NOAA Technical Report NMFS 83

Cephalopods from the

Stomachs of Sperm Whales

taken off California

Clifford H. Fiscus

Dale W. Rice

Allen A. Wolman

u.s. Department of Commerce

December 1989
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48. Widow rockfish: Proceedings of a workshop, Tiburon, California, December

11-12,1980, by William H. Lenarz and Donald R. Gunderson (editors). January 1987,

57 p.

49. Reproduction, movements, and population dynamics of the southern kingfish,

Menlieirrhus americanus, in the northwestern Gulf of Mexico, by Stephen M. Harding

and Mark E. Chittenden, Jr. March 1987,21 p.

50. Preparation of acetate peels of valves from the ocean quahog, Arclica islandica,

for age determinations, by John W. Ropes. March 1987, 5 p.

51. Status, biology, and ecology of fur seals: Proceedings of an international

workshop, Cambridge, England, 23-27 April 1984, by John P. Croxall and Roger

L. Gentry (editors). June 1987,212 p.

52. Limited access alternatives for the Pacific groundfish fishery, by Daniel D.

Huppert (editor). May 1987, 45 p.

53. Ecology of east Florida sea turtles: Proceedings of the Cape Canaveral, Florida,

sea turtle workshop, Miami, Florida, February 26-27, 1985, by Wayne N. Witzell

(convener and editor). May 1987, 80 p.

54. Proximate and fatty acid composition of 40 southeastern U.S. finfish species,

by Janet A. Gooch, Malcolm B. Hale, Thomas Brown, Jr., James C. Bonnet, Cheryl

G. Brand, and Lloyd W. Reiger. June 1987, 23 p.

55. Proximate composition, energy, fatty acid, sodium, and cholesterol contcnt of

finfish, shellfish, and their products, by Judith Krzynowek and Jenny Murphy. July

1987, 53 p.

56. Some aspects of the ecology of the leatherback turtle Dermoehelys eoriaeea at

Laguna Jolova, Costa Rica, by Harold F. Hirth and Larry H. Ogren. July 1987, 14 p.

57. Food habits and dietary variability of pelagic nekton off Oregon and Washington,

1979-1984, by Richard D. Brodeur, Harriet V. Lorz, and William G. Pearcy. July

1987, 32 p.

58. Stock assessment ofthe Gulf menhaden, Brevoonia patronus, fishery, by Douglas

S. Vaughan. September 1987, 18 p.

59. Atlantic menhaden, Brevoonia tyrallnus, purse seine fishery, 1972-84, with a

brief discussion of age and size composition of the landings, by Joseph W. Smith,

William R. Nicholson, Douglas S. Vaughan, Donnie L. Dudley, and Ethel A. Hall.

September 1987, 23 p.

60. Gulf menhaden, Brevooniapatronus, purse seine fishery, 1974-85, with a brief

discussion of age and size composition of the landings, by Joseph W. Smith, Eldon

J. Levi, Douglas S. Vaughan, and Ethen A. Hall. December 1987, 8 p.

61. Manual for starch gel electrophoresis: A method for the detection of genetic variation, by Paul B. Aebersold, Gary A. Winans, David J. Teel, George B. Milner, and

Fred M. Utter. December \987, 19 p.

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Cynthia S. Martin, Shelley E. Arenas, Jacki A. Guffey, and Joni M. Packard.

December 1987, 149 p.

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Douglas S. Vaughan and Joseph W. Smith. January 1988, 18 p.

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Farfante. April 1988, 32 p.

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Mitchell. March 1988, 28 p.

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Pacific, by M. James Allen and Gary B. Smith. April 1988, 151 p.

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application to the Pacific coast trawl fleet, by Dale Squires. July 1988, 34 p.

68. Annotated bibliography II of the hard clam Mereenaria mereenaria, by J.L.

McHugh and Marjorie W. Sumner. September 1988, 59 p.

69. Environmental quality and aquaculture systems: Proceedings of the thirteenth

U.S.-Japan meeting on aquaculture, Mie, Japan, October 24-25, 1984, edited by Carl

J. Sindermann. October 1988, 50 p.

70. New and innovative advances in biology/engineering with potential for use in

aquaculture: Proceedings of the fourteenth U.S.-Japan meeting on aquaculture, Woods

Hole, Massachusetts, October 16-17, 1985, edited by Albert K. Sparks. November

1988, 69 p.

71. Greenland turbot Reinhardtius hippoglossoides of the eastern Bering Sea and

Aleutian Islands region, by Miles S. Alton, Richard G. Bakkala, Gary E. Walters,

and Peter T. Munro. December 1988,31 p.

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and Louise M. Dery. December 1988, 135 p.

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by Michael Vecchione, Clyde F.E. Roper, and Michael J. Sweeney. February 1989,

23 p.

74. Proximate composition and fatty acid and cholesterol content of 22 species of

northwest Atlantic finfish, by Judith Krzynowek, Jenny Murphy, Richard S. Maney,

and Laurie J. Panunzio. May 1989,35 p.

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Jeffrey Stein, Kevin T. Herbinson, Robert H. Moore, Michael Mullin, and John S.

Stephens, Jr. April 1989, 16 p.

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78. Survey of fishes and water properties of south San Francisco Bay, California,

1973-82, by Donald E. Pearson. August 1989, 21 p.

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habitat off the southeastern United States, by Charles A. Wenner and George R.

Sedberry. July 1989, 49 p.

80. Laboratory guide to early life history stages of northeast Pacific fishes, by Ann

C. Matarese, Arthur W. Kendall, Jr., Deborah M. Blood, and Beverly M. Vinter.

October 1989,651 p.
NOAA Technical Report NMFS 83

Cephalopods from the

Stomachs of Sperm Whales

taken off California

Clifford H. Fiscus

Dale W. Rice

Allen A. Wolman

December 1989

U.S. DEPARTMENT OF COMMERCE

Robert Mosbacher, Secretary

National Oceanic and Atmospheric Administration

John A. Knauss, Under Secretary for Oceans and Atmosphere

National Marine Fisheries Service

James Brennan, Assistant Administrator for Fisheries
COVER: Drawing of sperm whale by Nancy Williams-Nelson.

The National Marine Fisheries Service (NMFS) does not approve, recommend or endorse any proprietary product or proprietary material mentioned

in this publication. No reference shall be made to NMFS, or to this publication furnished by NMFS, in any advertising or sales promotion which would

indicate or imply that NMFS approves, recommends or endorses any proprietary product or proprietary material mentioned berein, or which has as

its purpose an intent to cause directly or indirectly the advertised product to

be used or purchased because of this NMFS publication.

ii
CONTENTS

Acknowledgments iv

Introduction 1

Materials and Methods 2

Species Accounts 2

Loliginidae 2

Loligo opalescens Berry, 1911 2

Enoploteuthidae 2

Abraliopsisfelis McGowan and Okutani 1968 2

Octopoteuthidae 4

Octopoteuthis deletron Young, 1972 4

Taningia danae Joubin, 1931 4

Onychoteuthidae 4

Onychoteuthis borealijaponica Okada, 1927 4

Moroteuthis robusta (Verrill, 1876) 4

Gonatidae 5

Gonatus spp. 5

Gonatus pyros Young, 1972 5

Gonatus berryi Naef, 1923 5

Gonatus sp.

