RICHARD EDWARD YOU The Systematics

r

 and Areal Distribution

of Pelagic Cephalopods

from the Seas

off Southern California

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY NUMBER 97
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INTERNATIONAL MOK YEAR • 1*72
SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 97

Richard Edward Young Th e SyStematlCS

and Areal Distribution

of Pelagic Cephalopods

from the Seas

off Southern California

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON

1972
ABSTRACT

Young, Richard Edward. The Systematics and Areal Distribution of Pelagic Cephalopods from the Seas off Southern California. Smithsonian Contributions to Zoology,

number 97, 159 pages, 15 figures, 38 plates, 1972.—The mid-water cephalopods from

off the coast of southern California and adjacent areas are described and their areal

distribution is discussed. Forty-two species of pelagic cephalopods are now known

from this area including ten new species. Off southern California the fauna is part

of the transitional and subarctic fauna to the north, while primarily tropical species

are found off northern Baja California.

Official publication date is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year.

UNITED STATES GOVERNMENT PRINTING OFFICE

WASHINGTON : 1972

For sale by the Superintendent of Documents. U.S. Government Printing Office

Washington, D.C. 20402 - Price $2.00 (paper cover)
Contents

Page

Introduction 1

Material and Methods 1

Acknowledgments 3

Description of Study Area 4

Zoogeography 5

Material Examined 7

Key to the Species Described in this Report 11

Systematics 13

Records of Additional Pelagic Species from off Southern California 98

Checklist of Cephalopods Definitely Known to Occur in Southern Californian or Neighboring Waters 99

Summary 100

Systematics 100

Distribution 101

Literature Cited 101

Appendix 1 106

Appendix 2 107

Plates 109

Richard Edward Young Th e SyStematlCS

and Areal Distribution

of Pelagic Cephalopods

from the Seas

off Southern California

Introduction

The pelagic cephalopod fauna of the eastern North

Pacific is poorly known. The single major paper

treating cephalopod systematics of this area (Berry

1912b) dealt with only a small portion of the pelagic

species. Material for this study was collected by the

University of Southern California over a six-year

span from the fall of 1960 to the fall of 1966. During

this period, 445 mid-water tows were taken with an

Isaacs-Kidd mid-water trawl. This represents an estimated 911 hours of "fishing" at discrete depths from

the surface to about 1200 meters. This program

yielded over 5,000 specimens of pelagic cephalopods

belonging to 40 species of which 33 are described in

this report.

The present paper attempts to delineate the pelagic

cephalopod fauna occurring in the waters off southern California and to discuss the zoogeographic relationships of this fauna. Subsequent papers will treat

the vertical and seasonal distribution of the most

abundant species.

Materials and Methods

Specimens were collected with an Isaacs-Kidd midwater trawl with a mouth 10 feet in diameter. The

net was operated from the University of Southern

California's R/V VELERO IV. Comparative material was available from the large collections

Richard Edward Young, Department of Oceanography,

University of Hawaii, Honolulu, Hawaii 96822.

gathered by G. L. Voss at the University of Miami's

Institute of Marine and Atmospheric Sciences. The

University of Southern California trawling program initially studied the waters over the most accessible inshore basin, the San Pedro Basin, which lies

between the mainland and Santa Catalina Island.

The program subsequently was extended to include the area of the Santa Caltalina Basin to

the west of Santa Catalina Island. The most

intensive sampling (over 200 stations) has occurred in this area. While never completely abandoning these two basins, the program was extended to

include San Nicolas Basin, San Clemente Basin, the

waters just west of the continental slope, and the

waters to the south as far as the vicinity of Guadalupe Island. The total area studied is well within a

region extending from 28°N to 34°N. While the

region sampled is rather small, it lies within a transitional faunal region and, as a result, distinct differences were noted in the species composition between

the northern and southern ends of this area. To

demonstrate such geographic differences, I have divided the area of study into seven zones (Figure 2)

so that each includes a major area of sampling.

In this paper, the following abbreviations are used:

Sta. no. number of station where specimen was

captured

M.L. mantle length

M.W. mantle width

P.L. pen length

P.W. greatest width of pen

F.L. fin length
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

34°

FIGURE 1.—Localities of trawling stations.

Dots correspond to the midpoints of the trawling transects.

120° 116°

H.W. head width

H.L. head length measured from the anterior end

of the nuchal cartilage of the base of arms I

Tail L. tail length

Arm L. I length of arm I measured from the arm

base, between arms I (where the muscular portion

of the arms join) to the arm tip

Arm L. II length of arm II measured from the arm

base between arms II and III to the arm tip

Arm L. Ill length of arm III measured from the

arm base between arms II and III to the arm tip

Arm L. IV length of arm IV measured from the

arm base between arms IV to the arm tip

A.C. I-IV arm count (Total number of suckers

and/or hooks on the indicated arm)

l

/i A.C. I-IV total number of suckers and/or hooks

on the proximal 50 percent of the arm length of

the indicated arm measured from the basal sucker

A.C. Dist. S. I-IV number of suckers distal to arm

hooks on the indicated arm

A.C. Prox. S. I-IV number of suckers proximal to

arm hooks on the indicated arm

A.C. Hooks I-IV number of hooks on the indicated

arm

Sucker D. diameter of largest arm sucker

+ accurate count or measurement greater than

given figure due to damaged arm.

Club L. length of tentacular club

Club H. number of hooks on club

Club H. F. club hook formula (number of hooks in

the dorsal medial row over the number in the ventral medial row)

Club S. number of suckers on club

Club S. D. diameter (long axis) of largest club

sucker

Carp. S. number of suckers in carpal cluster

Carp. K. number of knobs in carpal cluster

Carp. F. carpal formula (number of carpal knobs

over the number of carpal suckers)

Dact. S. number of suckers on the dactylus of the

club

Dact. T.S. number of suckers in the sucker pad at

the tip of the dactylus

Tent. L. tentacle length
NUMBER 9 7

FIGURE 2.—Breakdown of area studied into

zones corresponding to major areas of sampling.

28°

120° 118°

Stalk S. number of suckers on the tentacular stalk

D.S.S. number of suckers in the dorsal row on the

tentacular stalk

V.S.S. number of suckers in the ventral row on the

tentacular stalk

M.S.S. number of suckers in median portion of tentacular stalk

R. right

L. left

Tub. L. length of cartilagenous strip bearing tubercules

Tubercles number of cartilagenous tubercules at

funnel-mantle fusion

Gill L. gill length

G.C. gill count (total number of primary gill

lamellae)

Gill W. gill width

Lam. L. length of longest lamella

Eye D. eye diameter

Egg D. egg diameter

Ov. G. Diam. diameter of oviducal gland.

Unless otherwise indicated in the tables, measurements and counts of structures on the right and left

sides have been averaged to give single figures. Measurements are in mm and have been rounded to the

nearest whole number in most cases.

Acknowledgments

I am indebted to many people who assisted me in

the present work. J. S. Savage, J. S. Garth, R.

Lavenberg, D. Bright, and R. Given associated

with the University of Southern California provided specimens and data. F. Hochberg kindly

supplied specimens taken by the R/V SWAN. G.

Y. Hendrix of the University of Miami provided

a considerable amount of help in sorting and caring for the collection and in making preliminary

identifications during the early stages of this work.

Diane McGinnis helped with sorting when time

was becoming a critical factor.

Drs. Garth and Savage also provided funds for

a trip to the University of Southern California

during the course of the study. I am also indebted

to the personnel of the R/V VELERO IV and

the students and staff at the University of South-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ern California that participated in the cruises.

This study was accomplished under the supervision of G. L. Voss, whom I thank for his assistance in innumerable ways and for his support,

encouragement, and help with the manuscript.

The comparative material, which was invaluable

to this study, was drawn from the large collections

gathered by Dr. Voss at the University of Miami's

Institute of Marine and Atmospheric Sciences.

I am grateful to D. DeSylva and R. Hurly for

reading the manuscript and offering valuable suggestions.

I am also indebted to C. F. E. Roper for supplying type material and much valuable information concerning specimens in the collections of the

National Museum of Natural History. Dr. Roper

has added considerably to the present work

through numerous discussions on systematic and

other biological problems. E. S. McSweeny also

provided aid through discussions of many of the

problems encountered.

The great majority of the illustrations were done

by Mrs. Constance McSweeny. Mrs. McSweeny,

a biologist as well as an excellent artist, on many

occasions detected important systematic characters

that had previously gone unnoticed. I thank Mrs.

McSweeny for her excellent illustrations and also

for her help with the systematic aspects of the

paper. Mrs. McSweeny deserves double thanks for

her contribution. After all the plates were completed, they were destroyed in a fire at Miami and

had to be redone.

During the course of this study, a number of

visits were made to Redlands, California where S.

Stillman Berry and I were able to have long discussions on many aspects of cephalopod biology. I

thank Dr. Berry for his interest and enthusiasm.

Finally, I wish to thank my wife, Shelia, for assisting with the typing, compiling data, and for

supplying ecouragement.

This research was partially supported by grants

G-10691 and G-23467 from the National Science

Foundation to the University of Southern California, and by grants GM-5729X, GA-0709, and

GA-1493 from the National Science Foundation

to the Univrsity of Miami, and by the National

Geographic Society-University of Miami DeepSea Biology Program.

This work was done in partial fulfillment of the

Ph.D. degree at the Institute of Marine and Atmospheric Sciences of the University of Miami.

Description of Study Area

The ocean floor off southern California between the

shoreline and the continental slope, a distance of

50 to 160 miles, is unusually complex. Shepard and

Emery (1941) designated this region the continental

borderland to distinguish it from typical continental

shelves. This area contains a dozen complete basins

ranging from about 500 to 2,100 meters in depth,

several troughs, numerous banks, and eight islands.

At the western edge of the borderland, the continental slope drops rapidly to a depth of about 3,500

meters.

The waters over the continental borderland are derived from the complex interaction of several water

masses. The California Current is the eastern component of the great anticyclonic circulation of the

North Pacific and, as such, it is a continuation of the

North Pacific Drift (sensu lato). The waters of the

northern portion of the California Current are derived largely from the Subarctic water. As this

cold current extends southward, it warms through insolation and by increased mixing with waters from

Central water to the west. At about 25° N, it

begins to turn westward and eventually merges into

the North Equatorial Current. Off southern California a nearly permanent eddy is present, creating

north-flowing currents between San Clemente and

Santa Catalina islands.

The surface waters over the borderland are modified to some extent by upwelling. Prevailing winds

along the southern California coast, which are from

the north and northwest, are strongest in May and

June. Surface waters, under the influence of these

winds and the earth's rotation, are driven offshore

and replaced by nutrient-rich waters which upwell

from below to a maximum depth of 200 to 300

meters. Entrainment of surface water by the California Current over the shallow Santa Rosa-Cortes

Ridge may also play a role in the upwelling.

During the winter an inshore surface countercurrent is present along most of the coast from the

southern tip of Baja California to at least 45° N. This

current seems to be somewhat variable off Baja California and off southern California, but is well defined north of Point Conception where it is known

as the Davidson Current.
NUMBER 9 7

Below 150 to 200 meters a deep counter-current of

Equatorial Pacific Water flows northward along the

coast to at least 40° N. and progressively mixes with

the lower layers of the California Current. Off southern California the major flow of the deep countercurrent, which has become transitional in composition at

this latitude, occurs just seaward of the continental

slope, although portions flow over the borderlands apparently filling the basins and forming the water of

the intermediate depths.

Thus, over the borderland, the influence of the

California Current, the surface countercurrent, and

upwelling predominate in about the upper 200 meters

while below 200 to 300 meters the influence of the

southern water predominates. (The above discussion

is based primarily on Reid, Roden, and Wyllie 1958,

and Emery 1960.)

Surface temperature ranges are shown in Figures 3

and 4.

Zoogeography

The present study area lies within an oceanographic

and faunistic transitional region. The faunal changes

that occur within this area have been demonstrated

by a number of authors. The faunal boundries seem

to apply equally well to the littoral (Ekman 1953),

deep benthic (Parker 1964), epipelagic (McGowan

1968), and mid-water (Brinton 1962, Lavenberg and

Ebeling 1967) faunas.

Brinton (1962) gave an excellent account of the

distribution of euphausiids in the northeastern Pacific. His chart showing the faunal relationships of

euphausiids in this area for a 2-month period (August—September) is reproduced here (Figure 5). A

more generalized chart (Figure 6) is given by McGowan (1968) based on the distribution of pteropods

and heteropods.

From the limited information available, it appears

FIGURE 3.—Extremes of temperature regression curves in the study area based on data

taken from 1950 to 1962. (Reproduced from

R. J. Lynn, 1967.)

REGRESSION CURVE MINIMA

TEMPERATURE CO
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

DEGRESSION CURVE MAXIMA

TEMPERATURE C O

28" -

•Jr

FIOURE 4.—Extremes of temperature regression curves in the study area based on data

taken from 1950 to 1962. (Reproduced from

R. J. Lynn, 1967.)

that cephalopods in this area exhibit general faunal

distributions similar to the euphausiids, pteropods,

and heteropods. Many cephalopod species during the

period studied were absent or only sparcely represented in the southern zones, but were abundant in

the northern zones; the reverse trend was also apparent. In many cases species can be confidently related to the established faunal assemblages of the

North Pacific, while for other species only indications

of their faunal relationships can be determined.

Subarctic and Transitional species. Due to our

present lack of information, it is difficult in many

cases to determine whether a species occurs primarily

in Subarctic, Transitional, or Subarctic and Transitional waters. Therefore, the placement of certain

species within one category or the other must be considered tentative.

Moroteuthis robusta appears to be the species most

closely associated with Subarctic water which has

been reported off southern California. It has only

rarely been collected from these waters (Phillips

1961). It is known from off northern California

(Phillips 1966), Oregon (van Hyning and Magill

1964), Canada, the Aleutian Islands, and northern

Japan (Clarke 1966).

Two other species, Berryteuthis ma&ster and B.

anonychus, probably are distributed primarily in Subarctic waters and are not known from California.

Berryteuthis magister (Berry 1913) has been recorded

from Japan, Kuril region, coastal waters of the Aleutian Islands, the Gulf of Alaska, Puget Sound, and

off Oregon (Clarke 1966). Berryteuthis anonychus

(Pearcy and Voss 1963) is known only from the

coast of Oregon.

Gonatopsis borealis is also known from Subarctic

waters, but is more abundant in the southern California waters than Moroteuthis robusta and has a

southern limit of distribution off northern Baja Cali-
NUMBER 9 7

Material Examined

Species

Abraliopsis falco

Abraliopsis felis

Pyroteuthis addolux

Pterygioteuthis gemmata

Ommastrephes bartramii

Symplectoteuthis luminosa

Neoteuthis sp.

Histioteuthis heteropsis

Histioteuthis dofleini

Octopoteuthis deletron

Gonatus onyx

Gonatus berryi

Gonatus pyros

Gonatus californiensis

Gonatopsis borealis

Onychoteuthis borealijaponicus

Mastigoteuthis pyrodes

Chiroteuthis calyx

Valbyteuthis oligobessa

Valbyteuthis danae

Grimalditeuthis bomplandii

Bathyteuthis berryi

Cranchia scabra

Leachia dislocata

Galiteuthis phyllura

Galiteuthis pacific a

Helicocranchia pfefferi

Japetella heathi & /. sp.

Ocythoe tuberculata

Alloposus mollis

Octopus sp.

Vampyroteuthis infernalis

Total Specimens

0-

10

187

1

9

26

1

11

—

69

—

—

6

31

1

27

14

5

26

333 +

14

n20

395

18

47

128

1

12

330+

123

41 +

29 +

63

47

4

1

11

3

65

24

29

33

2

100 +

131

21-

30

210

1

99

62

22

633

71

101

50

53

8

19

6

3

1

33

30

24

26

2

56

31-

40

6

118

13

20

46

225

29

41

25

12

3

2

38

6

1

19

37

1

25

12

40

41-

50

4

20

3

9

88

10

4

13

2

2

46

3

1

1

1

19

20

1

17

9

43

Mantle

51-

60

3

7

24

2

4

1

24

1

9

8

1

7

1

1

50

6/-

70

4

5

6

1

1

3

5

2

1

10

3

1

8

3

16

Length

71-

80

3

1

1

1

1

1

7

5

4

1

21

81-

90

1

1

1

1

1

1

4

1

8

5

1

10

in mm

91-

100

1

2

1

1

3

1

1

2

6

1

6

101-

110

1

1

3

7

3

3

HI- 121-131- 141- 15t

120

1

5

2

1

130

3

1

4

3

140

1

2

1

2

7

5

150

1

1

3

4

+

5

1

5

5

13

) Total

Specimens

10

930

20

168

5

2

1

251

3

116

1309 +

308

187 +

128 +

141

95

27

137

19

1

1

16

8

225

182

4

115

116

4

1

432 +

390

5,352 +

fornia. It has been recorded from California, Oregon,

the Bering Sea, Kamchatka, and nothern Japan.

Onychoteuthis borealijaponicus is known from off

California, Oregon, Canada, and northern Japan.

Probable records also occur in the Bering Sea.

If all northern Pacific Galiteuthis (except G. pacific a) belong to G. phyllura, then this species would

be known from the Kurile Islands and the Bering

Sea as well as off California and Oregon; it seems

likely that this will prove to be the case.

Loligo opalescens may extend as far south as the

tip of Baja California and extends north to the southern end of British Columbia (Fields 1965).

A number of additional species are believed to have

the southern limit of their distribution off northern

Baja California and are known to extend north to

the middle Oregon coast. These are: Gonatus onyx,

Chiroteuthis calyx, Octopoteuthis deletron, and Abraliopsis felis. It seems likely that these four species will

prove to be distributed primarily in Transitional

waters, although Abraliopsis felis has tentatively been

recorded off Hawaii. Octopoteuthis deletron is also

known from off northern Peru and may be an antitropical species.

Histioteuthis heteropsis is known from the southern

California area north to Monterey Bay. Its southern
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

160*

140* 130* MONORTHEASTERN PACIFIC EUPHAUSHOS

ASSOCIATIONS OF SPECIES

Subarctic species

100%

>50% (<50% Transition

species)

Transition species

100%

>50%

<50%

FIGURE 5.—Faunistic zones of the northeastern Pacific based on the distribution of

euphausiids. (Reproduced from Brinton, 1962).

limit is thought to lie off northern Baja California.

It has also been reported off Chile and is also considered to be antitropical in distribution.

Other species which, at present, are thought to

have southern limits off northern Baja California and

are known only from Transitional waters are Gonatus

berryi and Gonatus pyros. Gonatus californiensis extends somewhat farther south along Baja California,

and there is a single record of a larva from the Bay

of Panama that apparently belongs to this species.

This is, however, considered tentatively to be outside

the normal range for this species.

Central species. Two species, Leachia dislocata

and Abraliopsis falco, show definite preference for

the southern zones of the study area and have also

been found off Hawaii and therefore probably belong
NUMBER 9 7

GENERAL FAUN 1STIC ZONES

OF THE PLANKTON OF THE

CALIFORNIA CURRENT -

FIGURE 6.—Faunistic zones off California and

Baja California based primarily on the distribution of pteropods and heteropods. (Reproduced from McGowan, 1968.)

to the Central Water Mass fauna.

Equatorial species of the eastern Pacific. Two

species, Galiteuthis pacifica and Valbyteuthis danae,

are know from the eastern Pacific off Panama where

they are thought to be common. Only a few specimens have been taken in the southern California

area.

Occasionally, Dosidicus gigas is found off southern

California. A major invasion of this species took

place from 1934 to 1937. This squid first became

abundant in 1935 when 12 tons were landed at San

Pedro Harbor during August and September. The

squid varied from 2 to 2/2 feet in length. Many individuals were taken north of Monterey Bay during

the fall and winter of 1935 and January of 1936.

Most of these specimens were between 3 and 4 feet

long. In January 1936, 400 pounds were taken in a

purse seine. The individuals averaged close to 4 feet

in length (Clark and Phillips 1936). Croker (1937)

states that in 1934 and 1935 specimens weighed about

5 pounds, while those caught in the early part of

1937 were averaging over 20 pounds.

This information indicates that D. gigas was not

breeding in the southern California waters but invaded, grew, and disappeared after 1937. Apparently this species has not been present in any numbers

since 1937. Young D. gigas are common in the Bay of

Panama (personal observation) while only large

specimens are known from the coast of Chile, and

these disappear during part of the year (P. GarciaTello, personal communication). It seems likely that

this species is an equatorial form which migrates
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

away from its spawning grounds into cold water to

feed. If this is so, it is strange that this species does

not occur more commonly off southern California.

Argonauta pacifica has been reported from the

Galapagos Islands to Monterey Bay (Berry 1912b).

Its occurrence off southern California seems to be

sporadic. On occasion, it has been reported off southern California in considerable numbers (Berry

1912b).

Cosmopolitan tropical and subtropical species.

Species known from California that have been recorded as having a cosmopolitan distribution in

tropical and subtropical waters are: Pterygioteuthis

gemmata, P. giardi, Ommastrephes bartramii, Alloposus mollis, Ocythoe tuberculata, Histioteuthis dofleini. Two additional species, Cranchia scabra and

Vampyroteuthis infernalis, have a similar distribution

but also occur in somewhat higher latitudes. Symplectoteuthis luminosa appears to be broadly distributed

within warm waters of the Pacific Ocean only.

Species with insufficient distributional data. Mastigoteuthis pyrodes is found in greater abundance

within the southern zones of the study area, but is

not known from either the waters off Panama or off

Hawaii. Valbyteuthis obligobessa and Bathyteuthis

berryi are known only from the present trawling program and show no preference for the southern or

northern latitudes within the study area. Pyroteuthis

addolux also exhibits no latitudinal preference in the

study area, but is clearly more abundant in the more

oceanic zones. This species is also known from as far

west as longitude 147° W. Systematic problems or insufficient data prevent discussion of the zoogeography

of the following species: Neoteuthis sp., Japatella diaphana, Japetella sp., Grimalditeuthis bomplandii, and

Heliocranchia pfefferi.

Inshore and offshore species. Some species that

occur along the coast show preference for inshore

waters over the continental borderland, while others

show a preference for the more offshore waters seaward of the continental slope. In the present study,

Gonatus onyx, the most abundant species taken,

seemed to prefer near-shore waters as do G. californiensis and G. pryos, while Onychoteuthis borealijaponicus and Pyroteuthis addolux showed preference for offshore waters. McGowan and Okutani

(1968) found that larvae of Abraliopsis felis were

taken most frequently in offshore waters. A similar

trend was found in the present study in specimens 20

mm M.L. or less, but not in specimens larger than

20 mm. Valbyteuthis obligobessa is also more abundant in the offshore zones; this appears to be correlated with its deep-water habitat.

Faunal affinities. It is apparent that in the northern zones off Los Angeles the species with northern

affinities (i.e., Transitional and Subarctic species)

TABLE 1.—Distribution frequencies (specimens per hour of trawling) of most

abundant species. Frequencies are corrected for estimated depth of capture and for

contamination resulting from the use of open-net trawls.

Species

AU r >. • i r \ > 20 mm M.L. ..

Abraliopsis fehs 11 < l\ mm M.L. ..

Pterygioteuthis gemmata

Histioteuthis heteropsis

Octopoteuthis deletron

Gonatus onyx

Gonatus berryi

Gonatus pyros

Gonatus califoriensis

Gonatopsis borealis

Onychoteuthis borealijaponicus

Chiroteuthis calyx

Leachia dislocata

Galiteuthis phyllura

Helicocranchia pfefferi

Vampyroteuthis infernalis

Japetella sp

Zones

1 2 3 4 5 6 7

0.66 1.13 0.03 0.00 0.50 0.63 0.22

0.61 1.07 0.12 0.75 2.93 1.63 0.23

0.15 0.08 0.30 0.00 0.13 1.71 1.34

0.22 0.52 0.37 0.14 0.18 0.31 0.03

0.24 0.18 0.26 0.05 0.36 0.29 0.00

3.12 2.80 1.34 0.05 0.00 0.12 0.00

0.32 0.57 0.59 0.19 0.25 0.45 0.00

0.16 0.24 0.07 0.11 0.03 0.02 0.15

0.28 0.47 0.16 0.00 0.03 0.00 0.00

0.06 0.40 0.05 0.00 0.21 0.08 0.01

0.05 0.09 0.05 0.00 1.49 3.18 0.18

0.19 0.16 0.26 0.05 0.19 0.52 0.00

0.02 0.07 0.13 0.12 0.31 2.28 1.25

0.11 0.26 0.28 0.04 0.29 0.29 0.02

0.11 2.28 0.53 0.07 0.04 0.12 1.63

0.23 0.57 0.52 0.70 0.89 1.19 0.49

0.09 0.15 0.09 0.25 0.22 0.30 0.26
NUMBER 9 7 11

predominated during the study period from 1960-

1966. The 14 more abundant species (disregarding

Helicocranchia and Japetella due to systematic complications) taken in this survey give a total frequency

of capture (specimens/hour) of 9.41 in zone 2 (Catalina Basin—the most heavily sampled area), of which

8.59 specimens per hour represent species with northern affinities. In zone 7 (the most southernly zone)

these same 14 species have a total catch frequency of

3.92 specimens per hour, of which 3.08 specimens per

hour represent species with southern affinities (I.e.,

species of tropical waters including the North Pacific

Central waters). These figures also suggest a greater

abundance of individuals in zone 2 compared to the

more oceanic zone 7.

The predominant affinity with a more northern

fauna can be further emphasized by the total counts

of individuals. Almost 80% of the total specimens

taken in the survey belong to species with northern

affinities, while only slightly more than 11% of the

total specimens belong to species with southern affiinities. (The remaining 9% represent species whose

affinities could not be determined for various

reasons.)

Of the 42 species of pelagic cephalopods recorded

off southern California and nearby waters 8 are unidentified at the specific level as a result of their damaged condition, larval state, or merely insufficient

numbers. Of the remaining 36 species, 10 also occur

in the Atlantic Ocean. Available material was sufficient to allow a detailed comparison between Atlantic and Pacific populations of only four of these

species: Pterygioteuthis gemmata, Ommastrephes

rtramii, Cranchia scabra, and Vampyroteuthis in-

'lis. All four showed distinct differences between

o oceans. The differences in local specimens of

P. gemmata are probably a result of geographical

variation. The differences in the local specimens of O.

bartramii, C. scabra, and V. infernalis are presently

interpreted as resulting from geographical variation,

but this will require confirmation. There was abundant material of a fifth species, Heliocranchia pfefjeri, available, but comparative material was inadequate; however, a slight difference in eye size was

noted between specimens from the different oceans.

It is felt that mature specimens as well as more adequate comparative material must be examined before the identification of this species can be confirmed.

Although direct comparisons were made, material

of Alloposus mollis, Ocythoe, tuberculata, and Grimalditeuthis bomplandii was insufficient to allow a

careful evaluation of variations in specimens from

different oceans.

Comparisons of populations of Histioteuthis dofleini have been made by N. Voss. She informs me

that this widely distributed species shows considerable

geographical variation.

I have not examined specimens of the final cosmopolitan species, Pterygioteuthis giardi, which was recently reported from southern Californian waters by

McGowan.

It should be emphasized that all ten of these presumed cosmopolitan species are primarily distributed

in tropical and subtropical waters and in those species

that have been carefully examined, Pacific specimens

exhibit distinct differences from those of the Atlantic

Ocean.

Of the 26 species known only from the Pacific

Ocean, four, Onychoteuthis borealijaponicus, Moroteuthis robusta, Gonatopsis borealis, and Symplectoteuthis luminosa, are know from the waters around

Japan. It is possible, however, that Octopoteuthis

deletron and Galiteuthis phyllura are identical to

similar described forms from Japan (i.e., Octopodoteuthis sicula and Galiteuthis armata).

Only three species, Leachia dislocata, Abraliopsis

falco, and Abraliopsis felis, of the 26 species restricted

to the Pacific Ocean have been found off Hawaii, and

it seems likely that the juvenile A. felis (if this is a

correct identification) seen from these waters are

outside their area of primary abundance.

Key to the Species Described in this Report

This key is designed for the identification of only adult and subadult specimens. Species (other

than Octopus) known from southern Californian waters which are not encountered in the present

study or only poorly represented in the collections are not included in the key; these are briefly

mentioned at the end of this report where they are distinguished from the following species.

1. a. Suckers with chitinous sucker rings 2

b. Suckers fleshy, without chitinous rings 28

2. a. Hooks present on arms I—III 3
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to the Species Described in this Report—Continued

b. Only suckers present on arms I—III **

3. a. Armature in four series on arms *

b. Armature in two series on arms "

4. a. Tentacles absent Gonatopsis borealis

b. Tentacles present ^

5. a. A photophore present on ventral surface of eacH

 e

y

e

 Gonatus pyros

b. Eyes without photophores "

6. a. A median row of suckers and hooks present on tentacular club proximal to large central

hook. Single hook present distal to large central hook 7

b. A median row of only suckers present on tentacular club proximal to large central hook.

Single hook rarely present distal to large central hook Gonatus onyx

7. a. Oral face of tentacular stalk with numerous sucfcers between the marginal rows. Median

row on tentacular club with hooks at end proximal to large central hook

Gonatus califoriensis

b. Oral face of tentacular stalk without suckers or with only one or two between marginal

rows. Median row on tentacular club with suckers at end proximal to large central

hook Gonatus berryi

8. a. Tentacles absent Octopoteuthis deletron

b. Tentacles present 9

9. a. Fins sub terminal, with posterior margins convex * 0

b. Fins terminal, with posterior margins concave 1 *

10. a. Tentacular clubs with 3-5 hooks. Arms I—III with hooks in two rows (biserial)

Pyroteuthis addolux

b. Tentacular clubs with suckers only. Arms I-III with hooks only in ventral rows

Pterygioteuthis gemmata

11. a. Photophores on ventral surface of head randomly dispersed Abraliopsis felis

b. Photophores on head in distinct longitudinal rdws Abraliopsis falco

12. a. Mantle fused to the head dorsally 13

b. Mantle articulated with the head dorsally by cartilaginous pads bearing grooves and

ridges 17

13. a. Hooks present on tentacular clubs. Fins lanceolate 14

b. Only suckers present on tentacular clubs. Fins no t lanceolate 15

14. a. Cartilaginous tubercules present at points of funpel-mantle fusion Galiteuthis phyllura

b. Cartilaginous tubercules lacking at points of funnel-mantle fusion Galiteuthis pacifica

15. a. Cartilaginous tubercules present over entire surface of mantle Cranchia scabra

b. Cartilaginous tubercles not present over entire surface of mantle 16

16. a. Cartilaginous tubercules extend in a line posterior to each point of funnel-mantle fusion

Lechia dislocate

b. Cartilaginous tubercules absent Helicocranchia pfefferi

17. a. Funnel locking-cartilage with an inverted T-shaped groove 18

b. Funnel locking-cartilage without an inverted T-shaped groove 19

18. a. A pair of long luminous strips present on the ventral surface of the mantle

Sytnplectoteuthis luminosa

b. A pair of luminous strips not present on the ventral surface of the mantle

Otnmastrephes bartramii

19. a. Funnel and mantle locking-cartilages fused. Tentacles absent

Grimalditeuthis bomplandii

b. Funnel and mantle locking-cartilages not fused Tentacles present 20

20. a. Hooks present on tentacular clubs Onychoteuthis borealijaponicus

b. Only suckers present on tentacular clubs 21

21. a. Funnel locking-cartilage with an oval depression bearing one or two inward projecting

knobs 22

b. Funnel locking-cartilage with a straight groove and without knobs 25

22. a. Funnel locking-cartilage with only a posterior knob projecting into the oval depression.

Funnel valve absent 23

b. Funnel locking-cartilage with a medial knob afld with or without a posterior knob projecting into the oval depression. Funnel valve5

 present 24

23. a. Arms IV each with 2-4 suckers Valbyteuthis obligobessa

b. Arms IV each with 13-15 suckers Valbyteuthis danae
NUMBER 9 7 13

24. a.

b.

25.

26.

27.

28.

29.

30.

31.

32.

a.

b.

a.

b.

a.

b.

a.

b.

a.

b.

a.

b.

a.

b.

a.

b.

Tentacular clubs whiplike, unexpanded, with 18-22 longitudinal rows of small suckers.

No photophores on ink sac Mastigoteuthis pyrodes

Tentacular clubs expanded, with 4 longitudinal rows of suckers. Two photophores present

on ink sac Chiroteuthis calyx

Mantle covered with large photophores 26

Mantle without photophores 27

Base of arms IV with 8-10 photophores in a transverse series .... Histioteuthis heteropsis

Base of arms IV with 3 photophores in a transverse series Histioteuthis dofleini

Fins more than

 l/i of the mantle length. Arm suckers in two series Neoteuthis sp.

Fins less than J4 of the mantle length. Arm suckers in 3-4 series on distal half of arms

I—III Bathyteuthis berryi

Fins present. Numerous photophores present Vampyroteuthis infernalis

Fins absent. Photophores absent 29

Arms with biserial suckers throughout 30

Arms without biserial suckers or with biserial suckers only on distal half of arms 31

Arms II and III considerably shorter than arms I and IV Ocythoe tuberculata

Arms subequal Octopus sp.

Arms with uniserial suckers throughout 32

Arms with uniserial suckers proximally and biserial suckers distally Alloposus mottis

Eyes and liver with a brilliant, silvery, iridescent layer Japetella heathi

Eyes and liver without a silvery iridescent layer Japatella sp.

SYSTEMATICS

Class CEPHALOPODA

Order TEUTHOIDEA

Family ENOPLOTEUTHIDAE Pfeffer, 1900

DIAGNOSIS.—Arms with biserial armature (occasionally modified at arm tips) consisting both of

hooks and suckers. Tetraserial armature on the tentacular clubs, although marginal rows may be lost

with growth in some species. Hooks usually present on

clubs. Eight buccal lappets; buccal connectives attach

to the dorsal margins of arms IV. Photophores

present.

Subfamily ENOPLOTEUTHINAE Pfeffer, 1912

DIAGNOSIS.—Nidamental glands absent. Numerous

small photophores present on the surface of the mantle, head, and arms, but not in the tentacles or on

the viscera. Posterior margins of fins concave.

Genus Abraliopsis Joubin, 1896

DIAGNOSIS.—Arms IV with 2-4 very large photophores at the arm tips. Tentacular clubs with 2 rows

of hooks on manus.

TYPE SPECIES.—Enoploteuthis hoylei Pfeffer, 1884.

Abraliopsis falco, new species

PLATES 1A,B; 2A-G

DESCRIPTION.—The mantle is cylindrical anteriorly and tapers to a blunt point posteriorly. The

mantle musculature terminates at the end of the pen

which is in advance of the posterior tip of the body.

This broad posterior tip appears to be composed of

vacuolated connective tissue which continues laterally, for a short distance, up the sides of the muscular

portion of the mantle. The anterior margin of the

mantle projects slightly in the areas of the dorsal and

ventrolateral cartilages. The mantle is moderately

thick and muscular.

The large, muscular fins are terminal; they are

about 70-80% of the mantle in length and about

90-100% in width. Their lateral apex forms a

rounded point. The posterior margin is drawn-out

along the mantle to its apex. Anterior lobes are free.

The funnel is short and muscular. Each funnel

locking-cartilage has a straight groove which opposes

a straight ridge on the mantle. The dorsal pad of the

funnel organ has an inverted V-shape. A large ridge

extends diagonally across each lateral limb of the

organ and an anterior papilla is present. The ventral

pads are oval. A funnel valve is present. The funnel

is free laterally.
14

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The head bears large eyes that occupy its entire

lateral portions. A large sinus is present in the anterior margin of each eyelid, and distinct dorsal and

ventral windows lie above and below each eye. A

nuchal crest with 4 folds on either side of the head is

present; the ventral fold on each side is thick and

partially embrances the funnel; the second fold from

the funnel bears the "olfactory" organ; the dorsalmost 2 folds join posteriorly on each side forming

crescent-shape ridges.

The arms are large and muscular; they are considerably longer in the male. The arm formula is:

4 = 2>3>1. Arms I-IH all have well developed

aboral keels and arms IV have well-developed lateral

keels. Protective membranes are nearly absent proximally on the dorsal borders of arms I where they are

represented primarily by isolated trabeculae; the

membranes are very low distally. The dorsal borders

of arms II have a few trabeculae proximally, but

otherwise lack trabeculae and protective membranes

except near the sucker-bearing tip where a trabeculate protective membrane is present. The dorsal

borders of arms III show only a trace of a few proximal trabeculae except for a small trabeculate protective membrane near the distal, sucker-bearing tip.

The dorsal borders of arms IV have flat, rounded

trabeculae with low membranes in females. Arms I—

III all have large, trabeculate, protective membranes

on their ventral borders. These are present, but not as

well developed on arms IV in females.

Each arm I—III possesses a double series of hooks

which is replaced by a double row of suckers at the

arm tip. The number of distal suckers is difficult to

count with certainty. One female had about 34 on

each arm I, 23 on each arm II, and 15 on each arm

III. The most proximal suckers in this series have

broad outer rings; their inner rings have about 10-12

broad teeth which are rounded to truncate on the

distal margin. The more distal suckers are globular;

they lack outer rings and have reduced apertures

without teeth. All suckers are globular and lack

outer rings on the dorsal pair of arms in males. Arms

IV possess hooks, but lack distal suckers.

The arms in male specimens are longer and have

more hooks than do the arms of females. The increased number of hooks in males results from both

the greater length of the arms and the smaller size

of the hooks. The following chart comparing averages for 4 adult females and 5 adult males demonstrates these relationships.

Both arms IV are hectocotylized in the males;

however, the left arm is much more highly modified.

The trabeculate protective membrane is greatly enlarged on the ventral margin of this arm and is

heavily pigmented with dark brownish purple chromatophores. Small conical knobs line the oral margins

of the large trabeculae. On the dorsal margin, trabeculae are very prominent proximally but diminish

in size distally, while the protective membrane is low

and nearly absent proximally; these trabeculae are

also lined with conical papillae on their oral margins.

The proximal 3-4 hooks on the ventral margin are

much larger than the proximal 2-3 hooks on the

dorsal margin. The fourth or fifth hook on the ventral margin and the third or fourth hook on the dorsal margin are comparable in size. Distally the hooks

on the dorsal margin increase in size, while those of

the ventral margin abruptly decrease. Shortly, the

hooks of the ventral margin begin to increase in size,

while those of the dorsal margin begin a gradual decrease. Approximately the fifteenth hook of the ventral margin approaches in size the adjacent hooks of

the dorsal margin. The hooks of the ventral margin

then decrease more rapidly in size than those of the

dorsal margin up to the end of the series. The fleshy

bases from which the hooks arise in the dorsal row

bear small conical papillae; there are also a few

papillae on some of the hook bases of the ventral row.

The oral face of the arm proximal to the first hook

also possesses numerous papillae.

The right arm IV shows a lesser degree of modification. On the ventral border, from the second hook

to about the middle of the arm, are 4—5 enlarged

Am

No.

No.

A m

No.

No.

CHARACTERS

I length/mantle length

of hooks

of hooks/arm length ....

CHARACTERS

I length/mantle length

of hooks

of hooks/arms length

I

0.46

17

0.83

I II

0.63 0.77

27

1.20 1.

32

17

FEMALES

Arrm

II

0.55

21

0.89

MALES

Arms

III

0.68

26

1.15

r

III IV

0.50 0.56

19 24

0.87 0.97

IV

0.79

42 (left)

28 (right)

1.51 (left)

1.02 (right)
NUMBER 9 7 15

trabeculae, bearing conical papillae, with a low membrane between them; trabeculae are lacking on the

distal half of the arm. Along the distal third of the

ventral border, there are two semilunar flaps. The

dorsal border has trabeculae that are less prominent,

but also bear papillae and have very low protective

membranes. A small flap, present distally on the dorsal border, opposes the distal flap on the ventral

border. The proximal 3^4 hooks in the dorsal row are

considerably smaller than the first 4-5 in the ventral

row. Distally the hooks in the ventral row decrease

abruptly in size, while those of the dorsal row increase

abruptly in size so that the fifth or sixth hook of the

ventral margin is much smaller than the fifth one of

the dorsal margin. Each series terminates at the

proximal end of the distal flaps on each margin. The

tips of the ventral arms bear large photophores identical to those of the female. Both modified arms IV

are about the same length.

In the male, arms I—III show a slight amount of

dimorphism. The trabeculae on the dorsal borders of

arms I are considerably larger than in the females.

They lack a protective membrane proximally, but

one is well developed distally. The oral margins of

the trabeculae on both dorsal and ventral borders are

lined by conical papillae. In males, papillae are also

present to some extent on the trabeculae of arms

III and to a lesser degree on arms II. All arms in

males generally have 5—20 papillae proximal to the

first pair of hooks. In males, the proximal hook on

the ventral margin of each arm III is greatly enlarged over the succeeding hook and is approximately

equal to (or larger than) the proximal hook of the

dorsal margin. The second through fourth hooks of

the ventral margin are distinctly smaller than those

opposing them on the dorsal margin; farther distally,

they are approximately equal in size. Hooks on arms

I—III are smaller in males than in females.

The tentacles are short and robust. The tentacular

club is short and not expanded. The manus bears

hooks in 2 rows; those of the ventral row are about

twice the height of those in the dorsal row in females, while the difference may be less in males.

Hooks of the dorsal row vary from Z-A, while the

smaller hooks of the ventral row vary from 2-4.

Suckers alternate with the hooks in the ventral row.

Only the proximal sucker is present in the dorsal

row, and it is occasionally absent. The dactylus has 4

rows of suckers that extend to a circle of suckers at

the tip of the tentacle. A random count of 8 clubs

shows a variation in dactyl suckers between 48 and

60. A distinct carpal cluster is present which possesses 4 or occasionally 5 suckers and pads. No membrane or flap lies adjacent to the carpus or the manus.

A low, thick protective membrane lines the ventral

margin of the dactylus. There is no trace of a keel.

Numerous photophores of varying sizes are embedded in the mantle in a rather irregular arrangement, although some indication of a pattern is discernible. In the ventral midline, there is a narrow

strip that is generally free of photophores; however,

this strip is sometimes difficult to detect and often

does not extend to the anterior margin or to the

posterior end of the mantle. On either side of this

strip is a diffuse longitudinal band of photophores.

Lateral to these bands, the number of photophores

gradually diminishes to only a few scattered organs

situated dorsally. Superimposed on this pattern is a

single line of photophores that extends from the anterior mantle margin at the projection of the mantle

locking-cartilage, posteriorly for about half of the

mantle length. Between this series and the midventral

strip is another single line of photophores that can

be traced about half of the mantle length from the

anterior margin. Both series are not always readily

apparent.

