of Puget Sound (Tomilin, 1967). Pebbles and grand particles in the stomach suggest that the minke as well as the humpbacks feeds atthe bottom occasionally. The balaenids, the right whale group, as known by theirenormously long baleen plates with finest filter meshes but no ventral grooves perform a really way of skimming type of feeding (Scoresby, 1820; Watkins andSchevill, 1976). (Table 3).
The difference in feeding types of whales may characterize the group and/or
the species of food organisms in the order of more preferable feeding. Generally
speaking, the swallowing type of feeding performed by the balaenopterids is considered more suitable for harvesting the larger and faster swimming prey organisms
while the skimming in the near surface water by largely opened mouth would allow to entrap a really small and weak swimmers of planktonic organisms such as
ctenophores, pteropods, chaetognaths and copepods. (Scoresby, 1820; Hjort andRuud, 1929; Watkins and Schevill, 1976). Table 4 gives one of such differences
as having been proposed by Nemoto (1970) where he calls these trends as the orderof selection of the food in baleen whales. These, however, must be regarded as alargely generalized idea, which sometimes seems to be hardly applicable in actual
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cases mostly by the difference of ecologically heterogenious ocean systems to which
the whales come to concentrate and feed. One of typical examples of local differences may be found in the dietary habits of the balaenopterids demonstrated between the Antarctic and the North Pacific feeding grounds (Gaskin, 1976; Nemoto
and Kawamura, 1977).
KIND OF FOOD ORGANISMS
One of well known whales food is the krill, Euphausia superba Dana in the Antarctic Ocean. E. superba predominates over the southern oceans, and is not only
the food of baleen whales but also the food of almost all larger animals, seals, seabirds, fish, squids and perhaps of some benthic animals occurring on the shelf andunderwater ridges. One of comparative study relating this suqject was given by
Salinikov (1953 ). E. superba that is called as the Antarctic krill, certainly build up
those gigantic blue whale of weighing hundred tons and keeps growing the enormous biomass of various whale stocks. Undoubtedly, E. superba can be literally
regarded as the key species in the Antarctic marine ecosystems. (Marr, 1962;
Laws, 1977).
There are, however, many evidences on the food items of baleen whales occurring over the world oceans although the place of events somewhat biassed by
localities since our knowledge on the diet of larger whales largely depends upon thewhaling operations through which we can examine the carcasses of whales, but this isalso restricted by the whaling regulations for locality, seasons and whale species inclusive. Being due to these limited source for the informations, there are vast sea
regions such as Arabian Sea left under complete lack of knowledge concerning thewhales food. It seems, therefore, to be reasonable to describe the evidence on thefood habits of whales by the localities where the whaling of an appreciable extent
have had been and/or have been took place.
Southern Seasa.

Antarctic and Subantarctic

During earlier days of the Antarctic whaling in the South Georgian waters
there had been reported only Euphausia superba and some ' shrimp ' as the whales
food (Hinton, 1925). It might be simply due to the whale species hunted in those
years of the 1920s'-1930s', when the blue whale, Balaenoptera musculus and fin whale,
B. physalus were the main target of whaling. However, several years later, Peters(1938) found fish, Palarepis coregonoides and squid, Onichoteuthis banksii as food of blueand fin whales in addition to E. superba, and more later Peters (1955) added the following species for the Antarctic balaenopterids being based on the German whaling expeditions during 1936/37-1938/39; Euphausia recurva, Thysanoessa macrura, Cyllopus spezialis, Parathemisto gaudichaudii, Eusirus antarcticus, Calanus propinquus, and C.
( = Calanoides) acutus.
In accordance with the change of main harvestable whale species since theSci. Rep. Whales Res. Inst.,
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REVIEW OF FOOD OF BALAENOPTERID WHALES

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TABLE 5. STOMACH CONTENTS OF BALEEN WHALES CAUGHT BYJAPANESE PELAGIC WHALING FROM 1961 TO 1965 IN
THE ANTARCTIC (Nemoto, 1970)
Whale species
Food species
Euphausiacea
Euphausiacea & others
Copepoda
Amphipoda
Munida (Decapoda)
Pisces
Cephalopoda (squid)
VacantNo. of whales examined

Blue 1l

Fin

Sei2l

Humpback

Minke

517
4
2
6

16158
18

5936
4
2472
1514
75
31
5
16145
26182

7

88

2
9

10
93

9
76

674
1203

18878
35139

1) mainly subspecies Balaenoptera musculus brevicauda distributed in the lower Antarctic.
2) catch for 1966 season is included.

TABLE 6. FOOD ORGANISMS FOUND IN THE STOMACH OF BALAENOPTERIDWHALES IN THE SOUTHERN OCEANS (Abe, 1957; Nemoto, 1959, 1962, 1970;
Brown, 1968; Kawamura, 1970, 1974; Budylenko, 1978)
Euphausiacea :Copepoda:
Amphipoda:
Decapoda:
Pisces:

Cephalopoda :Pteropoda:
Chaetognatha :Tunicata :Euphausia superba*, E. crystallorophias, E. vallentini*, E. lucens, E. similis*, E.
recurva, E. frigida, E. spinifera, Thysanoessa gregaria,
T. macrura*, T. vicina, Thysanopoda actifrons
Calanus tonsus *, C. simillimus *, Clausocalanus laticeps *, Drepanopus pectinatus *,C. propinquus, C. acutus Calinocalanus macrocarinatus
Parathemisto gaudichaudii f. compressa*, f. bispinosa*, f. intermediate
Munida gregaria*, Penaeus sp.
Gymnospelus nicholsi, Myctophum subasperus, M. punctatum, Scomberesox saurus*,
Notolepis coatsi*, Vinciguerria attenuata*, Notothenia ramsayi, Paralepis sp.,
Tetroganurus curvieri, Lepidotus candatus, Protomyctophum normani, Xenocyttus
nemotoi, Engraulis australis*, Scomber sp.
Octopoda sp., Oegopsida sp., Onychoteuthis banksii
Clione sulcata, C. antarctica
Eukrohnia hamata
Species name is not stated* Important food species.
commencement of modern whaling in the Antarctic onward (blue-fin-humpbacksei/fin-sei-sei/minke-minke) (Kawamura, 1974), a considerable amount of knowledge on whales food have been accumulated. Table 5 is one of the summerized
figures which covers blue, fin, sei, humpback, and minke whales. It may clearly ·be noticed that the blue, fin, and minke whales as typical swallowing type feeder
mainly feed upon euphausiids while sei whale alone shows a stronger preference for
copepods and amphipods in addition to euphausiids, i. e., the former is consideredreally stenophagous and the latter is euryphagous at least in the Antarctic feeding
grounds.
Finding no significant differences in the structure of baleen plate of the North
Atlantic sei whale to that of the Antarctic, Tomilin (1967) considered that the sei
Sci. Rep. Whales Res. Inst.,
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KAWAMURA

168

whale firstly evolved as the microplanktophagous filter-feeder in the North Atlantic since there had been known only E. superba as the food of sei whale from the Antarctic, and he (Tomilin, 1967) concluded that the sei must be a species appeared
recently in the Antarctic regions. A part of this consideration, however, it does
not seem to be agreeable because the sei actually feeds on so many kind of minutecrustaceans in the Antarctic as well as in the North Atlantic although the evidence
from the palaeozoogeography supports the origin of balaenopterids somewhere in
the North Atlantic regions (Gaskin, 1976).
TABLE 7. KIND OF FOOD ORGANISMS OCCURRED IN THE STOMACHOF SEI, FIN AND PYGMY BLUE WHALES TAKEN IN THE
ENVIRONS OF CROZET ISLANDS (Pervushin, 1968).
Kind of food organisms

Sei

Euphausia frigida
Euphausia vallentini
Calanus propinquus
Calanus acutus
Calanus simillimus
Myctophum punctatum
Onychoteuthis banksii *

+
+
+
+
+
+
+

Fin

Pygmy blue

+

+
+

+

+
+

*Identification by Yu. A. Filippova.
TABLE 8.

FOOD SPECIES OF SEI WHALES IN THE ANTARCTIC (Doi et al., 1967)

SeasonEuphausiacea

Copepoda

MunidaAmphipoda

Pisces1964/65
1965/663688 (5)767 (7)192173 (7)1065

109 (2)1136 (2)

13 (3)
2

Cephalopoda(Squids)5

Number of concurrent food in brackets.