(cf. G. fabricii [Lichtenstein, 1818]) 5

Gonatopsis borealis Sasaki, 1923 5

Unidentifiable Gonatids 6

Architeuthidae 6

Architeuthis japonica Pfeffer, 1912 6

Histioteuthidae 6

Histioteuthis spp. 6

Histioteuthis dofleini (Pfeffer, 1912) 7

Histioteuthis heteropsis (Berry, 1913) 7

Ommastrephidae 7

Dosidicus gigas (d'Orbigny, 1835) 7

Unidentifiable Ommastrephid 7

Chiroteuthidae 7

Chiroteuthis calyx Young, 1972 7

Mastigoteuthidae 7

Mastigoteuthis spp. 7

Cranchiidae 8

Taonius pavo (Lesueur, 1821) 8

Megalocranchia spp. 8

Galiteuthis spp. 8

Galiteuthis phyllura Berry, 1911 8

Galiteuthis pacifica Robson, 1948 8

Mesonychoteuthis hamiltoni Robson, 1925 8

Unidentifiable Cranchiids 9

iii

Vampyroteuthidae 9

Vampyroteuthis infernalis Chun, 1903 9

Octopodidae 9

Octopus dofleini (Wulker, 1910) 9

Alloposidae 9

Alloposus moWs Verrill, 1880 9

Unidentifiable Beaks 9

Discussion 10

Citations 10
Acknowledgments _

We wish to thank the following authorities for their identification of whole

cephalopods or cephalopod beaks: M.R. Clarke, F.G. Hochberg, T.K.

Kristiansen, T. Kubodera, and C.F.E. Roper. Clarke, Hochberg, and Roper

also reviewed a draft of this manuscript and provided many helpful suggestions and comments.

iv
Cephalopods from the

Stomachs of Sperm Whales

taken off California

CLIFFORD H. FISCUS

DALE W. RICE

ALLEN A. WOLMAN

National Marine Mammal Laboratory

National Marine Fisheries Service, NOAA

7600 Sand Point Way N. E., Bldg. 4

Seattle, Washington 98115-0070

ABSTRACT

Cephalopod remains (beaks, bodies, and parts of bodies) were

collected from the stomachs of 157 sperm whales (Physeter

macrocephalus) taken off central California (lat. 37°-39°N). At

least 24 species representing 14 families were identified. Frequencies of occurrence of the six most numerous taxa were

Moroteuthis robusta 72.0%, Gonatopsis borealis 66.2%, Histioteuthis dofleini 36.9%, Galiteuthis spp. (including G. phyllura

and G. pacifica) 36.3%, Octopoteuthis deletron 35.0%, and Vampyroteuthis infernalis 27.4%. One find of two Mesonychoteuthis

hamiltoni beaks strongly suggests transequatorial migration by

one large male sperm whale.

Introduction _

Sperm whales (Physeter macrocephalus) feed mainly on

medium to large cephalopods, which they apparently capture at depths ranging from several hundred meters to perhaps

as great as 3000 meters (Rice 1989). The stomachs of sperm

whales thus provide samples of a portion of the cephalopod

fauna that can seldom be sampled by other methods. During

the course of research on the life history and ecology of the

great whales in the eastern North Pacific, Rice and Wolman

(1970) examined 637 sperm whales taken in commercial

shore-based whaling operations off central California (lat.

37°-39°N) during 1959-70. Rice (1963a,b) described the

whaling operations and area off central California and illustrated the bathymetry of the ocean floor.

Initial examination ofthe cephalopod specimens collected

from the stomachs of sperm whales began in the early 1960s.

Rice (1963b) listed Moroteuthis robusta, Gonatopsis borealis, Onychoteuthis sp., Octopus sp., and unidentifiable

squids. Rice and Wolman (1970) added an additional seven

genera to the published record. Fiscus and Rice (1974)

published a note on the finding of Architeuthis sp. in sperm

whale stomachs. Fiscus completed the initial examination of

the material in 1972, and a list of cephalopod taxa was

prepared but not published.

During this period, selected material was sent to other

cephalopod specialists for identification or confirmation of

identification. These specialists are acknowledged in the individual species accounts. Fiscus took samples of squid beaks

to the workshop on identification of cephalopod beaks held

1-12 June 1981 at the laboratory of the Marine Biological

Association of the United Kingdom at Plymouth, England

(Clarke 1986a). As a result of this workshop and discussions

with the participants, most of the remaining unidentified

beaks were identified. The entire collection was reexamined

by Fiscus between 1983 and 1988; the results ofthis examination are presented in Table 1 and in the following species

accounts.

In this report, we present our data on taxonomy and distribution of the taxa of cephalopods found in sperm whale

stomachs. An analysis of the sperm whale's prey-which

includes fishes and crustaceans as well as cephalopods-in

relation to feeding ecology, sex, age, and social behavior

of the whales, will be included in a forthcoming report by

Rice and Wolman on the biology of the sperm whale in the

eastern North Pacific.
Materials and Methods _

Sperm whales were landed at the whaling stations of the Del

Monte Fishing Company and the Golden Gate Fishing Company at Point San Pablo, Richmond, California. Whales were

taken during all months except January, but the majority were

taken May to November. Contents of the stomachs of 483

sperm whales were examined at least cursorily; however,

because ofthe rapidity of the flensing operations and the need

to gather other biological data, we were able to collect

samples of cephalopod beaks from only 157 stomachs. All

stomachs contained at least a few cephalopod beaks. The

number of beaks collected from each stomach ranged from

1 to 57, and the total number of beaks was 2060 (not including beaks from whole specimens, crowns, and buccal

masses). All of the remains collected represent a minimum

of 1700 individual cephalopods.

These samples of whale stomach contents were fixed in

10% formalin or 70% ethanol or isopropanol, and brought

back to the Marine Mammal Biological Laboratory (now the

National Marine Mammal Laboratory, NMML) in Seattle.

Fiscus began the examination of cephalopod material in the

early 1960s. The samples were mostly represented by beaks,

but fortunately some relatively intact specimens or crowns

(heads with arms and tentacles) were included in the samples.

After the whole bodies and heads were identified, the beaks

were removed and compared with other beaks in the samples.