The funnel has 6 separate groups of photophores,

none of which extends to the exhalant opening. One

group is located on either side of a clear median

strip on the ventral surface of the funnel. A second

group is found along each lateral margin of the funnel, and a third group is located on either side of the

dorsal surface of the funnel and extends onto the

bases of the bridles.

A triangular patch of photophores is located on the

ventral surface of the head dorsal to the funnel. This

patch continues forward over the ventral surface of

the head in the midline as a narrow, well-defined strip

to the junction of arms IV where the series divides

into a row that runs along the inner margin of each

arm IV for about half of the arm length. A second

narrow strip on each side of the head runs anteriorly

from the first nuchal fold, passes medial to the ventral "window" of the eye, and continues onto the

lateral margin of arm IV at the base of the lateral

keel. This series continues to the tip of the arm, but

is broken at 4—5 spots by short areas that lack photophores. Between these first 2 bands, near the base of
16

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

the ventral arms, a few scattered photophores are occasionally present.

A third strip extends anteriorly from each second

nuchal fold, passes on either side of the head lateral

to the ventral "window" and joins an area of scattered photophores anterior to each "window." Each

patch continues onto the margin of the lateral keel of

each arm IV as a single row of photophores which

extends along the proximal third of the arm. A final

series extends around the margin of each eyelid. In

the 3 males which allowed accurate counts, the number of photophores in this series varied from 48-54,

while in 4 females the number varied from 52-63.

This series extends anteriorly on either side onto arm

III at the base of the aboral keel and continues the

entire length of the arm except for a few breaks in

the series in the distal half of the arm.

On the ventral surface of each eye is a series of 5

light organs of which the first and last members are

considerably larger than the middle 3.

Distally the tips of arms IV bear 3 large, darkpurple photophores and 2-3 small photophores.

Within the mantle cavity of females on the inner

surface of the dorsal mantle wall, immediately anterior to the stellate ganglion, is a highly folded, apparently glandular, pocket. The pleats of this pocket

also extend beneath the collar in the nuchal region.

Several sperm reservoirs (discharged spermatophores)

were found attached within the pocket. The pocket

has a dark-purple pigmentation which can be seen

through the dorsal surface of the mantle beginning

about one-eighth of the mantle length from the anterior end of the pen.

The buccal membrane has 8 supports and bears

numerous long fleshy papillae on its brownish purple

oral surface. The buccal membrane connectives attach to the dorsal borders of arms I, II, and IV, and

to the ventral borders of arms III.

The specimens are covered with numerous small,

reddish brown chromatophores.

TYPE LOCALITY.—28°54' N, 118°08' W, eastern

North Pacific Ocean.

LOCATION OF TYPE.—University of Southern California, U.S.C. Hancock collections, AHF Cephalopod Type No. 1.

DISCUSSION.—The genus Abraliopsis has been in

an extremely confused state due to many inadequate

early descriptions, often based on larvae, followed by

numerous misinterpretations by subsequent authors.

Fortunately, the genus has been monographed by

Voss (in manuscript) and most of the problems have

been resolved.

The genus Abraliopsis is usually divided into two

subgenera: Abraliopsis, with a diffuse pattern of

photophores on the ventral surface of the head, and

Micrabralia, with photophores on the ventral surface

of the head arranged in distinct rows (Voss 1967).

A. falco clearly falls into Micrabralia which, according to Voss (in manuscript), contains only two other

named species: Abraliopsis affinis (Pfeffer 1912) and

Abraliopsis gilchristi (Robson 1924). Pfeffer based A.

affinis on Hoyle's (1904) description of 4 specimens

taken from the eastern tropical Pacific between 2° 34'

N, 82°29' W and 14°46' N, 98°40' W. The specimens were only partially described; however, specimens of A. affinis in the collections of the Institute of

Marine and Atmospheric Sciences which were captured at 7° 19' N, 79°44' W have allowed adequate

comparisons to be made with A. falco.

Abraliopsis falco differs most distinctly from A.

affinis on 2 characters: A. affinis has a distinct membrane on the ventral margin of the tentacle that extends from the proximal end of the carpus to the

base of the manus; no such membrane is present in

A. falco. A. affinis has a large keel on the club that

extends from the tip of the dactylus to about the

midpoint of the manus; a keel is lacking in A. falco.

The specific value of these features within the genus

has been emphasized by Voss (in manuscript). Several other characters are also of value in separating

these species. In A. falco, the first hook in the ventral

row of each arm III in males is usually equal to or

larger than the first hook in the dorsal row of the

same arm (this feature was valid in 9 out of 10

cases). In A. affinis the first hook in this same position

in males is distinctly smaller than the first hook in

the dorsal row (this feature was valid in 35 out of 36

cases). The midventral strip on the mantle in A.

affinis which lacks photophores is very broad and

generally expands at its posterior termination less

than two-thirds of the mantle length from the anterior end. In A. falco, this strip is very narrow

throughout and seldom reaches either the anterior or

posterior end of the mantle. The arms are considerably longer in A. affinis, but further study on more

material is necessary to quantify this feature. There

are a number of other features that seem to show

significant differences such as the relative sizes of the
NUMBER 9 7 17

TABLE 2.—Measurements in mm of Abraliopsis falco, new species.

Characters

Sex

M.L

M.W

H.W

F.L

F.W

Arm L. I

II

Ill

IV

A.C. Hooks I

II

Ill

IV

Tent. L

Club L

Club H.F

Carp. S

Dact. S

Station Number of VELERO

9899 9899 9899 9899 10679 10679 10679 10677 9901

9 9 9496 >4<J6 >

(Holotype)

43 46 41 35 44 36 37 36 37

13 15 12 12 11 10 11 11 12

15 — 13 13 15 11 11 14 11

31 33 29 25 32 27 29 28 26

39 44 37 34 39 34 35 36 37

20 23 18 22 20 21 22 24 26

24 25 21 26 25 26 26 — 32

23 23 20 23 22 23 23 27 27

26 25 22 28 25 26 25 30 33

L R L R L R L R L R L R L R L R L R

17 18 16 16 16 16 28 27 16 17 29 28 26 26 26 25 27 27

22 21 22 23 20 20 30 29 20 21 34 34 32 32 — — — —

19 20 19 19 19 18 24 26 18 19 27 26 26 26 — — 30 —

26 24 26 26 22 22 40 26 21 22 44 31 42 29 39 26 45 29

50 69 41 45 43 — 38 40 —

67556—5 6 —

L R L R L R L R L R L R L R L R L R

3/3 3/2 _ 3/3 3/4 — 3/4 3/3 3/3 — — 4/3 3/3 — 3/4 —

44_ 5 4_4 4 4 — — 4 4 4 4 —

46 55 — 50 54 — 51 — 60 — — 48 50 — — —

club hooks and the number of proximal arm papillae; however, the characters listed are sufficient to

separate the species.

Since Abraliopsis falco appears to be a warm-water

species, it is possible that the ranges of A. falco and

A. affinis overlap. Although the original description

of A. affinis is incomplete, there can be no question of

the true identity of this species since a club was illustrated by Hoyle (1904, plate 8, figure 1) that clearly

shows the presence of a keel and another illustration

(plate 10, figure 1) shows the midventral clear strip

on the mantle with the appearance characteristic of

specimens discussed in this paper as A. affinis.

Abraliopsis gilchristi was described by Robson

(1924) from specimens taken off South Africa and

has recently been redescribed by Voss (1967). The

most obvious feature that separates this species from

A. falco is the persence of 2 distinctly separate rows

of photophores on the midventral surface of the head

in A. gilchristi, each of which is continuous with the

medial series of the nearest arm IV. In A. falco, there

is only a single series in the midventral line of the

head. There are many other features (including the

presence of a semicircular membrane on the ventral

border of the tentacle between the carpus and manus

in A. gilchristi) which separates these 2 species, but

need not be discussed here.

There is another but unnamed species in the Atlantic Ocean which is treated fully by Voss (in manuscript) , and a comparison between this species and

A. falco will be made in this paper. For the present,

it is only necessary to state that they are distinct.

DISTRIBUTION.—The 10 specimens of A. falco

taken by the VELERO all came from zone 7. An additional specimen (M.L., 20 mm) was taken by the

SWAN near Hawaii at approximately 24° N and

151° W. These data indicate that this species is a

warm-water form and probably a mid-oceanic

species.

Abraliopsis felts McGowan and Okutani, 1968

PLATES 1C-D; 2H-N

Abraliopsis sp., Pearcy, 1965, p.261.

Abraliopsis felis McGowan and Okutani, 1968, p.72.

DESCRIPTION.—The mantle is long and slender; its

width is about one-fourth of its length. It is cylin-
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

drical anteriorly, but tapers posteriorly to end in a

blunt point. The muscular portion of the mantle

terminates at the end of the pen which is in advance

of the posterior tip of the body. This broad posterior

tip appears to be composed of vacuolated connective

tissue which continues for a short distance laterally

along the sides of the muscular portion of the mantle. The rhachis of the pen is visible in the dorsal

midline. The free margin of the mantle projects in

the nuchal region and in the regions of the mantle

locking-cartilages.

The fins are large, 62-68% of the mantle in length,

and broad, 70-83% of the mantle in width. The

lateral angles of the fins are rounded. Much of the

posterior fin margin forms a nearly straight line at

about 45 degrees to the body axis; however, as the

edge approaches the mantle, it turns posteriorly and

terminates at the tip of the mantle. Anterior lobes

are large and free.

The funnel, which is free laterally, is small and

muscular. The funnel locking-cartilage bears a

straight groove. The dorsal pad of the funnel organ

has an inverted V-shape and bears an anterior papilla

and lateral ridges. The ventral pads are oval. A large

funnel valve is present.

The head width is approximately equal to the

mantle width. The eyes are large and occupy the

entire lateral portions of the head. A small but distinct anterior sinus is present on each eyelid. Four

large nuchal folds are present on each side of the

head, the dorsal 2 of which are joined posteriorly.

The second fold from the funnel bears the "olfactory"

organ, and the fold nearest the funnel forms the posterolateral edge of the funnel groove and bears photophores on its ventral surface. The nuchal cartilage is

elongate and has a prominent median ridge that

bears a central groove.

The arms are relatively short (arms IV are 55-

60% of the mantle in length), and in the order

4>3 = 2>1. All arms I-III have well-developed

aboral keels; arms IV have large lateral keels. Welldeveloped trabeculate protective membranes are present on the ventral border of arms I-III, but are absent from the dorsal borders of these arms. Arms IV

lack protective membranes on both borders.

Arms I-III each bear 2 alternating rows of hooks

which give way abruptly to 2 rows of suckers near the

arm tip. Each arm has approximately 15-20 hooks

and about 20-26 distal suckers. These suckers have

roughly 7-10 slender teeth along the distal half of

their inner chitinous rings. Arms IV have 13-15

hooks and lack distal suckers; these hooks are smaller

than those of the other arms. The tips of arms IV

have usually 3 (sometimes 2 or 4) very large photophores encased in dark-purple pigment followed by

2-4 small photophores of similar appearance.

In the males the right arm IV is hectocotylized. It

carries 10-12 small hooks and near the distal end of

this hook-bearing portion a bilobed membrane,

broadest in its proximal half, is present on the ventral

border of the arm. Slightly more distal to the hooks,

a second membrane arises on the dorsal border. The

area beyond the membranes lacks hooks or suckers.

McGowan and Okutani (1968) have shown that

rarely both ventral arms are hectocotylized. The terminal photophores are the same as in the non-hectocotylized ventral arms.

The tentacles are robust and have short unexpanded clubs which lack protective membranes. The

manus has from 3-4 hooks (rarely 2) in each of 2

rows, with those of the ventral row being much

larger than those of the dorsal row. The latter generally alternate with small marginal suckers, although the full compliment is not always present.

The dactylus consists of approximately 14 transverse

rows of 4 suckers each. The inner chitinous ring of

each of these suckers is nearly smooth, but may have

a few irregular teeth on the distal margin of the first

few suckers. A distinct carpal cluster is present which

has 4-5 suckers and knobs. Membranes are lacking

from the clubs and stalks except for a slight indication of a dorsal keel along the distal half of the dactylus. This keel, when detectable, is very low and appears as only a slender ridge.

The buccal membrane is purple and 8-parted. The

buccal membrane connectives attach to the dorsal

borders of arms I, II, and IV and to the ventral

borders of arms III.

In females, there is a dark patch on either side of

the anterior end of the nuchal cartilage which marks

the position of the spermatophore receptacles. The

inner side of the collar and the outer surface of the

body wall adjacent to the nuchal cartilage form a

glandular pleated pocket which receives sperm reservoirs. Mature eggs are roughly 1.5 mm in diameter.

Nidamental glands are absent and the oviducal

glands are greatly enlarged.

The mantle is covered with numerous photophores
NUMBER 9 7 19

of varying sizes. The light organs have the greatest

concentration on the ventral surface, considerably less

on the lateral surfaces, and only a few scattered

organs are present on the dorsal surface. These organs

from a diffuse pattern over the mantle; there is, however, a narrow strip extending the full length of the

mantle along the midventral line which lacks photophores, and there is a somewhat regular line of photophores that extends around the ventral two-thirds of

the mantle along its free margin. Photophores on

either side of the mantle are not strictly bilaterally

arranged.

The funnel has 6 separate patches of photophores,

none of which extends to the exhalent opening. In

fully mature specimens, one patch, located on either

side of the midline on the ventral surface of the funnel, has between 16-23 photophores. A second patch

is found along each lateral margin of the funnel and

usually has 5 or occasionally 6 photophores. The

other patches, each of which has about 13-15 photophores, are located on either side of the dorsal surface of the funnel and extend onto the bases of the

bridles.

Photophores are abundant on the ventral surface

of the head, but have a diffuse pattern with the following exceptions: 2-3 photophores in a single line

extend along the medial nuchal fold. From this

series, extending dorsally along the posterolateral

margin of the head, is a single row of usually 4 photophores (occasionally up to 6) that reaches to a point

nearly level with the center of the lens. From usually

the second photophore from the ventral end of this

series, a single line of broadly separated photophores

extends anteriorly along the head (occasionally one

or two photophores fall above or below the main

line). This series runs lateral to the ventral "window"

of the eye. Between this line and the eye opening,

there are no photophores except for a line of organs

which lie along the margin of the eyelid. The number

of photophores in the latter series is relatively constant. A count on 19 eyelids gave 2 with 29 photophores, 3 with 30, 11 with 31, one each with 32, 33,

and 34 photophores. There are no photophores over

the ventral "window" of the eye. Medial to this area

the ventral surface of the head is covered with numerous scattered photophores, diffusely arranged but

with a nearly bilaterally symmetrical pattern. Anterior to each "window" there are a few scattered

photophores. In the ventral midline, the most posterior photophore lies dorsal to the funnel and immediately in front of the point of attachment of the

converging bridles. Immediately anterior to this

photophore is a distinct oval area that is usually devoid of photophores.

Arms IV each bear three series of photophores.

One series runs along the ventromedial margin of the

arm for about two-thirds of the arm length. The

middle series runs along the ventrolateral edge of the

arm. At the base of the arm this series is arranged in

2-3 irregular rows which shortly converge into a

single row that continues to the tip of the arm. Near

the arm tip, 2-4 photophores are greatly enlarged

and covered with a dark-purple pigment. These are

followed by 2-4 small photophores. The final series

runs along the edge of the lateral membrane and

terminates roughly at the same level as the ventromedial series.

A single row of photophores extends along each

arm III at the base of the aboral keel for slightly

more than half of the arm length. This series is

aligned with 2 photophores of the eyelid series that

are situated on the dorsal edge of the eyelid sinus;

however, a distant gap separates these 2 series (Plate

ID).

The number of photophores increases with the

size of the animal. Therefore, the above discussion

applies primarily to mature specimens although many

of the features mentioned can also be detected in

young specimens.

This squid is covered with numerous reddish brown

chromatophores.

TYPE LOCALITY.—31 °59' N, 122°24' W. Eastern

North Pacific Ocean.

LOCATION OF TYPE.—U.S. National Museum, catalog number 678792.

DISCUSSION.—A. felis belongs to the subgenus

Abraliopsis which contains only 2 other species:

Abraliopsis (Abraliopsis) pfefferi Joubin, 1896, from

the Mediterranean; and Abraliopsis (Abraliopsis)

hoylei (Pfeffer 1884), from the Mascarene Islands,

Indian Ocean (Voss 1967).

A. pfefferi was originally taken off Villefranchesur-Mer, but has since been found to be distributed

throughout much of the Atlantic Ocean and into the

eastern Indian Ocean (Voss 1967). This species is

easily distinguished from the Californian species by a

number of characters. A. pfefferi has a semicircular

membrane on the ventral margin of the tentacle that
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

originates opposite the carpal cluster and extends distally just past the first hook on the manus. This

membrane is lacking in A. felis. A low, but distinct

keel on the club in A. pfefferi extends from about the

middle of the manus to the dactylus. In A. felis an

indistinct keel is present only on the distal half of

the dactylus. There are usually between 10-12 hooks

on the tentacular clubs in A. pfefferi, while A. felis

has usually 6-8. In A. pfefferi there are roughly 44—

46 photophores surrounding the eyelid, while A. felis

has usually 30-31 photophores. In A. pfefferi, lateral

to the ventral "window" of each eye, there is a patch

of photophores consisting of at least 4 irregular longitudinal rows. In A. felis there is only a single row of

photophores in this position. In A. pfefferi, the series

of photophores surrounding the eyelid continues onto

arms III. In A felis a distinct gap separates the 2

series. In A. felis there is an oval patch devoid of

photophores near the midposterior portion of the

ventral surface of the head which is lacking in A.

pfefferi. There are other distinctive characters such

as the fin length and the number of arm hooks (see

also discussion of A. hoylei); however, the features

mentioned should be sufficient for the present.

A. hoylei has been the subject of considerable discussion since its original discovery. Hoyle (1904) ascribed several specimens from the Pacific Ocean near

Panama to this species. Pfeffer (1912), recognizing

the distinctive features of Hoyle's specimens, placed

these forms in a new species A. affinis (a member of

the subgenus Micrabralia). Robson (1948) rejected

Pfeffer's name and assigned specimens that he examined from Panama to A. hoylei which he considered to be a polymorphic species. There can be no

doubt, however, that A. affinis and A. hoylei are distinct species and even belong in separate subgenera

(Voss, in manuscript). Both Berry (1913, 1914) and

Allan (1945) ascribed specimens from the South Pacific to A. hoylei, although with some hesitation.

These specimens were either too small or too inadequately described for their identity to be ascertained

with certainty; however, it is unlikely that they belong to A. hoylei. Therefore, the only valid descriptions of A. hoylei are those of the type specimen

(Pfeffer 1884, 1912).

A. hoylei is much more difficult to separate from A.

felis than is A. pfefferi. This is not unexpected since

one must rely solely on Pfeffer's (1884, 1912) descriptions of the type as the type specimen has been destroyed (Voss, in manscript), and there are no other

specimens available. It is, however, possible to detect

the following differences: in the single specimen of

A. hoylei, the fins are 78% of the mantle in length

and 95% of the mantle in width; the arms (apparently Pfeffer refers to arms IV which are the longest) are 75% of the mantle length. In A. felis the

fins are 62-68% of the mantle in length and 70-

83% of the mantle in width. Arms IV are 55-60%

of the mantle length. The first few distal suckers in

A. hoylei are stated to have pointed teeth on the

inner rings while the rest have smooth rings. In A.

felis the distal suckers have long slender teeth in at

least the first 10 suckers. Pfeffer could not describe

the tentacular membranes in A. hoyei since the tentacles had undergone some desiccation; however, his

illustration (plate 17, figure 9) shows a keel that

originates near the middle of the manus. This differs

from the previously mentioned conditioned in A.

felis.

A number of differences can be detected in the

light-organ patterns. Pfeffer described a ventral field

of photophores on the ventral surface of the mantle

which has a median bare strip lacking photophores on

the posterior two-thirds of the mantle. The illustration shows this clear band extending somewhat

farther anteriorly, but clearly not approaching the

anterior margin of the mantle. In A. felis this clear

band extends to the anterior margin or sometimes is

separated from it only by a single line of photophores. The ventral field in A. hoylei is bounded

laterally by an area which "fast den Eindruck eines

leeren Langstreifens macht." Lateral to this zone is

another area of more densely packed photophores

which Pfeffer called the laterodorsal field. In A. felis

a relatively clear band separating the laterodorsal

field is lacking; instead, the number of photophores

gradually diminishes laterally. On the ventral surface

of the head, Pfeffer described a row of photophores

that corresponds to the row immediately lateral to

each ventral "window" of the eye of A. felis. Lateral

to this row, he describes "eine weitere, unbedeutende

kleine Reihe." This additional series is lacking in A.

felis.

One might be somewhat dubious of the small differences here indicated in the photophore pattern

between A. felis and A. Hoylei; however, A. pfefferi

differs from A. felis in exactly the same features and

in the case of A. pfefferi there can be no question
NUMBER 9 7 21

concerning the validity of these characters. In fact,

the other characters of A. hoylei mentioned here,

which separate it from A. felis, are also shared by A.

pfefferi. This condition suggests that A. hoylei and A.

pfefferi may be synonymous; however, characters such

as the number of hooks on the tentacular clubs and

other features make this relationship somewhat uncertain. At any rate, it is possible to separate A. felis

from both A. hoylei and A. pfefferi and thus verify

the specific status of A. felis.

McGowan and Okutani (1968) reduced Watasenia

to the status of a subgenus of Abraliopsis and included in it Abraliopsis (Watasenia) scintillans and

Abraliopsis (Watasenia) felis. In doing this, these

authors relied greatly on the sexual modifications of

the ventral arms in members of Watasenia and Abraliopsis.

Descriptions of the males of A. pfefferi and A.

hoylei have never been published; therefore, McGowan and Okutani were unaware of the nature of

the hectocotylus in these species. I have examined

two males of A. pfefferi from the Indian Ocean. The

right ventral arm is hectocotylized in each specimen

and bears an elongate membrane distally on the ventral margin of the arm. A much smaller membrane is

present on the dorsal margin in one specimen, but

not the other. The left ventral arm in each specimen

is not modified. It is assumed that the very similiar

A. hoylei would have a comparable condition.

Therefore, within the genera Abraliopsis and Watasenia all male members seem to have similarly modified right ventral arms although varying slightly between species. In the genus Abraliopsis two species, A.

affinis and A. falco, have highly modified trabeculate

protective membranes on the left ventral arm in

males; a third species, A. gilchristi has the left ventral

arm slightly modified by the presence of large

trabeculae, but with a reduced or absent protective membrane (Voss 1967). These three species

are placed in the subgenus Micrabralia. In A. felis,

A. pfefferi, presumably A. hoylei (all subgenus Abraliopsis), and W. scintillans the left ventral is not

modified (rarely, in A. felis both ventral arms are

similarly modified). Therefore, the sexual modification of the ventral arms in W. scintillans is similar

to these species of Abraliopsis; however, these should

not be included in the same subgenus for the following reason: All species of Abraliopsis have two rows

of hooks on each tentacular club. This is an extremely

important feature which is of generic value (all

species of Abralia have a single row of hooks, all

species of Enoploteuthis have two rows of hooks). W.

scintillans has a single row of hooks on each tentacular club and, therefore, is clearly separable from

all members of the genus Abraliopsis.

The tips of both ventral arms in W. scintillans bear

several greatly enlarged photophores. This is a highly

characteristic feature which is shared by all members

TABLE

Characters

Sex

M.L

M.W

F.L.

F.W.

Arm L. I

II

III

IV

A.C. Hooks I

II

III

IV

Tent. L.

Club L.

Club H.F.

3.—Measurements in mm for Abraliopsis

8024

9

29

8

18

22

12

14

13

17

16

16

17

13

30

6

3/3 3/3

8024

31

9

20

22

13

17

16

18

17

18

18

L R

13 11

36

7

3/2 3/3

felis McGowan and Okutani,

Station Numbers of the

9249

S

41

12

28

34

18

21

22

23

16

18

17

L R

14 —

32

8

3/3 3/3

7280

9

41

13

27

33

19

23

22

23

44

9

7221

9

42

11

26

34

18

21

21

23

16

18

18

15

48

8

3/3 —

VELERC

7221

9

45

13

28

34

18

22

21

25

17

19

18

15

—

)

7221

IS

40

11

25

30

21

21

24

15

17

L R

15 10

43

3/3 3/3

1968.

7221

$

40

11

26

28

18

19

20

22

L R

12 10

37

5

3/3 3/3

7221

3

40

10

26

29

18

20

21

22

16

L R

14 12

42

6

3/4 3/3
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

of Abraliopsis, but no other groups. Therefore, at

present, the status of Watasenia as a separate genus

or a subgenus of Abraliopsis rests purely on personal

preference, but it is clear that Watasenia scintillans

remains in an isolated position and that A. felis belongs in the subgenus Abraliopsis of the genus Abraliopsis.

DISTRIBUTION.—Specimens of Abraliopsis felis over

20 mm M.L. are well represented from zones 1, 2, 5,

and 6, having respective frequencies of 0.66, 1.13,

0.50, and 0.63 specimens/hour. In zone 7 the frequence of captures is significantly less, being 0.22

specimens/hour. Specimens less than 20 mm M.L.

have a similar occurrence, but have a greater frequency of capture in the more oceanic zones 5 and 6.

McGowan & Okutani (1968) record A. felis from

about 43° N to about 27° N along the coast of Oregon, California, and Baja California. Apparently, the

southern limit for the species is off Baja California.

Juvenile specimens taken by the SWAN, which appear to belong to this species, indicate that the

species may extend westward to at least 147° W at

26° N.

Subfamily PYROTEUTHINAE Pfeffer, 1912

DIAGNOSIS.—Nidamental glands present. Mantle,

head, and arms without photophores. Photophores

present on viscera and embedded in tentacular stalks.

Posterior margin of fins convex.

Genus Pyroteuthis Hoyle, 1904

DIAGNOSIS.—Left oviduct present, but reduced.

Hooks present on tentacular clubs.

TYPE SPECIES.—Enoploteuthis margaritifera Riippell, 1844.

Pyroteuthis addolux, new species

PLATES 3B; 4I-Q,S

DESCRIPTION.—The mantle is conical and tapers

rapidly to a posterior point. The tip of the mantle

consists of little more than an integumentary layer

which covers the pointed conus of the pen. The

mantle wall, for most of its length, is thick and muscular. The free margin projects in the area of the

ventral locking-cartilages and in the nuchal region.

The fins are separate and each is nearly circular

in outline, having broad, free anterior and posterior

lobes. The fin length is about 40-50% of the mantle

length, and the combined fin width is about 70-80%

of the mantle length.

The funnel is short; it reaches anteriorly to approximately the level of the midpoint of the lenses.

The funnel locking-cartilage has a straight groove.

The dorsal pad of the funnel organ has in inverted

V-shaped pad which bears median ridges on each of

the lateral arms and a very small anterior papilla.

The ventral pads are oval and extremely large. A

funnel-valve is present.

The head is large; its width is greater than that

of the mantle. The large eyes occupy the entire lateral

sides of the head. A distinct anterior sinus on the eyelid is lacking. A thick "olfactory" papilla is present

on either side of the head at the posterior end just

lateral to the funnel; the papilla curves forward in an

anteromedial direction. A nuchal crest and nuchal

folds are lacking. The nuchal cartilage is oval and

bears a median ridge with a central groove.

The arms are robust; they are about 50-60% of

the mantle in length and in the order of 4 = 3 = 2>1.

Large trabeculate protective membranes are present

on the ventral borders of arms I—III, but are greatly

reduced on the dorsal borders. Arms IV have low

membranes on their dorsal borders, but none on their

ventral borders. Arms I—III all have well-developed

aboral keels; arms IV have large, lateral keels. The

armature is biserial throughout and consists of hooks

over most of the arm length, although there are

suckers basally and distally. The hooks of arms IV

are considerably smaller than those of the other

arms. The hooks have no trace of an aperature and

a small secondary cusp is present beneath the major

one. This secondary cusp is greatly elaborated in the

proximal hooks of the hectocotylus. There appears to

be considerable sexual dimorphism in the armature;

however, since only a single male was examined, this

will need confirmation. The male lacks distal suckers

on arms II and III, while females have approximately 4-9 suckers. The male also has a slightly

greater number of hooks (21-23 hooks) on arms II

and III than do females (14-17) and a lesser number of proximal suckers. The structure of the suckers

appears to be similar on all arms except the dorsal

pair. The first 2-6 suckers in the distal series of arms

I are typical, but the following suckers have lost the

outer chitinous ring and are more globular. The first
NUMBER 9 7 23

few of these latter suckers are slightly enlarged over

the adjacent typical suckers. The globular suckers

have smooth rings. The proximal suckers from arms

IV have approximately 10 teeth around the margin

of the inner chitinous ring. The teeth of the distal

margin are rounded and separate from one another,

while those of the proximal margin are truncate and

closely adjacent, almost fused to one another.

The right arm IV of the male is hectocotylized by

the presence of a short semicircular membrane on the

ventral border of the arm about one-third of the arm

length from the tip of the arm. The 2 rows of hooks

proximal to the membrane are more widely separated than in the other arms.

The tentacles are moderately long and bear short,

unexpanded clubs. The club, on its proximal twothirds, is bounded by very low, trabeculate, protective

membranes. The manus has a single row of 3-5 hooks

dorsal to which is a slightly irregular row of relatively large suckers, and ventral to it is a row of

small suckers. The remainder of the club distally

bears 4 rows of suckers that gradually diminish in

size. There is a distinct carpal cluster that consists of

3 suckers and an equal number of pads. The largest

suckers of the club each have about 13 broadly separated, blunt teeth around the entire margin of the

inner chitinous ring, with those of the proximal margin distinctly larger than those of the distal margin.

There are no photophores present on the surface of

the mantle; however, a number of light organs are

situated on the visceral sac. A large photophore is

located on either side of the anus. A transverse series

of 3 photophores is located on the midline between

the gills; the median member is much the largest. A

large round photophore is located at the base of each

gill. Posterior to the median, ventral mantle artery

lies a single round photophore that is followed farther

posteriorly by another median photophore. The latter

2 photophores are often bilobed. There is a final

median, round photophore located near the tip of the

body at approximately the level of the most posterior

edge of the fin.

Twelve photophores are present on the bulbus of

the eye: 3 are situated on the anterior margin near

the bases of arms II and III, another is immediately

posterior to the lens. On the anteroventral margin

of the bulbus, there is a series of 5 large photophores

that are somewhat staggered in position. The second

photophore from the posterior end of the series is

the largest. Anteriorly and medially from this photophore is an additional series of 3 very small lightorgans.

A series of 7 photophores is imbedded in the tentacular stalk. Near the base of the tentacle, the stalk

is bent sharply. This bend is easy to locate and is a

permanent feature of all members of the subfamily.

At the distal end of this area is a round photophore

that is deeply buried in the stalk. Shortly distal to this

is a smaller round photophore that lies more superficially near the aboral surface of the stalk. Near the

proximal end of the carpal cluster is another round

photophore located near the aboral surface of the

stalk. Approximately evenly spaced between these

proximal and distal photophores are 4 elongate light

organs near the aboral surface of the stalk.

The dorsal surface of the mantle, head, and arms

is covered with large, reddish brown chromatophores;

smaller chromatophores of the same color are present

on the ventral surfaces. Most chromatophores overlie

a silvery, iridescent layer.

The buccal connectives attach to the dorsal borders

of arm IV.

The eggs of the gravid females measure 1.0 mm in

in diameter. The spermatophores' of the male are

short, compact, and lack coils in the ejaculatory

apparatus.

TYPE LOCALITY.—31 °39' N, 133° 16' W, eastern

North Pacific Ocean.

LOCATION OF TYPE.—University of Southern California, U.S.C. Hancock collections, AHF Cephalopod

Type No. 2.

DISCUSSION.—In addition to Pyroteuthis addolux,

there is only one other species, P. margaritifera (Riippell 1844), in the genus. Hoyle (1886) assigned 2

specimens captured in the South Pacific (4°21'S,

129°7'E, and 0°33'S, 151°34'W) to P. margaritifera.

He did not describe the animal, but merely illustrated

the club. Pfeffer (1912) felt that these specimens belonged to a new subspecies, P. m. oceanica, since the

illustration of the club showed only 3 hooks on the

manus and only 2 suckers in the carpal cluster. I examined specimens from the South Pacific which agree

with P. margaritifera from the Atlantic in the numbers of hooks on the manus and the number of suckers in the carpal cluster and in all other discemable

features. The craracters Pfeffer referred to are unusual for P. margaritifera, but almost certainly are

within the range of variation for this species. There-
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

fore, it does not seem advisable to retain P. m.

oceanica as a separate taxon.

Joubin (1924) described a form of P. margaritifera

from the Atlantic Ocean which he named P. m. aurantiaca which supposedly had 2 more occular photophores than does the typical P. margaritifera. Joubin's illustration, however, shows only typical photophores. Voss (personal communication) has examined the type which is deposited in the Monaco Museum and informs me that the eyes are indeed normal

for the species and that P. m. aurantiaca has no taxonomic validity.

The specimens of P. addolux are very similar to P.

margaritifera, but differ in a number of important

features. The most distinctive character is the presence of an additional photophore on the tentacular

stalk in P. addolux, giving a total of 7 photophores

compared with 6 in P. margaritifera. The long fleshy

pad situated on the hectocotylus of P. margaritifera

is quite different from the shorter, more membranous

pad of P. addolux. The hooks at the proximal end of

the hectoctylus are much larger and of a different

shape in P. margaritifera (see Plate 4R, S) . These

hooks are also more numerous in P. margaritifera. I

have counted the hooks proximal to the pad on 12

males from the Atlantic and South Pacific oceans and

find the following variation in P. margaritifera:

Number of hooks 13 14 15 16

Number of specimens 1 7 3 1

TABLE 4.—Measurements in mm for Pyroteuthis addolux, new species.

Characters

Ships and Station Numbers

SWAN VELERO

27A-66

Sex

M.L

M.W

H.W

F.L

F.W

Arm L. I

II

HI

IV

Tent. L

Club L

Club H

Carp. S

A.C. Dist. S. I

II

III

IV

A.C.Prox.S. I

II

III

IV

A.C. Hooks I

II

III

IV

(Holotype)

L

3

3

10

0

0

12

4

5

4

0

18

22

23

25

30

8

10

14

22

14

17

17

16

40

5

R

3

3

9

0

0

6

4

5

3

0

18

21

22

25

13AS6

L

4

3

18

8

6

21

8

8

4

2

17

15

15

20

9

45

14

17

19

34

22

26

26

26

59

10

R

5

3

22

9

8

22

9

7

4

1

16

15

17

21

8238

9

20

6

8

9

14

10

11

12

11

28

4

3

3

10+

5 +

3 +

15

11

9

5

4

15

15

16

18

R

4

3

12

3 +

4

16

9

9

5

3

15

15

17

19

10838

9

23

8

8

10

16

11

12

12

12

5 +

—

9

8

4

2

15

15

15

10

7

—

2

14

14

19 17
NUMBER 9 7 25

the single male of P. addolux has only 10 hooks. At

the tip of the hectocotylus, P. addolux has 6 small

suckers while P. margaritifera usually has none, although a maximum of 2 small suckers was observed

on the 12 males examined. The spermatophores in P.

addolux also seem to be fairly distinctive, but this

cannot be certain until more material can be studied

since spermatophores are sometimes subject to considerable variation.

DISTRIBUTION.—Only 20 specimens of P. addolux

have been taken during the VELERO program. Of

these 20, however, only 1 specimen came from the

inner 2 basins. Nine specimens were captured in zone

3, 1 specimen in zone 5, 7 specimens in zone 6, and 2

specimens in zone 7. In addition to these, 2 specimens, including the type, were captured by the R/V

SWAN between southern California and the Hawaiian Islands at 147° W, 26° N and 133° W,

32° N.

The spotty occurrence off California, the nearly

complete absence of P. addolux from the inner basins,

and the capture of specimens from the Eastern North

Pacific Central Water indicates that this species is

probably mid-oceanic and is only rarely present in the

inshore waters over the continental borderland.

Genus Pterygioteuthis Fischer, 1896

DIAGNOSIS.—Left oviduct absent. Hooks not present on tentacular clubs.

TYPE SPECIES.—Pterygioteuthis giardi Fischer,

1896.

Pterygioteuthis gemmata Chun, 1908

PLATES 3A; 4A-H

Pterygioteuthis gemmata Chun, 1908, p.87; 1910 p.108,

pl.13, fig.3; pl.14, figs.4,5,9; pl.15, figs.2,3,6-12; pl.16, figs.

1,2,5,7-19.—Pfeffer, 1912, p. 194.—Thiele, 1921.—Voss,

1967.

DESCRIPTION.—The mantle is cylindrical anteriorly, but conical posteriorly. The mantle wall is thick

and muscular. The pen extends posterior to the fins

and terminates in a sharp point. The free margin of

the mantle project distinctly in the nuchal region and

at the mantle locking-cartilages.

Each fin is large and nearly circular in outline and

has broad, free anterior and posterior lobes. The fin

length is about 40-50% of the mantle length, and the

combined width is about 70-80% of the mantle

length.

The funnel is moderate in size, reaching to the level

of the midpoint of the eyes. Each funnel lockingcartilages bears a straight median groove. The dorsal

portion of funnel organ is an inverted V-shaped pad

with a small anterior papilla and a low ridge on each

lateral arm. The ventral pads are roughly oval in

shape. A large funnel valve is present.

The head is large and has enormous eyes which occupy the entire lateral portions of the head. The eyelid has only a slight indication of an anterior sinus.

An "olfactory" papilla is present at the posteroventral

edge of each eye as a triangular flap which is swollen at the tip. The olfactory nerve passes along the

posterior edge of the flap.

The arms are short and robust. The arm formula is

3>4=2>1 . Extremely well-developed trabeculate

protective membranes are present on the ventral

borders of arms I—III and very weakly developed

ones are present on the dorsal borders. Arms IV in

females have low, fleshy, protective membranes that

are particularly thick near the bases of the arms.

Large aboral keels are present on arms I—III, but

are best developed on arms III. Arms IV have large

lateral keels. The arms have biserial armature consisting mostly of suckers, but with 3-7 hooks on the

midportion of the ventral row of each arm I—III.

Generally there are 2-4 hooks on each arm I, 4—5 on

each arm II, and 5-7 on each arm III, although the

maximum range is slightly greater than this (Figure

7). The hooks are considerably larger in males. The

suckers of arms IV are much smaller than those of

the other arms. The proximal suckers are very small

but are gradually larger in size over the first two

thirds of the arm, then are progressively smaller to the

tip. The larger suckers on all arms possess approximately 6-8 long, slender, truncate teeth on the distal

margin of the inner chitinous rings. The hooks are

low with broad bases and possess a rather large but

irregular secondary cusp beneath the primary one. No

remnant of the sucker aperture remains.

The left arm IV of the male is hectocotylized. It

bears 2 large fleshy longitudinal ridges (the distal one

is somewhat bilobed) between which is located a

pocket with pleated lateral sides that contains a

toothed plate. The dentition of this plate is slightly

variable, but usually has a proximal portion with 5

cusps, then a large single cusp, and a distal set of
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

b

3 4 5

No. of hooks

4 5 6 " 7

No. of hooks

Total no. of hooks

FIGURE 7.—Comparison of numbers of arm hooks between specimens of Pterygioteuthis gemmat a from the North Atlantic Ocean (white) and from the North Pacific Ocean off California

(black). Crosshatched area represents overlap. Comparisons shown: a, Arms II; b, Arms III;

c, Arms II and III.

2-3 cusps. The distal one-third of the hectocotylus

bears low, protective membranes which embrace

small, paired suckers in the last 15-20% of the arm.

These suckers are fragile and easily lost during capture and preservation. A large lateral keel is present

along the length of the arm.

The tentacles are short and the tentacular stalks

are naked. The club is only slightly expanded, and

possesses moderately developed trabeculate protective membranes; the dorsal membrane is slightly the

larger. A distinct keel is lacking. Each carpal cluster

consists typically of 3 suckers and 3 pads; occasionally

(only 2 suckers may be present. Suckers on the manus

and dactylus are in 4 rows. The suckers, in a tranverse series on the manus, grade in size from the

largest dorsally to the smallest ventrally. The suckers

of the ventral row are roughly half the diameter of

those of the dorsal row. The suckers of the dactylus

are small and equal in size in a transverse series.

Dentition of the inner chitinous ring of each of the

club suckers consists of approximately 10-14 short,

slender, pointed teeth on the distal margin. The

proximal margin is smooth or slightly knobby.

The buccal membrane is very long, purple, and has

8 lappets. The inner lining of the membrane is produced into numerous, pale, globular papillae. The

primary buccal membrane connectives attach to the

dorsal borders of arms I, II, and IV, and to the ventral border of arms III. Secondary connectives attach

to the ventral borders of arms I and II.

Each eye possesses 14 photophores on the surface

of the bulbus. On the anterior margin of the eye

there is a series of 3 large photophores followed posteriorly by a slightly smaller series of 4 photophores,

the third of which has a distinctly different structure.

Just posterior to the lens is a rather large, single
NUMBER 9 7 27

photophore. Along the ventral medial margin of the

eye are 2 large well-separated light organs and a more

anterior medial series of 4 very small light organs.

Most light organs on the eyes are spectacular even in

preservation, reflecting any color of the rainbow depending on the angle of incident light. Each tentacle

contains 4 light organs buried deep in the aboral tissue of the stalk. One of these photophores lies beside

the carpal cluster; 2 lie close together about onethird of the tentacle length from its proximal end,

and the other organ lies at the base of the tentacle.

Several light organs of various sizes lie within the

mantle cavity. A large photophore lies on either side

of the rectum. A medium-size photophore lies in the

midline on the viscera between the bases of the gills,

while 2 very large ones lie just posterior to the branchial hearts. Farther posteriorly on the midline, beginning at approximately the level of the midpoint of

the fins, there is a series of 3 small photophores of

which the anterior 2 are sometimes completely divided into paired organs; however, more often the

organs are simply bilobed. The most posterior organ

lies near the tip of the pen approximately at the level

of the most posterior point on the fins.

Large, paired nidamental glands are present. Only

the right oviduct, which has a well-developed oviducal gland, is present.

The body, head, and arms are covered by large,

reddish brown chromatophores. There are distinct

silvery iridescent areas on the body and head, although these are rendered nearly invisible by preservation. The sides of the mantle anterior to the fins

seem to have been silvery in life although overlain by

chromatophores. Similar iridescent spots are present

on the posterior-dorsal margin of the head, the posterior-ventral corners of the head, anterior-dorsal

corners of the eyes, and the bases of arms II, III and

IV and the tentacles.

TYPE LOCALITY.—South Atlantic Ocean.

LOCATION OF TYPE.—Berlin Museum.