The species of food organisms known to date from the Antarctic region are
as given in Table 6. Looking at the table, we are surprised at the greater
diversity in the composition of diet in balaenopterid whales, and item of foodorganisms turns more diversified features when the adjacent waters to the Antarctic are included. Actually, Budylenko (1978) shown a total of eighty-two foodspecies having been found in the southern sei whales. However, it must be took
in mind that the majority of them are found in sei whale alone, and also it is only
a few food species that actually occurs largely with significant nutritional importance in each individual stomach. The very rare food species are also included in
the table. The subspecies of blue whale, B. musculus brevicauda (pygmy blue whale)(Omura et al., 1970) occurring in the Kerguelen/Crozet waters feeds solely upon
Euphausia vallentini (Nemoto, 1962), but Pervushin (1968) reported E. frigida andsome another organisms (Table 7). However, the food habits of this animal shifts
to E. recurva and/or E. diomedeae in the South African waters (Bannister and Baker,1967).
At present, minke whale, Balaenoptera acutorostrata bonaerensis (Rice, 1977) is
only the harvestable whalebone whales in the Antarctic waters. The stomach conSci. Rep. Whales Res. Inst.,

No. 32, 1980

REVIEW OF FOOD OF BALAENOPTERID WHALES

169

TABLE 9. COMPOSITION OF FOOD ORGANISMS IN THE STOMACHS
OF SOUTHERN SE! AND FIN WHALES DURING THE
ANTARCTIC SEASON, 1969/70 (Kawamura, 1974)
Calanus
tonsusII

ParaCalanus
Drepa- Euphausia Euphausia Euphausia Euphausia themisto
simillimus p notpust
lucensvallentini superha diomedeae gaudiec ina uschaudii----- ----- ----- ----- -----,_....__----- -----

Notolepis
coatsi,_....__~fu~fu~fu~fu~fu~fu~fu~fu~fu

C. tonsus
C. simillimus
Th. vicina
Th. gregaria
Th. sp.
E. lucens
E. vallentini
E. superba
E. similis
E. diomedeae
Th. actifrons
P. gaudichaudii
Penaeus sp.
S. saurus
V. attenuata
N. coatsi

203 1 > 3
52>
2

3

2
10

38

8

1
19
4

8

20

2

I & II : Order of dominancy.
1) Including a mixture with Pseudochirella sp. in the Order II.
2) Including a mixture with a few individuals of P. gaudichaudii, E. vallentini, Th. gregaria and E. hamata
in the Order II.
Note: Finding a few specimens of Parathemisto gaudichaudii at South Georgia, Mackintosh and Wheeler(1929) considered that this species were eaten almost accidentally along with E. superba. More later,
Mackintosh (1942) discussed the food habits of South Georgian baleen whales and stated that P.
gaudichaudii consisted a wholly insignificant part of the diet, and Brown (1968) confirmed this again
in sei whale. But he (Brown, 1968) found two species of myctophids, Electrona (Elampa) subasper andElectrona (Protomyctophum) normani. Table 9, however, strongly demonstrates that the importance of P.
gaudichaudii, copepods and euphausiids other than E. superba in the subantarctic waters.

tents of this animal were consisted of Euphausia superba, E. spinifera and Calanus tonsus(Ohsumi et al., 1970). However, more later study suggests that E. superba may beconsidered to be solely responsible food organisms of the minke whale (Ohsumi,
1979b), and Kawamura and Kikuno (1980) found but a single occurrence of
Thysanoessa macrura out of 381 minke whale stomachs examined.
Among the balaenopterid whales the Bryde's whale alone does not seem toenter into the so-called Antarctic region, and none of information is available atthe present state.
Although there are considerable number of food species in Table 6, the occurrence of different food species largely depends upon the characteristics of their own
zoogeography, and this may be noticed by comparing Tables 7 and 8. For thefilter-feeders one of the important aspects of ecological characteristics is that the
prey organisms should be an aggregate or school forming animals. This has been
Sci. Rep. Whales Res. Inst.,
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KAWAMURA

deduced from the finding that the stomach contents of each animal are really monotonously or even monospecifically compositioned (Table 9). As it is noticed inTable 9, the stomach contents of whales are usually composed of one or two predominant food species, and those with no asteriks in Table 6 are mostly considered
as a temporal migrants or occasional contaminants when the major food species
were preyed. To look at Tables 6 and 9, it is considered that the most importantand staple food of the southern balaenopterids counts only a several numbers ofplanktonic crustacean species, most of which are the herbivorous and/or omnivorous
creatures. At the sametime, it can be considered from the ecological viewpointthat the marine production and the energetic flows in the Antarctic ecosystem are
structured under relatively simple pathways, a very short circuited flux between
primary production and higher consumers.
TABLE 10. INCIDENCE OF WHALES WITH FOOD IN THE STOMACH IN THE
SOUTH AFRICAN AND ANTARCTIC WATERS (Gambell, 1968)
Fin

Sei

Durban 1l

Cape Province•>
South Georgia
Antarctic pelagic(Area II)

1962
1963
1965
1962
1963
1960-1 to
1964-V
1961-11 &1962-III

Food
present

Empty

33
3
13
122
351

56
79
24
138
281
167
149
16

{M 164F 276
186%
Tota1 feeding
89
82
37
260
632
331
425
202

Food
present Empty

37
4
35
47
56

45
16
5

50}
65
92

%
Total feeding

13

97
129
7
13
30

142
145
12
24
43

32
11
42
46
30

390

653

1043

37

336

72

408

82

11

1) Bannister & Baker (1967).
2) Best (1967).
M and F in the table indicate male and female respectively.

b.

South African waters
The balaenopterid whales visit the South African waters and its environs on
their way to and from the Antarctic feeding grounds. Usually those temperate to
subtropical waters are considered only the migratory passages for the baleen whales
and consequently, very little feeding activity is expected (Mackintosh and Wheeler,
1929). However, in some regions of under the influence by the Agulhas and Benguela Current systems where relatively rich marine production or standing stocksof both planktonic and nektonic animals are expected to form a fishing grounds,
the migrating whales may stay and feed to some extent (Table 10). Because ofits geographical and oceanographical characteristics, even the temperate/tropicalwhale species, the Bryde's may come and concentrate in those offshore and inshore
waters (Best, 1960, 1967). As it is suggested by the annual catching season in Saldanha Bay during the austral winter (chiefly from May to October), the stomachof whales shows what are the nutritional sources during the 'off season' of the Antarctic whaling.
Sci. Rep. Whales Res. Inst.,
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REVIEW OF FOOD OF BALAENOPTERID WHALES

171

There are rather few available data concerning the dietary habits of whalesin the South African waters except one fine piece by Best (1967), in which a very
comprehensive results are given being based on the investigations during 1962-1963.
By examining a total of 1085 stomachs, Best (1967) demonstrated that about anhalf of the stomachs of baleen whales visited the South African waters of lying in
mid latitudes (36°30'-32°40' S) was repleted considerably with many kind of foodorganisms (Table 11 ). The species of prey by a taxonomical groups found in thewhales stomach were given in Table 12. It is clearly shown in the Table 12 that
TABLE 11. NUMBER OF WHALE STOMACHS EXAMINED AND FEEDING
CONDITION IN fHE SOUTH AFRICAN WATERS (Best, 1967)
1962

Sei
Fin
Bryde'sBlue
Humpback
Minke

1963

A

B

A

B

260
24
30
0

47
45.8
100
0
0
0

632
43
89
2
3

55.4
30.2
83.1

0

O*
0
0

A: Number of whale stomachs examined.
B : Number of stomachs with food in percent figures.
* a few mega!opa larvae was found.
TABLE 12. FREQUENCY OF OCCURRENCE OF ORGANISMS IN BALEENWHALE STOMACHS, DONKERGA T 1962 AND 1963 (combined)(Bannister and Baker, 1967 ; Best, !967)1l
SpeciesEuphausiacea
Euphausia lucens
Euphausia recurva
Euphausia diomedeae
Thysanoessa gregaria
Euphausia spinifera
Nematoscelis microps?
Nematoscelis megalops?
Nyctiphanes capensis
Copepoda
Calanus tonsus
Calanoides carinatus
Clausocalanus arcuicornis forma major
Nannocalanus minor
Euchirella ro strata
Centropages chierchiae
Oncaea media
Corycaeus sp.
Corycaeus speciosus

Sei

Fin

Bryde's

54
33

3
9

18
5

7
3
I

2
2

Pygmy
blue3 >

Humpback3 >

} I

}

72
59
56
14
14
10
3
4

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

172

KAWAMURA
TABLE 12.
Species

Candacia bipinnata
Centropages brachiatus
Scolecithrix danae
Undeuchaeta major
Eucalanus attenuatus
Metridia lucens
Temora turbinata
Pleuromamma borealis
Oncaea venusta?
Paracalanus parvus?
Calanus tenuicornis?
Pisces" Mackerel " 2 lScomberesox saurus
Hygophum hygomi
Myctophum humboldi
Engraulis capensis
Trachurus trachurusSardinops ocellata
Maurolicus muelleri
Lestidium sp.
Scombrid sp.
Amphipoda
Hyperia macrophthalma
Vibilia armata
Brachyscelus rapacoides
B. crusculum
Primno macropa
Plaryscelus serratulus
Phronima semilunata?
Pteropoda
Cavolinia gibbosa
I anthina ianthina
lanthina globosa?
Sryliola subula
Diacria trispinosa
Megalopa larvae
Velella sp.