Papers by Akimushkin (1965), Berry (1912), Clarke (1966),

Pearcy (1965), Phillips (1933, 1961), and Verrill (1882) were

most useful during initial stages of the study. Papers by

Okutani and McGowan (1969) and Young (1972) were important sources of information on species of cephalopods

present in the California Current system. Anderson (1978)

and Talmadge (1967) listed species of cephalopods from

Monterey Bay and northern California. Jefferts (1983, 1988)

provided important new information on cephalopod distribution in the northeastern Pacific Ocean.

Cephalopod beaks were identified by comparison with

vouchered beaks in the NMML collection and through the

use of beak descriptions and illustrations (Clarke 1986a, Iverson and Pinkas 1971, Wolff 1984). Lower beaks are most

useful in determining species of cephalopods; however, some

upper beaks are characteristic of certain species or genera,

and were used when possible.

2

Species Accounts _

Classification and sequence offamilies and genera of cephalopods follows Clarke and Trueman (1988), who updated the

list by Voss (1977). We identified 24 species representing

14 families of squids and octopods from the 157 stomach

samples examined. For each, we have calculated frequency

of occurrence and number of individuals (Table 1).

The following abbreviations are used throughout: GL = gladius length; LRL = lower rostral length; ML = dorsal

mantle length; NMML = National Marine Mammal

Laboratory.

Loliginidae

Loligo opalescens Berry, 1911 The occurrence of a single

upper beak in the stomach of a sperm whale almost certainly

represents a secondary occurrence, i.e., it was initially consumed by a larger cephalopod or fish which was in tum eaten

by the whale. Loligo opalescens is a neritic species that occurs off central California throughout the year (Fields 1965).

It is a schooling species, and we have found that it usually

constitutes the bulk of the contents when it occurs in marine

mammal stomachs. Hixon (1983) provided a detailed life

history of this species, which moves to_shallow inshore

waters to spawn. In the commercial catch in Monterey Bay,

California, the average ML of mature males was 150 mm

and of mature females 140 mm. Large mature specimens in

the NMML collection are in the size range 180-206 ML.

This species is of major importance as prey to several species

ofshallow diving marine mammals found off central California, such as the northern fur seal (Cailorhinus ursinus),

Pacific white-sided dolphin (Lagenorhynchus obliquidens),

and Dall's porpoise (Phocoenoides daili) (Stroud et al. 1981,

Kajimura 1984, Clarke 1986b); however, L. opalescens is

not usually found at the depths at which sperm whales feed.

Enoploteuthidae

Abraliopsis felis McGowan and Okutani 1968 The presence of a single upper beak of this species in the stomach

of a sperm whale must certainly represent a secondary occurrence. Abraliopsisfelis is a schooling species, and we have

noted that it is usually a major component of the contents

when it occurs in marine mammal stomachs. This species

is found in the waters ofthe California Current system, where

it is mesopelagic to epipelagic (Jefferts 1983). Mature

specimens may measure 60 mm ML. The species forms a

regular part of the diet of the shallow diving marine mammals (Kajimura 1984, Stroud et al. 1981).
Table 1

Frequency of occurrence and number of individuals of cephalopods in the stomachs of sperm whales collected off central California.

Frequency of occurrence Number of individuals'

FamilyZ Species Family Species

Species N % N % N % N %

Teuthida

Loliginidae 0.6 0.07

£Oligo opalescens 3 0.6 0.07

Enoploteuthidae 0.6 0.07

Abraliopsis felis l 0.6 0.07

Octopoteuthidae 56 35.7 125 9.33

Octopoteuthis deletron 55 35.0 124 9.25

Taningia danae 1 0.6 I 0.07

Onychoteuthidae 119 75.8 334 24.93

Onychoteuthis borealijaponica 15 9.6 22 1.64

Moroteuthis robusta 113 72.0 312 23.28

Gonatidae 108 68.8 421 31.42

Gonatus spp. 5 3.2 5 0.37

Gonatus pyros 3 1.9 5 0.37

Gonatus berryi 16 10.2 28 2.09

Gonatus sp. (cf. G. fabricii) 19 12.1 25 1.87

Gonatopsis borealis 104 66.2 352 26.27

Unidentifiable gonatids 4 2.5 6 0.45

Architeuthidae II 7.0 II 0.82

Architeuthis japonica 11 7.0 II 0.82

Histioteuthidae 73 46.5 207 15.45

Histioteuthis spp. 13 8.3 34 2.54

Histioteuthis dojleini 58 36.9 164 12.24

Histioteuthis heteropsis 6 3.8 9 0.67

Ommastrephidae 5 3.2 7 0.52

Dosidicus gigas 4 2.5 4 0.30

Unidentifiable ommastrephids 1 0.6 3 0.22

Chiroteuthidae 5 3.2 6 0.45

Chiroteuthis calyx 5 3.2 6 0.45

Mastigoteuthidae 3 1.9 3 0.22

Mastigoteuthis spp. 3 1.9 3 0.22

Cranchiidae 68 43.3 133 9.93

Taonius pavo 2 1.0 2 0.15

Megalocranchia spp. 8 5.1 9 0.67

Galiteuthis spp. 39 24.8 72 5.37

Galiteuthis phyllura 12 7.6 18 1.34

Galiteuthis pacifica II 7.0 15 1.12

Mesonychoteuthis hamiltoni I 0.6 2 0.15

Unidentifiable cranchiids 9 5.7 15 1.12

Vampyromorpha

Vampyroteuthidae 43 27.4 62 4.63

Vampyroteuthis infernalis 43 27.4 62 4.63

Octopoda

Octopodidae II 7.0 26 1.94

Octopus dojleini II 7.0 26 1.94

Alloposidae 2 1.3 3 0.22

Alloposus mol/is 2 1.3 3 0.22

Total identified individuals' 1340

Unidentifiable

Histioteuthidae or Cranchiidae 67 42.7 318

Megalocranchia or Taningia 4 2.5 7

Cephalopoda 19 12.1 35

Total unidentified individuals 360

Total individuals 1700

Total stomachs 157

IMinimum number necessary to account for the remains from each stomach; for example, 3 upper beaks and 2 lower beaks are counted as 3 individuals.

2Family totals may be less than sum of species totals because some stomachs contained more than one species

3probably secondary food item from stomach of sperm whale's prey species.

'Identified at least to family.

3
Figure 1

Lower rostral length-frequencies of 59 Moroteuthis robusta beaks from

the stomachs of sperm whales taken off California.

Moroteuthis robusta (Verrill, 1876) This large squid occurred more frequently (72 %) than any other species in our

survey. Most samples contained beaks of 1-7 squids; however, one sample contained beaks from 12 and another from

26 individuals. The characteristic cartilaginous end-cones of

the gladii of this species are a conspicuous component of the

stomach contents of most California sperm whales. Two

whole squids were collected from sperm whale stomachs,

one measuring 363 mm ML and the other C\J 1000 mm ML.