DISCUSSION.—At present the following 3 species of

Pterygioteuthis are established in the literature: P.

giardi Fischer, 1896; P. gemmata Chun, 1908; and P.

microlampas Berry, 1909. P. giardi is very distinctive

and easily identified by the presence of hooks in both

the dorsal and ventral rows of arms I—III (P. gemmata and P. microlampas have hooks only in the

ventral rows). It can also be separated by the presence of 15 photophores on each eye compared to 14

in the other 2 species, and by the lack of suckers on

arms IV.

Pterygioteuthis microlampas poses a problem. It is

known from only 2 specimens captured near the Hawaiian Islands, both of which are apparently no

longer extant.

In establishing P. microlampas, Berry (1914) listed

4 features that distinguished his species from P gemmata. The slight difference in the arrangement of the

ocular photophores noted by Berry is probably only a

result of the slightly damaged condition of his specimens. The small number of hooks on arms II and III

(3 on each) which was previously thought to fall outside the range of variation of P. gemmata is now

known to be well within the limits of the latter species, although it is somewhat unusual for a Pacific

Ocean specimen. Berry could not locate suckers on

the proximal half of arms IV and considered this a

distinctive feature of P. microlampas; however, the

proximal suckers are often very difficult to observe

on P. gemmata since they are very small and are frequently covered by the fleshy protective membranes.

It seems doubtful that there are any valid differences concerning this feature. The final character

concerns the presence of only 2 suckers in the carpal

cluster of P. microlampas. This feature is unusual,

but is also occasionally shared by P. gemmata. Therefore, the laterature provides no basis for separating

P. microlampas from P. gemmata. The examination

of specimens from Hawaii, however, is necessary before synonymizing the two species.

I examined a number of specimens of P. gemmata

from the North Atlantic and find that they have

fewer hooks on arms II and III than do the California specimens. The total number of hooks on all 4

lateral arms shows almost no overlap (Figure 7c).

This is the only feature, however, that seems to differ.

I recently examined 17 specimens from the South

Pacific and find the hook counts fall bomewhat between the specimens from the North Atlantic and

North Pacific. The mean number of hooks on arm II

is 3.3 for specimens from the North Atlantic, 3.6 for

specimens from the South Pacific, and 4.5 for the

specimens from the North Pacific. The values for arm

III are 4.2, 5.0, and 6.1 for the three areas, and the

mean values for the total counts of all 4 lateral arms

are 15.2, 17.3, and 21.1. The different hook counts,

therefore, seem to reflect geographical variation.

DISTRIBUTION.—P. gemmata has previously been
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 5.—Measurements in mm for Pterygioteuthis gemmata Chun, 1908.

Characters

Sex

M.L

M.W

H.W

F.L

F.W

Arm L. I

II

Ill

IV

Tent. L

ClubL *.

A.C. Hooks I

II

Ill

9603

$

28

8

10

13

22

10

11

13

12

20

4

L R

3 2

4 4

6 6

Station

9603

9

23

7

10

18

8

9

10

9

22

4

L R

3 3

5 4

6 6

Numbers

9605

9

30

9

10

14

22

10

12

13

12

20

5

L R

— 2

4 4

6 5

of the VELERO

9605

S

24

8

8

11

18

9

23

4

L R

2 2

4 4

— 5

9602

9

32

9

11

15

24

12

13

14

12

19

5

L R

4 4

5 5

7 7

8026

S

25

7

9

9

14

9

9

11

10

26

4

L R

3 3

4 4

6 6

recorded from the equatorial Atlantic (Thiele 1921),

from the South Atlantic (Chun 1910; Thiele 1921)

from the Indian Ocean (Voss 1967) and off Hawaii

(Berry 1914, P. microlampas). I have examined

numerous specimens captured by the DANA and

PILLSBURY from areas throughout much of the

north Atlantic and specimens captured by the ELTANIN in the region of the subtropical convergence

in the South Pacific. The species is worldwide in distribution, occurring primarily in warmer waters. For

more details concerning the distribution of P. gemmata see Voss (in manuscript).

In the present study P. gemmata had relatively low

frequencies of capture in the more northern zones of

the area under study (i.e., 0.15, 0.08, and 0.30 specimens/hour for zones 1, 2, and 3, respectively) and

relatively high frequencies of capture in the more

southern zones (1.07 and 1.34 specimens/hour for

zones 6 and 7, respectively). This species clearly

shows a preference for warm water.

Family OMMASTREPHIDAE Steenstrup, 1857

DIAGNOSIS.—Arms with biserial suckers. Tentacular clubs with tetraserial suckers on the manus and

usually on the dactylus. Buccal connectives attach to

the dorsal borders of arms IV. Funnel locking-cartilage with an inverted T-shaped groove. A muscular

bridge anterior to the funnel locking-cartilage passes

from the funnel to the ventral surface of the head.

Genus Ommastrephes Orbigny, 1835

DIAGNOSIS.—An enlarged tooth present in each

quadrant of the largest tentacular suckers. Protective

membranes on lateral arms greatly enlarged, particularly on arms III. Arm tips not attenuate; arms

containing 50-70 suckers. Funnel and mantle cartilages not fused.

TYPE SPECIES.—Loligo Bartramii Lesueur, 1821

Ommastrephes bartramii (Lesueur, 1821)

PLATES 5B, 6C; 7J-R

Loligo Bartramii Lesueur, 1821, p.90, pi. 7.

Ommastrephes Bartramii.—Orbigny, 1839, p.347, in F6russac

and Orbigny, Loligo pl.2; pl.21, fig.5; Ommastrephes pl.2,

figs.11,12.—Berry, 1912b, p.298; pl.47; pl.50, figs.4-5;

text figs. 7-8.

Ommastrephes Bartrami.—Steenstrup, 1880, p.73, fig.2; p.79,

fig.3; p.81.

Stenoteuthis bartrami.—Pfeffer, 1912, p.465; pls.35-36;

pl.39, figs. 1,2.—Sasaki, 1929, p.289, pl.l, fig.8; pl.24,

figs. 1-3; text fig. 139.
NUMBER 9 7 29

DESCRIPTION.—The mantle is long and slender

(width about 20% of length). Anteriorly it is nearly

cylindrical, but at the level of the anterior edge of

the fins, it begins to taper rapidly and terminates in a

blunt point. A median ventral keel extends from the

level of the conus forward for about 20% of the

mantle length. The free anterior margin of the mantle

has no distinct projections. The mantle wall is extremely thick and muscular.

The fins are about 40%-45% of the mantle in

length and about 70%-75% of the mantle in width.

They are rather sharply angled laterally and have

free anterior lobes.

The funnel is very muscular and rather short. The

funnel locking-cartilage is complex. The groove has

roughly the shape of an inverted T. The dorsal pad of

the funnel organ is very large and has an inverted Vshaped appearance. A small anterior papilla can

sometimes be detected. The ventral pads are large

and oval. A large funnel valve is present. The funnel

is fused to the head forward of each locking-cartilage by a muscular bridge.

The head, approximately the same width as the

mantle, has large eyes which occupy almost its entire

lateral sides. Each eyelid has a large anterior sinus. A

nuchal crest with 4 folds on each side is present; the

most medial folds embrance the funnel. Posteriorly

the folds all converge. A low oblique ridge on the posterior extension of the second fold from the funnel is

apparently the "olfactory" organ. A small "window"

is situated ventral to each eye, but there is only a

slight trace of a dorsal "window." The funnel groove

contains a large foveola at its anterior end. Seven

folds are present within the foveola, and 2-4 side

pockets are located lateral to it on either side.

The arms are short (30% to 40% of the M.L.) and

powerful. Arms I—III all have well-developed aboral

swimming keels. In addition, arms I possess accessory

lateral keels. Large lateral keels are present on arms

IV. All arms have well-developed trabeculate protective membranes. The membranes on the ventral margins of arms I—III are greatly enlarged, but are most

extensive on arms III. The elongate trabeculae extend to the margins of the membranes.

The arms have biserial suckers. Arms I—III bear

from 50-55 suckers while arms IV (excepting the

hectocotylus) have about 65 suckers. The dentition

of the suckers is somewhat variable. The basal suckers

from arms III have about 16-18 pointed teeth (some

teeth have irregular shapes) on the distal threefourths of the ring; the proximal margin is smooth or

finely toothed. By the fifth transverse row of suckers

from the base of the arm, the teeth are more regularly

shaped and the entire ring is clearly lined with about

28-32 teeth. The largest teeth occur on the distal

margin; however, these teeth frequently alternate

with smaller teeth. The outer ring is made up of

roughly 75 elongate slats; each, with only a few exceptions, extends unbroken from the inner to the

outer margin of the ring. The slats on the distal portion of the ring bear small knobs at their bases. At the

fourteenth row the proximal portion of the inner ring

is smooth, and the distal portion bears about 7 large,

pointed teeth that may or may not alternate with

very small teeth. The largest suckers of arms IV are

just slightly smaller than those of the first arms which,

in turn, are slightly smaller than those of arms II and

III.

The left arm IV is usually hectocotylized in males,

although occasionally the right arm is modified instead. The largest specimen available (303 mm

M.L.) is immature and the hectocotylus is probably

only partially formed. All specimens available show

some signs of hectocotylus formation which begins

with the loss of suckers at the extreme tip of the arm.

The largest specimen has 27 suckers on the proximal

part of the arm which is followed by a suckerless area

which would normally carry 7 suckers. Distal to this

is a single row of 6 suckers on the dorsal margin, and

the remaining portion of the arm lacks suckers. Distal

to the proximal series of suckers, the protective membranes do not appear to have trabeculae. It is likely

that this is not a completely mature hectocotylus.

The tentacles bear an aboral keel which runs along

the tentacular stalk and is continuous with the dorsoaboral keel of the club. Trabeculate protective membranes occur along both margins of the oral face of

the tentacular stalk for most of its length; however,

the membrane of the dorsal margin is much more extensive. These membranes extend to the tip of the

club along the lateral margins. The clubs are expanded and elongate. The suckers of the manus are

arranged in 4 longitudinal rows except at the proximal end where they are irregularly aligned and extend onto the tentacular stalk for a short distance.

The suckers on the dactylus are arranged in 4 rows

which are continuous with those of the manus. The

dactylus suckers grade in size from the largest on the
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ventral margin to the smallest on the dorsal margin.

At the tip of the dactylus, there is a pad of 17

smooth-ringed suckers. At the proximal end of the

manus, along the ventral margin, is a row of 2-4

smooth-ringed carpal suckers alternating with 2-4

pads. Proximal to this series lie 5-6 toothed suckers.

The suckers of the medial rows on the manus are

nearly 3 times the diameter of the marginal suckers.

Numerous fingerlike papillae arise from the fleshy

outer edges of these medial suckers. These papillae

vary in size and thickness and are often joined at

their bases in groups of 4-5, each set looking like a

human hand. The papillae are also present on the

marginal suckers but are located in only 2 tufts, one

on each dorsolateral corner with the biggest tuft

nearest the margin of the club. Papillae are not present on the dactylus suckers. On 2 of the largest suckers of the manus 123 and 128 slats were counted in

the outer ring. These slats have small knobs at their

bases.

The inner rings of the largest suckers of the manus

have 4 enlarged teeth, one in each quadrant. Occasionally an additional enlarged tooth may be observed. There are generally 7-10 smaller teeth, which

may vary considerably in size, in each quadrant between the large teeth. The marginal suckers have

about 25-30 teeth with those of the distal border

being much larger than those of the proximal border.

Generally, minute teeth may be observed alternating

with the distal teeth. The suckers of the dactylus have

the same dentition as the marginal suckers on the

manus.

The buccal membrane has 7 lappets. The buccal

connectives attach to the dorsal borders of arms I, II,

and IV, and to the ventral borders of arms III.

The animal is speckled with numerous, small, reddish brown chromatophores. The ventral surface of

the mantle bears an elongate, sharply delineated,

silvery strip that extends from the anterior edge of

the mantle posteriorly to the level of the anterior edge

of the fins. The ventral surface of the head also has a

silvery background that extends down arms IV. The

dorsal surface of the mantle and the dorsal midportion of the fins have a deep-purple coloration that

terminates very abruptly laterally. The dorsal surface

of the head is similarly colored, as are the dorsalmost

portions of arms I—III and tentacles. The lateral

sides of the head and mantle have a silvery, iridescent

layer benath the chromatophores.

Numerous photophores are imbedded in the musculature near the ventral surface of the mantle. These

photophores are not visible in preserved specimens

unless the skin is removed, and often it is necessary

to chemically macerate the muscle tissue. The photophores are small, closely packed, and extremely irregular in shape. Along the ventral midline of the

mantle, there is a strip extending from the conus of

the pen to nearly the free margin of the mantle in

which the number of photophores is reduced. In the

posterior quarter of this strip photophores are lacking.

The concentration of photophores is greatest near

the free margin of the mantle. This group of photophores extends along the lateral sides of the mantle

margin becoming progressively less numerous. There

are no photophores dorsally.

An irregular patch of photogenic tissue is situated

on either side of the head at the point of attachment

of the muscular bands which pass from the ventral

surface of the head to the funnel near the lockingcartilages. At least several photophores are located in

the integument and connective tissue immediately anterior to the foveola. Unfortunately, in the specimen

examined, the integument was stripped away from

most of this area; therefore, there may be more

photophores in an intact specimen. Another extremely irregular patch of photogenic tissue is situated

on the head near the base of each ventral arm. There

is also a long band of photogenic tissue that extends

from the base of each ventral arm nearly to the arm

tip within the base of the lateral keel.

Within the funnel there is a patch of photophores,

clustered approximately into a V-shape, that is located above (dorsal to) the dorsal pad of the funnel

organ.

With growth, the patches of photophores on the

ventral surface of the head increase in size and

change somewhat in shape. The specimen described

and illustrated here (Plate 6c) has a M.L. of only

210 mm; therefore, the size of the patches on the

head, at least, can be expected to be more extensive

in mature specimens.

DISCUSSION.—Three species are presently recognied in the genus Ommastreph.es: O. bartramii (Lesueur 1821), O. pteropus (Steenstrup 1855), and O.

caroli Furtado, 1887. The latter two are known only

from the Atlantic Ocean, whereas O. bartramii has

been reported from the Atlantic, Pacific, and Indian

oceans. The broad distribution of O. bartramii indicated that it might possibly represent a species complex. I have carefully compared the present speci-
NUMBER 9 7 31

mens against specimens of O. bartramii captured off

Florida and find that the eastern Pacific specimens

unquestionably differ from those of the Atlantic. The

tentacular clubs of the Pacific specimens are distinctly broader than those from the Atlantic. The

suckers of the Pacific specimens are about 20% larger

than those from the Atlantic specimens of the same

mantle length. A specimen of 303 mm M.L. from the

Pacific was immature; the hectocotylus was only partially formed and there were no spermatophores

present in the genital tract. A specimen of 302 mm

M.L. from the Atlantic had a completely formed

hectocotylus and spermatophores. There were several

minor differences in the structure of the beaks, but

it is not known whether these fall within the normal

variation for either form.

It is possible that a more complete comparison of

all stages of development will reveal additional differences. At present, however, the differences do not

seem sufficient to indicate anything more than geographical variation.

There is considerable confusion concerning the relationship between O. bartramii and O. caroli. It is

even possible that they represent different growth

stages of the same species. If this is the case, it is

curious that the distributions of O. caroli and O. bartramii do not more closely coincide. O. bartramii can

be distinguished from O. pteropus by the presence of

4-6 suckers on the tentacular stalk proximal to the

first knob of the locking apparatus compared to 0-2

suckers in O. pteropus. The silvery strip on the ventral surface of the mantle is distinctive of O. bartramii, as is the photophore pattern. The photophore

pattern in O. pteropus was described by Roper

(1963) and Clarke (1965). The most obvious distinguishing features of the photophore pattern in this

latter species are the presence of a large oval patch

composed of densely packed photogenic spherules

TABLE 6.—Measurements in mm of Ommastrephes bartramii (Lesueur, 1821).

Characters

Sex

M.L.

M.W.

H.W.

F.L.

F.W.

Arm L. I

II

Ill

IV

A.C. I

II

Ill

rv

Tent. L.

Club. L.

Club. S.

Dact. T.S.

Carpal F

$

211

44

42

91

158

66

83

81

78

51

55

55

L R

66» 68

131

72

143

16

L R

2/3 3/2

Stations

Near Guadalupe Island

S

217

44

47

90

157

63

81

78

77

50

54

57

L R

67 51*

123

70

136

16

L R

4/4 3/3

$

240

51

54

108

178

77

96

96

95

51

53

54

L R

48* 66

190

92

137

17

L R

2/2 2/2

$

303

65

137

220

%

123

114

L

117*

49

55

53

L

33*

230

119

131

17

L

2/2

R

121

R

67

R

2/2

Atlantic

off Florida

*

302

65

70

146

220

95

119

117

L R

141* 131

51

57

58

L R

19* 73

200

110

140

17

L R

2/3 3/2
32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

situated on the dorsal surface of the mantle beneath

the integument near the nuchal region, and the presence of small, spherical photophores, which are

broadly spaced, on the ventral surface of the mantle.

DISTRIBUTION.—There are a number of records of

O. bartramii from cold water (Pfeffer 1912, Grimpe

1925, Berry 1912b) ; however, as there has been considerable confusion regarding the identification of

this species these records retain an element of doubt.

O. bartramii is common in tropical and subtropical

waters, and I consider that this is the usual habitat

for the species, although, being powerful swimmers, it

would not be unexpected to find stragglers occasionally in high latitudes. In the present study, 5 specimens were captured at one night-light station off

Guadalupe Island.

Genus Symplectoteuthis Pfeffer, 1900

DIAGNOSIS.—Funnel locking-cartilage with typical

ommastrephid inverted T-shape, but fused in its midportion to the mantle locking-cartilage.

TYPE SPECIES.—Loligo oualaniensis Lesson, 1830.

Symplectoteuthis luminosa Sasaki, 1915

PLATES 5A; 6A,B; 7A-I

Symplectoteuthis oualaniensis.—Watase, 1906, p. 195.

Symplectoteuthis luminosa Sasaki, 1915, p. 144, text fig. 4, pl.4,

figs. 7-13; 1916, p.106; 1929, p.293, text fig. 140, pl.24,

figs. 4-5.

Eucleoteuthis luminosa.—Berry, 1916, p.60.—Okada, 1927,

p.4.

DESCRIPTION.—The mantle is long and slender

(width 15-20% of length); anterior to the fins it is

nearly cylindrical, but posteriorly it tapers to a point.

The free margin is only slightly produced near the

ventral locking-cartilages and not at all in the nuchal

region. The mantle wall is extremely thick and

muscular.

The fins are about 45-50% of the mantle in length

and about 55-60% in total width. They are muscular

and attenuate posteriorly. The fins extend to the posterior tip of the mantle. The anterior lobes are free.

The funnel is short and muscular. It is fused to the

ventroposterior side of the head by a narrow muscular

band extending forward anterior to the funnel locking-cartilage. The funnel locking-cartilage has an inverted T-shaped groove which is fused with the mantle locking-cartilage in the posterior portion of the

longitudinal groove. The dorsal pad of the funnel

organ has an inverted V-shape with only a slight trace

of an anterior papilla. The lateral arms are very long

and extend onto the funnel retractor muscles. The

ventral pads are rather slender and extremely elongate. The anterior and posterior extent of the ventral

and dorsal pads coincides. A large funnel valve is

present.

The head is approximately the same diameter as

the mantle. The eyes are large and occupy almost the

entire lateral sides of the head. A large anterior sinus

is present on each eyelid, and there is a small "window" in the skin above and below each eye. A nuchal

crest bears 4 folds which are connected posteriorly.

An oblique ridge, the "olfactory" organ, lies on the

posterior extension of the second fold from the funnel. A distinct funnel groove is present. The foveola

has 8 low ridges on its anterior wall. Only a single,

small, side pocket can be found on each side of the

foveola. The nuchal cartilage broadens at the anterior

end and has a median ridge with a central groove.

The arms are short (30-40% of the M.L.) and in

the order 2 = 3 = 4>1. Arms I-III all have welldeveloped aboral keels. An additional keel is present

along the lateral side of each arm I. Large lateral

keels are present on arms IV.

Trabeculate protective membranes are well developed on all arms. On arms I-III, the membranes

on the ventral margins are broader than on the dorsal

margins. These are most fully developed on arms III

where they are wider than the greatest width of the

aboral keel of the same arm. The long, slender, supporting trabeculae extend to the margin of the membrane. The trabeculate membranes of arms IV are

equally developed on both margins.

The suckers of the arms are biserially arranged and

total 43-48 on each of arms I-III and 56-58 on

arms IV. The basal few suckers on each arm generally have 3-5 broad, truncate teeth on the upper

margin of the chitinous ring and a smooth lower

margin. Each broad tooth is the homologue of 2

pointed teeth on the more distal suckers. At rows 5

and 6 on the arms, the teeth have approximately 9-12

sharply pointed teeth on the distal margin that merge

with 1-3 broad, generally truncate, teeth on each

lateral margin; the proximal margin is smooth. The

teeth are more or less of a uniform size, except for an

occasional small tooth situated near a larger tooth, al-
NUMBER 9 7 33

though more frequently these are fused, forming a

single, somewhat lopsided tooth. The number of

teeth is somewhat irregular; some of the large suckers

of arms II may have small broadly spaced teeth on

the proximal half of the sucker ring. This appears to

be a variable feature. Suckers of the seventeenth and

eighteenth rows each have 5-6 long pointed teeth on

the distal half of their inner rings. The largest suckers

of arms IV are roughly half the diameter of the

largest suckers of arms II. The largest suckers of arms

I and III are only slightly smaller than those of arms

II.

The tentacles are short and muscular.. An aboral

keel extends the full length of the tentacular stalk

and onto the club where it is continuous with the

dorsoaboral keel of the dactylus. The clubs are only

slightly expanded and bear roughly 115-123 suckers.

The suckers of the manus are arranged in 4 longitudinal rows except at the proximal end where the

number of suckers rapidly diminishes to a few irregularly arranged suckers that extend onto the tentacular stalk. There are no carpal suckers although a

single carpal knob was found near the base of the

manus close to the dorsal border of both clubs in one

specimen, but not in the other. The suckers in the 2

median rows on the manus are greatly enlarged, the

largest being 2/2 to 3 times the diameter of the marginal suckers. These suckers each bear a single enlarged tooth on the distal margin of the sucker ring.

The remaining margin has low jagged edges which

form teeth lacking a regular pattern or shape. The

dentition becomes more regular on the smaller suckers. The marginal suckers bear about 30-35 teeth

around the entire circumference of the chitinous ring.

The teeth of the distal margin are slender, pointed,

and roughly alternate between long and short sizes.

The teeth on the proximal margin are all short and

uniform. The dactylus has 4 longitudinal rows of

suckers which are continuous with those of the

manus. The suckers grade in size in a transverse series from the largest in the ventral row to the smallest

in the dorsal row. The sucker dentition is the same as

in the marginal suckers of the manus, although the

number of teeth decreases as the size of the suckers

diminishes near the tip of the dactylus. There is a

small pad of about 7-8 smooth ringed suckers at the

tip of the dactylus.

The buccal membrane has 7 lappets. The buccal

connectives attach to the dorsal borders of arms I, II,

IV, and to the ventral borders of arms III.

This species has a very distinctive photophore pattern. Two broad luminous strips extend almost the

full length of the mantle on either side of the ventral

midline. Each strip consists of 3 separate parts: a

small anterior segment is situated at the free margin

of the mantle; a short gap separates this from the

second part which extends to approximately the level

of the lateral angles of the fins; and this is barely

separated from the third portion which continues to

approximately the level of the conus of the pen. Near

the anterior edge of the mantle are two small patches,

one situated on either side of the ventral midline, but

medial to the anterior portion of the long strips. Also

near the anterior margin, but lateral to each of the

long strips, is a medium-size photophore. There is a

large photophore on the ventral surface of the head

at the base of each arm IV, and another is located on

midventral surface of each arm IV.

Many small photophores, generally invisible without dissection, are scattered over the mantle, funnel,

head, and arms. These form almost a complete line

along the anterodorsal margin of the mantle. They

are scattered irregularly over the ventral surface of

the mantle, but are absent from its dorsolateral surTABLE 7.—Measurements in mm of Sympletocteuthis

luminosa Sasaki, 1915.

Characters

Sex

M.L.

M.W.

H.W.

F.L.

F.W.

Arm L. I

II

III

IV

A.C. I

II

Ill

IV

Tent. L.

Club L.

Club S.

Station Numbers of the VELERO

10673

9

163

33

34

75

97

49

61

57

60

47

44

48

56

78

35

123

10673

9

136

26

30

67

79

42

50

48

48

47

43

48

58

61

29

115
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

face. They are scattered along the dorsal midline

from the anterior mantle margin to the fins. The

inner surface of the funnel has a single photophore

on each dorsolateral side at approximately the level of

the emergence of the cephalic vein. There is a photophore at the funnel attachment of each of the 2

bridles and 2 small photophores are imbedded in the

funnel valve. On the head, a few photophores are

scattered lateral to the base of the funnel. I have

found only a single photophore in the integument immediately in front of the foveola; however, since

most of the tissue there has been torn away, it is quite

likely that at least several others are present in intact

specimens. A small distinct photophore is present at

the medial base of each ventral arm very near the

point where the arms unite. A line of irregularly

shaped photophores extends along the lateral edge

of each ventral arm at the base of the keel in the

proximal half of the arm. There is also a series of

small photophores along the surface of the axial nerve

of each tentacle.

There can be no assurance that all photophores

have been located, since they are extremely difficult

to see. For example, Okada (1927) in a histological

examination found photophores in the fins, but I

have not been able to find them by gross dissection.

The larger specimen examined is a gravid female

with greatly enlarged nidamental glands. The mature

eggs measure just under 1 mm.

The animal is covered with numerous, small, reddish brown chromatophores. The dorsal surface is

darker than the ventral surface, and the lateral sides

of the animal have a distinct silvery appearance.

TYPE LOCALITY.—Off Misaki, Japan.

LOCATION OF TYPE.—In Tokyo Imperial University.

DISCUSSION.—Within the genus Symplectoteuthis

there are, presently, only 2 recognized species: S. luminosa Sasaki, 1915, and S. oualaniensis (Lesson

1830). (A third species very similar to S. oualaniensis has recently been reported (Clarke 1965), but

not described.) The long, luminous strips on the ventral surface of the mande are present on the former,

but not the latter, and easily allow their separation.

The present specimens agree with Sasaki's description except for some differences concerning dentition

of the arm and club suckers, the number of side pockets in the funnel groove, and the number of carpal

knobs. These features, however, are probably all variable. The small photophores buried within and beneath the integument were first reported by Okada

(1927).

DISTRIBUTION.—S. luminosa is known only from

the waters off Japan (Sasaki 1929), as possible record

from the Kermadec Islands (Berry 1916), and from

the California Current (Okutani 1967). It is here

recorded off Guadalupe Island at 29°37' N, 188°49'

W.

Family NEOTEUTHIDAE Naef, 1921

DIAGNOSIS.—Arms with biserial suckers; tentacular

club with 4 rows of suckers on the distal part of the

manus and dactylus and numerous small smuckers at

the proximal end of the manus. Buccal connectives

attach to the dorsal borders of arms IV. Funnel locking-cartilage with a straight, simple groove. Anterior

fin lobes lacking; posterior fin lobes free. Photophores

absent.

Genus Neoteuthis Naef, 1921

DIAGNOSIS.—Fins more than 60% of the mantle

length. Club length more than 60% of the mantle

length. Most of manus bearing many hundreds of

minute suckers.

TYPE SPECIES.—Neoteuthis thielei Naef, 1921.

Neoteuthis species

PLATE 8A-C

DESCRIPTION.—The mantle is cylindrical anteriorly

and gradually tapers to a point posteriorly. The

mantle wall is thick and moderately muscular; anteriorly it is produced slightly at the nuchal and

funnel locks. The pen is visible in the dorsal midline.

The fins are peculiar: they are very elongate (66%

of M.L.), but very narrow (29% of M.L.) ; and they

lack anterior lobes, but possess small posterior lobes.

The posterior attachment of the fins is slightly in advance of the pointed tip of the mantle. The fins are

attached to the lateral sides of the mantle and are,

thus, broadly separated from one another in the

dorsal midline.

The funnel is small: it reaches just past the level

of the posterior margin of the eyes. The funnel locking-cartilage has a simple, straight groove which opposes a straight ridge on the mantle. The dorsal pad
NUMBER 9 7 35

of the funnel organ has roughly an inverted V-shape.

A small papilla is present near the rounded apex. On

each lateral arm of the dorsal pad a ridge is present

that slopes from the lateral margin toward the midline, then gradually back to the lateral margin. The

lateral arms are very long and extend onto the funnel

retractor muscles. The ventral pads are also very

large and have a teardrop shape with the more

pointed end posterior. The funnel is free from the

head laterally.

The head is rounded and bears large eyes which

occupy almost the entire lateral sides of the head. A

sinus is present on the anterior border of each eyelid,

but is extremely small. There is no indication of nuchal folds or a nuchal crest. An "olfactory" papilla is

present on either side of the head immediately lateral

to the funnel. The nuchal cartilage is broader anteriorly and bears a median ridge with a central groove.

The long arms (60-80% of MX.) are rather brittle in preservation, and in the order of 4>3>?2>1.

Arms I—III bear low, broad, gelatinous protective

membranes with gelatinous trabeculae that project

well beyond the membrane. Protective membranes

are barely detectable on arms IV. The arms have

been rubbed, and no evidence of aboral keels remains. Arms IV have low, gelatinous lateral keels.

All arms bear biserial suckers. The suckers are large

except for those on the ventral arms, the largest of

which is less than half the diameter of the largest

suckers of arms III. On arms I—III the suckers are

largest in the middle of the arms, but on arms IV the

suckers are largest near the base of the arms. Although the sucker rings have almost been destroyed

by the fixative, it is still possible to detect about 13-15

apparently semilunar or truncate teeth on the distal

two-thirds of each ring; the proximal portion seems

to be smooth.

The tentacles are long (64% of the M.L.) and

bear very large clubs. The clubs are expanded and

the manus, which tapers gradually onto the stalk,

bears many hundreds of minute suckers arranged in

numerous irregular rows. Approximately 35 suckers

are present in a transverse series at about two-fifths

of the club length from the proximal end. The most

distal of the small manus suckers have very broad

outer rings. Each inner ring seems to bear triangular

teeth around its entire margin, with those of the distal

margin enlarged. The suckers in the middle of the

manus have teeth which are all nearly equal in size.

The suckers decrease in size at the proximal and lateral margins of the manus, but at the distal end the

suckers increase abruptly in size, the largest being

about 4 times the diameter of the smaller suckers. Except for the area where the manus and dactylus blend

together, the dactylus has suckers arranged in 4 longitudinal rows with the suckers of the ventral margin

2 to 3 times as large as the suckers of the dorsal

margin. The medial suckers grade in size between the

marginal rows. Except for an irregular circle of

smooth-ringed suckers at the tip of the dactylus, all

suckers of the dactylus have roughly 26-28 sharply

pointed teeth around the entire margin of each inner

ring. There is a large dorsoaboral keel on the dactylus that extends onto the distal end of the manus.

The buccal membrane connectives attach to the

dorsal borders of arms I, II, and IV, and to the ventral borders of arms III.

DISCUSSION.—The genus Neoteuthis and the family Neoteuthidae were erected by Naef (1921:48).

The only diagnosis, given in a footnote, was based on

3 young specimens which were not given a specific

name. Later in the same year, Naef named Neoteuthis thielei form a description that Thiele (1921) had

given of a small squid from Antarctic waters identified as Octopodoteuthis sp.; therefore, Neoteuthis

thielei becomes the type species of the genus by

subsequent monotypy. The diagnosis given by Naef

and the description given by Thiele greatly lack in

detail. Although Naef s diagnosis states the club has

4 rows of suckers, it seems likely that he merely overlooked the manus suckers since in young specimens

the tentacles are disproportionately small and the

manus suckers are extremely difficult to see. No new

information has appeared on this group since Naef s

work. Fortunately, at Miami there are specimens captured in Antarctic waters by the USNS ELTANIN

that look very similar to Thiele's sketch. Except for

this material, it probably would have been impossible

to correctly identify the California form.

Although Neoteuthis has a very distinctive appearance, it is closely related to the genus Alluroteuthis,

the sole genus of the family Alluroteuthidae. The similarities are: in both, the buccal membrane connectives attach to the dorsal borders of arms IV. Both

genera have very distinctive fins which arise from

the lateral sides of the mantle and are broadly separated from one another in the dorsal midline. The

fins are also slightly subterminal, lack anterior lobes,
36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

and have free posterior lobes. The fins in the 2 genera

are identical, except that they are much longer and

more slender in Neoteuthis. Likewise, the clubs differ only in degree. Alluroteuthis has a cluster of

minute suckers at the proximal portion of the manus

which corresponds to the much more elaborate

manus of Neoteuthis. The appearance of a large

broad head is similar in both groups as in the armature of the arms, including the occurrence of reduced suckers on the ventral arms.

These features are sufficient to indicate the close

relationship between Neoteuthis and Alluroteuthis.

Their families must therefore be considered synonymous, and the more recent family Alluroteuthidae

Odhner, 1923, must be submerged into the older Neoteuthidae.

The present species is very close to N. thielei. The

specimens of N. thielei that are available for comparison, however, are fragmentary and the original description offers little of specific value. Thus, I am unable to determine whether the present form is conspecific with N. thielei and, therefore, must refrain

from giving it a specific name.

Data from the single specimen of Neoteuthis sp.

here described are: Sex, o" ; M.L., 83; M.W., 18;

H.W., 23; F.L., 55; F.W., 24; Arms I, 49; II, 56+ ;

III, 64; IV, 68; Tent. L., 118; Club L., 53.

DISTRIBUTION.—The genus Neoteuthis has been reported from Antarctic waters (Thiele 1921), and

from the Mediterranean Sea (Naef 1921). The

specimen described here came from 30°23' N, 118°

29' W (VELERO Station 10840). Several specimens

are present in the collections at Miami which were

taken at the following localities: 3°30' N, 4°41' E,

Gulf of Guinea (PILLSBURY Station 306); 0°31' S,

ll°02' W and 13°31' S, 18°03' W, South Atlantic

(DANA collections); and 46°30' S, 170°9' W, South

Pacific (ELTANIN collections).

This broad distribution suggests that more than

one species may be present in the genus.

Family HISTIOTEUTHIDAE Verrill, 1881

DIAGNOSIS.—Arms with biserial suckers. Tentacular clubs with suckers generally arranged in more

than 4 rows. Large, anteriorly directed photophores

over the surface of the mantle, head and arms. Left

eye considerably the larger. Buccal connectives attach to the dorsal borders of arms IV. Funnel locking-cartilage with an approximately straight, somewhat broad groove.

Genus Histioteuthis Orbigny, 1839

DIAGNOSIS.—The characters coincide with those of

the family.

TYPE SPECIES.—Cranchia bonnellii Ferussac, 1835.

Histioteuthis heteropsis (Berry 1913)

PLATES 9A ; IOIHO

Melagroteuthis hoylei—Berry, 1912b, p.305, text figs. 13-

16; pl.50, figs. 1-3; pl.51, figs. 1-5; pl.52, figs. 6-7.

Calliteuthis (Meleagroteuthis) heteropsis Berry, 1913, p.75.

Histioteuthis heteropsis.—You, 1969, p.815.

DESCRIPTION.—The mantle is short and broad; it

tapers gradually posteriorly to a blunt tip. The mantle

margin projects slightly in the region of the dorsal

and ventral locking-cartilages. The mantle wall is

very thick, but moderately gelatinous.

The fins are short, about 35-40% of the mantle in

length and about 50-55% of the mantle in total

width. Each fin is roughly semicircular and has free

anterior and posterior lobes. Medially, the fins touch

one another only at the posterior end where they extend slightly past the tip of the mantle. Anteriorly,

the fin insertions are broadly separated from one

another.

The funnel is short. The funnel locking-cartilage

has a broad, simple, generally straight groove that is

deepest in the midportion. This structure opposes a

straight or slightly curved ridge on the mantle, the

anterior end of which is flanked on either side by

additional short broad ridges. The dorsal pad of the

funnel organ has an inverted V-shape with a small

papilla at the anterior apex. The ventral pads are

oval. A funnel valve is present.

The head is broad and asymmetrical due primarily

to the unequal development of the eyes; the left eye

has almost l2/s the diameter of the right. An anterior

sinus is present on each eyelid. A nuchal crest and

nuchal folds are not clearly developed. The nuchal

cartilage is short and much broader anteriorly than

posteriorly.

The arms are broad and short, although slightly

greater than the mantle in length. Aboral keels are

well developed only on the midportion of arms III;

they are represented by faint ridges on arms I and II.
NUMBER 9 7 37

The lateral keels are well developed on arms IV.

Low, thick, gelatinous protective membranes, lacking distinct trabeculae, are present on all arms. The

arms have biserial suckers. The suckers are globular

and have relatively narrow outer rings, and inner

rings with about 8-10 rounded to truncate teeth on

the distal margin. All arms have low inner and outer

webs.

The tentacles are short and have small (30-40%

of MX.) expanded clubs. The manus has 7 irregular

longitudinal rows of suckers. The third and fourth

rows from the dorsal margin bear the largest suckers;

the remaining suckers become progressively smaller

toward the club margins. Sucker rows near the

ventral margin of the club extend slightly farther

proximally than the others. Proximally, the dactylus

has 7-8 irregular longitudinal rows of suckers; however, these are not all continuous with the rows on

the manus. At the base of the dactylus, a new row

begins along the dorsal margin giving a total of 8

rows for a short distance; however, rows that have

continued on from the manus begin terminating, and

approximately at the midpoint of the dactylus the

number has dropped to 6 rows, then shortly to 5, and

finally to 4 rows at the tip of the dactylus. The extreme tip is occupied by a circular pad of small

suckers. The largest suckers of the manus have large,

pointed or slightly truncated teeth around the entire

margin of the inner chitinous ring. The teeth of the

distal margin are only slightly larger than those of the

proximal margin in large specimens. In smaller specimens, the size difference is much greater. A trabeculate protective membrane extends along the entire

ventral margin of their club, but dorsally it disappears

about the middle of the dactylus. A dorsoaboral keel

is present on the dactylus.

The carpal locking-apparatus begins in an oblique

line of 3 suckers with alternating pads along the

dorsal-basal part of the manus. Three to 4 suckers

follow with alternating pads that extend proximally

in a single line along the dorsal margin of the club

and onto the tentacular stalk. Medial to the end of

this series is a single small suckers and pad aligned

longitudinally and situated slightly to the ventral side

of the midline. The major series of suckers and pads

crosses from the dorsal margin of the stalk to the

ventral margin. Elements transversing the stalk may

consist either of 2 suckers, or 2 pads, or 2 suckers and

2 pads. Along the ventral margin of the tentacular

stalk, the locking-apparatus consists of a single line of

pairs of suckers alternating with pairs of pads. There

are 2-3 sets of suckers in this series. The last element

in the series is a single sucker set some distance proximal to the alternating suckers and pads.

Histioteuthis heteropsis is covered with numerous

small, anteriorly directed photophores. On the anterior portion of the ventral surface of the mantle,

photophores are aligned roughly in closely packed

transverse rows. There are approximately 40—4-5

photophores along the free edge of the mantle. On

the posterior portion, the photophores are irregularly

distributed. The number of photophores decreases

laterally; on the dorsal surface of the mantle they are

sparse and smaller in size. Photophores are absent

from the ventral surface of the fins, but occur in

small numbers on the dorsal surfaces. The funnel

lacks photophores. The ventral surface of the head is

covered with many irregularly distributed photophores. About 19-21 photophores are situated at the

edge of the circular eyelid of the right eye and only

a few, small, scattered light organs are present near

the eye opening of the right eye and none of these lies

on the edge of the eyelid. Large photophores are

scattered over the right side of the head, but the left

side has smaller and less numerous photophores. The

dorsal surface of the head has small, scattered photoTABLE 8.—Measurements in mm of Histioteuthis

heteropsis (Berry 1913).

Characters

Sex

M.L

M.W

H.W

F.L

F.W

Arm L. I

II

Ill

IV

Vi. A.C. I

II

Ill

IV

Tent. L

ClubL

Station Numbers of the VELERO

— 8018 8114 8025 10260

9 9 9 — —

75 65 33 35 22

38 32 18 23 13

41 — 21 26 15

26 — 12 13 9

38 — 17 18 12

88 86 35 42 26

94 87 36 45 29

97 87 36 43 29

97 87 35 42 28

30 29 27 23 27

29 29 30 28 29

31 32 30 29 28

28 29 28 27 34

139 158 47 65 45

24 23 11 14 9
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

phores. At the base of the ventral arms there are 8-9

longitudinal rows of light organs. These decrease to 7

about halfway along the arm length. Arms III have

4 rows near the arm base, but only the ventral row extends to the arm tip. Arms I and II each has a distinct row along its ventral border which extends to

the arm tip. Other portions of these arms bear small,

scattered photophores.

The buccal membrane has 7 lappets, and the primary buccal connectives attach to the dorsal borders

of arms I, II, and IV and to the ventral border of

arms III. An accessory connective attaches to the

ventral border of each arm I.

This animal has a uniform purplish red coloration.

TYPE LOCALITY.—Monterey Bay, California, eastern North Pacific Ocean.

LOCATION OF TYPES.—United States National Museum and personal collection of S. S. Berry, numbers

108, 110.

DISCUSSION.—H. heteropsis can be separated from

all other members of he family by the numerous small

photophores on the body, head, and arms (i.e., 8-9

rows across the base of each ventral arm), combined

with a lack of cartilaginous tubercules on the arms

(Voss 1969).

DISTRIBUTION.—H. heteropsis is a common species

off southern California. The northernmost record is

from Monterey Bay (Berry 1912b) while the southernmost record in this area is from off Baja California

at 27°34' N. (Voss 1969). Within the area of the

VELERO survey, the highest frequency of capture,

0.52 specimens/hour of trawling, occurred in zone 2

(Santa Catalina Basin). The other major zones also

had a relatively high frequency (.22 to .37 specimens/

hour) with the exception of zone 7. In this latter

zone only 3 specimens were taken, giving the low frequency of 0.03 specimens/hour. Although not entirely

convincing, this data suggests that H. heteropsis is

primarily a cold-water form, and that its southern

limit occurs somewhere off Baja California.

Voss (1969) also recorded a specimen, captured by

the USNS ELTANIN, belonging to this species from

off the west coast of South America at latitude 33° S.

Although the data are sparse, it is possible that H.

heteropsis is an antitropical species with disjunct

populations, one in the temperate northeastern Pacific and the other in the temperate southeastern

Pacific.