Sei

Continued.
Fin

Bryde's

Pygmy
blue3l

Humpback 3l

2
2
2
1

2

24
10
10
3
3

2
2

2

14

I

I) Retabulated (data by Best (1967) and Bannister & Baker (1967)).
2) These fish were not examined by a biologist.
3) Data at Durban, 1962-1963 by Bannister & Baker (1967).

the diet of whales is extremely variable in species composition as the evidence foundfrom such a spacially small scale feeding grounds. Especially, the compositions for
copepods, amphipods, and pteropods are of great complexity. The sei feeds mainly upon copepods as well as found in the Antarctic waters while fin and Bryde's feed mostly upon euphausiids though the latter shows more stronger trends of preferSci. Rep. Whales Res. Inst.,
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REVIEW OF FOOD OF BALAENOPTERID WHALES173

TABLE 13. PERCENT BY VOLUME OF DIET COMPONENTS IN SEI, BRYDE'S,
AND FIN WHALES IN THE SOUTH AFRICAN WATERS (Best, 1967)
1962

Euphausiacea
Copepoda
Megalopa larvae
PiscesOther organisms
Amphipoda

1963

Sei

Bryde's

Sei

Bryde's

Fin*

64.0
31.0
4.6
0.2

2.5

73.0
23.5
3.0
0.2
0.2

53

94.1
3.8

97.5

47
2.0

"' 1962 and 1963 combined.

TABLE 14.

SPECIES COMPOSITION OF DIET OF BRYDE'S
WHALES OFF DONKERGAT (Best, 1977)

Species

Inshore form
(number of stomachs)

Offshore form
(number of stomachs)

Engraulis capensis
Trachurus trachurusSardinops ocellata
Maurolicus muelleri
Lestidium sp.
Scomberesox saurus
Scombrid sp.
Euphausiacea
Euphausia lucens
Euphausia recurva
Nyctiphanes capensis
Tlo•sanoessa gregaria

25
11
10
0
0

0
0
0
3
3

Pisces

0
0
0
0
1
0

18
5

ing fish diet (Table 13).
Finding new rorqual species, B. brydei, Olsen (1913) reported the two instancesof very unusual food items where sharks up to 2 feet long was found in a stomach,and on another occasion there was found no less than 15 Jackass Penguins, Spheniscus demersus and one Cape Gannet "malagass ", Sula capensis. These are, however,considered to be taken accidentally while the whale was feeding on fish school,
and those stomach contents are presumably spit out later.
More recently, Best (1977) studied more about the Bryde's whale occurringin the same locality and showd a obviously different dietary habits between two
possible allopatric forms; the offshore and inshore forms. His data demonstratethat there exists stronger ichthyophagous form with more coarser baleen filter,
and the other is planktonophagous form .with finer, more sei-whale-like baleenfilter. The former corresponds to :the inshore form and the latter to the offshore
form. Their dietary characteristics which are chiefly due to the difference in occurring water masses are clearly observed in Table 14.
Because of very little whaling operatons in pelagic waters through the tropics,
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KAWAMURA

information around the dietary habits of baleen whales seem to scarcely found.
Land based whaling off Angola, the west coast of Africa was once reported but
there does not seem to exist any appreciable data about the stomach contents ofwhales. Along the Somalian coast both Bryde's and pygmy blue whales feed directly on fish, but species name is unknown (Yukhov, 1969). Another existing few
information suggests that the blue whale along the African coast feed euphausiids,
Nyctiphanes africanus and Euphausia lucens (Mackintosh and Wheeler, 1929), and the
humpbacks may feed the lobster-krill, Munida gregaria (Matthews, 1937), although
there is an indication of non-feeding activity of whales on the whole (Ottestad andRuud, 1936).
The variety and diversity of whales food items around the South African waters
may lead to a consideration that the baleen whales may feed upon those variableorganisms which occur with a larger biomass if not largely but even spacially limited distribution in such a local waters of well outside from their main feeding grounds
in the Antarctic Ocean.

Australian and New Zealand waters
The humpbacks had been the main baleen whale species hunted in the Australian and New Zealand waters. According to Dawbin (1956) the humpbacks on
the northward migration through the New Zealand waters usually feed little. In
some localities, however, there occurs very dense swarms of neritic species of euphausiids such as Nyctiphanes australis, on which the humpbacks feed occasionally.
N. australis is an important food species largely fed by the squid, Nototodarus sloani
sloani around the southern New Zealand waters (Kawakami et al., 1973; Kawakami,1976). In the Cook Strait, New Zealand, humpbacks also take the late larval formof" Glimothea" of galatheid decapods, Munida gregaria, the same prey fed by thebalaenopterids in the Patagonian waters. The Bryde's whale visited Hauraki Gulf
off North Island of New Zealand has been reported to feed on mugilid fishes called'Mullet' but its scientific name is unknown (Gaskin, 1976). Sei whale, on theother hand, feed largely on Calanus tonsus, Clausocalanus laticeps and Parathemisto gaudichaudii around southern New Zealand toward Tasmania (Kawamura, 1974).
In the western Australian waters, the food of humpbacks changes to Euphausia
spinifera and E. hemi.gibba (Dall and Dunstan, 1957). However, Hollis (1939) reported the occurrence of Pseudeuphausia latifrons, herring-like fishes tentatively been
identified as Clupeafimbriata and a few larval stomatopods in the three out of fiftyfive stomach samples of humpback whales taken in Shark Bay off the west coast ofAustralia, although the latter two food items were found in the mouth cavity of afemale animal. The anchovy, Engraulis australis and unknown species of young
mackerel also have been known as main food of Bryde's whales off southern West
Australia. In these waters, however, the baleen whales can be considered feed little during June to October as Hollis (1939) notes, "apparently the whales were notfeeding in this locality ", and empty stomach usually contained only heavy green
flocculent substance or "chunks" of dark green gelatinous material from the squamous epithelium of alimentary tract (Hollis, 1939).
c.

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REVIEW OF FOOD OF BALAENOPTERID WHALES

175

South Pacific and its environs
According to Budylenko (1978), Smirnov (1935) reported that the sei whale
stomachs from the tropical zone of the Pacific contained " fish ". One of few available data is that on the food and feeding habits of the southern Bryde's whale caught
in the Coral Sea and South Pacific regions (Kawamura, 1977, 1980). According to
Kawamura (1977), a total of 120 Bryde's whales (7 in the Coral Sea, and 113 in theSouth Pacific between New Zealand and Fiji Islands) were caught during OctoberNovember, 1976 and found that 75% of the stomachs over the regions contained
food with various state. One of another pelagic catches of the Bryde's whale offsouthern coast of Madagascar, was found that 46.7% of stomachs examined out of
105 were also filled with food. The compotition of stomach contents was completely different from that expected, i. e., they were solely comprised of euphausiids,
E. diomedeae, E. recurva, and Thysanoessa gregaria in the South Pacific animals while
there were only first two species in the Madagascar animals. Table 15 shows oneof results found by Kawamura (1980). It is again noteworthy that the feeding ofbaleen whales may takes place wherever the appropriate prey are found, and sogoes even in the tropical waters of generally poor in the first two stages of productivity.
d.

Brazilian waters
Since the commencement of whaling off Costinha, Brazil in 1910 onward, a
total of about 11,243 balaenopterid whales have been caught until 1974 (William-e.

TABLE 15.

NUMBER OF STOMACHS BY THE KIND OF FOOD
ORGANISMS (Kawamura, 1980).
1976/77
S. Pacific Coral Sea

E. diomedeae
T. gregaria
Euphausia sp. *E. recurva+ T. gregaria
T. gregaria +E. recurva
E. recurva
E. sibogae
P. latifrons
P. latifrons +E. diomedeae
T hysanopoda tricuspidata
Euphausia sp. **E. diomedeae + V. nimbaria
Fish larva***+E. diomedeae
V. nimbaria
Gonostomatid fish

SW IndianNo. 32, 1980

1978/79s. Pacific

E. Indian

5

Total

7
29
2
2

11
38
8
11
9
109
7
29
2
2

4

4

6

10

28

7
8
1
38

51***** Probably E. recurva.
** Probably E. sibogae.
*"'* Larval forms of Myctophiformes.
**** Mixture with one individual of hatchet fish is included.
Sci. Rep. Whales Res. Inst.,

1977/78

3
8
20

1

176

KAWAMURA

son, 1975). The abundant species as found in catch statistics are minke, sei, andhumpbacks yet there is no existing data about the diet of these animals. The
minke whales which have currently been caught about 700 annually do not seem
feed extensively in the Brazilian waters, but a short description stating; "The stomachs of nearly all the whales are empty. A little krill is found in the stomachs ofabout 3% of the whales" (Williamson, 1975). Unfortunately, however, Williamson (1975) did not give the scientific name for this 'krill '. However, feeding ofbaleen whales off tropical Brazilian coast does not seem to take place to an appreciable extent (Paiva and Grangeiro, 1965, 1970).

Indo-PacificArabian and Malaysian seas
Although the Bryde's whale occurs and strands occasionally in the ArabianSea regions (e.g. Roberts, 1970) and Malaysian seas (e.g. Berry et al., 1973), there
seems to exist no information about the dietary habits of whales.
a.