Rice has examined (but could not preserve) many whole

specimens measuring up to 1300 mm ML. Several crow~s

(head with arms and buccal mass) are also represented In

the collection, as well as fragments of gladii and cartilaginous

end-cones of gladii. LRLs of 59 beaks were in the size range

5-14 mm (x 10.1 mm) (Fig. 1). The same LRL range was

found by Clarke and MacLeod (1980) in beaks they examined

from the stomachs of sperm whales taken off western

Canada. Roper and Young (1975), commenting on the depth

occurrence of this species, stated that the upper limit of its

distribution appears to be C\J 100 m. Most are taken in commercial bottom trawls at depths much deeper than this.

Hochberg (1974), in summarizing capture records, stated that

Moroteuthis lives near the bottom at depths of 200-600 m;

he published records ofsix specimens taken by bottom trawl

in the Santa Barbara channel at depths of 200-300 m. Jefferts

(1983) reports a maximum depth of capture of M. robusta

of 514 m. The species is found in slope waters from southern

California northward around the Pacific rim to Japan. It has

been reported from the stomachs of sperm whales from central California (Rice 1963b), western Canada (Clarke and

MacLeod 1980), the Gulf of Alaska and Aleutian Islands

(Okutani and Nemoto 1964), the western and central Aleutian Islands (Kodolov 1972), the Kuril Islands (Betesheva

and Akimushkin 1955), and Japanese waters (Kawakami

Octopoteuthidae

Octopoteuthis deletron Young, 1972 This species is represented mostly by beaks of 1-5 specimens per stomach;

however, beaks from 12 specimens were found in one

stomach. No flesh remains were found. LRL of 13 beaks

measured 6.0-9.5 mm (x 7.1 mm). The species is mesopelagic to epipelagic in habit (Jefferts 1983, Roper and Young

1975) and is endemic to the California Current system.

Clarke (1986a) stated that Octopoteuthis spp. are important

in the diet of sperm whales in all but the coldest waters of

the world.

Taningia danae Joubin, 1931 We found one lower beak

with an LRL of 12.0 mm. No wing darkening was apparent

even at this size. To our knowledge, no whole specimens

of T danae have been collected from the California Current system. The only published record for California is that

ofCondit and Le Boeuf (1984) who reported this species from

the stomach of one elephant seal (Mirounga angustirostris);

although not stated in their paper, M.R. Clarke (Mar. Biol.

Stn. U.K., Plymouth, England) identified their specimen.

Roper and Young (1975) stated that little is known of the

depth range of this species; however, they cited one record

from the stomach of a bottom-dwelling shark taken on hookand-line at 1246 m. Clarke (1986a) indicated that this species

is important in the diet of sperm whales in the Atlantic from

lat. 400 S to 53°N and in the Pacific from lat. 40 0 S to 37°N.

This species has been taken from the stomachs of sperm

whales captured off the coast of Honshu, Japan (Okutani

et al. 1976, Okutani and Satake 1978). Okutani (1974)

listed 16 specimens taken in micronekton tows during the

EASTROPAC expeditions from the eastern tropical Pacific.

Onychoteuthidae

Onychoteuthis borealijaponica Okada, 1927 This species

is represented by beaks from 1-3 squids per whale stomach.

No fleshy parts were present in the samples, but gladii were

identified with GL in the size range 303-320 mm. LRLs of

three beaks measured 3.0, 5.5, and 6.0 mm. The species

is abundant in slope waters off central California, as indicated

by its frequent occurrence in marine mammal stomachs

(Kajimura 1984, Stroud et al. 1981), and ranges across the

North Pacific from Baja California north and west to Japanese

waters. It is mesopelagic to epipelagic, and at least part of

the population migrates to the surface at night (Jefferts 1?8?).

It forms an important part of the diet of the shallow dIVing

marine mammals (Stroud et al. 1981, Kajimura 1984), and

also is taken by the deep diving elephant seal (Condit and

Le Boeuf 1984, Antonelis et al. 1987). Mead et al. (1982)

found this species in four of five stomachs of archbeak whales

(Mesoplodon carlhubbsi) that stranded on the southern

California coast.

4

16

14

>=' 12

z

w

<..> 10 '"

w

>- 8 <..>

z:

w

:J 6 0

W

'"

"- 4

:11 II

S 6 7 8 9 10 11 12

LOWER ROSTRAL LENGTH (MM)

1I

13 14
1980). Antonelis et aI. (1987) and Condit and Le Bouef

(1984) reported this species from the stomachs of elephant

seals from California waters.

Gonatidae

Members of this large family are found in the subarctic and

transitional waters of the North Pacific, the Bering and

Chukchi seas, the Arctic Ocean, the North Atlantic, and in

Antarctic waters. In the subarctic Pacific the family is

represented by 4 genera and 19 nominal species (Okutani

etal. 1988). Three ofthe four genera are present in the waters

off central California: the genus Gonatus is represented by

at least five species, Gonatopsis by one, and Berryteuthis by

one or possibly two species.

Gonatus spp. Four occurrences represented by single

beaks, and one by a whole specimen, are recognizable as

Gonatus, but presently cannot be identified to species.

Gonatus pyros Young, 1972 Two occurrences are represented by one lower beak each, and one by three lower beaks;

LRLs are 4.5 mm (four specimens) and 5.0 mm (one specimen). Gonatus pyros has been taken in demersal tows as deep

as 440 m, and in midwater opening-closing tows at depths

of750-790 m and 500-1000 m (Jefferts 1983). This species

is found in the waters of the California Current system and

northwestward to about 175°E in the Alaska Current.

Gonatus berryi Naef, 1923 Each of our samples consists

of 1-5 lower beaks. LRLs of five of the largest specimens

were in the size range 5.0-5.8 mm. Jefferts (1983) reported

G. berryi from opening-closing midwater trawls as deep as

760-830 m, and from demersal trawls at 375-450 m. This

species ranges from the California and Alaskan current

systems into the southern Bering Sea (Jefferts 1983). Gonatus

berryi has been reported from the stomach contents of

elephant seals from San Miguel Island, California (Antonelis

et aI. 1987).

Gonatus sp. (cf. G.fabricii (Lichtenstein, 1818]) Gonatus

fabricii has not been reliably identified in the North Pacific

or its adjacent seas. However, beaks similar to those of

G. fabricii have been reported from stomachs of marine

mammals taken from the California Current system, as

detailed below. We have beaks representing only one or two

squids per occurrence. In this species-group, the upper beak

can also be used for identification. LRLs are in the size range

6.2-8.8 mm, with a mean of7.9 mm. Young (1972) in his

description of G. californiensis mentioned that it is very

similar to G. fabricii specimens from the Atlantic and

G. antarcticus from Antarctic waters. Clarke (1986a) mentions the difficulty in separating the beaks of G. fabricii and

G. antarcticus. Our beaks are a reasonable match with those

of Gonatus sp. illustrated by Mead et aI. (1982: 12, fig. 9)

from the stomach of a Mesoplodon carlhubbsi taken off

California, and with those listed as G. fabricii by Clarke and

MacLeod (1980) from the stomachs of sperm whales taken

off western Canada. Fiscus took two of our beaks (NMML

5

50

40

>-

Z

t..J

U

a::

t..J 30

>-

u

z

t..J 20 ::>

a

t..J

a::

.....