Histioteuthis dofleini (Pfeffer, 1912)

PLATES 9B; 10A-C

Calliteuthis ocellata.—Chun, 1910, p. 147; pl.I, figs. 1,2; text

fig.23a,b.

Stigmatoteuthis Dofleini Pfeffer, 1912, p.288.

Stigmatoteuthis dofleini.—Sasaki, 1929, p.258; text fig. 126-

127; pl.22, figs. 1-3.

Histioteuthis dofleini.—Voss, 1967, p. 74; 1969, p. 784.

DESCRIPTION.—Th e mantle is thick walled and

moderately muscular. It is conical, but tapers to a

blunt point. The free margin projects at the lockingcartilages. Beneath the thin outer integument, the

mantle is covered with short, broad, wartlike papillae.

The fins are short (about one-third of the mantle

in length and one-half of the mantle in width). Each

fin is nearly semicircular in outline and has small,

free anterior and posterior lobes. The papillate tissue

of the mantle extends onto the medial portion of the

dorsal surface of the fins.

The funnel is short, and free from the head laterally. The dorsal pad of the funnel organ has an inverted V-shape with an inverted V-shaped central

ridge terminating in a small anterior papilla. The

ventral pads are oval. The funnel locking-cartilage

has a straight but deep and broad central groove

which opposes a large, slightly curved ridge on the

mantle.

The head is large and broad. The left eye is nearly

twice the size of the right. There appears to be 3

nuchal folds rising from the nuchal crest on either

side of the head. The second fold bears the "olfactory" organ. The nuchal cartilage is long and slender;

it bears a median ridge with a central groove. The

head and aboral surfaces of the arms have weltlike

papillae beneath the outer epidermis as on the

mantle.

The arms are very long (over 175% of the MX.);

they are broad at the bases and attenuate at the tips.

The arm formula is I = IV>II = III. Rather broad

aboral keels are present on the distal half of arms III,

but arms I and II have keels that are only slender

ridges. Arms IV have thick, fleshy lateral keels. Arms

I—III have large trabeculate protective membranes

on both the dorsal and ventral margins; these are

much less developed on arms IV. Arms I of the single

large female examined have suckers arranged in 3

rows at the arm tips (this is not typical of the species

according to N. Voss) ; otherwise, all arms have biserially arranged suckers. The suckers of arms I-III
NUMBER 9 7 39

are much larger than those of arms IV. The inner

chitinous rings have been destroyed by the fixative in

the largest specimen; however, in the 14 mm M.L.

specimen, about 7-10 triangular teeth are present on

the distal half of the inner ring; laterally and proximally these diminish to small knobs. An outer web

connecting all arms at their bases is relatively well

developed, being approximately 20-25% of the mantle in length.

The tentacles are long and muscular. The tentacular clubs are slender but expanded, and nearly 40%

of the M.L. The suckers of the manus are somewhat

irregularly arranged, but can be generally divided into

5 longitudinal series. There is a row of very large

suckers in the median portion of the club. Dorsal to

this row is a very irregular row of medium-size

suckers that alternate somewhat in size. Ventral to

the central series is another, more regular row of

medium-size suckers. Along both the dorsal and ventral margin is a row of very small suckers. Along the

ventral margin, this row may be incomplete. The

suckers of the dactylus also appear somewhat irregularly arranged. In a transverse line through the

dactylus there are 3-4 suckers. At the tip of the club

there is a small flap that carriers 7-9 suckers. The

inner rings of the largest suckers of the manus have as

many as 60 sharp teeth around the entire margin. A

large dorsoaboral keel is present on the dactylus. The

carpal locking-mechanism consists of a row of 2

suckers with alternating pads at the base of the

manus along the dorsal border. Proximal to the manus there is a median row of 3-4 alternating suckers

and knobs. A small sucker is located near the proximal member of this series. Proximal to the median

row is a series of 2-3 pairs of suckers alternating with

2-3 pairs of knobs. The first set in this series is situated obliquely on the stalk, but the remaining lie

along the ventral margin. This series terminates with

a single sucker or knob.

The buccal connectives attach to the dorsal borders

of arms I, II, and IV and to the ventral borders of

arms III.

The ventral surface of the mantle is covered with

large, well-separated photophores. A somewhat irregular row of smaller photophores extends along the

free margin of the mantle. Dorsally the photophores in

this series lie somewhat farther from the mantle margin and are very small. There are 22-23 photophores

in this series. On the dorsal surface of die mantle, the

photophores are small and scattered. I counted a total

of 103 photophores on the entire mantle; however,

since some are very difficult to locate in the present material, this number may not be exact. The photophores on the ventral surface of the mantle are arranged approximately in diagonal rows.

Photophores are arranged in diagonal rows on the

ventral surafce of the head, except for a transverse

series that extends along the posterior margin. Seventeen photophores, embedded in the margin of the eyelid, encircle the right eye. Photophores are not so

regularly arranged around the enlarged left eye; unfortunately, their exact arrangement cannot be determined in the material available. A few scattered

photophores are present on the dorsal surface of the

head. In the proximal half of the ventral arms,

photophores are arranged in 3 longitudinal rows. On

the third quarter, or slightly more, of the arm, photophores are arranged in 2 rows, and on the distal segment only a single row is present.

Arms I—III all have a single row of large photophores, extending the full length of the arms along the

ventral margin of the aboral surface. A second row,

near the dorsal margin, of very small photophores is

also present on all arms. It is possible that other

photophores are present, but not detectable in the

present material.

This animal has a uniform purplish red coloration.

The data for the largest specimen available are:

Sex, $ ; M.L., 150 mm; M.W., 65 mm; H.W., 75

mm; F.L., 54 mm; F.W., 77 mm; arm lengths for

arms I-IV respectively, 265 mm, 318 mm, 313 mm,

268 mm; Tent. L., 345 mm; Club L., 61 mm.

TYPE LOCALITY.—Sagami Bay, Japan.

LOCATION OF TYPE.—Not traced.

DISCUSSION.—The present material was identified

by N. Voss. She informs me that this species can be

separated from all others in the family by the presence of paired genitalia in males, and by the papillate

skin in some specimens (i.e., this is a variable character) . The short, conical mantle and the very long

arms and relatively few photophores separates this

species, at a glance, from H. heteropsis. A third species Histioteuthis corona, subspecies berryi has been

reported once (Voss 1969) near this area (i.e., about

29° N, 126° W). It is superficially simliar to H.

dofleini except for the somewhat shorter arms. This

species can be easily separated from H. dofleini by

the presence of four longitudinal rows of light organs
40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

along the proximal half of each ventral arm, compared to three rows in H. dofleini.

DISTRIBUTION.—H. dofleini occurs primarily in

tropical and subtropical waters in both the Atlantic

and Pacific oceans, although there are a few records

from colder waters (Voss 1969). In the present study,

only three specimens (M.L., 150 mm, 14 mm, 9 mm)

were taken; these were captured in zones 1 and 7 .

Family OCTOPOTEUTHIDAE Berry, 1912

Veranyidae Chun 1910, p. 139. [Family name inadmissable

because originally based on a generic name not currently

valid. See Roper, Young, and Voss, 1969.]

Octopoteuthidae Berry, 1912a, p. 645.

Octopodoteuthinae Pfeffer, 1912, p.212.

DIAGNOSIS.—Arms with biserial hooks that may be

replaced by suckers near the arm tips. Tentacles present only in larvae (rarely in juveniles). Buccal connectives attach to the ventral borders of arms IV.

Fins more than 70% of the mantle length. Funnel

locking-cartilage with a simple, approximately

straight but broad groove.

Genus Octopoteuthis Ruppell, 1844

DIAGNOSIS.—Photophores present at the tips of all

arms. At M.L., 15-25 mm, all trace of tentacles has

been lost.

TYPE SPECIES.—Octopoteuthis sicula Ruppell,

1844.

Octopoteuthis deletron, new species

PLATES 10H-M; 11; 12A-D

Octopodoteuthis sp.—McGowan, 1967, p.209.

Octopoteuthis sicula.—Pearcy, 1965, p.261.

DESCRIPTION.—The conical mantle gradually

tapers to a blunt point posteriorly. The tip of the

mantle is almost always contracted and pulled ventrally and anteriorly, giving the posterior end of the

body a very blunt appearance. The mantle extends

slightly posterior to the fins in the larger specimens,

forming a short, broad tail. The tail is not detectable

in young specimens, but increases allometrically with

growth in specimens larger than about 40 mm M. L.

up to maturity when the tail is about 18% of the

M.L.

The fins are very large and muscular. At their posterior ends, they are reduced to narrow strips that extend along each lateral edge of the tail and join at

the apex of the tail. Although their anterior and posterior lateral edges are slightly flattened, the fins

are nearly elliptical in outline, except for the posterior attenuation. Their width is 95% of the M.L.,

and their length (exclusive of the tail) is 86% of the

M.L. in the holotype. Small free, anterior lobes are

present. The fins fuse at the dorsal midline.

The funnel extends approximately to the level of

the midpoint of the eyes. It is free from the head

laterally. The funnel locking-cartilage is basically of

the simple, straight type, although posteriorly the

groove curves slightly and flares. The dorsal pad of

the funnel organ has roughly an inverted V-shape

with a prominent ridge and papilla at the apex and

short ridges on the arms; the organ, however, is somewhat variable in appearance. The ventral pads are

very large and oval; a funnel valve is present.

The head is large and bears large eyes which extend

the entire length of the head. A small optic sinus is

present at the anterior end of each circular eyelid.

Along the posterior margin of each eyelid is a large,

broad, crescentshape patch of muscular tissue. The

"olfactory" papillae lie at the posterior edge of the

head, lateral to the base of the funnel; each consists

of a white, flattened terminal bulb with a short, swollen stalk. Both the mantle and head are encased in a

thick layer of gelatinous tissue.

The arms are long (over 70% of the M.L. in the

holotype). The arms are brittle and almost always

broken at the tips; in addition, those that can be

measured show considerable variation. The arm

formula, therefore, is difficult to determine with certainty; in mature specimens they appear approximately subequal. All arms have gelatinous aboral

keels. Low, thick, gelatinous protective membranes

are present on all arms, but are only apparent in large

specimens. These gelatinous ridges are joined across

the midline of the arm by membranes which alternate

with the hooks; the ridges and membranes form the

walls of cup-shape depressions from which the arm

hooks arise. The arms bear 2 rows of small hooks

which generally are partially or entirely hidden in

thick-walled sheaths. Two small, but distinct, accessory cusps occur on each hook. These cusps are relatively more developed in large specimens. The hooks

occupy most of the arm lengths, but near the arm

tips they abruptly give way to approximately 3-11
NUMBER 9 7 41

pairs of small, globular suckers which have about 6-9

somewhat irregular, pointed teeth, including a large

central tooth in each of the more proximal suckers.

The number of suckers seems to depend somewhat on

size and stage of development, although in the largest

specimens they vary from 6-11 pairs. The extreme

distal ends of the arms are bare, swell slightly, and

have a small bulb at their tips. The swelling and bulb

are generally more pigmented than the rest of the arms

and are luminescent.

The tentacles are completely lacking in adult specimens; however, a specimen with a mantle length of

17 mm has large blunt stubs of the tentacles. Larval

forms have small tentacles with a few enlarged suckers at the tips, but no details can be given due to the

damaged state of the larvae available.

Octopoteuthis deletron has a considerable number

of photophores; however, most are inconspicuous. A

single large photophore is buried deep in the mantle

tissue in the ventral midline near the posterior end of

the body. This abdominal photophore is difficult to

locate in specimens less than 30 mm M.L., but very

careful examination will reveal its presence in specimens somewhat less than 20 mm M.L.

A large photophore is embedded deeply in the

gelatinous tissue of the head just posterior to each

eye. As described earlier, the tips of all the arms bear

elongate photophores. An elongate, obliquely oriented

photophore lies inconspicuously near the anterior

medial margin of each eye on the ventral portion of

the head. These latter photophores do not seem to be

developed in specimens less than about 30 mm M.L.

A series of photophores lies near the central core of

each of the ventral 4 arms interior to the longitudinal

muscles, and are often adjacent to the axial nerves.

The basal photophore on each arm is larger than the

following ones and is somewhat more isolated from

the others. The remaining arm photophores, numbering about 25 on each arm IV, are situated at regular intervals up to the level of the distal suckerbearing zone of the arms where the series ends. Arms

II have only the basal photophore and the arms I

lack any trace of this series. In preserved specimens,

it is necessary to pick away the muscular tissue of the

arms to see these light organs. Another pair of photophores lies near the ventral surface of the ink sac.

These photophores are overlain by a pair of muscular

bands.

In mature males the penis is large and appears

capable of extending well beyond the opening of the

mantle cavity. Gravid females have greatly enlarged

nidamental and oviducal glands. The mature eggs

measure about 2 mm in diameter.

Specimens are covered with reddish brown chromatophores. The external layer of chromatophores that

lie in the aboral gelatinous tissue on arms III and IV

are arranged in a peculiar manner, in that they lie in

parallel transverse planes relative to the axes of the

arms and are evenly spaced, giving the appearance of

a series of lamellae. The mature female has a uniform

dark-purple color.

Octopoteuthis deletron undergoes rather drastic developmental changes at a M.L. of about 30-40 mm.

This is the change from the juvenile to the adult

form, although maturity is not reached until a much

larger size. The most apparent changes include the

great increase in eye size, the thickening of the arms

and body, and the acquisition of greater pigmentation. These changes are illustrated in Plate 12A-D.

The radula has a tricuspid rachidian and a bicuspid

first lateral, unicuspid second and third laterals, and

marginal plaques.

The primary buccal connectives attach to the ventral margins of arms IV.

TYPE LOCALITY.—33° 15' N, 118°37' W, eastern

North Pacific Ocean.

LOCATION OF TYPE.—University of Southern California, U.S.C. Hancock collections, AHF Cephalopod

Type No. 3.

DISCUSSION.—In the genus Octopoteuthis there

are 7 nominal species: O. sicula Riippell, 1844, near

Messina, Mediterranean Sea; O. megaptera (Verrill

1885), off east coast of United States, 44 mm M.L.;

O. persica Naef, 1923, off south tip of Africa, 34°31'

S, 26° 0' E, 4.7 mm M.L.; O. indica Naef, 1923, Gulf

of Aden, Indian Ocean, 13°2' N, 46°41' E, 3.8 mm

M.L.; O. danae Joubin, 1931, near Bermuda, 33° 15'

N, 68°20' W, 31 mm M.L.; O. niebeni Robson, 1948,

near Cocos Island, Pacific Ocean; O. longiptera

Akimushkin, 1963, from sperm whale stomach, 20° S,

25° W, Atlantic Ocean.

O. persica is based on an illustration in Chun

(1910). The specimen shows distinct swellings near

the tips of arms II and extremely broad fins. Both

features are strongly suggestive of the genus Taningia,

and I think it safe to transfer this species from Octopoteuthis to Taningia.

All members of the genus seem to have the same
42

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

basic pattern of large photophores, with the exception

of the abdominal series which has either 1 median or

2 lateral photophores, depending on the species.

Joubin (1931) first noticed the presence of the larger

photophores, and this resulted in the establishment of

a new species, O. danae. O. danae seems to agree in

almost all respects with O. sicula; unfortunately, the

arrangement of abdominal photophores in specimens

of O. sicula from the Mediterranean is unknown.

Still, there would be little evidence to prevent synonymizing these 2 forms if an additional, similar

species did not occur in the eastern North Atlantic.

In contrast to O. danae which has 2 abdominal

photophores this species has only a single abdominal

photophore, and I shall henceforth refer to it as Octopoteuthis A.

The other common Atlantic species, O. megaptera,

from the western north Atlantic and Gulf of Mexico

can easily be distinguished by the greater size of its

"tail," by the presence of a 2 abdominal photophores,

and by large accessory cusps on the arm hooks even

in young stages.

O. longiptera was very inadequately described. The

only character of possible specific significance is the

relative dimensions of the fins; their length is 135% of

the width. (This measurement apparently includes

the tail.) This feature is probably only a result of the

large size of the specimen (590 mm total length).

The specimens from California at 30-40 mm M.L.

have fins whose lengths (including the tail) are approximately 55%-65% of their widths; at 109 mm

M.L. the fin length is 90% of its width. In the 167

mm male ( =300 mm total length) the percentage is

105, while in the 148 mm female the percentage is

120. G. Voss informs me that the type of O. longiptera is not extant. Since O. longiptera cannot be

clearly separated from any of the existing species, it

must be considered a species dubia.

Unfortunately, O. indica from the Indian Ocean

must be ignored for the present, since its small size

prevents adequate comparison to related forms.

Therefore, we are left with at least 3 distinct species

in the Atlantic Ocean: Octopoteuthis A, O. danae,

and O. megaptera. One of the former 2 species (most

likely O. danae) must certainly be a synonym of the

Mediterranean species O. sicula. This problem, however, cannot be resolved with certainty until the photophore pattern of O. sicula is determined. In making

comparisons with the Califomian species, I shall

refer only to the 3 Atlantic forms and assume that O.

sicula is included in one of them.

In the eastern Pacific, at least 2 additional species

are present: O. deletron from California and possibly

Peru (see discussion of distribution), which is characterized by a short tail and a single abdominal

photophore, and O. nielseni from the region of PanTABLE 9.—Measurements in mm of Octopoteuthis deletron, new species.

Characters

Station Numbers of the VELERO

7414 7414 7414 8025 8716 8878 11097

Sex

M.L

M.W....

H.W... .

F.L

F.W

TaUL...

Arm L. I

II

III

IV

Yi A.C. I

II

III

IV

33

13

12

26

39

35

17

14

28

43

30

27

27

38

17

12

28

51

25 — —

32

27

9 <? d1

(Holotypc)

39

12

12

45

37

33

29

28

28

109

37

33

94

104

13

84

79

28

29

167

50

42

124

140

22

99

148

50

45

91

107

30

116

105

114

115

27

29
NUMBER 91 43

ama. O. nielseni was only briefly described by Robson, and the critical abdominal photophore pattern

was not given. G. Voss has recently (1962) examined

the type in the British Museum and informs me that

the specimen is so damaged that it is not possible to

determine the photophore pattern. In the DANA collections, there is a specimen which was captured at

6°48' N, 80°33' W off Panama. It is probably safe to

assume that this species is O. nielseni. It is a shorttailed form, as Robson (1948) illustrated, and possesses 2 posteroabdominal photophores. This is a large

specimen (M.L., 112 mm) which, surprisingly, has

only a slight trace of accessory cusps on the arm

hooks. This latter feature would seem clearly to separate O. nielseni from O. danae, which possess distinct accessory cusps at least in the adult stage (Adam

1952:73, as O. sicula), but the value of this character is somewhat uncertain; I examined a small

specimen captured off the west coast of Mexico which

also appears to be referable to O. nielseni, but has distinct cusps. Also specimens of O. danae from the Atlantic can be divided into young forms with accessory

cusps and young forms without them. Since O. deletron is quite consistant in the presence of this feature,

it is possible but perhaps unlikely that these different

types represent distinct species. The reason for this

confusion and difficulty in resolving the specific status

of the different forms results from the presence of

only a few specific systematic characters within the

genus. This condition is partially the result of the almost invariable loss of the arm tips during capture.

If the difficulties concerning the accessory cusps on

the hooks can be explained by individual variation,

then it becomes very difficult to separate O. danae

and O. nielseni. I believe, however, that O. nielseni

will prove to be a valid species and, therefore, believe

that its name should be retained at least until detailed

data are available on all growth stages of both species.

O. deletron is thus separable from all other species

except Octopoteuthis A by the presence of only a

single abdominal photophore. It differs from Octopoteuthis A by having a much shorter tail at equivalent mantle lengths and by having well-developed accessory cusps on the arm hooks which are lacking in

the latter species.

DISTRIBUTION.—In the present study, O. deletron

had a fairly uniform frequency of capture in zones

1, 2, 3, 5, and 6 (i.e., fluctuates between 0.18 and

0.36 specimens/hour of trawling). No specimens were

captured in zone 7. McGowan (1967) has recorded

larvae identified as Octopodoteuthis sp, which undoubtedly belong to the present species, from 34° N

to 30° N in the southern California area. Pearcy

(1965) reported O. sicula from off the coast of Oregon; his specimens possess a single abdominal photophore (G. Voss, personal communication) and can,

almost certainly, be referred to O. deletron. This information suggests that O. deletron is a cold-water

species with its southern limit somewhere off Baja California; however, the ELTANIN has taken a large

specimen of Octopoteuthis from 7°45' S, 81°37' W

off northern Peru that appears to be identical with

O. deletron. If this is the same species, present evidence would favor the occurrence of two disjunct

populations rather than a continuous distribution

from Oregon to Peru.

Family GONATIDAE Hoyle, 1886

DIAGNOSIS.—Arms with tetraserial armature; arm

tips may be modified. Median 2 rows on arms I—III

consist of hooks except at the extreme base and tip

of each arm, with the exception of Berryteuthis anonychus which lacks hooks except at the bases of arms

I—III in females. Funnel locking-cartilage has a simple straight groove which may flare slightly at the

posterior end in some species. Buccal connectives attach to the ventral borders of arms IV. Nuchal folds

present.

Genus Gonatus Gray, 1849

DIAGNOSIS.—Five teeth present in a transverse series on the radula. Tentacular club with a large central hook and generally additional hooks in the median line. Each tentacular club bears a distinctive

locking apparatus on the dorsal border of the manus

which consists of a series of large, thick, transverse

pads with medial alternating oval pads and suckers.

TYPE SPECIES.—Onychoteuthis? amoena Moller,

1842.

Gonatus onyx, new species

PLATES 13A; 14A,C-I; 17J

Gonatus fabricii.—Berry, 1912b, [part], p.308, pl.52, fig.4.—

? Pearcy, 1965, p.262.

DESCRIPTION.—The mantle is long and slender;

posteriorly it tapers gradually to a pointed tip. The
44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

muscular portion of the mantle which is moderately

thick terminates at the edge of the conus. A muscular

cone with a gelatinous core forms a tail that extends

posteriorly from the apex of the conus. This structure

is not apparent in young specimens, but increases allometrically and becomes prominent in large specimens. The free margin of the mantle projects slightly

at the mantle locking-cartilages.

The fins have free anterior lobes. The posterior

margins are drawn out along the tail in large specimens. The lateral edges of the fins are rounded.

The funnel is free from the head laterally and

reaches to approximately the level of the lenses of the

eyes. Each funnel locking-cartilage bears a straight,

simple groove. The dorsal pad of the funnel organ has

the shape of an inverted V; the arms are broadest in

the posterior portions; the anterior portions each have

a median longitudinal ridge that extend to the apex.

Near the anterior apex, which is blunt and slightly indented, a slender papilla arises. The ventral pads are

oval. A large funnel valve is present.

The head is oval and bears large eyes that occupy

the entire lateral sides of the head. An anterior sinus

is present on each eyelid. Three pairs of nuchal folds

are present; the "olfactory" organ arises from the second pair. The nuchal cartilage is elongate and

rounded at the anterior end; it has a median ridge

with a central groove and marginal ridges.

Arms II and III are very muscular, nearly equal in

size, and 50-60% as long as the pen. The ventral

arms are the same length or only slightly less than the

lateral arms, but are more slender. Arms I are the

smallest; they reach about 40-50% of the P.L. Arms

I—III all have gelatinous aboral keels which are best

developed on arms III. Arms IV have large lateral

keels along their entire lengths. Well-developed protective membranes with long, slender trabeculae are

present on arms I—III; they are less developed on the

arms IV. The armature is in 4 longitudinal series. On

arms I—III the median rows consist primarily of large

hooks and the marginal rows of small suckers. The

suckers arise near the distal ends of the trabeculae. At

the extreme tips of the arms, the hooks are replaced

by suckers. A total of 25-30 suckers and hooks occur

on the proximal half of each arm I—III. Arms IV

lack hooks and have 4 series of small suckers. Specimens larger than 40 mm P.L. have 30-37 suckers on

the proximal half of arms IV. The arm suckers all

have similar dentition. Long, pointed teeth are present on the distal margin, but decrease in size laterally.

The proximal border is smooth. The total number of

teeth is generally about 12-13. The suckers of the

median rows on arms IV are the largest arm suckers,

and the marginal suckers of arms I—III are the

smallest. The marginal suckers of arms IV are intermediate in size.

The tentacles are robust and have small (approximately 20-25% of the P.L.), complex clubs. The

manus possesses the 4 sucker-bearing zones typical of

the genus: a ventral-marginal zone that extends the

length of the manus and terminates behind the large

central hook; a locking zone along the proximal half

of the dorsal margin; a dorsal-marginal zone which is

continuous with the dactylus distally and proximally

passes medial to the locking zone; and, finally, a

median zone consisting of a single row of suckers,

hooks, or both. The ventral-marginal zone in its central part has 4, rarely 5, suckers that arise from each

trabecula and are arranged generally in 4 longitudinal

rows. The suckers in the medial row are smaller, but

not less than half the diameter of the large suckers in

the marginal row. Proximally, the rows converge to

a single one which is continuous with the sucker row

that extends along the ventral margin of the tentacular stalk. The largest suckers of the ventral marginal

zone have 10-12 long, pointed teeth distally, which

diminish in size laterally. The teeth are more broadly

spaced than are those of the arm suckers.

The locking zone consists of 4-6 thick ridges, each

with a large smooth-ringed sucker placed at its proximal end. There is usually a sucker at the distal end of

the zone without the adjoining ridge. Alternating

with the suckers are large rounded knobs. Proximally,

this zone ends abruptly and this break is here used to

indicate the proximal limit of the club.

The dorsal-marginal zone contains an irregular

group of suckers that do not appear to be arranged in

definite rows; distally, however, the suckers merge

into 3, or occasionally 2, rows which pass lateral to the

large central hook and merge with the suckers of the

dactylus. Proximally, the number of suckers decreases

rapidly and disappears medial to the locking zone.

The median zone bears a large central hook and a

proximal row of suckers. These suckers are never

transformed into hooks. Usually, there is no member

of this zone distal to the large hook; however, occasionally an enlarged sucker lies behind the hook.

Sometimes this sucker may bear an enlarged tooth on
NUMBER 9 7 45

the distal margin of the inner ring; and, rarely, this is

developed into a small, but completely formed hook.

If such a hook is present, it is usually on only one club

of the specimen. At the proximal end of the dactylus,

suckers are irregularly arranged in 5-6 rows, but

quickly decrease in number distally to form 4 regular

longitudinal rows. Except for the basal few transverse

rows, there is very little difference in size between

members of the dorsal and ventral rows. Distally, the

dactylus ends in a circle of smooth-ringed suckers at

the tip of the club. The suckers from the middle of

the dactylus have dentition similar to those of the

manus. The total number of suckers on the dactylus

(excluding the circle at the tip), the ventral marginal

zone, and the dorsal marginal zone varies from 160-

200. A well-developed keel is present on the dorsoaboral surface of the dactylus. A narrow, trabeculate

protective membrane extends lateral to the suckers

along the entire length of the ventral border of the

club. On the dorsal margin, a similar membrane

terminates in the proximal half of the dactylus.

The tentacular stalk bears a single series of suckers

along either margin. Distally, the row along the dorsal

margin consists of alternating suckers and pads with

adjacent ridges that show faint resemblance to the

locking apparatus of the manus; traces of the fleshy

ridges accompany only the first few suckers, then disappear. The knobs are displaced medially, relative to

the suckers.

The row on the ventral margin is a continuation of

the ventral-margin zone on the manus. Most sucker

rings in this series, have nearly smooth inner rings.

The suckers that are situated one-fifth of the tentacle

length from the base of the clubs are nearly smooth,

but have the distal margin lightly scalloped. Both

marginal rows extend almost the entire length of the

tentacle. The number of suckers in the ventral row is

70-115% of the number in the dorsal row. A protective membrane, apparently lacking trabeculae, extends along the entire dorsal margin of the tentacular

stalk. A trabeculate protective membrane is present

distally on the ventral broder. Only a few, generally

less than 10 (maximum range is 0-27), suckers are

found between the rows on the oral face of the stalk.

The buccal lappets are indistinct. The buccal membrane connectives attach to the dorsal borders of arms

I and II and to the ventral borders of arms III and

IV.

The animal is covered with numerous reddish

brown chromatophores. This species seems to have a

iridescent silver sheen over the lateral surfaces of the

head and arms.

Hooks begin to develop on the arms between 26-28

mm P.L. and on the tentacular club between probably 17-24 mm. A spindle-shaped liver set obliquetly

to the body axis is present in the larvae.

TYPE LOCALITY.—33° 19' N, 118°45' W, eastern

North Pacific Ocean.

LOCATION OF TYPE.—University of Southern California, U.S.C. Hancock collections, AHF Cephalopod

Type No. 4.

DISCUSSION.—Berry (1912b) described a number

of specimens under the name Gonatus fabricii. One of

his illustrations (plate LII, figure 4) showing part of

a tentacular club probably refers to G. onyx.

This species is unique in generally possessing only a

single hook on the tentacular clubs. Occasionally, an

enlarged sucker is present distal to the large median

hook and sometimes this is transformed into a small

hook; however, this rarely happens on both clubs of

a single specimen. Suckers proximal to the large hook

are never transformed into hooks. Other characters

also separate it from some of its close relatives. The

low number of suckers (usually less than 10) on the

tentacular stalk between the marginal rows separates

it from all species except G. berryi. The size at which

the hooks develop on the arms and tentacles separates

it from all species except G. californiensis. The number of suckers on the tentacular clubs and the number

of hooks and suckers on the proximal half of all arms

clearly separates it from G. californiensis. (See also,

discussion of G. californiensis.) The unique and distinguishing characters listed for each of the other

species of Gonatus are found in the following text.

DISTRIBUTION.—Gonatus onyx is the most abundant species in the collections. In the area under survey, its distribution is limited almost exclusively to

the San Pedro, Santa Catalina, and San Nicolas

basins; out of a total of 1,236 specimens only 7 have

been captured outside of these 3 basins. It is apparent

that the species seldom penetrates to the area of

Guadalupe Island since there is not a single record

from this area despite an estimated 76 hours of

trawling in appropriate depths. The almost complete

absence from all of the most oceanic zones (i.e., 5, 6,

7), however, indicates that this is an "inshore" species which may extend farther south in the more in-
46 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 10.—Measurements in mm of Gonatus onyx, new species.

Characters

Station Numbers of the VELERO

7279 8117 7279 8117 8112 7394 7394 7394 7394 10399

Sex

P.L

M.W. ...

F.L

F.W ..

Arm L. I

II

III

IV

Yi A.C.II

I .

HI

IV

Tent. L..

ClubL..

ClubS...

M.S.S...

D.S.S....

V.S.S....

59

15

25

42

26

32

31

30

28

30

30

35

41

12

194

12

75

71

98

21

—

52

40

47

49

47

30

31

30

34

66

16

181

3

45

45

55

12

38

25

27

31

30

26

29

28

34

44

12

184

1

65

60

(Holotypc)

67

15

25

41

29

36

36

35

27

28

27

32

67

15

174

1

66

54

72

17

28

37

30

36

37

35

28

31

31

35

66

13

172

5

69

65

50

11

34

24

27

27

26

25

26

29

36

39

11

171

2

61

48

44

12

21

31

20

23

24

21

28

29

28

32

35

9

177

15

74

67

34

9

15

26

14

16

17

15

28

29

31

33

30

7

187

1

68

73

42

11

20

31

17

20

21

18

28

28

28

33

42

9

165

2

67

64

69

17

28

47

34

38

41

36

29

31

30

34

47

14

185

5

66

64

shore waters which were not sampled south of latitude 32° N.

The northern limits of distribution of this species

are unknown. There is a probable record by Berry

(1912b) from Monterey Bay. I examined several

specimens of Gonatus collected by Pearcy from off the

Oregon coast which were deposited in the Miami collections. These specimens clearly belong to G. onyx,

and it is probable that the G. fabricii listed by Pearcy

(1965) belongs to this species, at least in part. Therefore, the known range of G. onyx extends from about

latitude 30° N to at least 43° N.

Gonatus berryi Naef, 1923

PLATES 15A; 16A,C-I; 17A-C,L

Gonatus fabricii.—Berry, 1912b, [parts], p.308; pl.52, fig.3;

pl.55, fig.2.

Gonatus berryi Naef, 1923, p.245.

DESCRIPTION.—The mantle is muscular, but thin

walled. The mantle margin is produced at the ventral

locking-cartilages, but only slightly, if at all, dorsally.

The mantle terminates in a point posteriorly; the

muscular portion ends at the conus of the pen. A

muscular cone with a gelatinous core forms a tail that

extends posteriorly from the apex of the conus. The

tail is disproportionately small in young specimens.

The fins are rounded laterally and have free anterior lobes. Posteriorly the fin becomes attenuate as

it extends along the sides of the tail.

The muscular funnel, completely free from the

head laterally, reaches to the level of the midpoint of

the eyes. The funnel locking-cartilage bears a straight

groove which flares slightly posteriorly and opposes a

straight ridge, which broadens posteriorly, on the

mantle. The dorsal pad of the funnel organ has an

inverted V-shape with a blunt or slightly indented

apex. The arms are strongly angled laterally and bear

median ridges that extend anteriorly to the apex. A

small papilla is present near the apex. The ventral

pads are egg-shape. A large funnel valve is present.

The head is oval and bears large eyes that occupy

almost the entire lateral sides of the head. Each eyelid has a well-developed anterior sinus. There are 3

pairs of nuchal folds; the second fold from the funnel

on either side bears the "olfactory" organ. The nuchal

cartilage is elongate, rounded at its anterior end, and

bears a median ridge containing a central groove,

and lateral ridges.

Arms II and HI are moderately long and muscular.
NUMBER 9 7 47

In specimens greater than 35 mm P.L. they vary between 60-70% of the P.L. Arms I are slightly shorter

and vary between 50-60% of the P.L. Arms IV are

nearly equal to arms II and III in length, but are

much thinner.

The presence of aboral keels on arms I—III is uncertain due to damage to the specimens. Welldeveloped trabeculate protective membranes are

present on arms I—III; they are less developed on

arms IV. Armature is in 4 series: on arms I—III the

2 median rows have large hooks and the marginal 2

rows have small suckers; at the arm tips the hooks are

replaced by suckers. On arms IV, all 4 rows have only

suckers. The number of suckers on the proximal half

of each arm IV is approximately 40 at a P.L. of 50

mm and 45 at a P.L. of 119 mm. The average number of hooks and suckers on the proximal half of each

of arms I—III is about 25 at a P.L. of 50 mm and 33

at a P.L. of 119 mm. The suckers of the arms all

have similar dentition. Long, pointed teeth, which

diminish in size laterally, are present on the distal

margin of the inner sucker ring. The proximal margin

is smooth. The total number of teeth varies approximately from 10-14. The largest arm suckers occur in

the median rows of arms IV; these are nearly V/z

times larger than the largest marginal suckers of arms

III. The marginal suckers of arms IV are intermediate in size.

The tentacles are robust; the clubs are moderately

large, generally between 30-37% of the mantle

length. The ventral-marginal zone of the manus

typically consists of 3 longitudinal rows in its broadest part. Occasionally, 1-2 small suckers are present

which are the only indication of a fourth row. The

suckers of the median row (occasionally rows) are

minute and not more than one-fourth the diameter of

the large suckers in the marginal 2 rows. The distalmost suckers of this zone are also minute. The largest

suckers of the ventral-marginal zone have about 8-9

long, sharp teeth on the distal half of the inner ring

which are rather broadly spaced. These diminsh in

size laterally and are replaced on the proximal margin

by small knobs. These suckers are distinctly larger

than those of the median rows on arms IV. The locking zone consists of 5-6 broad, fleshy ridges with a

large smooth-ringed sucker at the medial end of each.

An additional sucker is commonly present at the distal end of the zone.

The dorsal-marginal zone has relatively few

suckers. Distally, it continues past the central hook in

1-2 irregular rows and merges with the suckers of the

dactylus. Proximally, the row terminates medial to the

locking apparatus. The median zone contains a large

central hook followed distally by a small hook and preceded proximally by a single series of suckers and

hooks. One or two suckers always lie nearest to the

large hook and are followed by 2-4 hooks; the largest

hook is never the closest one to the large central hook.

At the base of the dactylus along the dorsal border,

there are 2 rows of large suckers. Between these rows

and the ventral margin are scattered a few minute

suckers. The number of rows increases distally to 4

regular series that extend to the tip of the dactylus.

The suckers of the ventral row are much smaller than

the others. This is most noticeable in the proximal

portion of the dactylus and in large specimens. The

largest suckers in the middle of the dactylus have

similar dentition to the suckers of the manus except

that the teeth are disporportionately smaller. At the

tip of the dactylus, there is a small circle of smoothringed suckers. The total number of suckers on the

dactylus (excluding the distal circle), the median

zone, and the ventral-marginal zone varies from 159—

181. A dorsoaboral keel is present on the dactylus. A

small, trabeculate protective membrane is present

along the entire ventral margin of the club. A similar

membrane is present on the dorsal border of the

manus, but ends along the base of the dactylus.

A locking apparatus, consisting of a series of suckers

and knobs, extends almost the entire length of the

tentacular stalk along its dorsal margin. There is only

a slight trace of the fleshy ridges characteristic of the

locking apparatus of the manus adjacent to the most

distal suckers in the series. The knobs are displaced

medially from the suckers. Another series of suckers

extends, in a single line, along the ventral margin of

the tentacular stalk in its distal half; however, the

length is somewhat variable since occasionally scattered suckers extend farther down the stalk. Almost

all of these suckers bear pointed teeth along the distal

margins of their chitinous rings. Suckers in this series

located one-fifth of the tentacle length from the base

of the club have 4-6 distinct, pointed teeth on the distal margin of the inner rings. The number of suckers

along the ventral margin varies from 38-58% of the

number along the dorsal margin. The oral surface of

the tentacular stalk between the marginal rows is

completely free of suckers except for an occasional
48

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

one or two. A low, protective membrane, apparently

without trabeculae, is present along almost the entire

dorsal margin of the tentacular stalk. A trabecular

protective membrane extends along the distal half of

the tentacular stalk on the ventral margin.

The buccal mambrane bears 7 indistinct buccal lappets. The buccal membrane connectives attach to the

dorsal borders of arms I and II and to the ventral

borders of arms III and IV.

The larva first develops hooks on the arms between

8-10 mm P.L. The large central hook on the club develops at 12-17 mm P.L. The more distal hook develops between P.L. 19-28 mm, while the proximal

hooks begin to develop between 25-32 mm. P.L. A

spindle-shape, larval liver is never present in this

species.

Photophores are absent. Members of this species are

covered with numerous, reddish brown chromatophores. There is some evidence, at least in younger

specimens, of the presence of iridescent silvery tissue

on the sides of the head and arms.

TYPE LOCALITY.—Monterey Bay, California, eastern North Pacific Ocean.

LOCATION OF TYPE.—?In personal collection of S.

S. Berry, number 98.

DISCUSSION.—Berry (1912b) described a number

of specimens of squid which he identified as Gonatus

fabricii. Naef (1923), noticing that one of the young

specimens illustrated by Berry had fins that were unusually large for G. fabricii, established for it the new

species G. berryi. No further information has been

published on this species until now. The large size of

the club in the type illustration as well as the illustrated details of the club structure make it very easy

to connect this name with the second most abundant

species of the genus taken in the present study.

G. berryi is very distinct, and there should be no

difficulties in its recognition. Among its unique features are the following:

1. In the median zone of the club there are 1-2 suckers

which lie near the large central hook. These are followed

by several hooks that generally increase in size as the series

proceeds away from the central hook. The usual case is the

opposite arrangement.

2. The ventral-marginal zone of the club has 2 marginal

rows of large suckers and 1-2 rows of minute suckers

medially.

3. Suckers in the ventrar row on the dactylus are considerably smaller than those of the dorsal row.

4. The tentacular clubs are 30-37% of the pen length.

5. The ventral-marginal row on the tentacular stalk has only

about 30-60% of the number of suckers present in the

dorsal-marginal row.

6. The larvae develop hooks on the arms when they have

a P.L. of only 8—10 mm.

TABLE 11.—Measurements in mm for Gonatus berryi Naef, 1923.

Characters

Station Numbers of the VELERO

8117

49

17

24

38

29

35

34

32

25

25

25

38

43

17

173

1

86

32

8117

52

16

24

34

31

37

37

34

25

25

25

40

57

19

165

0

71

25

7394

42

13

18

29

22

26

27

26

23

24

22

35

38

13

166

0

77

41

7394

56

14

28

35

33

37

37

36

28

28

29

41

55

20

162

0

74

26

7394

34

10

14

24

19

22

23

20

23

21

22

33

30

11

163

0

76

31

7343

cf

119

35

65

72

65

73

77

76

33

34

34

45

85

30

168

1

83

36

7279

46

16

22

34

27

31

33

32

26

26

24

39

37

17

162

0

74

33

8112

38

10

—

29

22

25

26

24

23

24

26

38

35

13

170

0

85

33

8018

33

13

14

26

18

19

20

18

24

24

26

33

23

10

178

1

77

41

8795

23

9

9

17

9

11

10

9

20

21

20

31

10

5

158

1

—

Sex

P.L

M.W. . ..

F.L

F.W

Ann L. I

II

III

IV

Yi A.C. I

II

III

IV

Tent L. .

ClubL..

ClubS..

M.S.S. ..

D.S.S....

V.S.S....
NUMBER 9 7 49

7. A spindle-shaped larval liver is not present at any stage

of the life cycle.

In addition, many features distinguish this species

from one or more of its relatives. Among these are

the almost complete lack of suckers on the oral face of

the tentacular stalk between the marginal rows;

toothed suckers along the ventral margin of the tentacular stalk; the relative length of the arms; the arrangement of suckers on the dactylus; the number of

suckers on the tentacular club; the number of suckers

and hooks on the proximal halves of the arms; and

the P.L. at which the hooks on the club develop.

DISTRIBUTION.—This species occurs in all zones except 7. Its frequency of 0.5 specimens/hour of trawling in zone 6 is surprisingly high and compares favorably with its frequency of 0.57 in the Santa Catalina

Basin. Its absence from the vicinity of Guadalupe Island, however, is significant and suggests that this

area is near the southern limit of the species. The type

specimen, which is the only other record of the species, was taken by the ALBATROSS in Monterey

Bay. Thus, the known range of this species extends

from latitude 30° to 37° N off the coasts of southern

California and Baja California.

Gonatus pyros, new species

PLATES 13B; 14B,J-O.; 17G-I,K

DESCRIPTION.—The mantle is slender; in almost all

specimens examined, it is contracted and tapers gradually from a flaring margin to a posterior point. The

mantle margin projects ventrally in the areas of the

mantle locking-cartilages, but not dorsally. The

mantle wall is moderately thick and muscular.

The fins are broad. A posterior "tail" is only

slightly developed in the largest specimens. Free anterior lobes are present, and the lateral fin margins are

rounded. The fins are very muscular, but are almost

always damaged.