Northern Seas-North PacificNorth Pacific and Bering Sea
The baleen whales occurring in the North Pacific of higher than 50°N andBering Sea regions have been studied extensively along with that in the Antarctica.

Antarctic

North PacificFig. 5. Two deformed schemata for the Eltonian pyramid (Nemoto and Kawamura, 1977).

since there have been heavy exploitation of whale stocks by the factory ship operattions. Since the studies by Ponomareva (1949), Betesheva, (1954, 1955, 1961),
Nemoto (1957, 1959) and several more Russian workers as compiled in Tomilin(196/), the main whaling grounds shifted southward year by year far south down to
20°N in 1972 by an ammendments of whaling regulations and measures, from which
many kind of prey organisms have been introduced (Omori et al., 1972; Kawamura, 1973) In accordance with these general trends, it became rather that clear
an overall composition of whales diet shows considerable complexity covering various taxonomical groups of food organisms over the whole North Pacific and Bering
Sea regions. It can be stressed here that the relative importance of fish and/or
squid diet among many others in the northern seas is hardly comparable to thatin the southern hemisphere. When we see the structure of ecosystem through the
viewpoint of nature in food chain arrangements, for example, the shape of the ElSci. Rep. Whales Res. Inst.,
No. 32, 1980

REVIEW OF FOOD OF BALAENOPTERID WHALES

177

tonian pyramids could be expressed two schematically different types (Nemoto and
Kawamura, 1977) as shown in Fig. 5. The proposed schemata demonstrate thatmost of the principal food organisms in the North Pacific are consisted of both omnivorous and carnivorous species while the herbivores largely form main diet of baleen
whales in the southern oceans (Fig. 6). The feeding grounds of baleen whales, whenviewed from the structure of trophic levels in the sea are not substantially equivalent from one to the another.
An overall food items for the North Pacific balaenopterid whales are given in
Table 16. To see the Table 16 along with Fig. 6, it may be noticed that relatively higher occupation of fish and squid in the whales diet in the North Pacific is cha-N

• -/-)'f

40'~~;

•
•

,
tl

I

---

I

140"E

•

\

I

'

II

() 0

®©0 0 0 0 0@ \ID © ®O 0 0 ()@ @ ® (')

~'1-

---

Foshes,,..P.

/·~ Ii)~~ -~ Eup1.~s-----;>lITC

N' <'t\

'·

50

36

120°W

~···- -~~
't f>·(1.~~ecopods. ~~'11~@@~~~()000 f~ ~ ~·() ~ ~ () () () (l Cl @'.) () () ~

)

/

140"

160"

~ Jr-t,.

·".- ~ r--"v' ~~I,.._,

v

180°

160"

140"E

I

!40'

l\~30
26

.. --

'
lso·

160"

160"

12o·w

140"

Fig. 6. Approximate occurrence of food organisms of baleen whales in terms ofpercentage figures in the North Pacific (Nemoto and Kawamura, 1977).
TABLE 16. STOMACH CONTENTS OF BALEEN WHALES CAUGHT BY JAPANESEPELAGIC OPERATIONS FROM 1952 TO 1971* IN THE NORTHPACIFIC (Nemoto and Kawamura, 1977).
Food species
Euphausiacea
Euphausiacea & copepoda
Euphausiacea & others
Copepoda
Copepoda & others
PiscesPisces & others
Cephalopoda (Squids)
Empty
No. of whales examined* Exclusive of data in 1966.
** Including Sergestes similis.
Sci. Rep. Whales Res. Inst.,
No. 32, 1980

Blue
97 .6%**

1.1%
1.3%

504
971

Fin

Sei

Bryde's

64.1%**
3.4%
0.3%
25.5%
0.0%
5.0%

12.6%**
0.0%

88 .. 9%

1.7%
10064
29575

82.7%
0.1%
3.4%
0.0%
1.2%
9665
21713

Right

Humpback

100.0%

77.3%
0.6%
3.9%
0.6%

11.1%

82
109

17.2%

0
9

0.3%
150
458

178

KAWAMURA

racteristic yet the importance of both euphausiids and copepods as staple food is unchanged. As it was observed in the South African waters, and possibly in the Antarctic, both Bryde's and humpback whales prefer strongly to feed fish than any
other members of the Balaenopteridae. On the contrary, sei whale that performs
both skimming and swallowing types of feeding shows a considerably wider rangesof selecting the preferable prey organisms.
Each group of prey organisms which represent both the northern North Pacific and the Bering Sea is shown in Table 17.
TABLE 17. FOOD OF THE BALAENOPTERID WHALES IN THE HIGHERLATITUDES OF THE NORTHERN NORTH PACIFIC AND BERINGSEA (Sleptsov, 1955; Nemoto, 1957, 1959; Tomilin,
1967; Kawamura, 1973).
Euphausiacea :Copepoda:
Mysidacea:
Decapoda:
Pisces:

Cephalopoda (Squids):
Pteropoda:

Euphausia pacijica, Thysanoessa inermis, T. longipes, T. spinifera, T. raschii
Ca/anus cristatus, Ca/anus plumchrus, C. finmarchicus, Metridia lucens
Gnathophausia gigas (larva)
Panda/us borealis, Sergestes simillis
Clupea pallasi, Mallotus catervarius, Theragra charchogramma, Colorabis
saira, Pleurogrammus monopterigius, Gadus macrocephalus, Eleginus gracilis,
Sebastodes polispinis, Boreogadus saida
Ommastrephes sloani pacijicus, many others & larva
Limacina sp. Clione sp.

Of those food organisms given in Table 17, the most important euphausiid
species from their frequency occurrence are considered to be Thysanoessa inermis andT. longipes as well as their similar situation in the North Atlantic waters (see Table
25). It is noteworthy that the genus Thysanoessa in the North Pacific and Bering
Sea region is extremely important while the genus Euphausia represents largely inthe southern oceans. In connection with this, the geographically biassed distribution of food species must be considered: i.e., T. inermis predominates in thenorthern side of the Aleutian Chains of especially eastern half of the region ofunder consideration while T. longipes may predominate in the southern side of the
Aleutian Chains. However, the humpbacks in Glacier Bay, Lynn Canal andFrederick Sound, Southeast Alaska were found to feed mainly on Euphausia pacifica,
herring, Clupea /zerengus, and/or capelin, Ma/lotus villosus (Jurasz and Jurasz 1979).
Similar geographical changes may perhaps be found in the copepod food. In
general, the distribution of food organisms shows a considerable locally biassed
concentrations according to the zoogeographical characteristics in distribution
pattern by each prey species, and the idea may be extended further over the almost
all groups of food organisms.
In the coastal waters of higher latitudes in the Bering Sea the fish food becomes
more important than in the pelagic waters. The fin, humpbacks and minke B.
acutorostrata davidsoni (Rice, 1977) in the Gulf of Anadyr, Olyutorsky Bay, Kronotsky
Bay and the Peter the Great Bay, feed chiefly on herring, capelin, saffron cod, andT. inermis. Both humpbacks and finbacks have been reported to feed largely on
Thysanoessa spinifera and its egg masses, and the latter was especially abudant inSci. Rep. Whales Res. Inst.,

No. 32, 1980

REVIEW OF FOOD OF BALAENOPTERID WHALESTABLE 18.

Date

179

WEIGHT OF STOMACH CONTENTS OF FIN WHALES TAKEN IN
THE BERING SEA (constructed from Ponomareva, 1949, Table l).

Body
weight(ton)

Food species
Ti

Tr

TI

Cc

Ct

Weight ofstomach contents (kg)Index ofstomachfulness

170.0
212.5
255.0
425.0
127 .1
85.0
68.0
255.0
212.5
85.0
34.0
38.2
63.7
340.0

470
570
690
112
250
270
340
580
560
150
97
120
200
640

68.0
51.0
136.0
85.0
297.5
233.7
170.0
27.5
85.0
25.5
106.2
106.0
170.0
340.0

150
110
36
230
810
530
450
400
230
BO
340
200
450
640

Kommandorsky region

16 VI
16 VI
18 VI
21 VI
25 VI
26 VI
27 VI
28 VI
4 VII
4 VII
5 VII
10 VII
13 VIII
6X

36.0
37.2
36.6
37.B
50.25
31.14
25.76
43.56
37.B
55.38
36.0
28.86
28.86
52.5

14 VI
14 VI
15 VII
19 VII
20 VIII
20 VIII
20 VIII
20 VIII
20 VIII
20 VIII
B IX
B IX
9 IX
9 IX

42.9
43.76
37.2
37.1
36.6
43.56
37.2
31.32
36.6
31.86
30.60
42.24
37.80
52.50

+
+

+
+

+

+

+
+
+
+
+
+
+

+

+

+
+

+
+
+

+

+

+

01,yutorsky Bay

Ti : Thysanoessa inermis,
Calanus plumchrus).