10

oc-. ---'-----'---'

5 6 7 8 9 10 11 12 13 14

LOWER ROSTRAL LENGTH (MM)

Figure 2

Lower rostral length-frequencies of 130 Gonatopsis borealis beaks from

the stomachs of sperm whales taken off California.

136 and 137) to the Plymouth workshop in 1981. There they

were examined by M.R. Clarke (Mar. BioI. Stn. U.K.,

Plymouth, England) who stated that they closely resemble

the beaks of G. fabricii and G. antarcticus in the Marine

Biological Association collections. The precise identification

of these beaks must await the finding of comparably large

beaks from known specimens of G. californiensis. For the

present, most authorities are in agreement that G. fabricii,

G. antarcticus, and G. californiensis are very similar morphologically, so we believe that they probably also have

similar beaks.

Gonatopsis borealis Sasaki, 1923 This species ranked second in frequency of occurrence (66 %) and first in total

number of individuals. Most samples in which G. borealis

occurred contained beaks of 1-8 squids, two samples each

contained beaks of 10 squids, and three samples had beaks

representing 11, 14, and 18 squids, respectively. Five complete specimens of G. borealis were obtained from the samples; four ofthese measured 202,212,230 and 320 mm ML.

The crowns of7 specimens were readily identified, as were

the gladii of 19 specimens in the size range 255-318 mm

GL. Gladius length closely approximates mantle length.

Beaks removed from the whole specimens and crowns were

used to identify the remaining beaks. The LRL of 130 beaks

were in the size range 5.0-9.0 mm (x 7.9 mm) (Fig. 2).

Gonatopsis borealis ranges across the subarctic waters of the

North Pacific Ocean and Bering and Okhotsk seas, and in

California Current waters as far south as Guadalupe Island,

Mexico (Okutani et aI. 1988, Young 1972, Fiscus unpubI.

data). This species is mesopelagic to epipelagic, and some

individuals migrate to the surface at night (Fiscus and Mercer

1982). This species has been taken in demersal tows between

85 and 650 m, and midwater opening-closing tows to

635-700 and 500-1000 m throughout the California and
Alaska currents (Jefferts 1983). The genus Gonatopsis has

been reported in the stomachs of sperm whales arou.nd

the Pacific rim. In the eastern Pacific only one speCIes

(G. borealis) is known. Rice (1963b) listed this species from

central California. Gonatopsis is not listed from western

Canada by Clarke and MacLeod (1980), but see below.

Okutani and Nemoto (1964) listed G. borealis makko from

the western Aleutians. Kodolov (1972) listed G. borealis

from the western Aleutians and the Commander Islands.

Betesheva and Akimushkin (1955) listed the genus Gonatopsis from the Kuril Islands. Okutani et al. (1976) listed two

species, G. octopedatus and G. makko, from the Pacific

Ocean off Honshu, Japan; and Okutani and Satake (1978)

listed both G. borealis and G. makko taken from the same

area off Japan one year later. In summary, the genus is an

important component ofthe diet of sperm whales in the North

Pacific, Bering, and Okhotsk Seas.

The beaks of Gonatopsis borealis and Berryteuthis magister

are not separable except when very small (LRL <4.0 mm),

when differences in the degree of wing darkening can be used

in identifying some beaks. At the Plymouth workshop,

Kubodera, Kristiansen, Fiscus, and others tried without

success to find a means of differentiating between the two

species; M.R. Clarke (Mar. BioI. Stn. U.K., Plymouth,

England) likewise could not distinguish between the two.

Gonatopsis borealis and B. magister occupy much the same

range in subarctic waters; however, G. borealis ranges much

farther south in the California Current system. Mature

B. magister range south to Oregon waters; however, Jefferts

(1983) identified two larval specimens from collections off

central California. The smaller B. anonychus occupies much

the same range as the two aforementioned species and occurs south to Monterey Bay, California (Anderson 1978).

Berryteuthis anonychus beaks can be readily separated from

the beaks of the other two species at most stages of growth.

Kubodera and lefferts (1984) listed the MLs of the largest

known specimens ofthe three species: G. borealis, 480 mm;

B. magister, 335 mm; and B. anonychus, 150 mm. We justify

calling the central California beaks G. borealis because

31 positive identifications were made from 5 complete

specimens, 7 crowns, and 19 gladii, all from near-adult or

adult sized squids. At present, there are no specimens of adult

or near-adult sized B. magister from central California

waters; if mature B. magister are later identified from these

waters, we may have to reconsider our identification ofthese

beaks. The beaks listed as B. magister from the stomachs

of sperm whales taken off western Canada (Clarke and

MacLeod 1980) may include some G. borealis, based on the

results of our examination of beaks of both species at the

Plymouth workshop.

Unidentifiable Gonatids Four occurrences of single beaks

or beak fragments were recognizable only as gonatids.

6

Architeuthidae

Architeuthis japonica Pfeffer, 1912 Beaks representing

single squids were found in 11 stomach samples. These included seven upper beaks, three lower beaks, and one pair.

Original identification was verified by M.e. Mercer and F.~.

Aldrich (Fiscus and Rice 1974). (One occurrence-two paIrS

of beak fragments-originally listed as Architeuthis were later

found to represent Mesonychoteuthis hamiltoni.) LRLs of

the only three measurable lower beaks were 11.0, 12.0, and

12.5 mm. The known distribution of the giant squid in the

North Pacific through 1973 was summarized by Fiscus and

Rice (1974). The only record from California waters at that

time was that of Pinkas et al. (1971). Since 1973, a number

of Architeuthis have stranded or been captured in trawls along

the west coast of the United States from Santa Barbara,

California, north to Oregon (F.G. Hochberg, Santa Barbara

Nat. Mus. Hist., Santa Barbara CA 93105, pers. commun.

24 Aug. 1987). Hochberg stated that Architeuthis from

around the Pacific rim from California to Japan can all be

attributed to the species A. japonica, and we have listed our

specimens as such. Clarke (1986a) stated that this genus is

found in the Atlantic, Pacific, and Indian Oceans south to

the subtropical convergence. Roper and Boss (1982) presented a review of what is known of the biology and habits

of this genus. A. japonica has been identified from sperm

whale stomachs at various locations around the Pacific rim:

central California (Fiscus and Rice 1974), the western Aleutian and Commander islands (Kodolov 1972), the Kuril

Islands (Betesheva and Akimushkin 1955), and Honshu,

Japan (Okutani et al. 1976).