The funnel is free laterally and extends forward to

about the level of the posterior edge of the lens. The

funnel locking-cartilage has a long, straight sulcus

which opposes a straight ridge on the mantle. The

dorsal pad of the funnel organ has roughly the shape

of an inverted V; the lateral arms are long and angled

laterally. Small, median ridges are present near anterior ends of each of the lateral arms, and a small

papilla lies at the blunt anterior apex. The ventral

pads are roughly oval in shape.

The head is short and rounded. Its width generally

exceeds that of the mantle. The eyes are large, and

occupy the entire lateral sides of the head; an anterior sinus is present on each eyelid. Each "olfactory"

organ is flaplike and located at the posterodorsal end

of the second nuchal fold from the funnel. The 3

nuchal folds are very feeble and can be recognized

only in the most perfectly preserved specimens. The

nuchal cartilage is elongate and rounded at either

end. It bears 3 longitudinal ridges; 2 are near the

lateral edges of the cartilage, and one is in the median

line. The median ridge has a narrow, central sulcus

along its midline.

Arms II and III are moderately long and muscular; their length in specimens larger than 35 mm P.L.

varies between 60-70% of the P.L. Arms I are

shorter and vary between 50-60% of the P.L. Arms

I—III appear to have had membranous or gelatinous

swimming keels. Arms IV are nearly the same length

at the lateral arms, but are more slender. Thin, broad,

lateral keels are present along the entire lengths of

arms IV. Arms I—III all have well-developed trabeculate, protective membranes; trabeculae and membranes are less developed on arms IV. The armature

of the arms is in 4 series. On arms I—III, the marginal

rows consist of small suckers that arise near the ends

of the trabeculae; the medial 2 rows are composed of

large, alternating hooks which are replaced by suckers

at the arm tips. Arms IV have suckers instead of

hooks in the medial rows. All of the arm suckers have

similar dentition. On the distal part of the inner ring

are long, pointed teeth that grade into smaller teeth

laterally. The proximal margin is smooth. The total

number of teeth in each ring varies roughly from 6-9.

The marginal suckers of arms I—III are larger than

the medial suckers of arms IV, while the marginal

suckers of arms IV are smaller than the medial suckers. The usual number of suckers and hooks on the

proximal half of arms I—III varies from 29-33, while

on arms IV, in specimens larger than P.L. 34 mm,

the half-arm count varies from 36-41.

The tentacles are long and robust. The tentacular

clubs are short, expanded, and very complex. The

club length varies from 20-25% of the P.L. The

locking zone has 4-6 thick transverse lamellae; adjacent to each at the base of the medial end lies a large,

smooth-ringed sucker, and alternating with these are

rounded knobs. Proximally, this structure is marked

off from the tentacular stalk by a large gap. Distally,

the thick lamellae abruptly decrease in size; generally,
50 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

the last two are rudimentary, but have typical suckers.

Suckers of the dorsal-marginal zone converge into approximately 3 rows which pass around the central

hook to merge with the suckers of the dactylus. Suckers in this group usually are absent at the proximal

edge of the club, although occasionally they extend

onto the stalk. In the median zone, a single, large

hook lies near the center of the club and is followed

distally by a hook of intermediate size and proximally

by 3-4 small hooks. The distal hook is rarely joined

by an enlarged sucker with a large median tooth. The

small hooks decrease in size proximally and are occasionally preceded by a proximal series of suckers.

The ventral-marginal zone consists of 3 or sometimes

4 longitudinal rows of suckers in its midportion, all

of which are nearly the same size. This group of suckers continues onto the tentacular stalk not only as the

marginal row, but also as a more medial, sligthly irregular series. Distally, this zone terminates opposite

the large central hooks.

The largest suckers from the ventral-marginal zone

have large, rather broadly separated, pointed teeth

on the distal margin of the inner rings which grade

into smaller teeth laterally. The proximal border is

smooth. The total number of teeth in each ring is

roughly 9-10. These suckers are approximately the

same diameter as the suckers from the medial rows on

arms IV. The suckers of the middle of the dactylus

have similar dentition, but with sligthly fewer teeth.

The suckers at the proximal end of the dactylus are

numerous and not arranged in distinct rows. Slightly

distally, they become aligned in 4 longitudinal rows.

The suckers in a transverse series are nearly the same

size proximally, but distally the suckers of the dorsal

row are slightly smaller than those of the ventral row.

The tip of the dactylus has a circlet of small, smoothringed suckers. The total number of suckers on the

dactylus (excluding the circlet), the dorsal-marginal

zone, and the ventral-marginal zone varies from 151—

184.

The locking apparatus on the manus continues

along the dorsal margin of the tentacular stalk proximal to the gap, but much reduced. The large lamellae

on the manus are represented by relatively low

rounded lobes which soon disappear; but they are

much more apparent than in any of the other 3 local

species of Gonatus. The marginal suckers and alternating knobs decrease in size proximally, and extend

along, roughly, three-fourths of the stalk. The suckers

along the ventral margins continue for about half the

length of the stalk. The suckers in the latter series

situated one-fifth of the tentacle length from the base

of the club have a few broad, slightly pointed teeth

on the distal margin of the inner ring. Almost all

specimens have an additional series of suckers on the

tentacular stalks which are continuous with the suckers of the ventral-marginal zone of the manus. This

series closely parallels the suckers of the ventral

margin on the stalk. This series is variable, but usually

extends about three-fourths of the length of the marginal row, although it is most clearly present in the

distal part. These suckers generally are arranged in a

single row, an irregular single row, or occasionally an

irregular double row. Sometimes, this series lies very

close to the marginal row; more frequently, it is separated from the latter by a distinct gap and rarely

contains the only suckers on the tentacular stalks that

lie medial to the marginal rows. More commonly, additional suckers are scattered about the oral surface

of the stalk. The number of suckers in the ventral row

varies between 80-110% of those in the dorsal row.

The number of suckers on the stalk between the marginal rows varies betwen approximately 50-125.

The thin buccal membrane attaches far up on the

bases of the arms. It has 7 lappets and the connectives

attach to the dorsal borders of arms I and II and to

the ventral borders of arms III and IV.

A single, large, white photophore lies on the ventral

surface of each eye. Each photophore is somewhat

oval and covers the greater portion of the ventral surface of the eye. It is immediately distinguishable from

the silver tissue surrounding the eye by its well-defined

boundaries and its peculiar surface which seems to

have many small pores.

Although the integument is damaged on all specimens, some indication of its color remains. The sides of

the head and the aboral sides of arms II and III and,

to a lesser extent, arms I, are covered by a brilliant

silvery layer. This layer extends over the ventral surface of the head, but it is not as well developed there.

It is not possible to tell whether or not this layer was

present on the lateral surfaces of the mantle. All of

the head, arms, fins, and mantle (including the silvery

layers) appear to have been covered by brownish red

chromatophores. The buccal membrane lacks pigmentation except for a few chromatophores on its

outer surface.

Hooks on the arms are present at a P.L. of 17-22
NUMBER 9 7 51

TABLE 12.—Measurements in mm for Gonatus pyros, new species.

Characters

Station Numbers of the VELERO

7415 7415 7389 7412 8031 8028 8028 8025 8123 7409

Sex

P.L

M.W. .. .

H.W

F.L

F.W

Arm L. I

II

III

IV

Yi A.C. I

II

III

IV

Tent. L..

ClubL..

Club S..

M.S.S. ..

D.S.S....

V.S.S....

35

10

13

15

25

19

23

22

23

31

35

34

41

43

8

170

104

86

74

38

10

13

15

29

20

25

24

23

29

31

31

38

44

8

180

67

78

68

40

12

15

17

29

24

27

27

26

30

33

31

38

45

10

181

57

84

72

(Holotype)

39

10

14

14

27

22

27

26

24

29

32

30

38

43

—

168

85

73

68

42

11

14

—

30

23

27

26

25

30

34

34

38

67

10

174

91

82

85

31

9

12

12

21

15

17

18

18

27

33

31

33

24

7

180

70

66

56

33

8

12

12

21

15

16

17

16

29

29

30

36

34

7

175

58

77

69

25

—

—

—

—

9

11

12

11

30

32

30

35

—

—

167

—

—

39

11

14

16

26

22

26

27

25

29

33

32

40

52

10

170

82

82

77

38

10

13

—

26

20

24

24

—

31

33

33

—

44

8

159

79

77

73

mm; the distal hook on the club begins to develop at

a P.L. of 18-23 mm; the median hook at a P.L. of

15-18 mm; and the proximal hooks at a P.L. of 21 -

26 mm. A spindle-shape liver lying oblique to the

body axis is present in the larva.

TYPE LOCALITY.—33°37' N, 118°26' W, eastern

North Pacific Ocean.

LOCATION OF TYPE.—University of Southern California. U.S.C. Hancock collections, AHF Cephalopod

Type No. 5.

DISCUSSION.—G. pyros is separated from all other

members of the genus by a large photophore on the

ventral surface of each eye. Another unique feature is

the double series of suckers along the ventral margin

of each tentacular stalk. Also among the features

which separate G. pyros from other species are: size

at which the hooks develop; number of suckers and

hooks on the proximal half of the arms; number of

suckers on the tentacular clubs; and the length of

the arms.

DISTRIBUTION.—No specimens of G. pyros were

taken south of latitude 32° N, and it is assumed that

this was the southern limit of the species during the

period covered by this survey. In the Santa Catalina

zone G. pyros is moderately abundant; its frequency is

0.47 specimens/hour of trawling. This species is

known only from the present trawling program. The

northern extension of the species, therefore, coincides

with that of the program.

Gonatus californiensis, new species

PLATES 15B; 16B,J-P; 17D-F

DESCRIPTION.—The mantle is long, slender and

moderately thick and muscular; it tapers to a point

posteriorly. The free edge projects at the ventral,

mantle locking-cartilage, but only slightly dorsally.

The muscular portion terminates at the conus. Posterior to the conus, a fleshy column continues as a tail

which is best developed in large specimens.

The fins have free anterior lobes and are attenuate

posteriorly along the sides of the tail.

The funnel is short and reaches to the level of the

lenses of the eyes. The funnel locking-cartilage bears

a straight groove that opposes a straight ridge on the

mantle. The dorsal pad of the funnel organ has an inverted V-shape. The apex is slightly indented. Each

arm is angled on its lateral edge and bears a median

ridge on the anterior portion. A slender papilla is

present near the apex. The funnel is entirely free

from the head laterally.
52 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The head is oval. Two large eyes occupy almost

the entire lateral surface of the head. A distinct sinus

is present on the anterior edge of each eyelid. Three

weak nuchal folds are present lateral to the funnel on

either side, the second of which bears the "olfactory"

organ. The nuchal cartilage is elongate, rounded anteriorly and posteriorly, and has a median ridge with

a central groove and marginal ridges.

Arms II and III are muscular and moderately

long; generally they are 40-50% of the P.L. Arms IV

are approximately the same length as II and III, but

are a weaker construction. Arms I are the shortest,

being roughly 35-45% of the P.L. Arms I—III bear

gelatinous aboral keels. A lateral keel is present on

each arm IV and extends the full length of the arm.

Well-developed trabeculate protective membranes are

present on all arms, but are less developed on arms

IV. The armature is in 4 series: arms I—III have 2

medial rows of hooks and 2 marginal rows of small

suckers. The suckers arise from the tips of the slender

trabeculae. At the arm tips, the hooks are replaced by

suckers. Arms IV have 4 rows of suckers only. All arm

suckers have similar dentition. Long slender teeth are

present on the distal part of the inner ring which

grade into smaller teeth on the sides; the proximal

margin is smooth. The total number of teeth varies

roughly from 9-13. The suckers of the median rows

on arms IV are the largest, followed by the marginal

suckers of the same arms and finally by the marginal

suckers of arms III, but the size differential is not

great.

The tentacles are moderately long and slender. The

tentacular clubs are small (17-24% of the P.L.). The

ventral-marginal zone of the club has 4-5 rows of

suckers in its central part. Proximally, they blend into

a single row which continues onto the tentacular

stalk. Suckers of the 3 lateral rows are nearly the same

size, while the medial row (or rows) are about half

the diameter of the larger suckers. This discrepancy in

size only becomes apparent in large specimens and it

is somewhat variable in these. The largest suckers

from the ventral marginal zone on the manus have

roughly 8-10 teeth on the distal two-thirds of the

inner ring. The teeth are moderately long and

pointed on the distal part of the ring and grade into

smaller teeth on the lateral sides. These suckers are

clearly smaller than the larger suckers on the arms.

The proximal margin appears to be smooth. The

suckers of the dactylus have similar dentition, but apparently have slightly fewer teeth. The locking zone

has 4-6 large, fleshy ridges with smooth-ringed suckers at their medial ends. Generally a single sucker is

present at the distal end of the zone. The suckers alternate with rounded knobs.

The dorsal marginal zone passes lateral to the large

median hook in 2-3 irregular rows that blend distally

into the suckers of the dactylus. Proximally, the

suckers are irregularly arranged and pass medial to

the locking zone where they decrease in number until

all suckers disappear near the base of the manus.

The median zone contains a very large central hook

with a moderately large hook immediately distal to

this. Proximal to the central hook are usually 3-5

small hooks followed generally by 1-2 suckers. At the

base of the dactylus, suckers are arranged in approximately 7-8 irregular rows, but decrease to 4 rows

about halfway out on the dactylus. The suckers of the

dactylus are nearly equal in size in a transverse series.

The dactylus terminates in a small circle of suckers. A

swimming keel is present on the dorsoaboral border

of the dactylus. A narrow, trabeculate, protective

membrane lies lateral to the suckers along the entire

ventral margin of the club. A similar dorsal membrane ends partway out on the dactylus. Both membranes extend down the entire length of the stalks,

but are barely visible.

A series of alternating suckers and pads extends

along most of the length of the tentacular stalk on its

dorsal margin. At the distal end, a few suckers are accompanied by small, fleshy ridges similar to those on

the locking apparatus of the manus. A single row of

suckers is present along the ventral margin of the stalk

for most of its length. The number of suckers in the

ventral row varies roughly between 90-120% of the

number in the dorsal row. Suckers in this series situated one-fifth of the tentacle length from the base of

the club have smooth inner rings. Between these 2

rows there are about 40-80 scattered suckers.

Seven indistinct lappets are present on the buccal

membrane. The buccal membrane connectives attach

to the dorsal borders of arms I and II and to the

ventral borders of arms III and IV.

The species has numerous reddish brown chromatophores. There is some indication that in life the lateral

sides of the head and arms have an underlying silvery

iridescent layer.

TYPE LOCALITY.—33°32' N, 118°24' W, eastern

North Pacific Ocean.
NUMBER 9 7 53

LOCATION OF TYPE.—University of Southern California. U.S.C. Hancock collections, AHF Cephalopod

Type No. 6.

DISCUSSION.—Gonatus berryi and Gonatus pyros

have unique features which easily separate them from

other members of the genus. G. onyx and G. californiensis are not as easily separated from one another.

The arrangement of hooks on the tentacular clubs of

the 2 species differs considerably; in order to demonstrate clearly that these differences are not merely

types of variation within a single species, they have

been correlated with sucker counts on the tentacles,

on the proximal halves of the arms, and in the middle

of the tentacular stalks (Figures 8-10). The figures

260-

240'

0

O 220'

<A

O 200-

d

z 180-

160-

A

A

A

A A

A

A

* • •

. • # S • '

• f *

* *

"A

A

:

* *

20 40 60 80

Pen Length

100

FIGURE 8.—Club-sucker counts, excluding median series and circlet at tip of dactylus, for

Gonatus onyx (dots) and Gonatus californiensis (triangles).

60

Pen length

100

FIGURE 9.—One-half arm counts of arms IV for Gonatus onyx (dots) and Gonatus

Californiensis (triangles).
54 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

80-

6 0 '

•

40 •

20'

A

A

A

A A

A

A A

•

* •

A*

A

•

\

A

» *

A

• 5 «

*

A

•

20 40 60

Pen length

80 100

FIGURE 10.—Sucker counts of the medial suckers, excluding suckers of the dorsal and ventral

marginal series, on the tentacular stalks for Gonatus onyx (dots) and Gonatus californiensis

(triangles).

also point out several other interesting features of

these species. At about a P.L. of 40-45 mm in G. onyx

and about 60-65 mm in G. calif orniensis, the number of suckers on arms IV no longer increase. Therefore, although the arms will more than double in size

as the animal continues to grow (the maximum sizes

of these species are not known), the number of suckers

will remain constant.

A similar situation occurs with the number of

suckers on the tentacular clubs. After the sucker buds

have all appeared on the clubs between roughly 10-

20 mm P.L., no additional suckers are added. Thus,

at almost the first appearance of the definitive clubs

the number of suckers is fixed. These features of the

arms and tentacles indicate that sucker counts, although little used in the past, can provide excellent

taxonomic characters, at least in Gonatus, and should

be included in all future descriptions of species within

this group.

DISTRIBUTION.—Gonatus calif orniensis is the least

abundant member of the genus in the collections, yet

it is the only species that has been captured in zone 7

around Guadalupe Island. Eleven specimens have

been captured there, giving a catch frequency of 0.15

compared to 0.24 specimens/hour of trawling in zone

2. The indication is that this species, although primarily a northern form, extends farther to the south

than any of the other species of Gonatus of this area.

Since all known records of Gonatus are primarily

from temperate or colder waters, it would be expected

that G. calf orniensis does not extend too much farther

south than recorded here, although, McGowan

(1967) recorded larvae identified as Gonatus fabricii

as far south as 25° S off Baja California. Recently,

the PILLSBURY captured a larval Gonatus (P.L.,

17 mm) near the Bay of Panama (7° 19' N, 79°42'

W). This record is extremely surprising and if the

specimen belongs to one of the species found off California, as would seem likely, then it can be identified

as G. californiensis. The small size of the specimen

would suggest that it had been spawned in the vicinity

of Panama. I cannot adequately account for this rec-
NUMBER 9 7 55

Sex

P.L ... .

M W . .

F.L

FW

Arm L. I. .

II

III. .

IV

Vi A.C. I. .

II. .

Ill

IV..

Tent. L.. .

Club L.. .

ClubS...

M.S.S. .. .

D.S.S

V.S.S.. ..

TABLE 13.—Measurements in

Characters

8114

29

9

10

18

9

12

12

10

31

36

38

37

19

6

241

—

8116

52

15

20

36

20

24

25

23

30

36

38

50

43

10

269

45

66

80

mm

7411

48

13

20

30

20

24

24

23

30

35

35

44

47

10

235

59

71

80

for Gonatus californiensis, new

Station Numbers of the VELERO

8114

34

11

12

23

13

16

16

14

33

38

36

42

23

8

234

55

68

75

8114

38

11

13

25

14

18

18

17

31

36

36

41

31

9

225

59

72

67

9960

46

15

32

19

24

23

22

34

37

38

48

37

10

233

30

78

80

7157

50

13

19

31

18

23

23

22

35

38

39

50

10

217

48

65

67

species.

7157

52

14

21

32

19

24

24

25

35

40

39

51

38

10

236

22+

—

7410

cf

107

22

52

58

43

50

53

55

40

40

41

56

97

18

219

40+

—

7221

(Holotype)

112

21

49

60

47

56

58

62

44

47

45

57

80

19

259

57

82

90

ord, but believe it unlikely that the normal range of

this species extends so far south. This problem can

only be resolved by further trawling.

At present G. californiensis is definitely known to

extend from latitutde 34° N to 28° N off the coasts

of California and Baja California.

DISCUSSION OF THE GENUS GONATUS.—Within the

genus Gonatus, sensu strictu, there was previously

considered to be only a single valid species, G. fabricii

(Lichtenstein 1818), with a worldwide distribution.

The discovery of 4 sympatric species of Gonatus living

off southern California suggests that some of the

other forms thought to be synonyms of G. fabricii may

represent valid species. This is most likely true with

G. antarcticus (Lonnberg 1899) from Antarctic

waters and G. kamtschatica (Middendorff 1849)

from the northwestern Pacific. It is not known

whether or not there is more than a single species in

the North Atlantic. Until a study of the latter area is

undertaken, the true identity of G. fabricii and its accurate characterization must remain in question. Similarly, there is confusion concerning the correct identification of G. kamtschatica and G. antarcticus. Both

have been inadequately described. The collections at

the Institute of Marine Sciences indicate that at least

2 species are living in Antarctic waters and at least

2 additional species are living in the Bering Sea. Also,

Sasaki (1929) described an adult and a juvenile specimen of "G. fabricii" from the northwestern Pacific.

From his description and illustrations, it is apparent

that Sasaki was dealing with 2 species and that the

juvenile, at least, is different from any of the abovementioned species. The latter form recalls in general

appearance a larval gonatid described by Okutani

(1966, plate 3: figure 2) called Gonatidae y. Okutani

also considered that these belonged to the same

species. Therefore, at least 7 species of Gonatus are

present in the north Pacific, 2 in the Antarctic and 1

in the Atlantic, with only those from off southern

California adequately characterized.

With this unsatisfactory state of the genus, it is

nearly impossible to properly compare the California

species with those from other areas; however, G.

berryi, G. onyx, and G. pyros each has a number of

unique features that allow its separation from other

species. Gonatus californiensis is very similar to specimens that I have seen from the North Atlantic and

Antarctic oceans. It is with some hesitation that I

give this species a new name, since it is not possible

to separate it clearly from forms in these latter locali-
56

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ii

o

« i

'/////////A G. pyros

G. californiensis

G. onyx

G. berryi

'//A

10 15 20 25

Pen length, mm

30 35

FIGURE 11.—Size at which arm hooks develop for species of Gonatus. Empty squares indicate

only suckers present. Squares with Xs show intermediate condition (first hint of hook development). Solid squares indicate hooks present including small incompletely formed hooks. Crosshatched rectangles represent estimated size range over which hooks begin to develop.

ties due to lack of sufficient material. A few differences, however, that substantiate its specific ranking

have been noted. The clubs of Gonatus californiensis

are larger than those of the specimens of G. "fabricii"

examined from the North Atlantic, but are considerably smaller than those from specimens captured in

the Antarctic. There are also differences in the sizes

at which the large hooks develop on the tentacular

clubs between specimens of G. "fabricii" and Gonatus

californiensis.

Genus Gonatopsis Sasaki, 1920

DIAGNOSIS.—Seven teeth present in a transverse

series on the radula. Tentacles present only in larvae.

TYPE SPECIES.—Gonatopsis octopedatus Sasaki,

1920.

Gonatopsis borealis Sasaki, 1923

PLATES 18B; 19I-P; 36H

Gonatopsis borealis Sasaki, 1923, p.2O3, text-fig, p.205;

1929, p.274, pl.30, figs. 3-6.—Okutani, 1966, p.65, pl.II,
NUMBER 9 7 57

5

3

E 2

r

'///////A G. pyros

n

G. californiensls

v//////////////.

'/////////////A

G^ onyx

G. bcrryi

V////////A

20 25

Pen length, mm

30

FIGURE 12.—Size at which the central club hook in species of Gonatus develops.

figs. 3-4.—Pearcy, 1965, p.261.

DESCRIPTION.*—Anteriorly the mantle is cylindrical, but it begins tapering approximately at the level

of the anterior margin of the fins and terminates in a

blunt point just short of the posterior margin of the

fins. The mantle wall is extremely thick and muscular. The free anterior margin of the mantle projects

slightly at the funnel and nuchal locking cartilages.

The fins are broad (approximately 65-70% of the

mantle length), moderately long (over 40% of mantle length), and muscular. The fins are sharply angled

*K. N. Nesis (1971) has proposed that G. borealis be

placed in a new subgenus, Boreoteuthis. The work of M.

Okiyama (1969) and W. Fields and V. Gauley (in press)

further indicates that present generic definitions in the

Gonatidae are no longer adequate.

at their lateral extremities and taper posteriorly to

form a broadly rounded apex where their posterior

margins fuse. The anterior lobes are free.

The funnel is well developed and powerful. The

funnel locking-cartilage has a long straight groove

which broadens posteriorly and opposes a long ridge

that widens posteriorly on the mantle. The dorsal

member of the funnel organ is an inverted V-shaped

pad with a small anterior papilla and a low, short

ridge near the anterior end of each lateral arm. The

lateral arms are long and extend onto the end of the

funnel retractor muscles. The ventral pads are large

and oval. A funnel valve is present.

The head is cylindrical and has a well-defined, inverted U-shape funnel groove. A nuchal crest is present from which arise 4 nuchal folds on either side of
58

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

C

0

"o

8. «

* 3

*5 2 6

Q. pyros

n

G. californiensis

V/////////////.

G. berryi

v///////////////////,

15

Pen length, mm

FIGURE 13.—Size at which the distal club hook in species of Gonatus develops.

M

0

V//////////A G. pyros

n IT

6. californiensis

V////////////A

n i r

G. berryi

W////////////A

15 30 40 45

P*r\ length, mm

FIGURE 14.—Size at which the first proximal club hook in species of Gonatus develops.
NUMBER 9 7 59

mini

G. pyros

G. californiensis

mi

G. berryi

mum

iiiiiiiiiimimiiii

'////A

Illllllllllllllllllllllll

5 10' 15 20 25 30 35

'ft Arm hooks

^ C. hook

mi Dhook

— P. hooks

' 40

Pen length.mm

FIGURE 15.—Pattern of hook development for the four Californian species of Gonatus:

C. hook, central club hook; D. hook, distal hook; and P. hooks, proximal hooks.

the head. The 2 dorsal folds on each side join posteriorly. The second fold from the funnel curves dorsally and bears a small "olfactory" papilla. The

ventral-most fold is continuous with the margin of

the funnel groove. The eyes are very large and extend

nearly the entire length of the head. A large eye sinus

is present on the anterior-ventral margin of each

eyelid. The nuchal cartilage is large, slightly constricted in the middle of its length and bears a long,

straight median ridge with a central groove.

The arms are long, muscular, and in the order II =

III>I>IV. Arms II and III are roughly 40-50%

of the mantle in length. Aboral keels are well developed on arms III, but are less apparent on arms II

and form only a small ridge on arms I. Arms IV have

well-developed lateral keels. All arms have low, trabeculate protective membranes. Each arm I—III has

39^8 hooks arranged in 2 medial rows and 2 marginal rows of small suckers that arise from the trabeculae of the protective membranes. Near the tips of

the arms, the hooks are replaced by large suckers. The 4

rows of suckers thus formed extend to the tips of the

arms and usually consist of 56-63 suckers. At the base

of the arms, proximal to the hooks are 5-10 suckers.

Each arm IV has 4 rows of suckers with 2 median

rows somewhat enlarged. All suckers have approximately 12-15 sharply pointed teeth on the distal half of

the inner chitinous ring. The proximal margins are generally slightly irregular except on the suckers from

the tips of arms III, at least, where the proximal

border has distinct small knobs. Peculiarly, in the

smaller suckers it is the narrow edge of each tooth

which faces the exterior while the broad face lies

parallel to the adjacent teeth.

Tentacles are absent except in the larvae.

The largest male, although not fully mature, has a

long (95 mm), slender penis that reaches nearly to

the mantle opening. There is no evidence of hectocotylization. The female has paired oviducts and typical

nidamental glands.

There are 7 lappets on the buccal membrane. The

buccal membrane connectives attach to the dorsal

sides of arms I and II and to the ventral sides of arms

III and IV.

The color in preservation is provided by numerous

very small purple-brown chromatophores.

The juvenile at 34 mm M.L. is very similar in appearance to the adult; the major exceptions are that

only suckers are present on the arms and that the fins

are proportionally smaller.
60

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

In larvae the arms are proportionally shorter, much

broader, and bear 4 distinct rows of suckers. Tentacles are present. In the youngest forms they are

short and broad, but with pointed tips. The oral face

of the tentacle is flattened and bears 3-5 irregular

longitudinal rows of suckers. The rows break up

proximally and near the base of the tentacle lie 1-3

enlarged suckers. Although differing in the manner in

which they arise from the head, the tentacles are

very similar to the arms in size and shape.

In the more advanced larva the tentacles are

slenderer and much longer than the short arms IV,

but only slightly longer than arms III. The enlarged

suckers at the base of the club are very prominent. At

a later stage, the tentacle is much more slender than

any of the arms and is equal in length or slightly

shorter than the arms which are roughly subequal at

this stage. The enlarged proximal suckers are still

prominent. There is considerable variation in the size

of the larvae at which these stages of tentacle development occur. At roughly 8-11 mm M.L. the tentacles are lost abruptly.

TYPE LOCALITY.—Kushiro, Hokkaido.

LOCATION OF TYPE.—In Hokkaido Imperial University.

DISCUSSION.—The genus Gonatopsis was established by Sasaki (1920) to include a new species

Gonatopsis octopedatus from the western North Pacific. This gonatid was particularly distinctive in its

lack of tentacles and the possession of 8-12 rows of

minute suckers on the distal one-third of each arm.

In 1923 Sasaki described another member of the

genus, G. borealis, which also lacked tentacles, but

had only 4 rows of suckers at the tips of the arms.

Okutani and Nemoto (1964) have described a subspecies of the latter, Gonatopsis borealis makko, from

specimens taken from the stomachs of sperm whales

in the North Pacific. This form differs from G. b.

TABLE 14.—Measurements in mm of Gonatopsis borealis Sasaki, 1923.

Characters

Sex

M.L

M.W

H.W

F.L

F.W

ArmL. I

II

Ill

IV

A.C. Hooks I

II

Ill

A.C. Prox. S. I

II

Ill

A.C. Dist. S. I

II

Ill

A.C. IV

Station Numbers oj the VELERO

10402 8697 10401 10260 8022

<? 9 <? — —

200 240 257 34 30

44 61 60 12 10

41 49 — — —

85 107 110 13 10

137 155 164 26 22

99 104 101 14 11

108 110 113 15 13

107 113 110 14 12

88 93 99 14 11

L R L R L R L R L R

39 40 45 43 42 40 — — — —

39 40 48 44 44 42 — —

39 39 47 46 44 43 — —

7 7 10 — —

7 6 8 — —

6 6 8 — —

Total Arm S. Total Arm S.

57 54 60 129 129

59 61 61 137 139

58 59 57 131 131

L R L R L R

129 153 139 149 — 177 144 144
NUMBER 9 7 61

borealis primarily in having a slenderer mantle,

longer arms (two-thirds of the M.L. versus two-fifths

of the M.L.), a slightly different arm formula, and

smaller fins. These differences are sufficient to indicate that G. b. borealis and G. b. makko should be

considered as distinct species and not as subspecies.

The distinctness of G. makko is further enhanced by

the recent description (Okutani 1966: 66, plate III:

figure 1) of a larval squid, thought to belong to this

species, which differs greatly in shape from the larvae

of G. borealis.

The present material of G. borealis differs from

Sasaki's description in the slightly smaller size of arms

IV and in the slightly greater number of suckers and

hooks on the arms. At best, these differences can only

be considered to represent geographical variation.

DISTRIBUTION.—Previously, G. borealis has been

recorded off California (Rice 1963) from a whale

stomach, off Oregon (Pearcy 1965), the Bering Sea,

off Kamchatka (55°40' N, 180°00' and 49°50' N,

165°49' E) (Okutani 1966), and off Japan (Sasaki

1929).

Sasaki's comments (1929) on G. borealis are worth

repeating:

Specimens examined have mostly come from Kushiro, but

also include those from Nemuro and Tokachi. These localities cover the eastern coast of Hokkaido strongly influenced by a cold current 'Oyashio.' Most of the specimens

of Yamaguchi and Shirai referred to were caught on 28th

July, 15-30 miles off Kushiro, where the water temperature

showed 11.6°C. at the surface and 2.5°C. at the depth of

20 fathoms below the surface. According to the reports of

Mr. Yamaguchi and Mr. Shirai who made researches on the

habits of the species by angling from the experimental boat

"Sangyo-maru" 1922, whenever Ommastrephes sloani pacificus [=Todarodes pacificus] was abundant, Gonatopsis

borealis was very rare, the reverse being also the case, or

when both the species happened to be met with at the

same region of the sea, the latter species was always angled

from a much greater depth than the former. All these facts

point to the conclusion that Gonatopsis borealis is physiologically adapted to much colder water than Ommastrephes

sloani pacificus, which generally inhabits water above 10°C

in temperature. [Spelling has been corrected.]

In the present study the frequency of capture of

G. borealis in the Santa Catalina Basin was 0.40

specimens/hour of trawling. The adjacent basins

(zones 1 and 3) show surprisingly low frequencies

(0.06 and 0.05 specimens/hour, respectively). The reason for the discrepancy is uncertain; however, the

relatively high frequency of capture in the Santa Catalina Basin, the most intensively studied area, indicates

that this species is fairly abundant in the northern

sector of the area under study. Only 4 specimens have

been taken in zone 6 and only a single one in zone 7

(frequency 0.08 and 0.01 specimens/hour). Although

the data is somewhat confusing due to the low frequency of capture in zones 1 and 3, I think that it is

safe to assume that zone 7 roughly coincides with the

southern limits of this species in the eastern North

Pacific.

Family ONYCHOTEUTHIDAE Gray, 1849

DIAGNOSIS.—Arms with biserial suckers. Tentacular clubs with tetraserial armature of which the 2

marginal rows may disappear during growth. Clubs,

where known, bear hooks in the medial rows. Buccal

connectives attach to the ventral sides of arms IV.

Funnel locking-cartilage with a simple, straight

groove. Nuchal folds present.

Genus Onychoteuthis Lichtenstein, 1818

DIAGNOSIS.—Tentacular clubs with 2 rows of

hooks; marginal suckers absent except in early

juveniles. Arm suckers with smooth chitinous rings

(teeth lacking). Photophores present on viscera and

a single photophore present on the ventral bulbus of

each eye.

TYPE SPECIES.—Onychoteuthis bergii Lichtenstein,

1818.

Onychoteuthis borealijaponicus Okada, 1927

PLATES 18A; 19A-G

Onychoteuthis banksii.—Berry, 1912c, p.83, text figs.44-48;

1913, p.77.—PPfeffer, 1912, p.79, p.85.—Sasaki, 1916,

p.90; 1929, p.228, pl.20, fig. 12, pl.30, figs. 1-2.

Onychoteuthis borealijaponicus Okada, 1927, p.4.

Onychoteuthis banksi?—Clarke, 1962, p. 424.—Pearcy,

1965, p.262.

DESCRIPTION.—The mantle is long and slender

(width 22-27% of length) ; the anterior half is cylindrical, and the posterior portion tapers to a terminal

point. The mantle wall is thick and muscular. The

free margin projects in the nuchal region and at the

mantle cartilages. The muscular portion of the mantle

terminates at the conus. The pen is clearly visible the

entire length of the mantle in middorsal line. The rostrum of the pen extends posterior to the conus and is
62 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

easily visible in ventral view. The rostrum is surrounded by a cone of gelatinous connective tissue.

The fins are 54-60% of the M.L. in length and 70-

79% of the M.L. in width. The anterior lobes are

free. Posteriorly, the fins terminate at the tip of the

mantle.

The funnel is small; it extends to the level of the

middle of the eyes. The funnel cartilage is pointed

anteriorly, flares slightly posteriorly, and bears a

straight median groove. The dorsal pad of the funnel

organ has an inverted V-shape. The posterior portion

of the lateral limbs broaden abruptly; the anterior

portion bears a weak lateral ridge. A low, conical

papilla is present at the anterior apex. The ventral

pads are very large and oval. The funnel is free from

the head laterally and contains a large valve.

The head is relatively small, being equal to or less

than the mantle width. The large eyes occupy the

entire lateral sides of the head. Each eyelid has a

large anterior sinus. Distinct dorsal and ventral "windows" are present on either side of the head. A funnel

groove with well-defined margins is present; in the

median anterior line the margin turns forward to

form an acute angle. There are 9-10 folds on the

nuchal crest on either side of the head; the second

bears the "olfactory" papilla. The second to fifth or

sixth folds are joined posteriorly. The first fold connects with the midportion of the second. The nuchal

cartilage is elongate, constricted slightly laterally, and

bears a prominent median ridge with a central

groove.

The arms are short (longest arms of the largest

specimens are 40% of the M.L.) and in the order

2 = 3 = 4> 1. Aboral keels are well developed on arms

II and III, but are present only on the distal halves

of arms I. Lateral keels are present on arms IV. Low

trabeculate protective membranes are present on all

arms. The armature consists of biserial suckers

throughout the length of each arm. The inner chitinous rings of the suckers are smooth; the outer rings

are reduced. The suckers of arms IV are distinctly

smaller than those of the other arms.

The tentacles are short and robust. The tentacular

clubs measured to the proximal end of the carpus

are 24-27% of the M.L. The club is not clearly divided into manus and dactylus. It bears biserial hooks

and no suckers except for a small cluster of 15-16

smooth-ringed suckers at the tip of the club. Marginal

rows of small suckers, however, are present on the

club in juveniles up to a M.L. of at least 33 mm. The

chitinous rings of these suckers are smooth. The first

5 hooks of the ventral row increase gradually in size

distally and have symmetrical bases. The following

2-3 hooks increase greatly in size and have asymmetrical bases. The succeeding hooks gradually decrease in size. Each hook distal to the largest hook

frequently bears a small knob near the ventral

border of its base, although this is somewhat variable.

In the dorsal row, the second hook is usually slightly

larger than the first. The following hooks gradually

decrease in size to the fifth or sixth hook. The succeeding hooks increase slightly in size, then gradually diminish in size. With the possible exception of the first

few proximal hooks, the hooks of the dorsal row are

much smaller than those of the ventral row. The

carpus occupies an oval patch clearly marked off from

the surrounding tissue. It contains 8-10 suckers and

pads. The tentacular stalk is bare. A protective membrane is present along the full length of the ventral

margin of the club. The proximal portion is much

more expanded. A low membrane is present along the

dorsal border lateral to the 5 proximal hooks. A large

dorsoaboral keel is present. An additional separate

keel extends along the aboral surface of the tentacular

stalk.

The buccal membrane bears 7 lappets and the connectives attach to the dorsal border of arms I and II

and to the ventral borders of arms III and IV.

A large photophore is present over the ventral surface of each eye. An oval photophore is present on the

ventral surface of the ink sac and an ill-defined photophore is present on the ventral surface of the ink

duct just before the duct enters the intestine. Part of

the ink sac and the ink duct between the photophores

is covered by a band of tissue with an iridescent gold

sheen which Okada thought was also photogenic

tissue. O. banksii, a closely related form, has a very

large and well-defined anal and intestinal photophores with no iridescent tissue between them. The

animal is speckled with reddish brown chromatophores.

TYPE LOCALITY.—Tokachi, Hokkaido, Japan.

LOCATION OF TYPE.—In the collection of Hokkaido

Imperial University.

DISCUSSION.—In recent years Onychoteuthis

banksii has been considered the only valid member of

its genus. Throughout its worldwide distribution it

can easily be recognized by its robust appearance,
NUMBER 9 7 63

large hooks on the tentacular clubs, a large photophore under each eye, ancf 2 photophores next to the

intestine. In 1927, Okada described a new species,

O. borealijaponicus, from Japan based primarily on

the slightly different form and structure of the light

organs. Sasaki (1929), in his monograph of the Japanese cephalopods, placed this species in the synonymy

of O. banksii indicating that Okada's distinguishing

characters might be a result of developmental or seasonal differences. I have examined specimens from

12-100 mm M.L. and find that the appearance of the

photophores can easily separate the 2 species throughout this size range. O. banksii also is much more robust than is O. borealijaponicus.

I have attempted to find additional characters to

separate the 2 species, but have run into difficulties.

The "O. banksii" from Florida waters clearly fall into

2 separate groups and, therefore, the former worldwide species "O. banksii" now seems to consist of at

least three species: O. borealijaponicus from the

North Pacific and 2 species from Florida waters. It

seems likely that "O. banksii' will eventually be

found to consist of a number of additional species,

and until this complex can be worked out it will be

difficult to completely characterize O. borealijaponicus. O. borealijaponicus, however, can be separated

from the Florida specimens, by the light organs, and

by the tentacular clubs. I have counted the number

of hooks on 37 specimens of "O. banksiV from off the

coast of Florida and the results are as follows:

Number of club hooks

Number of clubs examined

19 20 21 22 23

4 28 28 13 1

Voss (personal communication) has examined the

type of O. banksii and finds that each club carries 21

hooks. The number of hooks on the clubs of the relatively few large specimens of O. borealijaponicus

available to me clearly fall outside of this range:

Number of club hooks

Number of clubs examined

25

2

26 27

4

In addition, the relative sizes of the fins, arms, and

tentacular clubs separate O. borealijaponicus from

each of the Florida species, but in different ways.

DISTRIBUTION.—O. borealijaponicus was taken

throughout the area under study, although most records (72 of 95 specimens) came from zones 5 and 6.

This species is a powerful swimmer and presumably

epipelagic, and is thus ill-suited for capture by the

TABLE 15.—Measurements in

Character

Sex

ML

MW

H W

FL

F.W

Arm L. I

II

Ill

IV

II

Ill

IV

Tent L

Club L

Club H

Carp S

Carp K

mm of Onychoteuthis borealijaponicus Okada, 1927.

Station Numbers of the VELERO

8237

81

18

18

47

61

27

32

31

32

23

24

21

26

47

20

L R

27 27

9 9

9 9

8121

63

14

13

35

48

20

25

23

23

19

21

20

25

35

15

L R

26 26

9 10

10 9

8114

50

11

10

27

36

15

19

18

17

18

23

22

26

32

13

L R

27 26

9 10

10 9

8121

48

13

11

26

38

14

17

16

16

17

20

18

23

28

13

L R

25 25

00 ONON 00

Pawlson Col-

- lection off

southern

California

9

100

23

20

60

70

32

41

39

40

23

23

21

26

73

26

L R

26 26

9 10

10 9
64 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Isaacs-Kidd mid-water trawl. The collection consists primarily of larval forms which probably are

taken incidentally when the trawl is being lowered

and retrieved. Considering this method of collection,

the results are rather difficult to interpret, but they

seem to indicate that the young, at least, are more

abundant in the more oceanic zones of the study.

Pearcy (1965) recorded O. banksii from off Oregon. I have examined one of his specimens now at

the Institute of Marine and Atmospheric Sciences,

Miami, and find that it can be referred to O. borealijaponicus. O. borealijaponicus is also known from

off the western coast of Canada (Roper, personal

communication) and the eastern coast of Hokkaido,

Japan. Other records of O. banksii from the high

latitudes in the North Pacific (Clarke 1966) probably

belong to this species. I examined a specimen of a

more typical "O. banksii" from 16° N, 123° W, and

Berry (1914) reported the more typical form from the

Hawaiian Islands. It appears that O. borealijaponicus

replaces "O. banksii" in the colder waters of the

North Pacific.