+
+
+
+

Tr: Th. raschii,

+
+
+
+
+
+
+
+
+
+

+
+
+
+

TI: Th. longipes,

Cc : Calanus cristatus,

Ct : Calanus tonsus ( =the humpback stomach (Hollis, 1939). He (Hollis, 1939) believes that over a shortperiod of time the euphausian egg may be of some importance as whales food.
The blue whale were found to feed T. inermis and Nematoscelis megalops both inAvachinskii Bay and Kronotsky Bay (Tomilin, 1967). Another food items which
are absent from the Chukchi Sea animals are sand lance (Ammodytes personatus),
amphipod (Anonyx nugax), euphausiid (Nematoscelis megalops) and copepod (Calanus
cristatus). Ponomareva (1949) reported that both Tftvsanoessa inermis and Th. longipes
were the dominant food assembly in the Olyutorsky and Commandorsky regions.
For these sea regions in 1947, Ponomareva (1949) gives following percentage figureswhere she suggests the increase of copepod food towards autumn:
June: Th. longipes (35%), Th. inermis (30%), Th. raschii (15%), Calanus crisSci. Rep. Whales Res. Inst.,

No. 32, 1980

180

KAWAMURA

tatus (10%) and Euphausia lanei (=E. pacijica) (10%)
July-October: Th. longipes (30%), Th. inermis (33%), Th. raschii (12%) andCalanus cristatus (25%)
In connection with relative occurrence of each food species, there exists one ofvery few indispensable data on the amount of stomach contents which enable us toconsider the actual importance of food species (Table 18).
The knowledge on the Chukchi Sea animal is very limited since there seem tohave been a sporadic whale catches, and the following prey organisms may be seenin the stomachs of humpbacks, but fin whales have been reported to feed T. raschii,
T. inermis, and Eualus gaimardi during August-September (Tomilin, 1967).
Food of the Chukchi Sea humpbacks:
Mysis oculata
Tf?ysanoessa longipes
Pandalus goniurus
Eualus gaimardi
Capelin (Mallotus villosus)
Saffron cod (Eleginus glacilis)
Arctic cod (Boreogadus saida)
The food item for the humpbacks includes considerable number of demersal
fish and crustaceans, which strongly suggests the animal may perform the bottom
feeding in the shallower coastal regions.

b. Far Eastern Seas
Because of the geographical and topographical complexities, there have beenreported a large variety of whales food species from many localities.
In the Kurile region the fin shows a greater variety of food items, and euphausiids, cephalopods and fish having been reported, i.e. Thysanoessa raschii, T. inermis andEuphausia pacijica, Podonema longipes, Ommastrephes sloani pacijica (squid), and fish, Pacific saury (Colorabis saira), anchovy (Engraulisjaponica), and walleye pollock (Theragra
charchogramma) (Betesheva, 1954, 1955), but later she (Betesheva, 1961) added
Calanus tonsus (=C. plumchrus) C. cristatus and Pleurogramma sp. as food of fin whalesin the Kurile region.
Sei whale, on the other hand, feeds largely on copepod, Calanus plumchrus but
feeds also on T. raschii, capelin (Mallotus villosus), sand lance (Ammodytes personatus)and sardine (Sardinella melanosticta) (Zenkovich, 1937). Japanese whaler says thatthe minke whale in the pelagic Okhotsk Sea regions feed exclusively upon the herring (Clupea pallasi) and walleye pollack (Theragra charchogramma). They says thatthe operation of herring fishing boat in those waters is the obvious indications forlocating a fishable concentrations of minke whale. The humpbacks may show asimilar diet to minke whale but have been reported Euphausia pacijica, walleye pollack and even the pink salmon (Oncorhynchus keta) (Tomilin, 1967), while the blue
whale feeds on Calanus cristatus and E. pacijica (Sleptsov, 1955 ).
Russian investigations (Sleptsov, 1955; Klumov, 1963 ), suggest both euphausiids and fish are the most important diet for the baleen whales occurring in the Far
Sci. Rep. Whales Res. Inst.,
No. 32, 1980

181

REVIEW OF FOOD OF BALAENOPTERID WHALESTABLE 19. FOOD OF BALEEN WHALES IN THE PELAGIC AND COASTAL
WATERS IN KURILE REGION DURING 1947-1954 (Sleptsov, 1955)
Whale
species
Fin
Sei
Blue
Minke
Humpback
Total

No. ofstomach Vacantexamined

196
42
15
12
12
277

12
8

20

Ca+
Eu

Eu+
Cep*

PiscesCep+
Pisces

52

23

18

3

7

2

4

2

65

30

64
5
6
6
4
85

10

2

Ca/anus Euphausiacea
2
6

8

18

Cephalopoda(Squids)

6

15
12

5
21

30

3

* Cephalopoda (squid).

TABLE 20. SPECIES COMPOSITION OF FISH DIET OF THE BALAENOPTERIDWHALES IN THE FAR EASTERN SEAS (Sleptsov, 1955; Tomilin, 1967)

Clupea harengus pallasi (herring)Eleginus navaga gracilis (saffron cod)
Osmerus eperlanus dentex (Arctic smelt)lHallotus villosus socialis (capelin)
Theragra chalcogramma (walleye pollock)Ammodites hexapterus hexapterus (A. penonatus ?) (sand lance)
Gadus morhua macrocephalus (Pacific cod)Sebastodes glaucus (rock fish)
Pleurogrammus monopterigius (Atka mackerel)
Sardinops melanosticta (Far eastern sardine)
Oncorhynchus keta (chum salmon)
Boreogadus saida (Arctic cod)
Cololabis saira (Pacific saury)Podonema longipes * (a morid fish)* Laemonema longipes (by K. Amaoka).

Eastern Seas, Okhotsk Sea, Kurile, off eastern Kamchatska and subarctic regionof the coastal Pacific (Table 19). Although Sleptzov (1955) did not mention thespecies name of food crustaceans in Table 19, it may be supposed that Thysanoessa
inermis, T. raschii, Calanus tonsus ( = plumchrus ), C. cristatus and C. pacijicus would represent the stomach of whales. Later investigation (Klumov, 1963), however, revealed the following prey organisms from the balaenopterid whales;
Copepods: Calanus glacialis, Eucalanu~ elongatus, Metridia ochotensis, M. pacijica
Fish: sardine (Sardinops sagax), anchovy (Engraulis mordax, and E. japonica),
capelin (Mallotus villosus)
Squid: Loligo opalescens and Gonatius fabricii
One of characteristics in the dietary habits of whales in the Far Eastern Seas
as suggested in Table 20 is the more extensive diversity in the species compositionof fish diet than in the pelagic waters of the North Pacific and Bering Sea. Occurrence of gregarious fish in the Far Eastern Seas is an important fact as the trophic
environment where even the planktonophagous fin and blue whales feed largely on
those fish schools. The balaenopterids known as the stenophagous animal show a
Sci. Rep. Whales Res. Inst.,
No. 32, 1980

182

KAWAMURA

distinct euryphagous food habits than any other feeding grounds.
TABLE 21. LATITUDINAL CHANGES OF THE DIET COMPOSITION FOR
THE BALAENOPTERID WHALES IN THE NORTH PACIFIC(Nemoto and Kawamura, 1977).
Occurrence of food item in percentage
Latitude (N)300-35°35°-400
400-45°45°-50°500-55°Euphausiacea

Copepoda

Pisces

Cephalopoda(Squid)

14
5
9
23
31.5

55
59
44
37

86
40
23
6.5
8.5

16
17

Decapoda

910.5
6Subarctic to Subtropics of the Pacific regions
As well as the case in the South Pacific the existing informations in the region
are again very scarce but one reports the occurrence of" small fish" from the sei
whale in the tropical Pacific (Smirnov, 1935 cited from Tomilin, 1967). As Fig. 6
demonstrates the diet of baleen whales in the mid to lower latitudes of the North
Pacific may largely be represented by a gregarious fish and copepods though a pelagic shrimp, Sergestes similis occurs largely in the stomachs of fin and sei whales, andthe sergestid shrimp is considered to make up the staple local food over the midlatitudes of the eastern North Pacific (Omori et al., 1973). An overall trends oflatitudinal changes in the dietary composition is given in Table 21. In the lower
latitudes, say, south of 40°N, there also exhibits the changes in occurring whale species from sei to Bryde's whales. In accordance with an increase of Bryde's whaleamong the balaenopterids toward the tropics, the preferable feed item also changesc.

TABLE 22. FOOD ORGANISMS OF SEI WHALE CAUGHT IN THE
SOUTHERN NORTH PACIFIC WHALING GROUND DURING
MAY-AUGUST 1972 (Kawamura, 1973).
Copepoda
Calanus cristatus
Calanus plumchrus
Calanus pacificus
Euphausiacea
Euphausia recurva
Euphausia pacijica
Euphausia diomedeae
Euphausia tenera
Thysanoessa inermis
Thysanoessa spinifera
Nematoscelis difjicilis
Nematoscelis gracilis ll

Decapoda
Sergestes similis
PiscesScomber japonicus
Sardinops melanosticta
Engraulis japonica
Cololabis saira
Maurolicus muelleri
Pseudopentaceros richardsonii
Cephalopoda
Gonatus sp.2l
Berryteuthis anonychus2l1) Identification is doubtful due to ill conditioned specimens.
2) Identification by T. Kubodera.