Histioteuthidae

Three species of histioteuthids have been described from

the eastern North Pacific: Histioteuthis corona berryi,

H. dojleini, and H. heteropsis (Voss 1969). Young (1972)

and Jefferts (1983) listed records from California waters. The

three published records of H. c. berryi are not from the

whaling area off central California (Voss 1969, lefferts

1983). Histioteuthids were identified from 73 of the 157

samples (46%) (in four samples more than one species was

identified). Histioteuthids have been reported from the

stomachs of sperm whales taken around the Pacific rim: off

western Canada (Clarke and MacLeod 1980), the western

Aleutian Islands (Okutani and Nemoto 1964), the western

Aleutian and Commander islands (Kodolov 1972), and

Honshu, Japan (Okutani et aI. 1976, Okutani and Satake

1978).

Histioteuthis spp. Beaks representing 1-7 squids per sample were identified. Twelve of the occurre~ces listed .as

Histioteuthis spp. are from 1970 samples WhICh were mISplaced and could not be reexamined in 1987-88. Beaks in

this category are most likely H. dojleini or H. heteropsis.
Histioteuthis dofleini (Pfeffer, 1912) One entire specimen

(NMML 59) was identified by C.F.E. Roper (NatI. Mus.

Nat. Hist., Smithson. Inst., Wash., D.C. 20560). In addition to this specimen, two crowns and a number of buccal

masses were identified. The remaining 53 occurrences are

represented by 1-12 beaks per sample. LRLs of 21 beaks

were in the size range 3.0-7.0 mm. Jefferts (1983) reported

taking this species in demersal tows to maximum depths of

100-200 m and 2700-3700 m, and in opening-closing midwater tows as deep as 500-600 m.

Histioteuthis heteropsis (Berry, 1913) In five samples

single beaks represented each occurrence, and in one sample beaks representing four squids were identified. Jefferts

(1983) reported taking this species in both midwater and

demersal tows at depths of 0-500 m. Identification and

separation ofthe beaks of H. heteropsis from H. dofleini were

based on descriptions and illustrations by Wolff (1984).

Ommastrephidae

Two large ommastrephids are present at least seasonally or

periodically off the California coast: Ommnstrephes bartrami

and Dosidicus gigas (Young 1972, Anderson 1978, Wolff

1984). Beak differences between the two genera are slight.

Dosidicus gigas (d'Orbigny, 1835) In comparing beaks

from the sperm whale stomachs with those from whole

specimens in the NMML collection, the four lower beaks

from whale stomachs are a better match with D. gigas

(NMML 92) than with Ommnstrephes bartrami. Anderson

(1978), citing other authors and his own observations, stated

that D. gigas specimens from Monterey Bay, California,

generally measured less than 350 mm ML. His northernmost

capture record was about 20 miles north of Santa Cruz,

California. Hochberg and Fields (1980) and Erhardt et aI.

(1983) presented additional life-history information on

D. gigas. LRLs of the four beaks which we consider to be

D. gigas were 2.7,3.0,3.0, and 5.5 mm. Wormuth (1976)

reported that specimens taken north of the Equator were

much smaller than those taken south of the Equator. Of more

than 800 northern specimens measured, only five exceeded

300 mm ML; whereas off Peru, specimens of 1.50 m ML

are not uncommon (Clarke et al. 1976). Clarke et al. (1976)

report beaks 10-30 mm LRL in the stomachs ofsperm whales

taken in the Humboldt Current off Chile and Peru, where

this species constitutes the bulk of the diet of sperm whales

(Clarke et al. 1988). Two specimens in the NMML collection taken north of the Equator measured 167 and 378 mm

ML, and 3.5 and 5.8 mm LRL, respectively.

Unidentifiable Ommastrephid One large upper beak can

be listed only as an ommastrephid. Around the Pacific rim,

Ommnstrephes bartrami has been reported in sperm whale

stomachs from western Canada (Clarke and MacLeod (1980)

and from Honshu, Japan (Okutani et aI. 1976, Okutani and

Satake 1978), and an unidentified ommastrephid has been

reported from the Kuril Islands (Tarasevich 1963).

7

Chiroteuthidae

Chiroteuthis calyx Young, 1972 Three species of this

family have been collected off California-Chiroteuthis

calyx, Valbyteuthis oligobessa, and V. danae-although the

latter is a tropical species that is only a straggler to southern

California waters (Young 1972, Anderson 1978). There are

some indications that Valbyteuthis lives at greater depths than

Chiroteuthis (Roper and Young 1975). No Valbyteuthis beaks

were available in the NMML collection to compare with

beaks from the sperm whale stomach samples. Beaks from

our samples are quite similar to chiroteuthid beaks from

elephant seal stomachs obtained at San Miguel Island,

California (Fiscus, in prep.). Although they resemble the

Valbyteuthis beaks illustrated by Clarke (1986a, fig. 86B),

they are not a good match for the Valbyteuthis beaks figured

by Roper and Young (1967, plate 4). They are in reasonable

agreement with the Chiroteuthis beaks illustrated by Clarke

(1986a, fig. 86(A)). The crest line and lateral wall in our

specimens appear to be longer than those of C. calyx in the

NMML collection, somewhat similar to Chiroteuthis sp. B

of Clarke et aI. (1976, fig. 5). Elsewhere around the Pacific

rim, chiroteuthids have been reported in the stomachs of

sperm whales taken off western Canada (Clarke and

MacLeod 1980) and off Honshu, Japan (Okutani and Satake

1978), and Chiroteuthis veranyi has been reported from the

western Aleutian and Commander islands (Kodolov 1972)

and from the Kuril Islands (Betesheva and Akimushkin

1955).

Mastigoteuthidae

Mastigoteuthis spp. Two species of this genus are known

from the eastern North Pacific, M. pyrodes (Young 1972)

and M. dentata (Jefferts 1983). Fiscus took an unidentified

lower beak (NMML 149) to the Plymouth workshop in 1981,

where it compared favorably with Mastigoteuthis specimens

in the Marine Biological Association collections. The beak

was later examined by M.R. Clarke (Mar. BioI. Stn. U.K.,

Plymouth, England) who stated it matched his ?Mastigoteuthis species B (see Clarke 1980). Three other lower beaks

from two different samples were later judged by Fiscus to

belong to this genus. LRLs for all four beaks are 5.5, 6.0,

6.5, and 8.0 mm. Mead et aI. (1982) reported finding seven

lower and eight upper beaks of M. pyrodes in the stomach

of a Mesoplodon carhubbsi which had stranded near San

Diego, California. The LRLs of these beaks measured

3.4-3.8 mm. Clarke and MacLeod (1980) reported the occurrence of one ?Mastigoteuthis sp. from the stomach of a

sperm whale taken off western Canada. The genus has not

been reported from sperm whale stomachs elsewhere around

the Pacific rim.
Cranchiidae

At least eight species of cranchiids, representing seven

genera, have been reported from the eastern North Pacific;

most of these are present in the California Current system.