Family MASTIGOTEUTHIDAE Verrill, 1881

DIAGNOSIS.—Arms with biserial suckers. Tentacles

whiplike, with long unexpanded clubs bearing many

thousands of minute suckers. Funnel locking-cartilage

with an oval depression, usually bearing 1 or 2 knobs

(tragus, antitragus). Buccal connectives attach to the

ventral borders of arms IV. Small light organs generally present on the surface of the mantle, head, and

arms.

Genus Mastigoteuthis Verrill, 1881

DIAGNOSIS.—Cartilaginous tubercules not present

on surface of mantle, head, funnel, and arms.

TYPE SPECIES.—Mastigoteuthis agassizii. Verrill,

1881b.

Mastigoteuthis pyrodes, new species

PLATES 25; 26I-Q

DESCRIPTION.—The mantle is cylindrical for most

of its length, but posteriorly it tapers rapidly, and the

muscular portion terminates in a blunt point just posterior to the anterior end of the fins. The mantle continues past this point as a narrow cylindrical cord of

tissue which merges into the tail. The tail extends for

a short distance posterior to the fins, but its exact extent is uncertain since it is invariably damaged. The

pen appears to extend to the tip of the tail. Anteriorly, the mantle projects strongly in the nuchal region, but less so in the areas of the mantle lockingcartilages.

The fins are large, about 60% (excluding the tail)

of the M.L., and nearly circular in outline. The anterior lobes are attached for the most part, but project

anteriorly very slightly.

The funnel reaches approximately to the level of

the posterior edges of the lenses. It contains a large

valve and a well-developed funnel organ with an inverted V-shaped dorsal pad bearing a small anterior

papilla, and 2 oval ventral pads. The funnel lockingcartilage has an oval depresion with a strong tragus

on the medial margin. Occasionally, a low antitragus

can be detected; however, it is usually not apparent.

The head is slightly broader than long. Its width

is nearly equal to the M.W., and its length is approximately 20% of the M.L. The eyes are very large, occupying almost the full length and depth of the head.

An eye sinus is not apparent on the eyelid. A crescentshape patch of tissue borders the posterior margin of

the eyelid, and a small "olfactory" papilla is located

posterior to each eye.

The arms are large, the ventral pair extremely so,

being comparable to the mantle in length. The arm

formula is IV>II>III>I. Although the arms are

generally damaged, they appear to contain rather

feeble protective membranes supported by flabby, triangular trabeculae. Arms IV have large lateral keels.

It is not possible to determine the presence or absence of swimming keels on the other arms due to

damage. The suckers are arranged biserially and have

slender stalks arising from large conelike bases which

are fused with the lateral trabeculae. The oral face of

each arm IV is very narrow and the suckers are

forced nearly into a single row, particularly in the

distal half of the arm. The number of suckers in the

proximal half of each arm in the largest female

(165 mm M.L.) is 36, 36, 39, and 38 for arms I

through IV, respectively. Each sucker ring bears

sharply pointed teeth on the distal margin and truncate teeth on the lateral margins. The total number of

teeth varies roughly between 10 and 15. The proximal edge of the chitinous ring is smooth. There is no

evidence of hectocotylization in males.
NUMBER 9 7 65

The tentacles are very long, over twice the mantle

length, and very slender. The club is unexpanded and

approximately 50% of the tentacle length. The club

possesses a microscopic protective membrane with tiny

trabeculae which defines the borders of the suckerbearing portion of the club. The proximal end of the

club is damaged in all specimens, so its exact structure cannot be determined with certainty; however,

it appears that the sucker-bearing area tapers rapidly

to an abrupt end. Distally, the suckers progressively

cover increasingly more of the club surface until they

completely surround the tentacle a short distance

from its tip. The smallest suckers occur at the tip of

the club. The largest suckers occur in a broad area

about one-third of the club length from its distal end

and are arranged in approximately 18-22 rows. The

suckers have long, slender stalks which result in a disorderly arrangement in preservation and makes an

exact determination of the number of rows difficult.

The club suckers are elongate in the proximal-distal

plane. The outer chitinous ring is broad and each

lateral margin of its inner edge has 2-3 greatly enlarged, blunt teeth that project into the aperature of

the sucker. The inner ring has 3 distinct blunt teeth

on the distal margin; the proximal and lateral margins are smooth. The largest suckers on the club

measure 0.28-0.32 mm in the long diameter of the

holotype (110 mm M.L.). The club in its middle

portion has a diameter about 7 times the width of

the largest suckers of this portion.

The numerous photophores present are almost exclusively in the outer integument and, therefore, are

easily lost with only a small amount of abrasion. As

a result of such injury to the present specimens, it is

impossible to determine the exact distribution of light

organs. The ventral surface of the mantle is covered

with small photophores which appear to diminish in

number laterally. On the ventral surface of each fin,

adjacent to the mantle, photophores are lacking,

while lateral to this area scattered light organs appear, but gradually diminish toward the lateral margins of the fins. Thus, there appears to be a somewhat

narrow patch of photophores down the ventral surface of each fin. Photophores are abundant on the

dorsal surface of the fins, particularly on the lateral

halves. Both the dorsal and ventral surfaces of tHe

funnel are sprinkled with photophores as are the

ventral and lateral surfaces of the head. A large light

organ is present deep in the tissue near the anteroventral margin of each eyelid. It is oval in shape and

may be completely hidden by an aggregation of dark

chromatophores. Numerous light organs are present

on the ventral surface of arms IV.

The buccal membrane attaches to the dorsal

borders of arms I and II and to the ventral borders

of arms III and IV.

The color in preservation consists of a reddish

purple background pigmentation upon which dark

reddish purple chromatophores occur in the outer

integument.

TYPE LOCALITY.—33°32' N, 118°23' W, eastern

North Pacific Ocean.

LOCATION OF TYPE.—University of Southern California, U.S.C. Hancock collections, AHF Cephalopod

Type No. 7.

DISCUSSION.—It is often impossible to identify with

certainty many species of Mastigoteuthis because of

several factors. These animals are soft and apparently

inhabit a rather deep zone in the ocean. As a result,

specimens are usually brought to the surface badly

damaged; their long tentacles are often lost along

with the delicate epidermis that frequently (if not

always) is covered with photophores. Although these

squid apparently are common inhabitants of the deep

sea, only rarely is more than 1 or 2 specimens taken

at a time. These difficulties, compounded by the

large number of species within the genus, have led to

the present systematic jumble. Many species are

known only from a single mutilated specimen, and

most species are inadequately described.

It is with some hesitation that I add a new name

to a group that is in such a state of instability; however, there appears to be no other course available.

In the following brief review of the 15 nominal

species, pertinent characters are listed only when they

are derived from the type material.

Mastigoteuthis agassizii Verrill, 1881b. Club suckers

are armed with 2-3 sharp teeth on one side. Arm

suckers have smooth inner rings. Fins are 60-65% of

the M.L. in length. The arm formula is IV>III =

II>I. At the proximal end of the club, the suckers are

in 2 rows. Light organs are distributed over the entire

body. Tragus and antitragus are both weakly developed. Type locality: Off east coast of United

States (about 34° N; 76° W). I examined the smaller

of VerrilFs syntypes which unfortunately lacks tentacles, and find that there is an enlarged photophore

imbedded in the anteroventral margin of each eyelid.
66 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

This species can be easily separated from M. pyrodes

by the lack of dentition on its arm suckers.

Mastigoteuthis grimaldii (Joubin 1895). Tentacles

are not known. Arm suckers have 13 pointed teeth on

the distal margin of the inner ring. The fins are 54%

of the M.L. in length. The arm formula is IV>III =

II>I. Light organs are distributed over the entire

body. No antitragus is present. Type locality: Near

Azores (39°43' N; 33°22' W). This species was described from a juvenile specimen that had a M.L. of

approximately 26 mm. Due to its lack of described

characters, it cannot be separated with any certainty

from 5 of the other species of the genus and, therefore, must be considered a species dubia.

Mastigoteuthis levimana Lonnberg, 1896. Tentacles

are not known. Arm suckers have broad, truncate

teeth around the entire circumference of the inner

ring. The fins are 38% of the M.L. The arm formula

is IV>III = II>I. Arms IV are more slender than

others and almost whiplike. Suckers are present only

on the proximal portions of arms IV and are arranged in intervals of 2 to 4 mm. Type locality:

North Atlantic (43° N, 24° W). The fragile structure

of the ventral arms with their peculiar arrangement

of suckers, the small fins, and the sucker dentition,

indicate that this species does not belong in Mastigoteuthis and is here placed in the genus Valbyteuthis

(Chiroteuthidae).

Mastigoteuthis dentata Hoyle, 1904. Tentacles are

not known. The arm suckers have 13 teeth on distal

half of the inner ring. Fins are 43-67% of the M.L.

in length. The arm formula is IV>III = II>I. Type

locality: Off Cape Mala, Panama (7°21' N, 79°35'

W), and east of the Galapagos Islands (0°36' S,

86°46' W). As can be seen from the preceding diagnosis, Hoyle gave very few characters of specific value

in the original description. Also, judging by the great

discrepancy in fin length between his 2 specimens, it

appears that he probably had 2 species. In order to

clarify this situation, I here designate his syntype from

off Cape Mala (number 7957, male, 72 mm M.L.)

as the lectotype. I examined this specimen which is

deposited in the United States National Museum (according to C. F. E. Roper, Associate Curator of Mollusks, United States National Museum, the other

syntype has been lost). Although the animal has

undergone complete desiccation, it is still possible to

determine from bits of integument still clinging to the

animal that the entire body is covered with photophores and that an enlarged photophore is imbedded

in the anteroventral margin of each eyelid.

Recently the PILLSBURY captured 2 specimens

from off Cape Mala which agree in all discernible

features with M. dentata. From these specimens,

which unfortunately were badly damaged, I can add

the following characters to the description of this

species: the clubs, although badly scraped in places,

appear to occupy about half of the tentacular length.

The largest suckers occur about one-third of the club

length from the distal end. This area of the club bears

suckers in roughly 35-40 rows. Suckers near the base

of the club are smaller and less thickly spaced, but

their precise arrangement cannot be determined.

Suckers also decrease in size near the lateral margins

of the club. The inner chitinous rings of the club

suckers are smooth (Plate 26T). The inner edge of

the outer ring has 12-14 small knobs around its entire margin that project slightly into the aperature of

the inner ring (Plate 26s). The club suckers are very

small. The long diameter across the outer ring of the

largest club suckers from a specimen with P.L. of 92

mm was from 0.13-0.16 mm. A minute trabeculate

protective membrane is present. The funnel lockingcartilage has a tragus and only a weakly developed

antitragus, although the posterior end of the cartilage

is well undercut. The 2 specimens had fin lengths of

66% and 67% of the M.L.

I had originally thought that the specimens of

Mastigoteuthis from southern California belonged to

this species; however, after examining the specimens

from Panama, there can be no doubt that the 2 forms

belong to separate species. The much smaller size of

the tentacular suckers as well as the differences in the

dentition of the inner and outer chitinous rings clearly

verifies their separate status.

Mastigoteuthis talismani (Fischer and Joubin

1906). Arm suckers are smooth (?). The fin is 89%

of the M.L. in length. Photophores are present on the

fins and probably over the entire body. There is no

distinct antitragus. Type locality: South of Azores

(34°46' N, 36° 11' W). The original and only specimen of this species was very badly multilated. The

club length is given as about one-tenth of the tentacle

length; however, this is almost certainly a misinterpretation resulting from the poor condition of the

animal and should not be considered a specific character. The larger size of the fins easily separates this

form from M. pyrodes.
NUMBER 9 7 67

Mastigoteuthis glaukopis Chun, 1908. The club is

seven-tenths of the tentacle length. Club suckers have

10-12 small teeth around the margin of the inner

ring. Club suckers measure 0.1 mm in a specimen

with a M.L. 37 mm. Arm suckers have 5—7 large

teeth on the distal portion of the inner ring and somewhat smaller teeth on the proximal side. The arm

formula is IV>II>III>I. The fins are 50% of the

M.L. in length. A single photophore is imbedded in

the anteroventral border of each eyelid. Other photophores are apparently absent. No distinct antitragus

is present. Type locality: Region of the east African

coast. Due to the small size of Chun's specimen, it is

not certain whether the dentition of the club suckers

refers to the inner or outer ring. The dentition of the

arm and club suckers, fin size, and possibly some of

the other characters separates this species from M.

pyrodes.

Mastigoteuthis flammea Chun, 1908. Tentacles are

not known. Arm suckers have 3-5 teeth on distal

margin of inner ring. The arm formula is: IV>

II>III—I. Tragus and antitragus are present. Photophores are distributed over the entire body. Eyes

are reduced in size. Fins are 50% of the M.L. Type

locality: Gulf of Guinea (0°25' N, 7°00' W; 1°14' N,

2° 10' W. This species can easily be distinguished

from all others now known by the small size of the

eyes.

Mastigoteuthis cordiformis Chun, 1908. The club

is two-thirds of the tentacle length. Club suckers have

numerous pointed teeth around the entire margin of

the inner ring with those of the distal border being

much larger. Diameter of the suckers is 0.5 mm in a

specimen of M.L. 83 mm. Arm suckers have about 30,

very small, rounded teeth on the distal two-thirds of

the inner ring. The fins are 78% of the M.L. in

length. A tragus is present and an antitragus is present, but small. The arm formula is: IV>II>III = I.

Proximal suckers of the club are arranged in 2 rows.

The middle portion of the club has approximately 25

rows of suckers. The whole body is covered with small

light organs. Type locality: South of Sumatra (0°15'

N, 98°8' E). This species is easily separated from M.

pyrodes by its large fins and by the dentition of the

club suckers. Descriptions by Chun (1910), Sasaki

(1929), Adam (1954), and Voss (1963) indicates

that this species exhibits considerable variation.

Mastigoteuthis famelica (Berry 1909). Tentacles

are not known. Arm suckers are "minutely toothed."

The arm formula is: IV>II>III—I. Fins are longer

than broad. Type locality: Vicinity of Kauai Island,

Hawaii. This species, based on a single specimen of

39 mm M.L., is inadequately known. It differs from

California specimens of the same mantle length by

having much shorter arms (each arm IV in M. famelica is 20 mm; in M. pyrodes of the same size each

arm IV is 35 mm). A redescription of M. famelica

is badly needed; unfortunately, the type, which was

deposited in the United States National Museum, has

been destroyed (Roper, personal communication).

Therefore, the further elucidation of this species depends on the capture of new material from the

Hawaiian Islands.

Mastigoteuthis magna Joubin, 1913. The club is

nine-tenths of the tentacle length. Club suckers are so

small that they are invisible to the naked eye. The

inner rings of the club suckers are smooth. Arm suckers have smooth inner rings. The fin is 65% of the

M.L. The arm formula is IV>II>III = I. The funnel locking-cartilage bears a straight groove anteriorly

that enters a bowl-shape depression posteriorly. Photophores are apparently absent. Type locality: Middle

of North Atlantic Ocean (31°45' N, 42°39' W).

This species is very distinctive and can easily be

separated from all other species by, among other

things, its peculiar funnel locking-cartilage.

Mastigoteuthis hjorti Chun, 1913. Tentacles have

suckers of 0.25 mm diameter at the proximal end of

the club in a specimen with a mantle length of 95

mm. Tentacular suckers are arranged in 18-20 rows.

Arm suckers have 9-10 teeth on the distal half of the

inner rings. Fins are equal to the M.L. The arm

formula is IV>III>II>I. Two large photophores

attached to the bulbus of each eye. Type locality:

North Atlantic Ocean (31°24' N, 34°47' W). This

species is easily separated from all others by its

enormous fins and by the presence of photophores on

the eyes.

Mastigoteuthis latipinna (Sasaki 1916). The club

is three-fifths of the tentacle length. The proximal

part of the club is slightly flattened and has broad

protective membranes with trabeculae. The proximal

suckers of the club are equal to large arm suckers in

size and bear 10 blunt, minute teeth on the distal

margin of the inner rings. Distally, the suckers

become smaller and more extensively toothed.

The arm suckers have smooth inner rings. The arm

formula is IV>II>III>I. The tragus and antitragus
68

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

are very faint. The fins are 84% of the M.L. The

left eye is much the larger. Type locality: Sagami

Sea. This species is very distinctive and can be distinguished from M. pyrodes by almost any of the characters mentioned above.

Mastigoteuthis schmidti Degner, 1925. The club

is three-fifths of the tentacular length. The club

suckers have smooth inner rings and lack inward

projecting knobs on the outer rings. Arm suckers have

13-14 slender teeth on the distal border and 11-12

blunt teeth on the proximal border. The arm formula is IV>III = II>I. The fin is 65% of the M.L.

Tragus and antitragus are present. Photophores are

distributed over the entire body. From 20-23 suckers

are present in transverse series in the distal third of

the club. Type locality: Eastern North Atlantic

(46°30' N, 7°00' W). This species can be easily separated from M. pyrodes by the dentition of the inner

and outer rings of the tentacular suckers.

Mastigoteuthis atlantica Joubin, 1933. Tentacular

suckers have 2 or 3 elongate knobs that project inward from the inner edge of the chitinous ring. The

inner chitinous ring is apparently smooth. Sucker diameter is 0.36 mm in a specimen with a mantle

length of 112 mm. Arm suckers have about 11

sharply pointed teeth on the distal border of the inner

ring; proximal borders are smooth except for a possible cleft in the median proximal line. The arm

formula is IV>II>III>I. A photophore is embedded

in the anteroventral margin of each eyelid. No antitragus is present. Fin is 71% of the mantle length.

Type locality: Near Bay of Biscay (46°28' N, 8°01'

W). Joubin's measurements state that the club is 31%

of the tentacle length; however, his illustration indicates that it is much longer. Joubin originally placed

this form as a subspecies of M. glaukopis, owing to

the presence of a large photophore embedded in each

eyelid in both forms. It is now apparent that this feature is widespread throughout the genus and that

M. glaukopis atlantica cannot remain at the subspecific level; it is here elevated to specific status. The

differences in sucker dentition and possibly fin size

are sufficient to verify that M. atlantica and M. glaukopis require separate specific status. This species is

very close to M. pyrodes, but differs primarily in the

dentition of the inner rings of the tentacular suckers

and in its locality.

Mastigoteuthis iselini MacDonald and Clench,

1934. Tentacles are not known. Arm suckers have 13

pointed teeth on the distal margin of the inner ring

TABLE 16.—Measurements in mm for Mastigoteuthis pyrodes, new species.

Characters

Station Numbers of the VELERO

7279 8934

10841

or

10844

11181 10844 11225 9905 9905 9905

Sex

M.L

M.W....

F.L.V. .

F.W

ArmL. I.

II.

Ill

IV.

Tent. L..

Club L. .

EyeD...

TailL...

Club S.D.

(Holotype)

110

27

64

61

42

55

52

99

233

121

15

17+

.28-.32

65

17

36

37

23

30

26

56

—

—

—

7

—

35

—

18

18

6

14

8

35

—

—

5.5

3

—

165

—

89

96

70

85

79

126

—

—

22

20 +

—

148

34

89

90

62

78

74

120

378

150

21

20+

.30-.35

86

—

50

50

33

47

44

79

188

90

14

12 +

.24-.26

94

20

55

58

52

41

85

15

—

77

16

44

45

29

41

31

70

12

7 +

84

18

47

45

26

37 +

31

70+

13

13

•Excluding tail length
NUMBER 9 7 69

which are replaced by 6 blunt teeth on each side of

the ring. The median proximal portion is slightly

crenulated. There is ho antitragus. Fins are 67% of

the mantle length. The arm formula is IV>III—

Type locality: 39°04' N, 71°29' W. This form is inadequately described and cannot be distinguished

with certainty from M. atlantica, M. dentata or M.

schmidti and must be considered a species dubia.

DISTRIBUTION.—M. pyrodes is known only from

the present trawling program. One specimen each was

taken from zones 1 and 2, none from zones 3 and 4,

4 from zone 5, 5 from zone 6, and 16 from zone 7.

Apparently this species is more abundant in the offshore zones and, in particular, the more southern

zones.

Family CHIROTEUTHIDAE Gray, 1849

DIAGNOSIS.—Arms with biserial suckers except occasionally at the arm tips. Tentacular clubs with

tetraserial suckers. Funnel locking-cartilage oval and

with 1 or 2 distinct knobs (tragus, antitragus) projecting toward the center of the cavity. Buccal connectives attach to the ventral borders of arms IV.

Genus Chiroteuthis Orbigny, 1845

DIAGNOSIS.—Arms IV greatly enlarged. Photophores present at least on eyes and arms IV. Funnel

locking-cartilage generally with both tragus and antitragus. Tentacles greatly elongated and with secondary clubs (larval clubs) lost.

TYPE SPECIES.—Loligopsis veranyi Ferussac, 1835.

Chiroteuthis calyx, new species

PLATES 20; 21A,B; 22A-K,N-Q

Chiroteuthis veranyi.—Berry, 1963, p. 128, text figs. 1-6.—

Pearcy, 1965, p.262.

DESCRIPTION.—The mantle is short and conical; its

musculature is weak and terminates in a rounded

point just past the anterior end of the fins. The pen

continues posterior to the fins, but has been broken

off in all specimens so that its full length is unknown.

The mantle margin protrudes in the middorsal line

and very slightly near the mantle locking-cartilages.

The fins are nearly circular in outline, 55-65% of

the M.L. in length, and have free anterior lobes. On

the posterior margin of the fins, where the pen

emerges, there is a slight indentation of the musculature covered by a transparent membrane.

The funnel is moderate in size, and is free from the

head laterally. The funnel locking-cartilage is oval

and bears both a tragus and an antitragus. The dorsal

pad of the funnel organ has an inverted V-shape with

short, wide diverging arms and a prominant anterior

papilla. The ventral pads are oval. A small, delicate

valve is present.

The head is elongate and cylindrical, although

swollen slightly in the region of the eyes. Nuchal folds

are lacking. The "olfactory" papillae which arise

from the ventral surface of the head lateral to the

exhalant opening of the funnel have long stalks and a

bumpy, terminal swelling. Each eye is large and the

eyelid bears only a slight indication of an anterior

sinus. The posterior margin of the eyelid consists of a

large muscular patch of tissue.

The arms are very large and in the order of IV>

III>II>I. Arms IV are extremely long, over V/t

times the mantle length, and broad. Arms I—III have

low protective membranes supported by fleshy trabeculae which fuse with the broad bases of the sucker

stalks. Protective membranes and trabeculae are absent from arms IV. Arms I—III possess thick, low,

keels. Arms IV have usually broad, lateral keels which

exceed the arm diameter in width. The suckers on

each arms I—III are biserial except at the arm tip.

The arm tip (i.e., approximately one-tenth of the

M.L. on each arm) bears small, closely packed suckers in 3 longitudinal rows. The suckers on each arm

IV are broadly separated and arranged in 2 alternating series which tend to converge distally into nearly

a single line at the arm tip. Very large globular suckers occur on the middle third of arms I and II, but

are somewhat more distally situated on arms III. The

suckers on arms IV are largest at the base of the

arms, then gradually diminish in size distally.

The enlarged suckers of arms I—III have nearly

smooth rings, but may show regular indentations in

the margin indicating that they have been formed by

the fusion of broad, truncate teeth. On the arm proximal to the enlarged suckers, the inner rings have

roughly 18-20 broad, truncate teeth which are separate from one another on the distal two-thirds of the

sucker margin; the proximal one-third is smooth. On

the arm distal to the enlarged suckers, individual

teeth on the suckers also can be recognized. Near the
70 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

tip of the arms, the suckers have relatively longer and

slenderer truncate teeth on the distal margins and

nearly smooth but slightly irregular proximal margins. Distinct truncate teeth also occur on all suckers

of the ventral arms. The largest suckers of arms III

are more than twice the diameter of the basal suckers

of arms IV and are more than 3 times the diameter

of the adjacent suckers on arms IV.

The tentacles are extremely long, as much as 5-6

times the mantle length; however, their length depends greatly on their state of contraction at the time

of fixation. The right and left tentacles in one specimen at hand differ in length by 10 cm, while those

of the holotype differ by 6 cm. Thick oval pads are

loosely adherent along the length of the tentacle

stalks, except near the club where they become embedded in the stalk and decrease in size. It is impossible to get an accurate count of their numbers since

many are invariably lost during capture and preservation. It is estimated, however, that there are over 50

of these organs on each tentacle in specimens of about

60 mm M.L.

The club is long (55-65% of the M.L.) and

bordered by protective membranes with large broad

trabeculae. The membrane is divided into a proximal

and a distal portion of nearly equal lengths by a slight

decrease in width and by a change in the structure of

the trabeculae. Each proximal trabecula is split into

2 or 3 parts in its outer half, except for the small

trabeculae in the initial 10% of the club. Each proximal trabecula is separated by a small gap from its

neighbors. The trabeculae of the distal portion of the

club are undivided and broadly separated from one another at their bases. The club is not differentiated

into a dactylus, manus, and carpus. The suckers of

the club are arranged in 4 longitudinal rows. Each

sucker is set on a long stalk composed of 2 portions.

A rather broad cylindrical basal portion terminates in

a dark purple, pleated "skirt" from which arises the

slender distal portion of the stalk that bears the

sucker. The basal portions of the marginal stalks are

nearly twice the size of the medial ones and each has

a narrow, transparent keel along its lateral edge

which runs onto the protective membrane between

the trabeculae. The number of suckers varies from

134-142 on the adult clubs in specimens examined.

The inner chitinous ring of each sucker bears a large

recurved median tooth on its distal margin along

with approximately 10-12 smaller lateral teeth. The

proximal margin is smooth.

Two large photophores lie embedded in the ink sac,

one to either side of the intestine. The ventral surface

of each eye is provided with 2 longitudinal strips of

luminous tissue which parallel each other on the inner

and outer eye margins. Lying between these strips at

either end is an oval luminous patch. An additional

small luminous organ is situated between the 4 other

organs and slightly in front of the posterior patch

(Plate 22N). The large pads on the tentacular stalks

may be photogenic. The tip of the club is occupied by

a very large photophore which faces aborally. Large

spherical photophores, each covered by a layer of

dark-brown chromatophores, line the lateral-oral

edges of arms IV along the base of the lateral membrane.

The buccal membrane attaches to the dorsal

borders of arms I and II, and to the ventral borders

of arms III and IV.

In preservation C. calyx is speckled with numerous

red-brown chromatophores.

Apparently all members of the genus possess a

larval stage that attains an unusually large size. C.

calyx is no exception; its larval form (it is a "doratopsis" larva) has a maximum size of about 60 mm

M.L. This larva has recently been described by Berry

(1963), and it is necessary to mention here only the

following points. The late doratopsis stage has enlarged suckers on the arms, the largest of which have

many closely set, slender, truncate teeth. The full

compliment of adult club suckers is present proximal

to the larval club. The larval club has approximately

63-65 suckers, tetraserially arranged and set on short

stalks. The suckers of the ventral row are considerably

larger than those of the other rows. At the tip of the

club is a small lobe which carries 5 suckers.

TYPE LOCALITY.—Santa Catalina Basin?, eastern

North Pacific Ocean.

LOCATION OF TYPE.—University of Southern California. U.S.C. Hancock collections, AHF Cephalopod

Type No. 8.

DISCUSSION.—Presently, there are 8 recognized

species belonging to the genus Chiroteuthis in addition

to C. calyx. Three species, C. imperator Chun, 1910;

C. picteti Joubin, 1894; and C. macrosoma Goodrich, 1896, are easily distinguished from C. calyx by

the presence of 3 rows of oval photophores on the

ventral surface of each eye (many other characters

separate them equally well). C. capensis Voss, 1967,
NUMBER 9 7 71

seems more closely related to the above 3 species although the eye photophore pattern, which is imperfectly known, seems to be more similar to the pattern

in C. calyx. At any rate, it can easily be separated

from the California species by the dentition of the

suckers (8-10 long, slender, sharp teeth on the upper

edge of the ring and about 20 low, blunt teeth on the

lower edge of the sucker ring), by the lack of photophores on the viscera, by the lack of a striped "skirt"

on the stalks of the club suckers, and by the different

proportional lengths of the arms, mantle, and clubs.

C. atlanticus (MacDonald and Clench 1934) is

known only from a single mutilated specimen; however, its elongate mantle clearly indicates its relationship to the above-mentioned species. C. joubini Voss,

1967, can easily be distinguished by the unique photophore pattern on the eye (10 oval photophores in 2

series) and by the presnce of a single, median visceral photophore.

The remaining 2 species, C. lacertosa Verrill, 1881,

and C. veranyi (Ferussac 1835), are much more

closely related to C. calyx than are the other 6 species.

C. lacertosa was considered a subspecies of C. veranyi

by Pfeffer (1912). These 2 forms are unquestionably

very close and even the validity of C. lacertosa still

has not been definitely established. The 2 forms differ

in the shape of the fins (they are round in C. veranyi,

but elongated—60 mm long by 40 mm wide—in C.

lacertosa) ; otherwise, the animals closely agree in

most characters. Unfortunately, the holotype of C.

lacertosa lacks tentacular clubs; however, if the clubs

found by Verrill on other occasions belong to this

species, as Verrill believed, then it would appear that

the species is more similar to the California species,

particularly regarding the arrangement of the lateral

trabeculae on the club, and that C. lacertosa is certainly a valid species. It is, however, necessary to examine more material from the type locality of C.

lacertosa (i.e., Brown's Bank off Nova Scotia) before

its relationships can be unquestionably resolved. At

present, C. calyx can be separated from C. lacertosa

by the differences in sucker dentition (sharply pointed

teeth in C. lacertosa and truncate or fused teeth in C.

calyx, different shape of the fins (circular in C.

calyx, elongate in C. lacertosa), and by the greater

size of the enlarged suckers on arms I—III in C. calyx.

Of the clearly established species, C. calyx is most

closely related to C. veranyi. Relative to the latter

species, the suckers in the middle section of arms IIII in C. calyx are considerably enlarged, while the

TABLE 17.—Measurements in mm for Chiroteuthis calyx, new species.

Characters

Station Numbers of the VELERO

9890 8292 10260 7280 9349?

Sex

M.L

M.W

H.L

H.W

F.L

F.W

Arm L. I

II

III

IV

Tent. L.

ClubL. .

ClubS..

EyeD..

Larva Larva

49

7

28

6

18

19

7

11

14

33

46

12

idult Larval

142 53

4

56

12

24

9

22

23

14

20

26

49

51

18

Adult Laval

140 63

7

55

17

21

16

23

28

38

51

62

87

190

30

137

11

60

25

20

24

29

61

74

91

126

40

142

13

(Holotype)

60

17

23

20

25

28

48

61

69

100

215

34

134

14
72

suckers of arms IV are relatively reduced in size. The

ratio of the diameter of suckers at the midsection of

arms III to adjacent suckers on arms IV is greater

than 3:1 in C. calyx, while it is less than 2:1 in C.

veranyi (Plate 22B, I, L, M). This difference between

the 2 species is large enough that it can be detected

at a glance. In C. calyx, the sucker rings have distinctly truncated teeth on the distal margin, which

on the largest arm suckers may all fuse giving a completely smooth ring. In C. veranyi, the suckers have

many sharply pointed teeth on the distal half of the

ring.

The final major differences between the 2 species

concern the protective membranes of the tentacular

club. In C. veranyi, the proximal portion of the membrane is much broader than in C. calyx. Also, in C.

veranyi, the individual supporting trabeculae in this

membrane can be distinguished only at their bases

since they are subdivided for most of their length;

these subdivisions are all closely aligned making it impossible to separate the subdivisions of one trabecula

from those of its neighbors (Plate 21c). In C. calyx,

the trabeculae are subdivided only in their distal half

and the trabeculae remain separated from their

neighbors by a distinct gap (Plate 21B).

It is worth noting that the arrangement of photophores is identical in C. veranyi, C. lacertosa, and C.

calyx.

DISTRIBUTION.—I examined a specimen of C. calyx

taken by Pearcy off Oregon at 44°40' N and 125° 16'

W, and his record of C. veranyi (1965) from Oregon

certainly can be referred to this species. The only

other record of C. calyx is from off southern California (Berry 1963, as C. veranyi).

In the present study, the frequency of capture was

fairly even in zones 1, 2, 3, and 5 (i.e., 0.16-0.26 specimens/hour). The highest frequency of 0.52 specimens/hour occurred in zone 6; however, this high

figure is largely the result of a single, unusual trawl

capture that yielded 15 larvae. If this trawl was

ignored, the catch frquency for zone 6 would be 0.24

specimens/hour. No specimens were captured in zone

7.

Although the data are somewhat contradictory, the

complete lack of specimens from zone 7 coupled with

the records from Oregon suggests that this is a coldwater species with a southern limit of distribution off

Baja California.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Genus Valbyteuthis Joubin, 1931

DIAGNOSIS.—Funnel locking-cartilage oval. Antitragus present, tragus absent. Tentacular club short,

compact and with suckers in 4 rows. Club suckers on

short stalks. Fins subterminal. Photophores absent.

Funnel valve absent.

TYPE SPECIES.—Valbyteuthis dana Joubin, 1931.

Valbyteuthis oligobessa, new species

PLATES 23A,B; 24H-N,Q,R

DESCRIPTION.—The thin, delicate mantle is saclike,

except posteriorly where it tapers to a point. The

anterior mantle margin is broad and only loosely surrounds the head and funnel.

The fins are small, about 23-33% of the M.L. in

length. Each is roughly semicircular and has only

slightly projecting anterior and posterior lobes. The

broad, tubular conus of the pen projects posteriorly

past the fins. It has been broken in all specimens so

that its full extent is unknown. The fins join in the

dorsal midline.

The funnel is very large; it reaches anteriorly to the

level of the anterior margins of the eyes in mature

specimens, but is smaller in immature forms. The

funnel locking-cartilage is distinctive of the genus;

the cartilage has an oval depression that deepens

posteriorly where it is overhung by a small antitragus.

The opposing mantle cartilage has the appearance of

a human nose with a single nostril; the "nostril" locks

with the antitragus. The dorsal member of the funnel

organ is very small relative to the large size of the

funnel. The organ is approximately shield-shape with

a rough or papillate surface and an anterior papilla.

This papilla, at its base, is slender, but distally it

broadens in the transverse plane and is slightly cupped. The ventral pads are somewhat variable, but

generally are elongate and oval and slant obliquely toward the midline. There is no funnel valve.

The head is elongate and cylindrical. The eyes are

small; in all specimens, they have been damaged. The

paired bridles and cephalic vein are prominent along

the ventral surface of the head beneath the funnel.

"Olfactory" papillae are present lateral to the funnel

and considerably posterior to the eyes. Each consists

of a very elongate stalk with a terminal swelling. The

nuchal cartilage is straight, slender, and elongate. A
NUMBER 9 7 73

central ridge with a median groove is barely detectable on its anterior half.

The arms are short, extremely delicate, and in the

order IV>III>II>I. In young specimens, arms IV

are greatly elongate (up to 130% of the M.L.), but in

the mature animal they are less than the length of the

mantle and only slightly longer than arms III. There

is only a slight indication of protective membranes,

and keels appear to be lacking from arms I—III, although lateral keels are present on arms IV. The

suckers on the arms are biserially arranged along the

entire length of each arm I—III. On arms IV, the

suckers are broadly separated and arranged in apparent single file. These are preset only on the proximal end of the arms and number only 2-A on each

arm.

The outer chitinous ring of each arm sucker consists of an outer row of parallel scales followed medially by 3-4 rows of scales with generally square bases.

The latter bear large knobs which have enlarged,

flattened ends in the most medial series of the distal

half of the ring. On the proximal side of the ring,

the knobs are replaced by hollow cylinders that have

a portion of one side missing (Plate 24R). The inner

chitinous ring has about 25-35 small, blunt teeth

around the distal three-fourths of the margin. The

proximal portion is nearly smooth.

The tentacles are relatively robust and long. Each

tentacular club is short, only slightly expanded, and is

not differentiated into a manus and a dactylus. Low,

protective membranes are present on both borders.

The suckers are arranged in 4 regular longitudinal

rows over most of the club, but become slightly irregular at the tip and at the base of the club. The

suckers of the ventral row are the largest. The suckers

become progressively smaller in a transverse series, although the suckers of the dorsal row are still only

slightly smaller than those of the ventral rows. Five or

six suckers, arranged in 2 alternating rows, are present

in the carpus. The larger of these is sligthly larger

than the suckers of the club.

The buccal membrane connectives attach to the

dorsal borders of arms I and II and to the ventral

borders of arms III and IV.

No photophores could be found. This animal is

covered with numerous reddish brown chromatophores.

The holotype is a gravid female. The mantle cavity

is occupied almost entirely by a large mass of mature

eggs and large nidamental glands. The ripe eggs are

about 1.5 mm in diameter. Several large sperm

reservoirs can be found in the enlarged ovary and,

surprisingly, one can be seen protruding through each

nephridiopore. Paired oviducts are present with enTABLE 18.—Measurements in mm for Valbyteuthis oligobessa, new species.

Characters

Sex

M.L

M.W

F.L

F.W

Ann L. I

II

III

IV

Tent L

Club L. *

ArmS. IV

10265

23

9

7

15

9

11

14

24+

L R

2 3

11101

33

11

87

6

Station Numbers of the

10976

34

11

7

12

12

15

18

46

67

6

L R

3

10265

34

11

8

11

15

16

41

55

4

L R

3 2

VELERO

10973

36

13

9

10

14

45

L R

3 3

11181

9

70

23

14

17

20

23

10976

9

(Holotype)

76

28

14

19

27

29

30

L R

2 3

*Does not include carpus
74 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

larged oviducal glands. This is the first record of a

mature specimen in the genus.

TYPE LOCALITY.—32° 17' N, 120° 12' W, eastern

North Pacific Ocean.

LOCATION OF TYPE.—University of Southern California. U.S.C. Hancock collections, AHF Cephalopod

Type No. 9.

DISCUSSION.—In addition to Valbyteuthis oligobessa there are, presently, only 2 other described species in the genus: V. danae Joubin 1931, from the

Pacific Ocean off Panama and V. levimana Lonnberg, 18%, from the North Atlantic; however, at

least 2 undescribed species are also present in the

Atlantic. It seems that this genus will eventually prove

to be rather speciose. V. oligobessa differs from V.

danae in a numbr of respects. The most obvious distinguishing feature is the much smaller size of the fins

in V. oligobessa (23-33% of the M.L. versus 50-55%

of the M.L. in V. danae). Other characters include

the number of suckers on the ventral arms {2-A in

V. oligobessa and at least 12-15 in V. danae), the

nnumber and shape of teeth on the inner chitous

rings of each arm sucker (25-35 narrow teeth in V.

oligobessa and 7-10 broad teeth in V. danae), the

relative size of the tentacular club suckers compared

to the size of the largest suckers of each arm (Plate

24B-F, H-L) and the size of the sucker orifice (Plate

24B-F, H-L) . This latter character is particularly

noticeable on the distal half of each arm in V. danae

where the size of the orifice relative to the size of the

sucker rapidly decreases, whereas the sucker orifice in

V. oligobessa show very little relative reduction. In

addition, the suckers of the ventral row on the tentacular clubs of V. oligobessa are slightly larger than

those of the dorsal row. In V. danae both rows have

equal-size suckers.

It is more difficult to separate the Californian species from V. levimana, since the latter is known only

from the type description which was based on two

mutilated specimens. The only definite characters

of specific value are the greater number of suckers

(6-8) on each arm IV in V. levimana and the dentition of the arm suckers, which is described in V.

levimana as consisting of broad, truncate teeth around

the entire aperture.

DISTRIBUTION.—V. oligobessa is known only from

the present trawling program. Four specimens were

taken in zone 2, 1 in zone 4, 5 in zone 5, 3 in zone 6,

and 5 in zone 7. If the number of trawling hours in

each zone is considered, it is apparent that this species is most abundant in the offshore zones. This distribution probably results from the circumstance that

Valbyteuthis is one of the "deepest living" species

taken in the program.

Valbyteuthis danae Joubin, 1931

PLATES 23C; 24A-G

Valbyteuthis danae Joubin, 1931, p. 188.—Roper and Young,

1967, p.l.

DESCRIPTION.—The mantle is cylindrical anteriorly,

but tapers rapidly posteriorly; the muscular portion

terminates at roughly the midpoint of the fins. The

pen is broken at the posterior end of the fins, so its

full length is unknown. The free margin projects

slightly at the nuchal and ventral mantle locks.

The fins are large (52% of the M.L. in length) and

muscular. Each is nearly semicircular and has free

anterior and posterior lobes.

The funnel reaches to approximately the level of

the middle of the eyes. The funnel locking-cartilage

has an oval depression and a distinct antitragus. This

opposes a cartilage on the mantle that is shaped like

a human nose with a single nostril; the latter locks

with the antitragus. The dorsal pad of the funnel

organ is nearly triangular, but has a slight posterior

indentation. The ventral pads are oval and very

large. There is no funnel valve.

The head is elongate and broadest posteriorly. The

bridles and cephalic vein are prominent on the ventral surface of the head up to the level of the middle

of the eyes. The head has been damaged; no other

details can be ascertained.

The arms are slender and delicate. The arm formula appears to be IV>III>II>I; however, all arms

have been damaged and it is impossible to get accurate measurments. All of the arms have low trabeculate protective membranes and biserial suckers.

On arms I—III, the suckers near each arm base have

relatively larger aperatures than the more distal

suckers. Near the arm tips the suckers become globular and have reduced aperatures and reduced outer

chitinous rings. Suckers are present only on the proximal portions of arms IV. These suckers are distinctly

biserial and number 12 on the left arm and 13 on

the right arm. The inner chitinous rings of the arm

suckers have about 7-9 broad, truncate teeth on the

distal margin; the proximal margin is smooth.
NUMBER 9 7 75

The tentacles are long and relatively robust; the

clubs are short and compact. Suckers are arranged

tetraserially and possess broad outer rings and inner

rings with narrow aperatures and smooth margins.

The club is not divided into a manus and dactylus.

The carpus has 6 suckers arranged in 2 alternating

rows. A thick protective membrane lines either side

of the club.

The buccal connectives attach to the dorsal borders

of arms I and II and to the ventral borders of arms

III and IV.

TYPE LOCALITY.—Off Panama, 7°3O' N, 79° 19' W.

LOCATION OF TYPE.—Copenhagen Museum.

DISCUSSION.—The above description is based on

a single, damaged specimen (o", 44 mm M.L.). It

agrees in all respects with specimens of V. danae

captured at the type locality off Panama. The illustrations used in this report are taken from Roper and

Young (1967) and are based on specimens taken

from the eastern tropical Pacific Ocean. For comparisons with the other species presently known, see

discussion of Valbyteuthis oligobessa.

DISTRIBUTION.—V. danae was previously known

only from the Pacific Ocean off Panama at about

latitude 7° N and off Peru at about latitude 8° S.