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

REVIEW OF FOOD OF BALAENOPTERID WHALES

183

to some extent. The food composition for sei whale over the subarctic to temperate waters between 30°-50°N of approximately 160°E-l 70°W is given in Table
22, and the recent unpublished data by Kawamura on the stomachs of Bryde's
whale in the lower latitudes revealed the followings as newly known organisms thatmust be added to the dietary list of the balaenopterids (mostly Bryde's) in thesouthern North Pacific regions (Kawamura, 1973);
Fish: Vinciguellia nimbaria
Gasterosteus acreatus acreatus
Scomber tapeinocephala
Pleurogrammus azonus
Tarletonbeania taylori
Ranzania laevis
Am phi pods: hyperiids (larvae)
Squids: Berryteuthis anonychus
Gonatus sp. (young)In the waters around Bonin Islands (about 25°N), the stomachs of sei and
Bryde's whales are mainly consisted of fish and euphausiids. From the investigations through February to May during 1947-1949 along with data from anothersources (Mizue, 1951), Nemoto (1959) reported the following food items:
Euphausiids: Euphausia similis
Euphausia recurva
Fish: Yarrella microcephala (Gonostomatidae)
Myctophum asperum (M yctophidae)
In addition to above mentioned food items, Ranzania rypus (Molidae) and twosternophychid fish, Argyropelecus and Polyipnus sp. were found occasionally (Nishimoto et al., 1952). In the pelagic waters, slender mola, Ranzania laevis is largely
found in stead of R. rypus (Kawamura, unpublished data). It is noteworthy that
there occurs no copepod food in the Bonin Island waters although a possible occurrence of copepod such as Candacia sp. as a constituents of whales food has been
pointed out but no evidence have been reported to date.
In Monterey Bay, California, fin whale was found to contain "enormous
quantity of codfish," and the blue whale in the waters off the port of San Quentin
pursued for the school of sardines and prawns (Scammon, 1874). After mentioning the animalculae, the ' right whale food ' or ' brit ' for bowhead whale in thenorthern waters, Scammon (1874) continues about the minke, "when roaming
about the inland waters of lower latitudes, they often shoot along the shallow borders of the bays in search of the myriads of small fry on which they mainly sustain
themselves ". In the eastern Pacific off Mexican coast, Pacific red crab, Pleuroncodes planipes (Galatheidae) is known as food of sei and blue whales (Matthews,
1938a), but the prey changes to Euphausia pacifica, Thysanoessa spinifera and possibly
Calanus plumchrus in the more northern Pacific coast. The fin whale off Vancouver
Island has been reported to feed Euphausia spinifera, herring, and squid, Gonatusfabricii (Tomilin, 1967). Kellog's description by citing Cornwall (1928) made us
confirm the occurrence of G.fabricii in these waters (Kellog, 1929).
Sci. Rep. Whales Res. Inst.,
No. 32, 1980

KAWAMURA184

The balaenopterids in the waters off Peruvian coast are believed to take anchovy, Engraulis ringens and some euphausiids but details are unknown (Gaskin,1976). There seems to exist little information concerning baleen whales food in
the Pacific coast along South America. Clarke and Aguayo (1965) found only
' fluid ' in the first and second stomachs of Bryde's whale taken at Caleta Molle,
lquique on the coast of Chile, but Budylenko (1978) suggested that sei whale feeds
but slightly along the Chilean coast.
As it was described in the South African waters, it is noticed in Table 22 that
there are very distinct similarity in the species composition of whales food, i. e., alargely diversed food composition over the tropical to temperate waters throughout the southern and northern hemispheres.

Coastal waters of Japan and East China Sea regions
The region includes the traditionally famous 'Japan Grounds' (Beale, 1839).
Today, it is called Sanriku region situated in the Pacific side of northern Japan, andis the most important fishing ground for the land based whaling. The informationof whales food, however, is somewhat scarce due to few comprehensive study worked out because the catches in coastal whaling are usually very sporadic. Mizue(1951), however, examined the stomachs of some balaenopterid whales as shown in
Table 23. The krill in this region is represented by Euphausia pacifica along withd.

TABLE 23.

Krill
SardineSaury
MackerelRock fish
Squid
Octopus
Empty

FOOD OF BALEEN WHALES IN THE SANRIKU WHALING GROUND,
OFF PACIFIC COAST OF NORTHERN JAPAN (Mizue, 1951).
Blue

Fin

Sei*

Humpback

16

43

253

2

103

11

41

25
1
2
10
1
547

4

* Sei+Bryde's.
less important two Thysanoessa species while it changes to T. inermis and T. longifJes
in far more northern regions adjacent to Hokkaido. Only Calanus pacificus forms
the main copepod food in the Sanriku region though it may change to Calanus
plumchrus in the more northern grounds. Squids are mostly composed of Ommastrephes sloani pacificus on which sei whale feed extensively. Sardine in Table 23 actually means the anchovy, Engraulis japonica. During April to May in the Sanriku
grounds there occurs large school of young sand lance, Ammotfytes personatus, andthe minke whale pursues for them. In accordance with their northward boundmigration along the Pacific coast of Japan, the minke also follows after them up to
the Hokkaido region. Both Engraulis japonicus and Ammotfytes personatus are the mostSci. Rep. Whales Res. Inst.,
No. 32, 1980

REVIEW OF FOOD OF BALAENOPTERID WHALES

185

important food items of minke whale in the coastal waters of Japan (Omura andSakiura, 1956), but Euphausia pacijica must be added in the waters along southwestern Japan Sea. However, the sei during summer largely feed on young mackerel,
Scomber j aponicus.
TABLE 24.

1955

1956

FOOD OF FIN WHALES IN THE EAST CHINA SEA
AND ITS ENVIRONS (Nemoto, 1959).

E. pacifica
Empty
Unknown
E. pacifica
Flying fishll
Blind eei 2>
Empty
Unknown

July

AugustSeptember

October

6
4

22
65
3
20
1

18
76
10
5

7
14

9
63

25
47

2

3
49

19

I) very little quantity.
2) Eptatretus burgeri.

In the Okhotsk Sea coast of Hokkaido, Nemoto (1959) describes both Euphausia pacifica and Thysanoessa inermis with occasional contamination by T. raschii forfin whales caught off Abashiri, Hokkaido. Later observations, however, revealed
that minke whale feeds on E. pacifica and squid, Ommastorephes sloani pacijicus and fin
whale caught far off shore region fed Thysanoessa inermis, which increases its importance gradually toward Sakhalin regions.
In the southern Japan around Kyushu the Bryde's whale frequents its offshore
waters and feeds Euphausia similis during May but gradually shifts to anchovy,
Engraulisjaponica during the summer season. (Nemoto, 1959).
Not so many are known for the East China Sea regions but the following brief
informations (Nemoto, 1959) (Table 24). The Bryde's whale which perhaps thenorthern counter part of the inshore form found in the South African waters (Omura, 1977), feeds mostly on anchovy, E. japonica and three kinds of horse mackerels, Trachurus japonicus, T. declivis, and T. argenteus though the first two are more
prefered by the whales.
In the adjacent waters of Korean coast, the Yellow Sea and Japan Sea, it has
been known that the fin and minke whales feed on euphausiids with occasional occurrence of squid. The corresponding euphausiid species has been believed to be
Euphausia pacijica. There is single record of the fish, Clupanodon punctatus from the fin
whale in the Yellow Sea. (Nemoto, 1959).
The humpbacks in the Ryukyuan waters seems to feed little but a few records
indicate Euphausia similis and Pseudoeuphausia latifrons to be the possible whales food(Nishiwaki, 1959 ).