Taonius pavo (Lesueur, 1821) This species is represented

by two lower beaks with LRLs of 7.2 and 8.5 mm. It has

been reported by Young (1972) and Jefferts (1983) in the

California Current system. Voss (1980) describes the distribution of the genus as circumglobal in subantarctic to

subarctic waters, mainly at depths greater than 200 m. In

California waters, Taonius pavo has been reported from the

stomachs of elephant seals at San Miguel Island, California

(Antonelis et al. 1987). Around the Pacific rim, Taonius spp.

have been reported from stomachs of sperm whales taken

off western Canada (Clarke and MacLeod 1980), in the

western Aleutian Island area (Okutani and Nemoto 1964),

in the western Aleutian and Commander Islands (Kodolov

1972), in the Kuril Islands (Betesheva and Akimuskin 1955),

and off the coast of Honshu, Japan (Okutani et al. 1976,

Okutani and Satake 1978).

Megalocranchia spp. M. speculator has been reported by

Jefferts (1983) from the central Pacific north ofthe Hawaiian

Islands. To our knowledge, there are no specimen records

of Megalocranchia spp. from the California Current system.

Voss (1980) reported a circumglobal distribution in tropical

and subtropical waters, and a depth distribution from subsurface to deeper than 1000 m. One of our lower beaks

(NMML 274) was taken to the Plymouth workshop. M.R.

Clarke (Mar. BioI. Stn. U.K., Plymouth, England) stated

that this beak is Megalocranchia; F.G. Hochberg (Santa

Barbara Nat. Mus. Hist., Santa Barbara, CA 93105) also

examined this specimen and is in agreement with Clarke.

Fiscus' identification of other beak samples is based on comparisons with this beak and with illustrations published by

Clarke (1986a). LRLs ofsix specimens were 6.5, 7.0, 14.0,

14.5, 15.0, and 15.5 mm. Clarke (1980), who at that time

called this genus Phasmatopsis, reported its occurrence in

the stomachs ofsperm whales taken at Durban and Donkergat, South Africa; Albany, western Australia; South Georgia;

and from Antarctic waters. It has not been previously

reported from sperm whale stomachs around the Pacific rim.

Galiteuthis spp. Two species of Galiteuthis occur in the

California Current system, G. phyllura and G. pacifica

(Young 1972, lefferts 1983, Voss 1980). Voss (1980) reported that the genus has a circumglobal distribution from

antarctic to subarctic waters, and that it occurs from subsurface waters to depths of 1500 m. We found Galiteuthis

specimens in 57 (36.3%) of the stomachs. Most could not

be identified to species. These included several crowns and

buccal masses, one gladius measuring 445 mm GL, and beaks

representing 1-5 squids per sample. LRL measurements of

seven of these beaks were in the size range 5.0-7.0 mm.

Around the Pacific rim, Galiteuthis spp. have been reported

from the stomachs of sperm whales from British Columbia

(Clarke and MacLeod 1980), off the western Aleutian Islands

8

(Okutani and Nemoto 1964), off the western Aleutian and

Commander Islands (Kodolov 1972), off the Kuril Islands

(Betesheva and Akimushkin 1955), and off Honshu, Japan

(Okutani and Satake 1978). They have also been reported

from elephant seal stomachs at San Miguel Island, California (Antonelis et al. 1987).

Galiteuthis phyllura Berry, 1911 Fiscus took two lower

beaks (NMML 215) to the Plymouth workshop. M.R. Clarke

(Mar. BioI. Stn. U.K., Plymouth, England) examined these

beaks and agreed that they were Galiteuthis, most probably

phyllura. These two beaks had LRLs of 5.8 and 6.0 mm.

The remaining eleven occurrences listed here are based on

comparisons of these two beaks and other Galiteuthis specimens in the samples. Roper and Young (1975) mention that

G. phyllura larvae move deeper as they develop in size, and

at approximately 60 mm ML larval development is complete

and they descend to 900 m or more. Galitenthis pacifica off

Hawaii exhibit similar characteristics, and most occupy

depths below 600 m both day and night (Roper and Young

1975). The species ranked second in abundance of the

cephalopods taken in 63 midwater trawl samples taken in

Monterey Bay, California (Anderson 1978).

Galiteuthis pacifica Robson, 1948 Fiscus took one pair of

beaks from a crown (NMML 127b) to Plymouth. M.R.

Clarke (Mar. BioI. Stn. U.K., Plymouth, England) examined

the lower beak and said it most closely resembled G. pacifica.

Beaks representing 1-3 squids per sample were identified

by Fiscus based on this specimen. LRLs of two beaks were

5.0 and 5.5 mm.

Mesonychoteuthis hamiltoni Robson, 1925 Two pairs of

beak fragments (NMML 121) had LRLs of 30.0 and 32.0

mm. They were taken from the stomach of a 16.2-m male

sperm whale, killed on 19 October 1970 and one of the

largest taken in the Richmond whaling operations. Rice and

Wolman (1970) estimate its age at 36 years on the basis of

annual growth layers counted iri a mandibular tooth. These

beaks were originally identified as Architeuthis sp. by Fiscus,

and were included in the samples described by Fiscus and

Rice (1974). These beaks have much larger LRLs than the

three lower beaks of Architeuthis sp. (11.0, 12.0, and 12.5

mm). Fiscus took these beaks to the Plymouth workshop

where they were examined by M.R. Clarke (Mar. BioI. Stn.

U.K., Plymouth, England) who stated, "After much careful

consideration I now feel quite certain that the lower jaws are

not Architeuthis but are definitely Mesonychoteuthis, this in

spite of the locality of collection. As far as we know this

species does not live north oflat. 400 S and these beaks certainly look as ifthey have been in the whale's stomach for a

very long time." These Mesonychoteuthis beaks strongly

suggest the movement of at least one sperm whale from a

southern- to a northern-hemisphere feeding ground. Ivashin

(1967) reported that a male sperm whale (body length 10.4

m) tagged at lat. 21 °33'N was recovered 4.25 years later at

33°20'S in the Atlantic Ocean. This is the only evidence to

date of long-distance transequatorial migration by sperm

whales.
Unidentifiable Beaks

Alloposidae

Alloposus mollis Verrill, 1880 One OCCl.!rrence of two

beaks and one represented by one beak. Measurements of

two lower beaks were:

The life history and range of the giant octopus have been

described by Hartwick (1983). Hochberg and Fields (1980)

report the eastern North Pacific subspecies o. d. martini

ranges from British Columbia south to Monterey, California, at depths to 100 m, and an unnamed subspecies from

off the California Channel Islands and San Diego. Anderson (1978) and Talmadge (1967) report this species from

Monterey Bay and northern California. Clarke and Macleod

(1980) report two octopods from the stomach of a sperm

whale taken off western Canada, with hood lengths of 12.5

and 13.3 mm, which no doubt represent O. dofleini. Betesheva and Akimushkin (1955) report this species (under the

name O. gibertianus) from sperm whales taken off the Kuril

Islands.