Within the Panamanian region, however, this species

appears to be fairly common. The single specimen

described here was taken in the Santa Catalina Basin

(VELERO Station 8120) at 32°20' N, 118°43' W.

There seems to be little doubt that this specimen is a

straggler from farther south.

Family GRIMALDITEUTHIDAE Pfeffer, 1900

DIAGNOSIS.—Arms with biserial suckers. Tentacles

absent. Buccal connectives attach to the ventral

borders of arm IV. Funnel and mantle lockingcartilages fused. Nuchal articulation free.

Genus Grimalditeuthis Joubin, 1898

DIAGNOSIS.—The characters coincide with those of

the family.

TYPE SPECIES.—Grimalditeuthis richardi Joubin,

1898.

Grimalditeuthis bomplandii (Verany, 1837)

PLATES 27B; 24O,P

Loligopsis Bomplandii Verany, 1837, p.99, pi. la.

Chiroteuthis Bonplandii.—F£russac and d'Orbigny, 1839, p.

326.—Gray 1849, p.44.

Grimalditeuthis Richardi Joubin, 1898a, p. 101, figs. 1-2;

1899, p.71; 1900, p.79, pl.4, figs. 1-2, pl.5, figs. 1-13,

pi. 10, figs. 1-2.

Grimalditeuthis Bonplandii.—Pfeffer, 1900, p. 188.—Chun,

1910, p.217; 1913, p.8.

Grimalditeuthis bonplandii.—Voss, 1956, p. 151, text fig. 13b.

DESCRIPTION.—Th e soft, gelatinous mantle is

slender (width is 16% of M.L.) and tapers gradually

toward the posterior end. Near the level of the anterior edges of the fin, the muscular layers of the

mantle diminish. The gelatinous portion of the mantle continues as a thick cylindrical mass of tissue

which is divided internally into a number of compartments giving the appearance of a honeycomb.

The funnel and mantle locking-cartilages are fused

to one another and thus form a permanent union between the mantle and funnel. The mantle is free in

the nuchal region.

The fin is large (46% of the M.L. in length and

6 1% of the M.L. in width) and relatively muscular.

It is subcircular in outline. The anterior lobes are

almost completely attached. Posterior to the fins, the

mantle and pen have been broken off. At this point,

the pen is very broad and undoubtedly continues for

some distance in the intact animal. Joubin (1900)

had an intact specimen in which the pen and mantle

continued posteriorly to support a second pair of fins.

The funnel is very small. No trace remains of a

funnel organ or funnel valve due to damage.

The head is cylindrical, elongate, and bears a pair

of small, lateral eyes. A pair of "olfactory" papillae

is present at the posterior edge of the head. Each

papilla lies immediately lateral to the funnel and

consists of a terminal swollen bulb set on a slender

stalk.

The arms are long and slender; arms IV have exceptionally attenuate tips. All arms lack protective

membranes, trabeculae, and swimming membranes.

Each sucker stalk arises from a large base from which

project 3 conical papillae. The largest suckers on

arms I—III occur at roughly two-thirds the distance

along the arm from its base. On arms IV, the largest

suckers are very near the base of the arm and are

smaller than the largest suckers of the other arms.

Near the arm bases, the suckers are globular and

have very small apertures. Distally, the apertures become relatively much larger. The suckers have about

12-13 slender, pointed teeth on the distal two-thirds
76 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

of the inner chitinous rings. The proximal margin is

smooth. The tips of all the arms are bare; however,

the length of these tips are longer in the ventral arms

than any of the others. The tips show no evidence of

being photogenic. A large sucker from the right arm

III has 13 sharply pointed teeth.

Tentacles are apparently lacking in this species.

They are represented by minute stubs located between

arms III and IV.

The buccal membrane has 7 lappets which attach

to the dorsal borders of arms I and II, and to the

ventral borders of arms III and IV.

The preserved specimen lacks pigmentation except

for a few small, scattered, brown chromatophores.

The data for the single specimen from off California are: Sex, $ ; M.L., 89 mm; M.W., 14 mm;

H.L., 23 mm; H.W., 16 mm; F.L., 41 mm; FW., 54

mm; Arm L. for arms I-IV respectively, 54 mm, 59

mm, 56 mm, 54 mm; J/4 A.G. for arms I-IV respectively, 40, 40, 42, 39.

TYPE LOCALITY.—29° N, 39° W, North Atlantic

Ocean.

LOCATION OF TYPE.—Not traced.

DISCUSSION.—Pfeffer (1912) synonymized the 2

nominal species in the family. Grimalditeuthis bomplandii (Verany, 1837) and Grimalditeuthis richardi

Joubin, 1898a. There has been some doubt concerning the validity of this action, since only G. richardi is

known to have photophores at the tips of the arms.

Judging from the present specimen, it seems likely

that this difference represents merely an ontogenetic

change. The VELERO specimen lacks photophores

at the tips of the arms, but all of the arm tips are

free of suckers. This condition is peculiar, but could

be easily explained if one assumed that the arm tips

were developing into photophores. I compared the

VELERO specimens with Atlantic material, but have

not been able to detect any differences.

This species is believed to be without tentacles;

however, delicate stubs of tentacles are present on all

specimens available to me from both oceans which

comprise a rather large size range. This indicates that

either extremely weak tentacles are present, but are

always lost during capture or, that the stubs represent rudimentary tentacles which persist, probably

due to their retention of some function which is

perhaps sensory. In this regard, it should be emphasized that in other species that lack tentacles

(i.e., Octopoteuthis, Gonatopsis) the stubs are present

in young forms, but are quickly lost with growth.

DISTRIBUTION.—This species is know only from a

very small number of specimens. It has been reported

from the north Atlantic at 29° N, 39° W (Verany

1837), 34°51' N, 33°1' W (Chun 1913), 38°55' N,

23°39' W (Joubin 1898a), from the Gulf of Mexico

(Voss 1956), and from the South Atlantic at 28° S,

28° W (Pfeffer 1912). The VELERO specimen was

taken from the Santa Catalina Basin at 33° 13' N,

118°28' W.

Family BATHYTEUTHIDAE Pfeffer, 1900

DIAGNOSIS.—Suckers on arms arranged in 2 rows

proximally, increasing to 4 irregular rows distally.

Tentacular clubs with minute suckers in approximately 8-10 rows. Minute suckers present on buccal

lappets. Fins small and separate with convex posterior

margins. Buccal connectives attach to dorsal borders

of arms IV.

Genus Bathyteuthis Hoyle, 1885

DIAGNOSIS.—The characters coincide with those of

the family.

TYPE SPECIES.—Bathyteuthis abyssicola Hoyle,

1885.

Bathyteuthis berryi Roper, 1968

PLATES 26A-H; 27A

Bathyteuthis berryi Roper, 1968, p. 169.

DESCRIPTION.—(Paraphrased from a manuscript

by C.F.E. Roper which was based on VELERO

specimens). The mantle is very plump and robust; its

width is 50% of its length. The mantle remains broad

for much of its length, then tapers and terminates

posteriorly in a broad, bluntly rounded tip.

The fins are short, rounded, and widely separated

from one another. Anterior and posterior fin lobes

are free.

The funnel is large; it extends to the level of the

anterior borders of the eyes. The posterior part of

the funnel is bound to the posteroventral surface of

the head with integument and gelatinous tissue. A

small pore lies in the ventral midline of the head near

the area where the funnel fuses with the head. The

funnel locking-cartilage has a simple, elongate

median sulcus. The dorsal member of the funnel
NUMBER 9 7 77

organ is an inverted, approximately Y-shape pad with

short, broad limbs. A spatulate papilla is present at

the anterior tip of the organ. The ventral pads are

approximately oval, but slightly pointed anteriorly. A

funnel valve is present.

The head is long and narrow. The eyes are large

and directed anterolaterally. The eyelids lack an

optic sinus. Small, stubby "olfactory" papillae are

present. Nuchal folds and crests are absent. The

nuchal cartilage is long and narrow and bears a

median and 2 lateral grooves.

The arms are short, but have attenuate tips. All

arms are subequal in length, giving the formula IV =

III = 11 = I. A moderately deep web joins the bases

of all arms except the 2 arms IV. The depth of the

web decreases from the dorsal to the ventral arms.

Low aboral keels are present on arms I—III; lateral

keels are present on arms IV. Protective membranes

occur on all arms. They are particularly well developed at the basal portion of the arms where they are

thick, fleshy, and ruffle-like. The thickened ruffles

quickly diminish distal to the bases of the arms, and

the protective membranes extend distally as low, even

keels. Protective membranes on arms IV are considerably less developed than on arms I—III. No distinct or separate trabeculae support the protective

membranes.

The arms bear numerous, small suckers. Proximally, the suckers originate in a single row, but soon

become biserial. About one-third of the way out on

the arms, the suckers become more closely packed and

the rows become irregularly arranged so that occasionally 3-4 suckers occur in a transverse series. On

the distal one-third to one-fourth of the arms, the

suckers become very closely packed and are exceedingly numerous. On arms IV, the sucker rows do not

become as irregular and the suckers are less numerous

than on arms I—III. Suckers extend to the extreme

distal tips of all arms. The suckers are largest proximally and gradually become smaller distally. About

275 suckers occur on each arm I—III of the holotype.

Arms I may have very slightly fewer, and each arm

IV has around 150 suckers. The inner sucker rings

from the arms bear 10-14 very low, small, rounded

or subtriangular, knob-like teeth. There is no hectocotylization.

A tentacle is known only from a single juvenile

(19 mm M.L.). This tentacle is long and robust. The

club is short, unexpanded, and simple; no keels or

membranes are present and no discrete divisions into

manus, carpus, or dactylus exist. About 7 or 8 rows

of very small suckers are distributed across the distal

half of the club; fewer rows occur proximally. Between 150 and 200 suckers are present at this stage.

The extreme distal portion of the club is reduced to

a small, papilla-like tip with a few minute bumps that

are probably precursors of future suckers. Dentition

on the inner rings varies from being nearly smooth

or slightly scalloped to having a few minute, low,

subtriangular teeth as shown in Plate 26H, although

the figure may exaggerate the size of the teeth.

The 7 buccal lappets of the buccal membrane are

long and bear 4-6 small suckers; the inner chitinous

rings of these suckers have about 10 small, low,

papilla-like teeth. The buccal membrane attaches to

the dorsal margin of arms I, II, and III, and to the

ventral margins of arms IV.

Most of the pigmentation is bleached out of the

holotype, but the smaller specimens exhibit the maroon coloration typical of Bathyteuthis.

A single, small, simple photophore is embedded in

the subcutaneous tisssue at the base of each arm IIII. The photophores in the holotype are not easily

seen, but the photophores in the juvenile and larval

specimens contrast more with the background pigmentation, are slightly raised, and are more readily

seen.

The largest specimen known, the holotype, has a

mantle length of 49 mm.

TYPE LOCALITY.—33° 18' N, 118°40' W, eastern

North Pacific Ocean.

LOCATION OF TYPE.—University of Southern California. U.S.C. Hancock collections, AHF Cephalopod

Type No. 10.

DISCUSSION.'—According to Roper (1968) the

genus Bathyteuthis includes 3 species: B. abyssicola

Hoyle, 1885, B. bacidifera Roper, 1968, and B. berryi

Roper, 1968. B. berryi can be separated from the

other 2 species by the much larger number of arm

suckers (i.e., over 250 compared to 150 in B. bacidifera and 100 in B. abyssicola). It can further be separated from B. bacidifera by the lack of free trabeculae at the bases of the arms, and from B. abyssicola

by the greater size of the grills.

DISTRIBUTION.—This species is known only from

the present trawling program. Eight specimens were

taken in zone 2, 1 each in zones 3 and 4, 3 in zone 5,

1 in zone 6, and 2 in zone 7.
78 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Family CRANCHIIDAE Prosch, 1847

DIAGNOSIS.—Mantle fused to head in nuchal region and to lateral corners of funnel. Buccal connectives attach to ventral borders of arms IV. Armature of clubs generally in 4 longitudinal rows. Arms

generally with biserial suckers.

Subfamily CRANCHIINAE Pfeffer, 1912

DIAGNOSIS.—Cartilaginous strips bearing tubercules

extend posteriorly on the mantle from the points of

the funnel-mantle fusions. Funnel fused laterally to

ventral surface of head in the adult stage.

Genus Cranchia Leach, 1817

DIAGNOSIS.—Mantle and fins covered with complex cartilaginous tubercules. Fins separate, each

nearly circular in outline.

TYPE SPECIES.—Cranchia scabra Leach, 1817.

Cranchia scabra Leach, 1817

PLATES 28B; 29J,Q

Cranchia scabra Leach, 1817, p. 140.—Hoyle, 1904, p.43,

pl.10, fig.ll.—Pfeffer, 1912, p.679, pl.48, figs. 22-28.—

Sasaki, 929, p.329, pl.26, figs. 13-15, text figs. 151-153.—

Voss, 1963, p. 142, text fig. 31.—Pearcy, 1965, p.261.

Cranchia tenuitentaculata Pfeffer, 1884, p.26, pl.3, fig. 36.

Cranchia hispida Pfeffer, 1884, p.27, pl.3, fig.37.

DESCRIPTION.—The mantle is saclike, although it

tapers abruptly to a point posteriorly. The mantle

wall is very thin, but muscular. The anterior edge of

the mantle is fused to the head in the nuchal region

and to lateral corners of the funnel. From each of

the latter points of fusion, two, short cartilaginous

strips diverge to form a V-pattem. The entire surface

of the mantle is covered with cartilaginous tubercules. The tubercules are somewhat variable, but

generally have 3-4 well-separated cusps whose tips are

usually subdivided into 2-3 additional cusps.

The fins are small and terminal (15-20% of the

M.L. in length and 25-30% of the M.L. in total

width). Each is nearly circular in outline and has

free anterior and posterior lobes. The dorsal surface

of the fins is covered with cartilaginous tubercules,

but the ventral surface appears to have only small,

rounded papillae.

The funnel reaches slightly past the bases of the

arms. The dorsal pad of the funnel organ has roughly

an inverted V-shape with a slender median ridge that

is larger in its posterior portion; a flap occurs at the

lateral edge of each diverging arm. The ventral pads

are large and kidney-shaped, with the concave margin

facing anteriorly. The surface of the funnel is covered

with simple papillae; those on the ventral surface, immediately anterior to the mantle, are interconnected

by low, narrow ridges that give a reticulated pattern.

A funnel valve is present. The funnel is fused laterally

to the head.

The head is short but broad and bears very large

eyes that occupy the entire lateral sides of the head.

Each eyelid has a small anterior sinus. Small, short

"olfactory" papillae are present on either side of the

head just lateral to the base of the funnel. Small, low

papillae are scattered over the surface of the head.

The arms are short and in the order of III>IV =

II>I. Arms I—III are joined basally by a web. The

arms all have broad, trabeculate protective membranes on both dorsal and ventral borders. Welldeveloped aboral keels occur on arms III, but are

barely detectable on arms I and II. Large lateral

keels are present on arms IV. The arms have biserially arranged suckers except near the tips of arms

III in males and on the hectocotylus. The suckers in

the proximal halves of the arms each have about 5—7

low, rounded, irregular teeth on the distal portion of

the inner chitinous ring. Distally, the broad teeth become more regular in arrangement and distinctly

truncate. The teeth of the inner sucker rings in females seem to be more regularly arranged throughout

the arm length, but are particularly so in the more

distal suckers which have slender truncate teeth. Each

arm III in males has the distal fifth modified by an

increase to 4-5 slightly irregular longitudinal rows of

suckers and by an abrupt decrease in the size of the

suckers. The suckers become biserial again at the extreme tips of these arms.

The right arm IV is hectocotylized in males. It is

considerably shorter than the left arm, and its tip is

swollen and curled laterally. Proximally, the suckers

of the hectocotylus are biserially arranged, but become tetraserial near the base of the curved tip. The

curved tip has biserial suckers which are considerably

enlarged proximally. Trabeculate protective membranes are present, but diminish on the curved portion. A large lateral keel extends to the tip of the

arm. Each sucker on the proximal segment of the
NUMBER 9 7 79

hectocotylus has about 6-7 broad truncate teeth on

the distal margins of the inner chitinous ring. The

largest suckers of the distal segment (curved portion)

each have about 13-15 rounded teeth around the

entire margin of the inner chitinous ring; the teeth

on the distal margin are the largest.

The tentacles are short and carry small, compact

clubs. Four longitudinal rows of suckers on the manus

grade into 4 rows on the dactylus. Suckers of both

manus and dactylus have a uniform size in a transverse series. There are 4-7 small, smooth-ringed

suckers at the tip of the dactylus. The dentition of the

inner rings of the manal suckers varies with the location of the sucker. The proximal suckers have 6-7

slender, pointed teeth on the distal half of the inner

ring. On the more distal suckers, the number progressively increases and smaller teeth also develop on

the proximal margin of the ring. At the distal end of

the manus, the number of teeth per ring approaches

30. Pairs of alternating suckers and pads extend down

the tentacular stalk for about two-thirds of its length;

the total number of suckers varies from 33-36 in the

2 specimens in which accurate counts were possible.

The buccal membrane attaches to the dorsal

borders of arms I and II and to the ventral borders

of arms III and IV.

In preservation, the specimens have small, brown

chromatophores scattered over the body, head, and

arms. In life, with chromatophores expanded and

head contracted into the mantle, the animal looks

like a large orange.

Photophores are present only on the surface of the

eyes. There is a proximal row consisting of 8 photophores in a U-shaped series passing from the anterior

to the medial section to the posterior surface of the

eye. A distal series of photophores lies near the lens

and has 4 photophores in a cescent-shape row ventral to the lens and 2 photophores dorsal to the lens.

The eye, therefore, carries a total of 14 photophores.

TYPE LOCALITY.—Off West Africa.

LOCATION OF TYPE.—British Museum (Natural

History).

DISCUSSION.—There is a single species, Cranchia

scabra Leach, 1817, presently recognized in the genus.

Distinct but minor differences exist in the pattern of

tuberculation in different specimens of C. scabra.

Pfeffer (1912) considerd C. scabra to consist of 2

subspecies, C. s. tenuitentaculata and C. s. hispida

which he separated primarily by the greater density

TABLE 19.—Measurements in mm for Cranchia scabra

Leach, 1817.

Characters

Sex

M.L

M.W

H.W

F.L

F.W

Arm L. I

II

Ill

IV

Tent. L

Club L

Club S

Stalk S

A.C. I

II

Ill

IV

Station Numbers of the

8025

c?

107

39

30

18

30

15

24

34

L R*

25 15

50

12

L R

92 96

36 36

.65

89

162

L R

88 98*

10897

9

78

38

18

15

24

9

14

19

15

31

9

L R

94 91

34 33

56

78

98

66

VELERO

10259

9

53

28

10

7

13

7

9

12

7

28

7

*hectocotylus

of tuberculation in the former. I have carefully compared the few specimens available from off California

with material from the Gulf of Mexico and find that

there is a distinct, but difficult to define, difference in

tuberculation between these forms. The Gulf form

has a greater number of large tubercules that gives

the mantle a rougher appearance than the California

form. The Atlantic specimens also have a much

thicker mantle wall. I also examined a specimen of

C. scabra from the Pacific Ocean off Panama that has

a thick mantle wall, but which has tuberculation that

is clearly less dense than the California or the Gulf

specimens. In other features, the specimens coincide rather closely.

It will not be surprising if eventually C. scabra is

shown to consist of a complex of species. An extensive series of specimens from each locality will be required to get necessary measurements, sucker counts,

and other data needed to establish the distinctness of

the forms since the differences will be small.
80 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

DISTRIBUTION.—C. scabra has been reported primarily from warm waters of the Atlantic, Pacific, and

Indian oceans between about 35° N and 37 S

(Clarke 1966). Pearcy (1965), however, recorded

C. scabra from off Oregon at about 44° N. Seven

VELERO specimens were taken from the Santa

Catalina Basin at 33° N and one from zone 6 at

31°42' N.

Genus Leachia Lesueur, 1821

DIAGNOSIS.—A single cartilagenous strip, bearing

tubercules, extends posteriorly from the point of each

funnel-mantle fusion. Arms are not attenuate. Arms

III of adult males have less than 100 suckers each.

(The latter two features separate Leachia from the

closely related Drechselia which has attenuate arms

with more than 200 suckers on each arm III. These

generic characters are only tentative at present, since

the adult stage is not known for most species of

Leachia.)

TYPE SPECIES.—Leachia cyclura Lesueur, 1821.

Leachia dislocata, new species

PLATES 30B; 31; 32

DESCRIPTION.—The mantle is long and slender

with an extremely thin but muscular wall. The pen is

clearly visible in the dorsal midline throughout the

length of the mantle and often separates from the

mantle during capture. The mantle is fused to the

head at the lateral sides of the funnel and in the

nuchal region. A line of cartilaginous tubercules, 13-

18% of the M.L. in length, extends from each

funnel-mantle fusion posteriorly along the ventral

surface of the mantle. The tubercules arise from a

single continous, cartilaginous band and are somewhat variable in form and arrangement; several basic

patterns are apparent. At the extreme anterior end

of the cartilaginous strip (i.e., the edge of the mantle), there is a large, complex tubrcule of variable

appearance. Closely posterior to this is another large,

complex tubercule also variable in appearance, but

clearly displaced toward the median line of the mantle. The position of this latter tubercule is invariable

and can easily be seen in all specimens beginning at

a mantle length of roughly 15 mm. Posterior to the

displaced tubercule, large, complex tubercules are

generally separated from each other by 2-3 small

tubercules usually with a single cusp. Generally, the

complex tubercules in the middle section of the strip

have a tripartite construction. The middle section has

3 cusps aligned in a row paralleling the body axis

with small additional cusps scattered sometimes at

various places. The lateral sections are distinctly

separated from the middle section and are quite variable in form. In the simplest cases, each appears as a

very large single cusp perpendicular to the median

complex. More commonly, the tip of this lateral section is formed into 2-4 small cusps. Generally 4-7 of

these complex, tripartite tubercules are found posterior to the displaced tubercule.

In the nuchal region, there is a cartilagenous pad

which overlies the anterior end of the pen. Two small

rounded tubercules arise from the anterior end of the

pad, one on either side, which project slightly past

the nuchal fusion. Often a low, flattened tubercle is

present lateral to each of these tubercles, and occasionally a small, median cusp may be present.

The fins are transversely oval in outline; they are

about 20-25% of the M.L. in length and about 30-

35% of the M.L. in width. Small, free anterior lobes

are present. The posterior margin of the fins is nearly

flat.

The funnel reaches approximately to the level of

the anterior end of the eyes. The funnel is fused to

the head laterally by narrow strips of tissue. The funnel organ has roughly an inverted U-shape dorsal pad

with a median ridge terminating in a long, slender

papilla. At each posterolateral corner, there is a long,

flattened papilla. Immediately anterior to each of

these there may or may not be a similar, but smaller

papilla. Sometimes one of these latter papillae will be

present on one side of the pad, but not on the other.

The ventral pads are elongate and often have an extended narrow anterior end. There is no funnel valve.

The head is broad and short. The eyes are large,

each occupying almost the entire lateral sides of the

head, and sessile in adults. A small anterior sinus is

present on each eyelid. An "olfactory" papilla is

situated on each side of the head, although the swollen tip of each barely rises above the surface of the

head.

The arms are short and stout. The arm formula is

III>II = IV>I. Arms III are extremely enlarged,

being approximately twice the size of arms II and

IV. All the arms bear short, broad trabeculae with

low protective membranes on both dorsal and ventral
NUMBER 9 7 81

borders, but there is no trace of aboral keels. All arms

have biserially arranged suckers. In males, the suckers

of the mid portions of all the arms (except the hectocotylus) have enlarged suckers with very small

apertures; in females, the suckers are smaller and

the apertures broader. The arm tips are not modified

except for arms III in females and the hectocotylus

in males. The inner chitinous rings have approximately 10-14 pointed teeth on the distal margin in

both sexes. In mature and nearly mature females (i.e.,

about 130-135 mm M.L.) the tips of arms III become modified by the loss of suckers and by the enlargement of the trabeculate protective membranes

into a spoonlike structure, generally with the edges

folded inward. The membranes are dark purple. The

oral surface bears a thick layer of highly convoluted

tissue that appears glandular. The modified arms III

have from 40-45 suckers. In males, arms III have

about 60-70 suckers in the large specimens.

The right arm IV is hectocotylized in males. No

mature males have been collected, and the hectocotylus described here from the holotype is probably

not completely developed. The tip of the arm is expanded and bears enlarged suckers with small apertures along the ventral margin and reduced suckers

along the dorsal margin. In the type, there is only a

slight indication of dentition in the distal suckers of

the ventral margin; in a slightly larger specimen

these suckers are distinctly toothed around their entire margins. A keel is present along the aboral surface of the arm. The right arm shows only very slight

hectocotylization at a M.L. of 110 mm and probably

cannot be detected at a smaller size.

The tentacular club is short, slightly expanded, and

bears suckers in 4 longitudinal rows. The suckers of

the median 2 rows on the manus are greatly enlarged.

The width of the inner chitinous ring is about 3

times that of the marginal suckers. The larger suckers

have about 30 teeth around the entire circumference

of the inner ring. Those of the distal margin are

longer and may be pointed; the others are all truncate. In the middle of the dactylus, the suckers are

largest in the ventral row, and grade in size to the

smallest in the dorsal row. There is a small ring of

suckers at the tip of the dactylus. Alternating suckers

and knobs in 2 rows extend down the tentacular stalk

for about half of the tentacle length. The suckers

number about 10-13.

The gravid females have undergone considerable

alternation in appearance. The funnel is enlarged and

extremely flaccid. All muscular tissue in the animal

appears to be more gelatinous than in nongravid

specimens, and the silvery iridescent tissue around

the eye has largely disappeared. The mature eggs are

approximately 1.2-1.4 mm in diameter.

In the adult animal 8 photophores are aligned in a

proximal series extending from the anterior end of

the eye and passing ventrally and medially along the

bulbus to the posterior margin. A second series of

photophores more closely encircles the lens. The ventral portion of this distal series consists of 3 large,

oval photophores with 2 interspersed small photophores giving the sequence; large, small, large, small,

large. The dorsal portion consists of 2 small photophores. There are no photophores on the viscera. It

is possible that the modified tips of arms III in females are photogenic.

The buccal membrane attaches to the dorsal

borders of arms I and II, and to the ventral borders

of arms III and IV.

The largest specimens appear to have been covered

by numerous, small brown chromatophores. The

young stages seem to lack chromatophores. At 5 mm

M.L., there is no trace of the cartilaginous tubercules

on the mantle; but at 10 mm M.L., they are present.

In some specimens of this size, it is possible to locate

the displaced tubercle. At 16 mm M.L., 2-3 dark

patches appear on the proximal face of the rostrum

of the eye, which are the first indications of developing photophores. The first 3 photophores seem to develop nearly simultaneously. At this size, the displaced

tubercle is clearly present. At 22 mm M.L., the first

3 photophores are well developed. At 25 mm M.L.,

the eyes have 4 photophores on the proximal face of

the rostrum. At 35 mm M.L., the fifth photophore has

developed and the photophores on the distal face of

the rostrum are beginning to form. The sixth photophore of the proximal series is present at 44 mm

M.L., and the seventh at 64 mm M.L. At this latter

size, the 3 large photophores of the distal series are

well developed. At 86 mm M.L., the eyes are still

borne on the long stalks as they have been throughout

the larval development, but by 110 mm M.L. the eyes

have become sessile and an eighth photophore (the

most posterior and last member of this series) has developed in the proximal series. This photophore does

not fall in perfect sequence with the other members

of the series since it is somewhat more distantly
82

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

placed. Also, at about 110 mm M.L., the right arm

IV in males begins to show the first evidence of modification into a hectocotylus. By 125 mm M.L., traces

of the small photophores of the distal series can be

detected. At 143 mm M.L., full maturity has been

reached, females are gravid, and the photophore pattern is completley developed.

TYPE LOCALITY.—32°35' N, 118°06/ W, eastern

North Pacific Ocean.

LOCATION OF TYPE.—University of Southern California, U.S.C. Hancock collections, AHF Cephalopod

Type No. 11.

DISCUSSION.—The differences between the genera

Leachia and Pyrogopsis depend almost entirely on

the size and position (i.e., stalked or sessile) of the

eyes and on the ocular photophore pattern. Because

specimens of Pyrgopsis generally have a small size

compared with those of Leachia, there has resulted

considerable debate concerning whether or not the

genus Pyropsis is based upon growth stages of members in the genus Leachia (Naef 1923; Adam 1960;

Voss 1967). The problem was not a simple one since

at least one adult Leachia has been recorded (Voss

1960) that is smaller than some specimens of

Pyrogopsis.

For the first time a complete series of growth

stages of a single species within this complex has been

available for study. This material indicates that the

genus Pyrgopsis Rochebrune, 1884, can no longer be

maintained and must be placed in the synonymy of

the older genus Leachia Lesueur, 1821. Specimens in

the present series from 10-35 mm M.L. exhibit all

the characters of the genus Pyrgopsis. Specimens from

35 to at least 86 mm M.L. have some features of both

genera, and specimens at 110 mm M.L. have all the

features of the genus Leachia. The fully mature females have features (i.e., photophore pattern) that

supersede Leachia and approach the genus Drechselia. The latter group, however, is sufficiently distinct to retain its generic standing pending information on the fully developed hectocotylus of Leachia

dislocata.

By synonymizing these two genera, I do not necessairly indicate that any of the present species of "Pyrgopsis" are necessarily the young of one of the known

species of Leachia. After examining many collections

of specimens belonging to this group, I am convinced

that it will prove to be very speciose. Therefore, it is

possible that all of the presently recognized species

could be valid; however, since many species are very

inadequately described, it may prove impossible to

incorporate their names into the future classification

of the group.

The genus Leachia as now constituted contains, in

addition to Leachia dislocata, the following species:

Leachia cyclura Lesueur 1821, south Indian Ocean

(37° S, 33° E) ; Leachia eschscholtzii (Rathke

1835), south Indian Ocean (28° S, 310° W);

Leachia zygaena (Verany 1851), near Messina, Mediterranean Sea; Leachia rhynchophorus (Rochebrune

1884), near Island of St. Paul, equatorial Atlantic

Ocean; Leachia schneehagenii (Pfeffer 1884) off

Chile; Leachia pacificus (Issel 1908) south Pacific

Ocean (14°32' S, 167°43' E) ; Leachia lemur (Berry

1920), east of Cape Hatteras (35°27' N, 73° 14' W) ;

Leachia atlantica (Degner 1925), eastern North Atlantic (36° 13' N, 9°44' W).

There is probably no genus in the Oegopsida that is

more systematically confused. Of the 8 species previously recognized, none can be presently identified

with any certainty. L. zygaena and L. rhynchophorus

are known only from very young specimens, 22 mm

M.L. and 17 mm M.L., respectively. Both species

lack enlarged arms III and have suckers extending

along the entire length of the tentacular stalks.

Neither of these features can be attributed to the

small size of the animals. If these features have been

correctly described, then there should be no difficulty

in separating them from the remaining members in

the genus; however, it seems unlikely that arms III

are as described. Both Pfeffer (1912) and Naef

(1923) have considered these 2 species as possible

synonyms. Eventually, both of these forms may have

to be considered as species dubia. L. schneehagenii,

L. pacificus, and L. lemur were all described from

specimens with a M.L. less than 35 mm and L. atlantica with a 50 mm M.L. In defining these species

Naef, Berry and Degner put considerable importance

on the shape of the fin. This character is of doubtful

value due to the very delicate nature of the fin and

particularly its anterior edge. I can find no valid

characters to separate these 4 species other than differences in their type localities.

Leachia cyclura and Leachia eschscholtzii are

known from mature or nearly mature specimens.

These 2 species cannot be separated from the preceding 4 species since any differences could be ascribed
NUMBER 9 7 83

Characters

Sex

ML

F.L

F.W

Arm L. I

II

IH

IV

Tent. L

Club L

Tub. L

TABLE 20.—Measurements in mm

10840

9

148

38

54

19

27

49

25

21

9905

9

140

29

39

16

21

46

21

21

9905

9

136

30

41

14

19

42

21

47

7

20

9905

9

130

30

45

15

21

36

19

18

for Leachia dislocata, new species.

Station Numbers of the

10985 11100

d"

137

"V)

42

15

74

48

L

?n

18

&

(Holotype)

125

30

42

15

23

47

R L R

16* 19 19*

42

7

19

VELERO

8025

9

87 +

26

35

9

13

29

12

48

7

20

8025

9

112

27

34

7

11

24

11

45

7

17

8116

9

76

19

25

3

6

17

5

36

5

12

8024

9

58

12

2

3

9

3

30

4

8

8881

cf?

50

11

15

2

2

9

3

23

3

7

8028

cf

110

23

31

7

11

25

10

34

6

19

*hectocotylus

to growth. Their type localities are, however, very

different.

Pfeffer (1912) identified a specimen captured off

the coast of Argentina (39° S, 53° W) as L. eschscholtzii. This specimen has an extremely long series

of cartilaginous tubercles which extend about half of

the M.L. This form probably does not belong to L.

eschscholtzii due to the differences in the lengths of

the tubercular strips. This identification rests almost

solely on the common occurrence of 3 light organs in

the distal series of the eye, a feature that is not

unique.

L. cyclura and L. eschscholtzii can be separated

from each other by the presence of a single photophore in the former species in the distal series near

the lens compared with 3 phorophores in this series

in the latter species. It appears now (personal observation) that a number of species which share these

photophore patterns have been lumped under these

2 specific names. It would appear that only a major

revision of the genus will stabilize this very confusing

group. It should also be mentioned that L. Boone

(1933) described a specimen identified as Pyrgopsis

schneehageni from the Pacific Ocean near Panama

that seems to have distinctive dentition on the tentacular clubs; however, if I am correct in assuming

that this has resulted from confusing the enlarged

third arms with the tentacles (this is possible if the

description was taken from the illustrations), then it

seems likely that the specimen may belong to Drechselia danae.

It would be nearly impossible to describe a new

species in this genus if it did not possess some extremely distinctive features. Fortunately, the California species has these. The displaced tubercle near

the anterior end of the cartilagenous strip on the

ventral surface of the mantle separates L. dislocata

from all other known species. In adult specimens, the

number of photophores in both the proximal and the

distal series is greater than in any other member of

the genus.

DISTRIBUTION.—L. dislocata is represented by 225

specimens from the present trawling program. The

frequencies of capture are lowest (0.02 and 0.07

specimens/hour) in the San Pedro and Santa Catalina basins and highest (2.28 and 1.25 specimens/

hour) in zones 6 and 7, respectively. Recently, the

SWAN captured a small specimen (17 mm M.L.( off

the Hawaiian Islands at approximately 22° N,

156° W.

The low occurrence of capture in the inshore basins

under study combined with the high occurrence in

warmer water, more oceanic zones, and its presence

off Hawaii indicates that L. dislocata may be primarily an inhabitant of the eastern North Pacific

Central Waters.
84

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Subfamily TAONIINAE Pfeffer, 1912

DIAGNOSIS.—Cartilagenous strips not present on

mantle. Funnel free from the head laterally in adults.

Genus Galiteuthis Joubin, 1898

DIAGNOSIS.—Larval forms have tetraserial suckers

on the tentacular clubs; the medial 2 series convert

into hooks in the juvenile stage. Fins are terminal

and lanceolate. Arms have biserial suckers throughout.

TYPE SPECIES.—Galiteuthis armata Joubin, 1898

Galiteuthis phyllura Berry, 1911

PLATES 33; 34A,B,D-I,L

Galiteuthis phyllura Berry, 1911, p.592; 1912, p.315, textfigs. 17-18, pl.46, figs. 1-3, pl.54, figs. 5-6, pl.56.

Galiteuthis armata [parts],—Pfeffer, 1912, p.731.—Pearcy,

1965, p.262.

DESCRIPTION.—The mantle is long and slender; its

muscular portion tapers to a point just past the anterior ends of the fins. The very thin but muscular

mantle is fused to the head at the nuchal region and

to the lateral corners of the funnel. Each of the latter

2 areas is marked by a fiat cartilaginous pad with 2-A

small tuebrcules at its anterior end.

The fins are lanceolate, extremely long (41—47% of

the M.L. in specimens greater than 100 mm M.L.),

and slender (width 28-35% of the M.L. in specimens

greater than 100 mm M.L.), and lack anterior or

posterior lobes. The fins become very antenuate posteriorly, but extend to the tip of the pen.

The funnel is moderate in size, reaching approximately to the level of the midpoint of the eyes. The

dorsal pad of the funnel organ has an inverted Ushape with rounded lateral arms. Three large, slightly

flattened papillae are present, one on each lateral arm

and one on the anterior midline. The dorsal pads are

nearly oval in shape. There is no funnel valve. The

funnel is free from the head laterally.

The head consists almost entirely of 2 very large,

bulging eyes which are directed slightly forward. The

eyes nearly touch each other in the ventral midline.

In specimens up to about 60-65 mm M.L., the eyes

are set on stalks. The eye opening is rather small and

has an anterior sinus. From the skin over the midposterior surface of each eye arises an "olfactory"

papilla which has a cup-shape terminal portion set

on a swollen stalk.

The arms are relatively short, but muscular. The

arm formula is IV>III>II>I. All arms possess

large trabeculate protective membranes which are

better developed on the ventral border of each arm.

Arms I—III appear to have low, gelatinous aboral

keels. Narrow lateral keels are present throughout the

length of arms IV. Arm suckers are biserially

arranged.

The suckers are globular, but have rather large

apertures. The largest suckers of arms IV occur in

the middle of the arms. The chitinous rings are

smooth. In the larger of the males, the tips of arms

I—II narrow abruptly and possess very small suckers

in 2 rows. This distal area gives the impression that

the arm tips have been lost and are in a state of regeneration; however, these tips are affected identically among opposite membrs of arms I and II and

in all of the larger males (3 specimens). An Atlantic

specimen of G. armata (also a male) shows the same

feature. Therefore, it appears that this modification

of the arm tips is a type of hectocotylization. A low

web is present which is most noticeable between arms

I-II.

The tentacles are short and muscular. The clubs

are only slightly expanded and have narrow, potective

membranes which extend the full length of the club

along both margins and contain very broad trabeculae. A minute keel is present at the extreme tip of

the club. The manus bears 2 rows of large hooks with

1-2 suckers in each marginal series at the proximal

end. In young specimens, the marginal series are

complete. Hooks first develop at approximately 40-50

mm M.L. The dactylus is short and bears from 22-32

small suckers arranged in 4 longitudinal rows. The

inner chitinous rings of the dactyl suckers have generally 4 slender, widely separated truncate teeth on

the distal margin, with 1-2 smaller teeth on each

lateral margin. The proximal margin is slightly irregular, but lacks definite teeth. A distinct carpal

cluster consists of 7-10 suckers and pads. Suckers are

arranged in pairs along about the distal 70% of each

tentacular stalk on the oral surface; the number of

these suckers varies generally from 36-44. The suckers

alternate with low, indistinct knobs.

The buccal membrane connectives attach to the

dorsal borders of arms I and II and to the ventral

borders of arms III and IV.
NUMBER 9 7 85

There are 2 photophores on each eye. The more

distal organ lies just ventral to the lens and is approximately bar-shape, but may curve slightly and is

divided into 2 sections. The larger photophore occupies almost the entire medial ventral surface of the

eye. It has a broad, crescent shape and is thickened

on its medial side.

In preservation, G. phyllura has many reddish

brown chromatophores distributed over the head,

arms, tentacles, and dorsal surface of the fins. The

mantle has only a few large scattered chromatophores

of the same color.

All specimens examined are immature.

TYPE LOCALITY.—Off Point Pinos, Monterey Bay,

California.

LOCATION OF TYPE.—United States National Museum 214325.

DISCUSSION.—Joubin, 1898b, established the genus

Galiteuthis based upon a new species, G. armata

taken from the Mediterranean Sea. Berry (1911)

briefly described a second species, Galiteuthis phyllura, from Monterey Bay, California, and subsequently (Berry 1912b) gave a more detailed description of the species; however, his species was already

relegated, apparently unknown to Berry, to the status

of a junior synonym of G. armata by Pfeffer in his

monograph of oegopsid cephalopods and has remained in that position ever since.

I directly compared this species to several specimens from the North Atlantic and believe that the

Californian form deserves the specific status that

Berry originally gave it.

My comparative material is inadequate to give

good morphometric comparisons. There appears,

however, to be a difference in the length of the "tail"

(i.e., longer in G. armata), although this feature is

difficult to use since it is apparently somewhat variable and is subject ot considerable alteration due to

damage. The specimens of G. armata available to me

have damaged eyes, and it is impossible to determine

their size. The literature indicates that the eyes of

G. armata are considerably smaller than those of G.

phyllura. This is readily apparent when one compares

my figure (pi. 33A) with Chun's illustrations from the

VALDIVIA reports (1910, plate 59, fibres 1, 2)

which show animals of nearly identical ' ly lengths.

TABLE 21.—Measurements in mm for Galiteuthis phyllura Berry, 1911.

Characters

Sex

ML

H.W

F.L

F.W

Arm L. I

II

Ill

IV

Tent. L

Club L ...

Tail L

Eye D

Club H . .

Carp. S

Stalk S

Dact S ...

Tubercles

8030

9

173

23

76

21

18

23

28

35

78

11

14

14

L R

11 11

10 10

36 39

22 26

3 2

9056

c?

211

30

91

32

25

30

38

43

79

14

28

18

L R

13 13

8 8

44 43

23 25

4 4

10265

c?

254

42

119

38

38

45

55

63

105

19

17

23

L R

13 13

8 8

39 37

26 24

3 3

Station Numbers of the VELERO

8030

85

12

31

10

8

11

14

16

43

7

8

6

L R

10 10

8 8

36 36

30 27

3 3

10259

60

S.E.*

19

7

5

7

9

11

35

7

5

5

L R

7 8

8 7

40 42

32 —

2 2

10265

108

16

44

13

12

14

18

23

55

9

14

9

L R

12 —

9 8

35 38

2 4

9661

45

S.E.*

12

7

4

5

6

7

26

5

L R

7 6

7 8

42 40

27 32

10265

60

S.E.*

18

6

5

7

8

10

33

5

5

L R

9 7

8 7

42 42

32 29

2 3

11183

c?

359

61

164

46

52

71

82

87

137

21

40

33

L R

14 14

8 8

36 36

21 21

4 3

10265

65

10

21

9

5

8

10

12

39

6

6

L R

9 8

8 7

38 38

27 26

3 3

•S.E., stalked eyes.
86

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The holotype of G. armata described by Joubin

(1898b) has a head width of 20 mm. The specimen

of G. phyllura of the same body length has a head

width of 30 mm. There also appears to be differences

in arm length, fin lengths, fin widths, sucker sizes,

and hook sizes. These differences, however, require

verification through the study of a large series of

specimens which are not presently available.