Sci. Rep. Whales Res. Inst.,

No. 32, 1980

186

KAWAMURA

Northern Seas-North Atlantic

Northeast Atlantic
According to Gaskin (1976), Allen (1916) earlily reported Thysanoessa inermis
as food of North Atlantic humpbacks. As Hjort and Ruud (1929) reviewed on
the fishing and whaling industries in the North Atlantic, the shelf region aroundthe Norwegian Sea and the Greenland Sea has been exposed to the extensive whaling of the blue, fin, sei, and humpback whales since the beginning of this century.
Norwegian whalers have been aware of the importance of food plankton (=krill)abundance in relation to their catch (Hjort and Ruud, 1929). The krill undoubtedly correspond to one of the largest euphausiid species, Meganyctiphanes norvegica
and several others such as Thysanoessa inermis, T. raschii, and T. longicaudata. Since
there also largely occurs copepods and two amphipod species, Themisto libellula andParathemisto oblivia in addition to euphausiids, the latter crustaceans group may also
· likely to be fed by the baleen whales.
Although Hjort and Ruud (1929) did not mention about the whales food in
the Greenland waters, they showed the following items as having been found in
the whales treated at Aukra Island, off the coast of More, Norwegian Sea during
1925-1928. A total of 619 baleen whales were examined (366 fin, 252 sei and 1blue) and found the enormous amount of plankton food among many animals during January-August.
The food items found were:
1. Herrings (especially ' large ' or spring herrings)
2. Euphausiids (krill)a. " Stor-krill" (large krill) corresponds to Meganyctiphanes norvegicab. " Smaa-krill" (small krill) mostly Tl!Jsanoessa inermis
3. Rod-aate " (copepods) mainly consisted of Calanus finmarchicus
According to Hjort and Ruud (1929), Sars (1874) states that the food of blue
whale in the waters of Finmarken was exclusively comprised of " loddle " or capelin, Mallotus villosus and Thysanopoda ( = Thysanoessa) inermis, while fin whales follow
after spawning shoals of capelin (M. villosus) off the coast of Finmark (Jonsgard,1966). The herring does not seem to be fed by the blue whale but both humpback
and minke take herring largely in this waters. In the Davis Strait, the fin feeds
herring and Thysanoessa inermis during winter (January-March) but it changes tosmaller euphausiids and Calanus finmarchicus in the spring (April-May) and thenshifts to Meganyctiphanes norvegica from May to August (Tomilin, 1967). In addition
to above mentioned food items, followings are the fish food of greater importancein the North Atlantic; cod, mackerel, sand lance, dog fish, whiting, and pollock.
Although the food item may varies with season and whale species, capelin, M. villosus must be an another important whales food in the Finmarken waters.
One of characteristics of the marine distribution in the North Atlantic is theoccurrence of very small copepod called " swamps " (Millais, 1973 ), Temora longicornis as food of baleen whales as well as the food of herrings in the Irelandic waters(Collet, 1886; Millais, 1906; Herdman, 1971), and in the Newfoundland watersa.

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187

REVIEW OF FOOD OF BALAENOPTERID WHALESTABLE 25. STOMACH CONDITIONS OF MINKE WHALE AROUND THE
NORWEGIAN WATERS (constructed from Jonsgard, 1951).
Stomach contents
RegionNo. of abs.

Area

ArcticNorwegianwatersBarentz Sea

Lofoten
Vestraalen
Helgoland

Empty

Fish 1>

94
105

2
333>

14
56

(753>+20)
(9)
(1)
199

(33 3>)(27 3>)

(73>)

35

70

(77)

Krill+ Copepod/Krill +Fish

Milk

78 2>
143>

13)

93

(73>)

(13>)

(16)

1•>

1) Cod, "skrei "2) Krill, Thysanoessa inermis
3) Vestfjord proper4) Calf, 16 feet 10 inches
TABLE 26.

RegionFOOD OF MINKE WHALE IN THE NORWEGIAN COASTAL
WATERS (constructed from Jonsgard, 195!1>)

Area

No.
of
abs.

Herring

Vestfjord

R0st

20

12

Vesteraalen

Outside99Arctic2 >Bear Island

HerCapering+!inothersHaddock

Cod

3+

Spitz berg en+

Barentz Sea

+

+
+

Other
species

Remarks

5

0-group herring"musse"I-group herring"bladsild"especially inautumnLarge herring,small codI-group herring" bladsild " large
haddock

1) Jonsgard notes that "the truth is evidently that minke whales prefer krill when this animal abounds,
but they resort to a great variety of shoaling fishes wherever these are met with."
2) No actual number of animals examined is stated.

where blue whale takes this species (Tomilin, 1967). T. longicornis have never beenfound as the whales food over any another whaling grounds in the world. In addition to crustaceans, there have been reported the pteropod molluscus which is considered to be Clione limacina and/or Limacina helicina. (Hjort and Ruud, 1929;
Peters, 1938).
To see these food items by each whale species, it can be said that the sei inthe coastal banks and slopes region feeds almost exclusively on Calanus .finmarchicus,
while the fin feeds herrings, Calanus .finmarchicus and two euphausiid species above
mentioned. The abundance of migrating whales into the whaling grounds in Moreregion is closely related to the abundance of these food organisms, i. e., there can be
expected richer whale migrations into the whaling ground in the years of moreabundant food distribution. The whales food as mentioned above may changes by
whale species and also by the seasons. The fin whales during January-March,
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KAWAMURA

for instance, feed either herrings or Thysanoessa species, while they feed Calanus finmarchicus and T. inermis in April towards early May. During August, the seasonof" summer fishing" (Hjort and Ruud, 1929) as they call, the fin survives exclusively on Meganyctiphanes norvegica.
According to Christensen (1975) the food items of the North Atlantic minke
whales B. acutorostrata acutorostrata (Rice, 1977) in the Barents Sea and off East
Greenland were largely consisted of capelin, but they changed to sandeels andkrill, Thysanoessa inermis in the waters off West Greenland. One of interesting
evidensces found in the North Atlantic to the Arctic waters is greater variabledietary items of minke whale by season and localitites as shown in Tables 25 and 26.
The minke in these waters, however, feeds generally on herring in spring along
the Norwegian coast, while it feeds mainly on capelin during summer in thenorthern waters along with gadoid fishes such as cod, whiting, and coalfish (Jonsgard, 1951). Although there may exist some confusion regarding the identity offood animals, Jonsgard (1951) gave fourteen possible fish species as the food ofminke whale; herring, sprat, mackerel, cod, coalfish, haddock, ling, pollock, whiting, Norway haddock, catfish, salmon, capelin and sand-eel (Table 27). He (1951)
TABLE 27. POSSIBLE FISH FOOD FOR THE NORTH ATLANTIC MINKE WHALES(Jonsgard, 1951). THE SCIENTIFIC NAME CORRESPONDING TO THE COMMONNAME WAS BASED ON WHEELER (1969), LYTHGOE (1971) and UNESCO (1973)
Common name
SalmonCapelin
Herring
Mackerel
Sand lance
Cod
Coal fish
Haddock
Norway haddock
Pollock
Whiting
Sprat
Ling
Catfish (Wolf fish)
Dog fish

Scientific name

Salmo salar
Mallotus villosus*
Clupea harengus*
Scomber scombrus
Gymnammoqytes cicerellus
Gadus morhua*
Pollachius virens ( = Gadus virens)
Melanogrammus aeglefinus (=G. aeglefinus)
Sebastes marinus
Pollachius pollachius
Merlangius merlangius*
Sprattus sprattus*
Molva molva
Anarhichas lupus*
S!J1liorhinus caniculus* UNESCO (1973) recommends trinominal nomenclature for these fishes.
also :suggested the occurrence of copepods (Calanus), krill (Euphausiacea), small
shrimp or ' aate ', and even mature cod ' skrei ' and big gadoid fishes. Gravel andstones up to the size of an egg are occasionally found (Jonsgard, 1951).
In the region of southern slopes of the Norwegian Sea-the Norway-Shetland-Faroe Island area-three euphausiids, M. norvegica, T. inermis and T. longicaudata occur with larger quantity. Although Hjort and Ruud (1929) did not show
the whales food in this region, it may be supposed that the fin whale occurring off
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REVIEW OF FOOD OF BALAENOPTERID WHALES

189

Faroe Island feeds above three euphausiids. The similar dietary situation could be
supposed to the northern slopes of the Norwegian waters.
In the Icelandic waters there has been reported an appreciable number of theblue, fin, sei and humpback whales come to concentrate and feed. The coincidence of their concentration with the pronounced abundance of krill distribution
suggests the possible whales food to be Thysanoessa inermis and Meganyctiphanes norvegica. Reviewing the fin whale fishery off the West Coast of Iceland, R0rvik et al.
(1976) described, "In all the investigated years, 1925-1928, the fin whales were
eating herring, small krill ( Thysanoessa inermis) or copepods, mainly Ca/anus finmarchicus, from the middle ofJanuary to the early part of May ". From May to the middle of August, " .... the fin were feeding exclusively on the large krill, Meganyctiphanes norvegica ". However, the greater part of the fin whales food seems to change
to capelin, M. villosus toward the eastern Greenland. (Jonsgard and Christensen,
1968).
Towards south closer to the temperate North Atlantic there seems to increasein the variety of fish food among many others, especially the fish belonging togadoids, clupeids and scombroids and they may become important particularly
m the coastal waters (Gaskin, 1976).

b.