The family Alloposidae is represented by this single

species. Young (1972) took one specimen from the Santa

Catalina basin ofsouthern California. F.G. Hochberg (Santa

Barbara Mus. Nat. Hist., Santa Barbara, CA 93105, pers.

commun. 21 Oct. 1988) mentioned several unpublished

California records from the Santa Barbara Natural History

Museum and Berry collections. Roper and Young (1975)

report this species from Hawaiian waters. Thore (1949) said

that this species is restricted to tropical and subtropical waters

throughout the world. Akimushkin (1965) reports it in the

Pacific between lat. 40 0

N and 40 oS, and lists it as bathypelagic abyssal in habit. Roper and Young (1975) report this

species has been taken from the surface to depths of3180 m.

Alloposus mollis has been reported from the stomachs of

sperm whales taken off the Kuril Islands (Betesheva and

Akimuskin 1955), off Durban and Donkergat, South Africa

(Clarke 1980), and from the Tasman Sea (Clarke and

MacLeod 1982).

Crest length

24.0 mm

27.0

Hood length

13.0 mm

13.0

Clarke (1986a) mentions that M. hamiltoni, the only

species of its genus, has a circumpolar Antarctic distribution, and that it forms a large portion of the sperm whale

diet in that region. He mentions beaks with LRL peaks at

12.0 and 25.0 mm, and offthe South Shetlands at 36.0 mm;

his largest specimen measured had an LRL of 48.0 mm. The

largest specimen caught in an opening-closing net was taken

at 2000 m measuring 1050 mm ML and 22.0 mm LRL.

Additional information on this species was given by Clarke

(1980).

Unidentifiable Cranchiids We list nine occurrences only

as cranchiids; two of these are represented by gladii only,

one by a lower beak distinguishable only as a cranchiid. The

remaining six occurrences are from the 1970 samples which

were not reexamined in 1987-88; they were most likely

Galiteuthis spp.

Vampyroteuthidae

Vampyroteuthis infemalis Chun, 1903 Beaks representing

1-4 specimens were found in 43 whale stomachs. Both

upper and lower beaks can be identified. Hood length

measurements from 10 lower beaks were 9.0. 11.0, 11.0,

11.5,12.0,12.5,13.0,13.5,14.0, and 14.0 mm. Vampyroteuthis infernalis is the only species in the family. The

vampire-squid is described as an inhabitant of the abyssopelagic zone by Voss (1967). Young (1972) and lefferts

(1983) report this species from California Current waters.

Roper and Young (1975) report the upper limit of most captures at 600 m and a lower limit of 1100-1200 m. The species

has been reported from the stomachs ofsperm whales taken

off western Canada (Clarke and Macleod 1980), but not

from other locations around the Pacific rim. It has been found

in the stomachs of elephant seals at San Miguel Island,

California (Antonelis et al. 1987). Clarke (1980) reported

specimens from the stomachs of sperm whales taken at

Durban and Donkergat, South Africa, with hood lengths

similar to our specimens.

Octopodidae

Octopus dofleini (Wulker, 1910) Eleven of our samples

each contained 1-5 beaks. Measurements of three lower

beaks were:

Hood length

18.0 mm

16.2

16.0

Crest length

41.5 mm

37.8

37.8

We have separated this category into three groups:

Group 1 Nineteen occurrences represented mostly by

unidentifiable fragments and upper beaks.

Group 2 Sixty-seven occurrences represented by upper

beaks which can be attributed to two families, Histioteuthidae

and Cranchiidae.

Group 3 Four occurrences represented by upper beaks

which can be attributed to either of two genera-Megalocranchia (family Cranchiidae) or Taningia (family

Octopoteuthidae).

9
Table 2

Families of cephalopods identified from stomachs of deep-diving marine mammals from the west coast of the United States (except Alaska)

and Canada.

Physeter macrocephalus Mirounga angustirostris Mesoplodon carlhubbsi

Cephalopod family

Loliginidae

Enoploteuthidae

Octopoteuthidae

Onychoteuthidae

Cycloteuthidae

Gonatidae

Architeuthidae

Histioteuthidae

Omrnastrephidae

Chiroteuthidae

Mastigoteuthidae

Cranchiidae

Vampyroteuthidae

Octopodidae

Ocythoidae

Alloposidae

1Present study

2Clarke and MacLeod 1980

)Antonelis et a!. 1987

'Condit and Le Boeuf 1984

SMead et a!. 1982

Central

California 1

x

X

X

X

X

X

X

X

X

X

X

X

X

X

British

Columbia2

X

X

X

X

X

X

X

X

X

X

X

Southern

California)

X

X

X

X

X

X

X

X

X

Central

California'

X

X

X

X

X

X

X

Southern

Californias

X

X

X

X

X

Discussion _

Berzin (1972), Kawakami (1980), and Clarke (1980) have

provided lists of cephalopods identified from the stomachs

of sperm whales from various locations around the world,

and Rice (1989) has cited the major regional studies ofsperm

whale food habits. Clarke (1980) corrected or updated older

records to conform to current nomenclature, making his

account particularly useful.

Around the world, the same cephalopod families contribute

to the sperm whale's diet, varying somewhat for each locality. With few exceptions, the larger, deeper-living species

contribute most to the diet. Since the above authors have

documented these, we do not add a similar compilation. We

have felt it useful to document published records of cephalopods identified from the stomachs of deep-diving marine

mammals along the west coast ofthe United States (excluding

Alaska) and Canada (Table 2).

Recent feeding by sperm whales captured off central

California is demonstrated by the presence of cephalopod

flesh in their stomachs. Families represented by the presence

of entire squids include Onychoteuthidae (Moroteuthis),

Gonatidae (Gonatopsis), and Histioteuthidae (Histioteuthis);

in addition, Cranchiidae (Galiteuthis) is represented by buccal

masses. Ranked by frequency of occurrence, Moroteuthis

(113) was identified in 72% of the 157 samples, Gonatopsis

(104) in 66%, Histioteuthis (73) in 47%, and Galiteuthis (62)

in 36%.

10

All the genera reported in this paper, with three exceptions (Taningia, Megalocranchia, and Mesonychoteuthis),

have been taken by various types of sampling nets or commercial trawl nets in the California Current system. Marine

mammals still remain the best sampling device for capturing adult and near-adult sized specimens.

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12