Northwest Atlantic
From temperate to subarctic waters along the eastern coast of North America
there exists one of two places where the large number of the balaenopterid whales
come to concentrate and feed during the warmer season. Composition of the whales
food in this region is generally comprised of similar animal communities to thatknown in the northeastern Atlantic waters, where copepods, euphausiids, and gregarious fish predominate in the whales stomachs. In the region of Bay of Fundy,
copepods, euphausiids, herring and mackerel are said to begin rise into the surface
during the summer, and the fin whale being attracted to those possible prey organisms begin to gather in the region (Gaskin, 1976). Although the diet of baleen
whales may generally go similar way over the whole North Atlantic, yet there seems to exist qualitative differences from localities and seasons.
The baleen whales food as revealed by the Canadian workers gives a comprehensive knowledge about whales food (Mitchell, 1974). By examining both fin andsei whales at Blandford, Nova Scotia, and sei at Dildo and Williamsport, Newfoundland between 1965 and 1972, Mitchell (1974) found the following food species: krill, copepods, sand lance, mackerel, herring, squid and ' fish ' for the Nova
Scotian animals, whilst the food in Newfoundland animals was composed of capelin, sand lance, lantern fish, krill, white fish, and cod. Although Mitchell. (1974)
did not give the scientific name for copepods and krill, they were perhaps comprised of Thysanoessa inermis, Meganyctiphanes norvegica, Ca/anus finmarchicus, and Temora
longicornis. One of distinct differences existing between two locations are the complete lack of capelin from the diet in the Nova Scotian animals. The capelin comprises the main diet of fin whales of mostly 80-100% through May to Novemberin the Newfoundland. On the other hand, the same whale species in the Nova
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KAWAMURA

Scotian waters feed largely on krill possibly be comprised of M. norvegica and several Thysanoessa euphausiids through May to November in more than 80% ofanimals, but the sei alone feeds largely on copepods (60-80%) between June andOctober. Brodie et al. (1978) reports that of 67 stomachs of fin whale taken in theNova Scotian waters 63 were full of krill, Meganyctiphanes norvegica, three contained
copepods and one was full of young herring. The importance of krill, M. norvegica
as food of fin whale is clear in the northwestern Atlantic region. To see the whales
food as shown in the two above localities, it is observed that both krill and copepods
are the most important food items in the Nova Scotian waters. Both sand lance
and mackerel are also fed by the fin whale but none of them by the sei whale.

Fin&

Bryde'sFig. 7. Schematic food web showing predation by "swallowing" and "skimming" baleen whales on main food sources (Mitchell, 1974).

Lantern fish and white fish may come to the next importance in the Newfoundland
especially in October through November. The humpback whales taken at Nova
Scotia (Blandford) and Newfoundland (Williams port and Dildo) were found to feed
capelin, sand lance, white fish, mackerel and krill (Mitchell, 1973). Of those prey
items, both capelin and krill are considered to be the main diet of humpbacks in
the Canadian Atlantic waters. Generally, it can be considered that there exists a
considerably different feeding grounds in the northwestern Atlantic region whereas
its expansion is spacially very limited. One of such feeding grounds may largely
depends upon the planktonic crustaceans in the formation of feeding ground while
the another one depends upon the small to medium sized gregarious fish species(Fig. 7).
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Summerizing addendum
In summerizing the review, it may be said that although the main subject of
this article was confined to deal with the diet of" balaenopterid whales ", the foodof the balaenids-right, bowhead, and pygmy right whales as stenophagous animal
is generally consisted of very similar food items with that found in the balaenopterids while the eschrichtiid whale alone shows quite different aspects from the above
two taxonomical groups. As mentioned in the morphological characteristics of thebaleen filter, the balaenid whales obviously perform "skimming" and feed preferably on smaller, weak powered swimmers of crustaceans such as copepods even insuch waters of rich euphausiid distributions as known in the the southern oceans,
where an another problem-competion for food between southern right and sei
whales may arise. (Kawamura, 1978b).
The bowhead, however, by their extraordinal biological, ecological, and zoogeographical distinctions, feeds rather complicated prey organisms. In his marverous book, "An Account of the Arctic Regions'', Scoresby (1820) demonstrated a fantastic plate showing the food organisms of bowhead whales, in which he introduced;
actiniae, cliones, sepiae, medusae, cancri and helices. He also added squllae or shrimp
from the different observations. To see Scoresby's plate XVI (Scoresby, 1820),
several additional food organisms may be observed; that is, two kinds of chaetognaths and a kind of both planktonic tunicates possibly belong to Oikopleura and gammarid amphipods. Although it is difficult to interprete each organisms from the
Scoresby's descriptions, Hjort and Ruud (1929) identified some groups of whales
food being based on the figures in the Scoresby's plate and found:1. The four ctenophores: Beroe cucumis, Pleurobrachia pileus, Mertensia ovum,
and Bolina septentrionalis
2. The two pteropods: Clione limacina and Limacia helicina
3. The two or three crustaceans: Hymenodora glacialis, Calanus hyperboreus,and possibly Themisto libellula
These organisms, however, seem to be unlikely as the staple food of bowhead,
and it was concluded later that their main food in the Atlantic Arctic region must
be a copepod, Calanus .finmarchicus as illustrated very likely in the Scoresby's plate(Ruud, 1937). Euphausiids may also be included in the diet of Atlantic bowead
whale. In the Alaskan waters the bowhead feeds copepods, Calanus glacialis andC. hyperboreus during their Bering Sea/Chukchi Sea migrations although the benthic
gammarids are likely to be taken supplementary. Very recently, Lowry et al.
(1978) examined two female bowhead whales caught in the vicinity of Point Barrow, Alaska. The stomach contents analysis revealed that the Alaskan bowhead
feeds mainly on Tfrysanoessa raschii, gammarid amphipods, Gammarus zaddachi, Acanthostepheia behringiensis, Monoculoides zernovi, and Rozinante fragilis, and hyperiid amphipod, Parathemisto libellula. Of those above mentioned food species, T. raschii
made up 90.3% of the total volume. Lowry et al. (1978) also found an unidentifiable carapace of shrimp and even a small pebble in the stomach. According toSergeant and Hoek (1974a), however, "nothing is yet known of the nature of thefood of bowheads " in the Franklin Bay region, Canadian Arctic, but they (SerSci. Rep. Whales Res. Inst.,
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KAWAMURA

geant and Hoek, 197 4b) suggested that Calanus hyperboreus, Parathemisto libellula,
Thysanoessa inermis and Th. raschii are the most possible food organisms of bowheads
in the Cape Lambton and Cape Parry regions in the Beaufort Sea.
Scoresby (1820) writes the way of feeding by the bowhead whale, the largest
skimmers ever existed as follows: " When the whale feeds, it swims with considerable velocity below the surface of the sea, with its jaws widely extended. A
stream of water consequently enters its capacious mouth, and along with it, large
quantities of water insects; the water escapes again at the sides; but the whalebone,which, from its compact arrangement, and the thick internal covering of hair, does
not allow a particle the size of the smallest grain to escape ".
Right and pygmy right whales are the dwellers of the subarctic and/or subantarctic regions where the most highest marine production is usually expected.
There have been reported that these animals as microplanktonophagi feed exclusively on copepods which predominate in theirown habitat: Calanus .finmarchicus,
Thysanoessa inermis and pteropods but lesser extent in the North Atlantic waters,
but this assemblage changes to Calanus plumchrus (Matuura and Maeda, 1942), C.
cristatus, and possibly Metridia sp. in the North Pacific and Okhotsk Sea. They
may also feed Calanus paci:ficus and Euphausia pacifica in the more southen waters from
northern Japan to the boreal regions of the North Pacific (Omura, 1958). The
southern counterpart, Eubalaena glacialis australis (=E. australis) feeds both euphausiids and copepods in the subantarctic waters. Their main prey animal, however,must be Calanus tonsus, Calanus simillimus and Euphausia vallentini as it is supposed by
the occupation ofthier main feeding (=whaling) grounds during the austral summer(Townsend, 1935; Kawamura, 1978b). There also have been reported that Euphausia superba is taken in the Antarctic proper, whereas the food species change to
Grimothea larva of Munida gregaria in the waters off Patagonian coast (Matthews,
1932; 1938b).
Very little is known for the pygmy right whale. It has been reported thatthe animal as their finer baleen fringes suggest was found containing several numberof Calanus species in the South Atlantic waters (Ivashin et al., 1972).
The composition of baleen whales food in the North Atlantic waters as has been reviewed in this article seems to be somewhat monotonous assemblages when it
was compared with that found in the North Pacific and its environs. It may be un-:likely to consider these differences simply due to the faunistic characteristics in thecommunity composition of food animals, but it was the result possibly due to thelimited material sources from the less exploitation of whale stocks in the pelagic
waters using the whaling factories in the North Atlantic regions. There is no reason therefore, to believe the greater difference in the biological environments concerning the diet of baleen whales between the North Pacific and the North Atlantic oceans.
The dietary habits of great whales as well as their general biology have long
been a mythological subject but the modern whaling accompanied with a extensive biological works gradually unvailed the mystery of whales one after anotherduring the past several decades and, perhaps some more will be found but gradually
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REVIEW OF FOOD OF BALAENOPTERID WHALES

193

in the next.
ACKNOWLEDGMENTS

The publish of this contribution became possible sorely by the courtesy of Dr Hideo
Omura, Director of the Whales Research Institute, Tokyo, and his kind offering
the space in this volume is greatly acknowledged. Several tables and figures as indicated in the text were able to be reproduced by the courtesy of Drs P. Best, E.
Mitchell, T. Nemoto, and Fishing News Books Ltd. Without their welltimed cooperation the article was not completed in a manner intended, and all these are
greatly acknowledged. Although I tried to get reproducing permit from Dr M. M.
Sleptzov, it was unable to obtain any response, and some of his tables were usedwithout permit.
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