The features mentioned here are not sufficient to

prove the separation of G. armata and G. phyllura. I

feel that when more material becomes available for

study, the differences between these species will be

substantiated; and I therefore retain the name G.

phyllura for the California specimens.

DISTRIBUTION.—G. phyllura has previously been

recorded only from Monterey Bay (Berry 1912). I

examined a specimen captured off Oregon by Pearcy

that belongs to this species as does, presumably, G.

armata listed by Pearcy (1965).

The frequency of capture in the present study is

almost constant for zones 2, 3, 5, and 6 (i.e., 0.26 to

0.29 specimens/hour). Zone 7, however, with over

120 hours of trawling has yielded only 2 specimens.

It seems likely that zone 7 is near the southern limit

for this species. Galiteuthis phyllura, therefore, has a

distribution that extends from approximately latitude

28° N to at least 44° N along the western coast of

North America. Sasaki (1929) recorded G. armata

from Japan; it is not certain whether his specimens

belong to G. phyllura.

Galiteuthis pacifica (Robson, 1948)

PLATES 30A; 34C,J-K,M

Taonidium pacificum Robson, 1948, p. 130.

DESCRIPTION.—The slender mantle, broadest in

the anterior half, tapers to a point posteriorly and has

an extremely thin but muscular wall. The mantle

margin is fused to the nuchal region and at the lateral

edges of the funnel. Cartilaginous tubercles are lacking at all of these points.

The fins are small and lanceolate; they are about

25% of the M.L. in length and 20% of the M.L. in

width. There are no anterior or posterior lobes. The

pen, which is visible along the middorsal line, extends

slightly posterior to the fins, but this may be an

artifact of preservation.

The funnel is small, but reaches nearly to the anterior margin of the eyes. The funnel is free from

the head laterally. The dorsal pad of the funnel organ

has roughly an inverted U-shape. Usually a small

papilla arises from the center of the anterior portion

of the pad. The ventral pads are small and oval.

There is no funnel valve.

The head is short and the eyes are relatively small.

The eyes are not clearly talked in any of the available specimens since they lack extended optic tracts,

although the eyes project at a 45° angle to the body

axis. A small olfactory papilla arises posterior to each

eye.

The arms are short and muscular. Low gelatinous

swimming keels are present on the distal portions of

arms I—III, and lateral keels are present on arms IV.

Low, trabeculate, protective membranes are present

on all of the arms. A low, interbrachial web exists

between arms I—II. The arm formula is IV>III>

II>I. The suckers are arranged biserially on each

arm. On the midportions of the arm, the suckers are

large, and the inner chitinous rings are smooth and

have broad apertures.

The tentacles are short and have only slightly expanded clubs. An ill-defined carpal locking-apparatus

is present at the base of the manus which seems to

include 7-9 suckers; the knobs are indistinct. Proximally, the carpal suckers merge with paired suckers

that extend along more than two-thirds of the tentacular stalk. These suckers alternate with indistinct

knobs and can therefore be considered as an extension

of the carpal cluster. There are 48—74 suckers on the

tentacular stalk. The median 2 rows on the manus

consist of apparently 5 hooks each, although the

clubs have been damaged so that the exact number is

uncertain. Lateral to the hooks on either side is a

marginal series of suckers. The dactylus is very short

and appears to have from 14-24 suckers. The inner

chitinous rings of the marginal and carpal suckers

each have a few short, truncate, and broadly separated teeth around the entire margin; those of the

distal margin are the largest.

On the ventral surface of each eye is a U-shaped

photophore that extends from the posterior margin

along the medial edge to the antrior margin of the

eye. A crescent-shape patch of tissue which is limited

medially by the photogenic band just described is apparently part of this photophore, although distinctly

different in appearance. In the largest specimen, the

more distal photophores, characteristic of most members of the subfamily, is just beginning to develop.
NUMBER 9 7 87

This photophore is also composed of 2 parts: a short

distal bar and a proximal oval patch. Combined,

these have a circular outline.

The specimens have scattered reddish brown

chromatophores.

The buccal connectives attach to the dorsal borders

of arm I and II and to the ventral borders of arms III

and IV.

TYPE LOCALITY.—Off Cocos Island, eastern tropical Pacific.

LOCATION OF TYPE.—Numbers 1948.7.8.1 and 2.

British Museum (Natural History).

DISCUSSION.—Galiteuthis pacifica can easily be

separated from the other 2 members of the genus, G.

phyllura and G. armata, by 2 distinctive features: the

points of the funnel-mantle fusions are smooth and

the distal photophore on each eye is oval. (This latter

feature can be observed only in specimens larger than

about 60-65 mm M.L.) Both G. phyllura and G.

armata have complex tubercles at each funnel-mantle

fusion and a bar-shape, distal photophore on each

eye. There are other, less obvious, features concerning

the shape and position of the fins, the stalked or sessile

condition of the larval eye at a given M.L., and the

dentition of the dactyl suckers that also distinguish

the species described here.

I examined a large specimen of Galiteuthis captured by the DANA and a small specimen taken by

the PILLSBURY from the tropical Pacific off Panama. Both specimens closely agree with those from

California and are certainly conspecific with them.

Robson (1948) described Taonidium pacificum

from 2 small specimens taken near Panama. The

specimens, due to their small size (the largest had

a M.L. of 29 mm), lacked hooks on the tentacular

clubs and therefore, their affinity to Galiteuthis was

not recognized. I here transfer Robson's species to

the genus Galiteuthis. The late Dr. W. J. Rees,

formerly Curator of Coelenterates at the British Museum, kindly examined the types of G. pacifica for

me and found that the funnel-mantle fusions

definitely lack tubercules. Therefore, there can be no

doubt that Galiteuthis pacifica (Robson) is indentical

with specimens here examined from off Panama and

southern California.

DISTRIBUTION.—G. pacifica was previously known

only from the type specimens captured off Cocos Island. The VELERO material was taken between

29° 12' N and 33°20' N. PILLSBURY and DANA

TABLE 22.—Measurements in mm of Galiteuthis

pacifica (Robson, 1948).

Characters

Sex

M.L...

M.W..

F.L....

F.W.. .

Arms I.

II.

III.

IV.

Tent. L

ClubL.

Stalk S.

Staton Numbers of the VELERO

8243 8018 11022 10906

36

11

9

7

4

6

6

7

28

5

72

42

12

10

8

4

6

7

8

25

4

56

52

15

13

11

6

8

10

11

24

5

49

65

—

17

11

8

10

12

13

—

—

have provided specimens from 7° 19' N, 79°42' W

and 6°40' N, 80°47' W. G. pacifica is very rare in

California waters, being represented in the collections by only 4 specimens. Since an equal number of

specimens are now known free from the Panama area

from only minimal trawling, it is safe to assume that

the species is common to these waters and only occasionally ranges as far north as the California area.

Genus Helicocranchia Massy, 1907

DIAGNOSIS.—Funnel-mantle without cartilaginous

pad. The mantle terminates in a blunt point, and the

pen rises free from the mantle shortly in advance of

the mantle tip. Fins pedunculate and attach to the

free pen except at their extreme anterior ends. Funnel large.

TYPE SPECIES.—Helicocranchia pfefferi Massy,

1907.

Helicocranchia pfefferi Massy, 1907

PLATES 28A; 29A-I

Helicocranchia pfefferi Massy, 1907, p.382; 1909, p.34, pl.3,

figs. 1_3.—Robson, 1948, p.130.

Teuthowenia (Helicocranchia) pfefferi.—Pfeffer, 1912, p.

740.—Grimpe, 1922, p.98.—Massy, 1928, p.36.

Desmoteuthis megalops [parts], Muus, 1956, p.9; 1962 p.9.

Megalocranchia (Helicocranchia) pfefferi.—Voss, 1960, p.

433.

DESCRIPTION.—The mantle is broadest in the midportion; posteriorly, it tapers to a blunt point. The
88 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

mantle wall is thin but muscular and often has a

leathery texture. It is fused to the head in the nuchal

region and to the lateral edges of the funnel; no cartilaginous tubercules or pads are present at these points.

The fins are separate, paddlelike in shape, and have

large posterior lobes. Except at the extreme anterior

ends, the fins are supported exclusively by the pen

which projects dorsally above the mantle. The small

posterior tip of the mantle, therefore, not only extends beneath fins without being attached to them,

but also lacks the supporting tip of the pen.

The funnel is extremely large; it extends well past

the bases of the arms and is very broad. The dorsal

pad of the funnel organ has roughly the shape of an

inverted V. A slender papilla arises from the midanterior portion of each lateral arm and also from

the midportion of the apical region. Each ventral

pad has an L-shaped appearance with the transverse

limb shorter and much broader than the longitudinal

one. A funnel valve is not present, and the funnel is

free from the head laterally.

The head is short and narrow. The eyes, small

relative to the size of the body, project from the head

and point forward at roughly a 45° angle to the body

axis. Each eye has a well-formed ventral rostrum with

an ill-defined photophore on its proximal face. A

small "olfactory" papilla arises from the tissue covering the lateral surface of each eye.

The arms are very small and slender. The arm formula is III>II> I = IV. All arms have low, protective

membranes without apparent trabeculae on both the

dorsal and ventral margins. Narrow aboral keels are

present on arms I—III, and lateral keels are present

on arms IV. All arms have biserial suckers over their

entire lengths except for arms I and II of males. Each

inner chitinous ring has about 7-10 low, rounded, or

sometimes pointed teeth on the distal margin; the

proximal margin is smooth. These teeth are very difficult to see under the microscope.

In males, approximately the distal one-fourth to

one-third of arms I and II are modified. On each

arm I, the size of the suckers and the diameter of

the arms decrease abruptly and the number of sucker

rows increases to about 4. On each arm II, the tips

are similarly modified except that the number of rows

increases to 8-9. These suckers have inner rings that

are disproportionally small and apparently lack teeth.

The tentacles are long and moderately robust. The

tentacular clubs are small but expanded and bear

suckers in 4 series. At the proximal end of the manus,

the suckers of the median rows are slightly enlarged

over those of the marginal rows. On the dactylus, the

suckers in a transverse series grade in size from the

largest in the ventral row to the smallest in the dorsal

row. At the tip of the dactylus, a small lobe bearing

3-4 tiny suckers is present. On the manus, the inner

chitinous rings of the suckers have about 8-10

slender, pointed teeth on the distal half of the margin.

Laterally, the teeth diminish rapidly in size and form

only small pointed knobs proximally. The proximal

2 suckers in the dorsal row of the manus are distinctly

reduced in size. A broad, trabeculate protective membrane is present along the entire ventral margin of

the club, but the dorsal membrane terminates along

the base of the dactylus. A short dorsoaboral keel is

present on the dactylus. About 13-17 pairs of small

suckers alternating with knobs extend along almost

the full length of the tentacular stalk. Buccal connectives attach to the ventral borders of arms IV.

H. pfefferi has a characteristic color pattern. The

mid-dorsal surface of the mantle has scattered elongate chromatophores. A strip of chromatophores extends along each dorsolateral side of the anterior twothirds of the mantle. The lateral sides have a series

of short, transverse bars, while the ventral surface has

scattered chromatophores with a small concentration

in the anterior midline.

TYPE LOCALITY.—Near Ireland, 51°54' N; 11°57'

W.

LOCATION OF TYPE.—British Museum (Natural

History).

DISCUSSION.—Helicocranchia was established by

Massy 1907 for H. pfefferi. Subsequently, there has

been considerable question concerning the validity of

the genus (Grimpe 1922; Massy 1928; Muus 1956,

1962; Voss 1960). The group is very distinctive,

however, and it is hoped that the above description

will remove all doubts concerning its independent status. Presently, the genus contains 2 additional species,

H. beebei Robson, 1948, from the region of Cocos

Island and the Galapagos Islands, and H. papillata

(Voss 1960) from 32°05' N, 64°38' W in the North

Atlantic. H. papillata can easily be separated from

the others by the presence of small, sharp papillae

over the surface of the funnel and mantle. H. beebei

is very similar to H. pfefferi, and it is possible that

they are synonymous. Some time ago, I examined
NUMBER 9 7 89

TABLE 23.—Measurements in

Characters

Sex

M.L

M.W

F.L

F.W

Arm L. I

II

Ill

IV

Tent. L

Club L

StalkS

mm for

9249

c?

52

20

8

14

9

12

13

8

43

6

L R

34 32

Helicocranchia pfefferi Massy,

Station Numbers of the VELERO

8030 8111

56

18

5

10

9

11

12

8

37

5

L R L

32 31

9

60

22

4

10

8

11

14

8

37

6

R

8111

65

22

4

9

11

14

15

11

L R

1907.

8027

9

79

23

8

17

13

16

19

12

69

7

L R

29 26

the type of H. beebei which is deposited in the British

Museum. The arms of the largest specimen were

considerably thicker and fleshier than the California

and Atlantic material of the same size. It seems likely

that H. beebei will prove to be a valid species. If this

is so, it places the identification of the California material somewhat in doubt, since it indicates a rather

peculiar geographical distribution for H. pfefferi.

Unfortunately, the only difference that I have been

able to detect between specimens of H. pfefferi from

the Atlantic Ocean and from off California is the

distinctly larger eyes in the Atlantic forms. This feature is, of course, insufficient by itself to require

separate status for the California specimens.

The California specimens, therefore, show slight

differences from H. pfefferi and H. beebei, but

these differences are small and since the validity of

H. beebei is in doubt, there is no recourse but to call

the California specimens H. pfefferi. Perhaps when

mature animals of all 3 forms become available for

study, it will be possible to determine their relationships more satisfactorly.

Voss (1962) described a new cranchiid genus,

Ascocranchia, based on a specimen with a number of

peculiar features, the most important of which is the

presence of 3 small oval photophores on the eye combined with the absence of cartilaginous strips and

tubercules on the mantle. This combination of characters cuts across formerly designated subfamilial

boundaries within the Cranchiidae and placed Ascocranchia in an uncertain position. I recently examined several damaged females belonging to this

genus that were captured by the PILLSBURY in the

Gulf of Guinea. The tentacles are missing, and the

eyes have been damaged in all of the specimens. They

exhibit, however, the basic features of Helicocranchia

in having a large funnel and paddlelike fins that are

supported by the tip of the pen which rises dorsally

from the surface of the mantle. The type was a male

and shows at least an indication of the peculiar hectocotylization seen in H. pfefferi and H. beebei, although it occurs on the arms III instead of arms I

and II. The light-organ pattern on the eyes of the

species in Helicocranchia are certainly not typical for

the subfamily. In addition, the photophores are

known only on the rostrum of the eye in juvenile

specimens. It is likely that the rostrum is absent in

mature specimens, and that there will be a corresponding alteration in the photophore pattern. Therefore, the light-organ pattern in Ascocoranchia may

not be as significant as it was thought at first. Until

the photophore pattern in mature specimens of Helicocranchia is known, the ultimate fate of Ascocranchia as an independent genus cannot be resolved

with certainty.

DISTRIBUTION.—In the present study, H. pfefferi

showed the greatest abundance in the northern zone 2

(2.28 specimens/hour), while the second highest

concentration (1.63 specimens/hour), occurred in the

most southernly zone, zone 7.
90

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Order OCTOPODA

Suborder INCIRRATA

Family BOLITAENIDAE, Chun 1911

DIAGNOSIS.—Rachidian and lateral teeth of radula

multicuspidate. Suckers uniserial. Mantle opening

broad. Liver and stomach in "normal" position.

Genus Japetella Hoyle, 1885

DIAGNOSIS.—Eyes are large. Right arm III in

males with many slightly enlarged suckers on the

distal two-thirds of the arm.

TYPE SPECIES.—Japetella prismatica Hoyle, 1885.

Japetella heathi (Berry, 1911)

PLATES 35B; 36D

Eledonella heathi Berry, 1911, p.589; 1912, p.276, pl.32, fig.

4, pl.33, figs. 2-4.

fapetella heathi.—Robson, 1932, p.335.—Thore, 1949, p.

7.—Pearcy, 1965, p.262.

DESCRIPTION.—The mantle is soft and sacular with

a gelatinous consistancy. The mantle opening is

broad, extending to at least the level of the midpoint of the eyes.

The funnel is large and embedded for most of its

length in the gelatinous tissue on the ventral surface

of the head. The tip of the funnel extends approximately to the level of the anterior edge of the eyes.

The funnel organ is an inverted V-shaped pad.

The broad head is encased in gelatinous tissue. The

eyes are large (13 mm in a specimen of 78 mm M.L.)

and elliptical in outline with the longest diameter

transverse to the body axis. The ventral-lateral surface of each eye has a strong silvery sheen. This

silvery patch also loops above the lens although it

may, in this area, be partially covered by reddish

brown chromatophores. The dorsal surfaces of the

eyes are black with a slight bluish iridescence. The

brain is about 4—5 mm in width while the optic stalk

is about 8 mm long in a specimen with a mantle

length of 71 mm. The peduncle lobe complex lies 3

mm from the optic lobe. A small "olfactory" papilla

is present posterior to each eye on the collar.

The arms are moderately long and each bears

globular suckers arranged in a single row. The

suckers are so closely packed that each is in contact

with its neighbor. The arms are thick but fragile, and

their tips are frequently broken off. The arms are arranged in order of III>IV = 11 = I. The suckers are

deeply embedded in the gelatinous tissue which surrounds the arms; this is particularly so near the bases

of the arms.

The web, although easily damaged or distorted, appears to be in the order of A>D>B = OE . The

right arm III in males is modified by an increase in

sucker size and perhaps a slight increase in length.

Usually the fifth sucker from the base of the arm is

the first enlarged sucker; the size of the enlarged

suckers decreases gradually to the tip of the arm.

The liver is usually oval and very large, although

its size varies greatly. Its ventral and lateral sides are

covered by a brilliant silvery tissue that extends over

the lateral sides of the crop. At its posterior end, the

liver laterally embraces the spiral caecum and the

anterior part of the stomach. The dorsal surface of

the liver is slightly concave where the large crop

presses against it. The crop in this species is very

capacious and is almost always in the expanded condition even if it is nearly empty. The crop is formed

by a large outpocketing of the esophagus. The crop

lies primarily to the left side of the esophagus. Large

posterior salivary glands lie on the right side of the

crop. The pancreas lies embedded in the dorsoposterior portion of the liver. It is slightly bilobed since it

follows the contours of this part of the liver. There

are 16-22 primarily gill lamellae. The ink sac is

small and embedded in the liver at the anterior end

of the pancreas.

The radula is ctenoglossan. The rachidian has a

large central cusp with 3-4 assymmetrically placed

cusps on either side. The admedian has 5-7 cusps,

the more medial of which is the largest. The second

lateral has 6-7 cusps with the medial cusp slightly

enlarged. The third laterals have a single elongate

cusp. Broad marginal plaques are present.

The eggs of the largest female (M.L., 90 mm) are

elongate and slightly swollen in the distal half. The

length is between 1.2 and 1.3 mm and the greatest

width is between 0.3 and 0.4 mm.

Data for 2 of the largest specimens are: from the

VELERO 8933, sex, tf; M.L., 78 mm; M.W., 42

mm; H.W., 38 mm; Arm L., I-IV, respectively, 55

mm, 60 mm, 77 mm, 60 mm; and from the

VELERO 8715, sex, d1; M.L., 72 mm; M.W., 45
NUMBER 9 7 91

mm; H.W., 36 mm; Arm L., I-IV, respectively, 39

mm, 42 mm, 57 mm, 40 mm.

TYPE LOCALITY.—Off Santa Catalina Island, California.

LOCATION OF TYPE.— United States National Museum 214318.

DISCUSSION.—At present only 2 species are recognized in the genus, /. diaphana, Hoyle, 1885 (type

locality, 0°42' S, 147° E) and /. heathi (Berry

1912b). The original description of /. heathi reveals

no definite features which could unquestionably relate

this name to one of the 2 .species present in these

waters. Berry's (1912b) statement that the eyes are

nearly black would seem to identify it with the species

referred to in this paper as Japetella sp.; however, it

seems more likely that the eyes had lost their silvery

iridescence due to preservation. The type, unfortunately, has been destroyed. I think the only solution

is to assume the most abundant species in this area

is /. heathi.

I compared specimens of /. heathi with specimens

of /. "diaphana" from the Atlantic Ocean off Florida

and find that-there is little to separate these forms;

however, considering the great similarity of the

sympatric species which occurs off California (see

discussion of next species) which allows the separation only of large specimens, one must be careful in

discussing worldwide relationships. For the present, I

think it best to retain the name /. heathi, at least

until we have a better understanding of speciation

within the genus.

DISTRIBUTION.—Since /. heathi has been imperfectly separated from Japetella sp. (see following

section), the distribution of the 2 species is considered

together. It should be emphasized, however, that the

vast majority of the specimens, belong to /. heathi.

Japetella is well represented throughout most of the

area studied, but did show a slight increase in frequency of capture in the southern zones. Zone 2 had

a capture frequency of 0.15 specimens/hour while

zone 6 and 7 had a frequency of 0.30 and 0.26

specimens/hour, respectively. /. heathi has also been

reported from off the coast of Oregon (Pearcy 1965).

Japetella species

PLATE 36A-C

DESCRIPTION.—The mantle is saclike and gelatinous. The mantle opening is broad and extends on

either side to at least the level of the midpoint of the

eye.

The funnel is well developed and reaches approximately the level of the anterior edge of the eyes. The

funnel organ is an inverted V-shaped pad situated on

the inner dorsal surface of the funnel. The funnel is

embedded in the gelatinous tissue of the head for

most of its length.

The head is broad and enveloped in gelatinous

tissue. The large eyes, nearly circular in outline, are

black although in live specimens they have a greenish

hue on the ventral half. The pupils of the eyes are

slitlike and oriented at approximately right angles to

the body axis. Each optic stalk, on a specimen with a

mantle length of 72 mm, is 8-9 mm long, while the

width of the brain is approximately 5 mm. The

peduncle lobe complex is situated 4 mm from the

optic ganglion. A small "olfactory" papilla is present

posterior to each eye on the collar.

The arms are moderately long and each bears a

single row of large, globular suckers. The suckers are

closely packed so that each is in contact with its

neighbor. The arm formula is HI>IV= I = ?II. The

suckers are very deeply embedded in the gelatinous

tissue that surrounds the arms. The right arm III of

males is hectocotylized. It is slightly longer than the

corresponding left arm and possesses enlarged suckers

beginning abruptly with usually the sixth sucker from

the base of the arm. The enlarged suckers decrease

gradually in size distally to the tip of the arm. The

web formula cannot be determined in the available

material.

The liver is small and slender. The intestine extends along the ventral surface of the liver well past

its anterior end. Posteriorly, the intestine passes ventral to the pancreas, between the hepatic ducts, and

enters the caecum. The bilobed pancreas is embedded in the ventral-posterior surface of the liver;

two large hepatic ducts arise from the pancreas, then

join to form a single duct before entering the caecum.

The caecum is embraced by the posteriorly directed

projections of the liver. The crop is a saclike outpocketing of the esophagus. Dorsal and lateral to the

crop on either side lies a posterior salivary gland.

These organs, although rather variable, are generally

the same size as the liver. The ink sac lies embedded

in the ventral surface of the liver at the anterior end

of the pancreas. There are about 18 primary gill

lamellae.
92

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The radula is ctenoglossan. The rachidian has a

large central cusp with 3-4 asymmetrically placed

cusps on either side. The admedian has 5-7 cusps,

the most medial of which is the largest. The second

lateral has 5-7 cusps with the medial cusp slightly

enlarged. The third laterals have a single elongate

cusp. Broad marginal plaques are present.

The measurements for the largest specimen available are: M.L., 72 mm; M.W., 46 mm; H.W., 40

mm; Arm L., I-IV, respectively, 62 mm, 48+ mm,

76 mm, 55 mm; length of posterior salivary gland,

16 mm; length of liver, 21 mm; width of liver, 6 mm;

eye diameter, 14 mm; height of largest sucker of hectocotylus, 6 mm.

DISCUSSION.—This species is very closely related

to /. heathi, but differs in several significant features.

The most apparent character is the lack of the brilliant silvery layer present on the eyes and liver of /.

heathi. This is very obvious in fresh material and

usually Formalin—preserved material, although the iridescence in /. heathi may be lost in some specimens

after preservation. This characters is difficult to use

in material preserved in alcohol. The testis and the

male reproductive tract are much larger than in /.

heathi at equivalent mantle lengths. Also, the viscera,

although showing considerable variation, are much

larger in /. heathi. In large fresh specimens these differences are readily seen, but I have been unable to

separate small specimens.

Although clearly different from /. heathi and /.

diaphana, it would be unwise to give this species a

name when it is only imperfectly separable from other

species. This must wait until better material is available for study and more detailed comparisons can be

made. Such a study, comparing this species with /.

heathi, would be extremely worthwhile for it could

set guidelines for the study of speciation within the

genus. Such guidelines are badly needed in this group

which at present seems to be a very confused state,

contrary to the conclusions of Thore (1949).

DISTRIBUTION.—See distribution for /. heathi.

Family OCYTHOIDAE Gray, 1849

DIAGNOSIS.—Web almost entirely absent. Right

arm III is hectocotylized and retained in a sac until

mature. Ventral surface of mantle covered with

tubercules and interconnecting ridges. Arms II and

III much shorter than arms I and IV. Arm suckers

biserial.

Genus Ocythoe, Rafinsque, 1814

DIAGNOSIS.—The characters of the genus are the

same as those of the family.

TYPE SPECIES.—Ocythoe tuberculata Rafinesque,

1814.

Ocythoe tuberculata Rafinesque, 1814

PLATE 37

Ocythoe tuberculata Rafinesque, 1814, p.29.—Berry, 1916,

p.l, fig. 1; 1955, p.177, fig. 1.—Naef, 1923, p.749, text

figs. 447-454.—Sasaki, 1929, p.26, pl.3, figs. 13,14, pl.8,

figs. 12-16, text fig.8.—Robson, 1932, p.201, text fig.27.

DESCRIPTION.—The thick, muscular mantle is

short and broad. The free edge extends to the level

of the dorsal margin of each eye. The ventral surface

of the mantle is covered with tubercules and inner

connecting ridges in females. The tubercules are more

closely spaced at the posterior end of the mantle,

along the ventral mantle margin, and along the

lateral sides.

The funnel is very large; it reaches anteriorly well

past the bases of the arms. Laterally, the basal portion

is fused to the head; the tubular portion is long and

free. Between the base of each arm IV and the

funnel is a large pore. The funnel organ is very large;

the dorsal pad has the shape of an inverted V. The

2 ventral pads are separate and elongate. The locking-apparatus on the funnel consists of a knob with

an anterior flaring lip that lies above a deep pit. The

structure locks with a corresponding groove and ridge

on the mantle. The "olfactory" organ consists of a

flattened pad which may either protrude from the

body surface or reside in a shallow pit and is located

near the dorsoposterior edge of each eye. The eyes are

moderately large. The arms have the characteristic

formula I = IV>II = III. There is no web connecting

the bases of the arms. The suckers, which are biserial

and project well above the surface of each arm, are

connected along either margin by a muscular membrane. The arms lack keels or at most are merely

angled aborally.

The males are much the smaller. The only male

in the present collections has a M.L. of 15 mm, but

is immature. The right arm III is hectocotylized and

enrolled in a membranous sac. This arm has 2 wellseparated rows of flattened suckers. The inner aperatures of the suckers are small and displaced toward

the medial end of the transversely oval sucking disk.
NUMBER 9 7 93

TABLE 24.—Measurements in mm for Ocythoe

tuberculata Rafinesque, 1814.

Characters

Sex

M.L

M.W

H.W

Arm L. I

II

Ill

IV

Catalina Island

9 9

60 80

45 38

27 28

123 148

83 97

80 90

132 141

Station numbers of

VELERO

8242 8238

9 <?

12 14

12 13

11 11

35 38

18 19

L R

16 18 12*

38 41

*hcctocotylus

The lateral margin of the suckers is fused to a marginal membrane which extends along both sides of

the arm. There are 66 of these large suckers on the

arm. The arm narrows abruptly at the distal end of

the sucker-bearing portion and continues as a slender

and attenuate filament which appears to be considerably longer than the sucker-bearing portion of the

arm. The proximal portion of the filament is enrolled in a separate sac which forms a bulb at the

tip of the sucker-bearing portion of the arm. There

are 2 small pads that appear to be reduced suckers on

the surface of this sac.

Females have approximately 34 gill filaments, and

the only male specimen has 21.

The radula has a tricuspid rachidian tooth, tricuspid first lateral, bicuspid second laterals, unicuspid

third laterals, and marginal plaques.

DISCUSSION.—At present only a single species,

Ocythoe tuberculata, is recognized in the family

Ocythoidae; however, there seems to be considerable

variation between described specimens. The number

of gill filaments varies approximately between 33 and

48. Most authors (Jatta 1896, Robson 1932) described the radula approximately as in the present

specimens, but Naef (1921, 1923) stated the second

lateral tooth is unicuspid. Both Sasaki (1929) and

Jatta (1896) indicate that the hectocotylus has about

100 suckers, but the California specimen has only 66.

Also, the California specimens lack the rather large

aboral keels of arms I and IV described by Naef

(1923) and Robson (1932), although this may be an

ontogenetic difference. At present, these variations

are assumed to be of little significance, but clearly

warrant a more detailed study.

DISTRIBUTION.—Only four specimens have been

avaiable for this study. The smaller two animals

were captured in the mid-water trawl in zone 2. The

two large specimens, collected by Dorothy Petersen,

washed ashore at the isthmus on Catalina Island on

two successive days (16, 17 January 1967).

Family ALLOPOSIDAE Verrill, 1882

DIAGNOSIS.—Rachidian tricuspid and without seriation. Right arm III in males hectocotylized and

coiled in a pocket in front of the right eye. Suckers

uniserial proximally and biserialy distally. Other than

hectocotylization, sexual dimorphism lacking. A funnel-mantle locking mechanism present.

Genus Alloposus Verrill, 1880

DIAGNOSIS.—Characters are the same as those of

the family.

TYPE SPECIES.—Alloposus mollis Verrill, 1880

Alloposus mollis Verrill, 1880

PLATE 38B

DESCRIPTION.—The mantle is short, gelatinous, and

broad (length is nearly equal to width). The mantle

opening is wide and reaches roughly to the level of

the dorsal edge of the eyes. On the inner surface of

the mantle, just medial to the gills and lateral to

the median pallial adductor muscle is a distinct transverse fold of the mantle which forms a deep groove.

This is the mantle component of the funnel-mantle

locking mechanism.

The funnel is very large. It extends far past the

eyes and past bases of the ventral arms. Except for a

short free tip, it is completely embedded in the gelatinous tissue of the head. The funnel organ is large

and W-shape. The posterior lateral edges of the funnel, near the point of attachment of the funnel retractor muscles, fold sharply forward to form the

funnel component of the funnel-mantle locking

mechanism.

The head is broad and bears large eyes. There is

no lateral or dorsal constriction between the mantle

and head. Posterior to each eye lies a low, flattened

knob which is the "olfactory" organ.
94

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The arms are very large, but gelatinous. They are

in the order of I>II>III>IV. The suckers begin in

a single but slightly irregular series near the base of

each arm and gradually become biserially arranged

proximal to the edge of the web.

The gills are huge and bear 18 lamellae, each of

which is robust and highly folded. The body wall

which covers the ventral surface of the viscera is surprisingly tough and opaque. Very small nephridial

papillae project from near the base of the gills. Posterior and lateral to the large median pallial adductor

muscle, a pair of oviducts opens through prominent

papillae. Each papilla is extremely large and muscular. The largest eggs in the ovary are about 1.5

mm long by 0.5 mm wide. The viscera are essentially

as illustrated by Thore (1949). The kidneys are very

large and extend to the posterior end of the visceral

mass. The crop is situated dorsal to the liver, and the

stomach and caecum are dorsal and partially posterior to the liver. The entire dorsal surface of the visceral mass is free from the overlying mantle. The

space between the visceral mass and the mantle communicates with the ventral mantle cavity laterally

around the visceral mass anterior to the funnelretractor muscles.

The rachidian tooth of the radula is tricuspid; the

first lateral is small and bicuspid, and the second and

third laterals are unicuspid. The marginals are simple

and broad.

The data for the single specimen captured are:

Sex, $ ; M.L., 115 mm; M.W., 126 mm; H.W., 120

mm; Arm L., I-IV, respectively, 360 mm, 260 mm,

240 mm, 200 mm; Eye D., 38 mm.

TYPE LOCALITY.—South of Newport, Rhode Island,

western north Atlantic.

LOCATION OF TYPE.—United States National Museum.

DISCUSSION.—Thore (1949) concluded that there

is only a single species in the Alloposidae. I have compared my large, but rather poorly preserved, specimen with material from the Atlantic Ocean and can

find no noticeable difference.

DISTRIBUTION.—The single specimen, captured in

August by the VELERO, came from the Santa Catalina Basin (33°23' N, 118°49' W). Thore (1949) reviewed the distribution of A. mollis and has shown

that it is restricted to tropical and subtropical waters

throughout the world.

Family OCTOPODIDAE Orbigny, 1845

DIAGNOSIS.—Benthic octopods with simple lateral

radular teeth and without a detachable hectocotylus.

Stomach and caecum posterior to liver.

Octopus Lamarck, 1798

DIAGNOSIS.—Arms with biserial suckers. Ink sac

normal, web moderate. Third right arm hectocotylized. Single penial diverticle. Without velar pouches

or finlike membranes. Mantle aperature wide.

TYPE SPECIES.—Octopus vulgaris Lamarck, 1798.

Octopus species

PLATE 38A

DESCRIPTION.—The mantle is short and broad; its

width is roughly 80-85% of its length. The mantle

opening extends to the ventral margin of the eyes.

The head is narrower than the mantle, and there is

only a slight constriction between them. The eyes are

small. The integument over the mantle and head appears to contain a definite pattern of papilla in the

larger specimens which is rather inconspicuous when

retracted. There is a simple median papilla on the

dorsal surface of the head at the level of the anterior

edge of the eyes. Dorsal to the posterior portion of

each eye, there is a complex occular cirrus. On the

mantle immediately posterior to each eye is a simple

papilla. Just slightly posterior to these, in the dorsal

midline is a single median papilla. At about 40% of

the mantle length from the anterior edge of the

mantle is a pair of simple papillae; each member is

situated on a dorsolateral edge of the mantle. At

about 25% of the M.L. from its posterior end a

similar pair of simple papillae is situated on the

dorsolateral areas of the mantle, but slightly ventral

to the more anterior pair. There are other papillae

that are smaller and often difficult to locate.

The entire surface of the mantle in the larger specimens is slightly granular.

The funnel extends to the level of the anterior

edges of the eyes. The funnel organ is W-shaped,

with the lateral arms at least as long as the median

ones.

The arms are short and robust. The arm formula

is somewhat variable, but in general the arms are

subequal. The right arm III of the largest male is
NUMBER 9 7

hectocotylized. The ligula is short and broad and incompletely formed; it is about 2% of the arm length.

A calamus is not present. Suckers are arranged in 2

rows on the arms, and there are no especially enlarged suckers. The web formula is somewhat irregular, but appears to be B = C=D> A = E. The

gills are large and have approximately 23-26 primary

lamellae.

Numerous reddish brown chromatophores are scattered over the animal, but are more concentrated on

the dorsal surface. In life the ventral surface has a

silvery, iridescent sheen.

DISCUSSION.—There are numerous specimens of

pelagic larval Octopus in the collections which appear to belong primarily to one species. Specimens

range from less than 2 mm M.L. up to 25 mm M.L.

This description refers to the larger specimens only.

The following survey briefly discusses possible relationship of the large specimens to the known species

of this area.

There are currently 9 species of Octopus recognized from southern California and adjacent shores.

These are: O. bimaculatus Verrill, 1883; O. bimaculoides Pickford and McConnaughey, 1949; O. leioderma (Berry 1911); O. californicus (Berry 1911);

O. rubescens Berry, 1953; O. dofleini (Wiilker 1910);

O. pricei (Berry 1913); O. micropyrsus Berry, 1953;

and O. penicillifer Berry, 1954.

Octopus bimaculatus and O. bimaculoides are sibling species that are easily distinguished from the

others by the presence of a pair of ocelli near the

bases of arms II and III. O. bimaculoides has a welldeveloped ocellus at a mantle length of 9 mm and

does not have a pelagic larval stage. O. bimaculatus

has small eggs and presumably pelagic larvae. Type

localities: O. bimaculatus, San Diego, California; O.

bimaculoides, Devil's Slide, La Jolla, California.

There are no ocelli present in the pelagic juveniles of

VELERO collections; also, the number of primary

gill lameallae (23-26) falls outside the maximum

range (14-20) for O. bimaculatus. This latter feature is sufficient to eliminate the inclusion of the

VELERO specimens in O. bimaculatus.

The adults of O. leioderma have a short, squat

body, smooth integument, a lateral keel-like fold

around the mantle, and large eyes. Also the dorsal

arms are longer than the other pairs. The smallest

specimens overlap the VELERO material in size

which, according to Berry, exhibit all characteristics

95

of the adults. Type locality: Shelikof Strait, Alaska;

also recorded by Berry (1912b) from Monterey Bay.

The arm formula, the keel on the mantle, the smooth

integument and the large eyes would seem to eliminate the VELERO specimens from this species.

O. calif ornicus can be distinguished by the presence

of two, norrowly separated V-shaped funnel organs.

In addition, the juveniles which overlap the

VELERO specimens in size have enormous eyes.

Clearly, the VELERO juveniles do not belong to this

species. Type locality: Vicinity of San Diego, California.

Berry (1953) states that O. rubescens seems to be

the commonest small offshore species of Octopus in

the waters of southern California and Baja California. In addition, he states the eggs are minute

(1.3 mm), and from this it may be assumed that the

larvae are pelagic. Berry described the integument as

"superficially smooth but under a lens seems to be

raised in a tessellate pattern of low blister-like welts,

with a few symmetrically disposed, rather small,

papillae-like cirri on the dorsum and sides." In the

largest of the VELERO specimens, there is some indication of "blister-like welts" and the papillae seem

similar to those in the present larvae. The holotype of

this species is a male with a body length (32 mm)

only slightly larger than the largest VELERO specimen. The much longer arms, long ligula, and a few

enlarged suckers on the arms indicated that this

species may be separate from the VELERO species,

although it is quite possible that these differences

would be resolved after the pelagic larvae adopt a

benthic habitat. Unfortunately, the gill count is unTABLE 25.—Measurements in mm of Octopus sp.

Characters

M.L

M.W

H.W

Arms I

II

Ill

IV

Eye D

GillF

Sucker D

Station Numbers of the VELERO

7393 7387 7412 7414

15 17 18 25

11 13 13 18

9 11 10 15

27 35 39 48

29 38 43 54

30 33 40 50

29 29 41 48

4 4.5 4 5

24 25 26 25

1.5 1.7 1.5 2.0
96

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

known. Type locality: 7-17 fathoms off South Coronado Island, Baja California, Mexico.

Pickford (1964) referred the giant octopus of

California to Octopus dofleini. The gap in size between described specimens of this species and the

VELERO specimens is great; however, O. dofleini

has eggs that are 6-8 mm long and, therefore, are

probably not pelagic. Also the number of primary

lamellae on the gills varies from 40-60. The number

of gill lamellae, at least in some species (Burgess

1966) do not seem to vary with the mantle length.

Although it is possible that the gill lamellae increase

in numbers upon settling, at our present state of

knowledge the great number of lamellae in O. dofleini

must be considered as evidence against relationship.

Other differences concern the greater length of the

arms and the very elongate ligula in O. dofleini. Both

features could be due to growth. The weight of evidence appears, however, to be against aligning the

VELERO specimens with O. dofleini. Type locality:

Hokkaido, Japan.

O. pricei has been described from specimens considerably smaller (18 mm M.L.) than the larger

VELERO specimens. Berry (1913) stated that this

species has very large eyes; this feature alone is

enough to separate it from the VELERO specimens,

which possess small eyes. Type locality: From

stomach of Onchorhynchus tschawytscha taken off

Point Pinos, Monterey Bay, California.

O. micropyrsus is considerably smaller (10 mm

M.L.) than larger VELERO specimens. It has more

slender arms, larger eyes, a few enlarged suckers, and

an enormous calamus which clearly separate it from

the VELERO specimens. Type locality: Off La

Jolla, California.

O. veligero is also a small species, although specimens have not been recorded which overlap VELERO specimens in size. Its tripartite funnel organ,

fewer gill lamellae and the velumlike prolongations

of the web along the ventral margins of the arms

separate it from the VELERO Octopus. Type locailty: 50 fathoms off San Juanico, Baja California,

Mexico.

The unfortunate conclusion resulting from this

brief survey is that the VELERO specimens are not

clearly referable to any of the known species. This

is a somewhat surprising result considering the great

abundance of these larvae in the plankton. Part of

the difficulty in identification results from the inadequate descriptions of known species. Certainly much

of the difficulty is due to their possession of larval

characteristics, in spite of their rather large size, and

that only after settling and subsequent metamorphosis will it be possible to definitely determine their

identity. The problem remains an interesting one for

future workers.

Order VAMPYROMORPHA

Family VAMPYROTEUTHIDAE Thiele, 1915

DIAGNOSIS.—Oral crown consists of 8 arms and 2

retractile, sensory filaments. Mantle fused to the head.

Fins are present. Suckers are arranged in single series

on the arms and lack chitinous rings. Photophores are

present.

Genus Vampyroteuthis Chun, 1903

DIAGNOSIS.—Characters are the same as those of

the family.

Vampyroteuthis infernalis Chun, 1903

PLATES 35A; 36E-O

Vampyroteuthis infernalis Chun, 1903, p.88, text fig. [no

number].—Thiele, 1915, p.534, text figs. 67-70, pl.40,

pl-41, figs. 1-5.—Robson, 1932, p.90.—Pickford, 1946,

p.l; 1949, p.l, pi. 1-9.

Cirroteuthis macrope.—Berry, 1912b, p.2733, text fig.l, pi.

32, figs. 1-3.

DESCRIPTION.—The short, broad mantle is extensively fused ot the head dorsally. The mantle wall is

thick, but very gelatinous. The fins which arise subterminally are large and paddlelike. The large funnel

is embedded in the gelatinous tissue of the head for

almost its entire length. The funnel organ consists of

only 2 small, oval pads. A funnel valve is present.

The head is broad and bears very large eyes. There

is no anterior sinus on the circular eyelid. A small "olfactory" papilla is present posterior to each eye. The

animal has 8 arms, each of which bears a single row

of suckers. The suckers, which lack chitinous rings,

alternate with lateral cirri along most of the arm

length except basally, where the arms are bare. About

12-18 primary cirri occur proximal to the first sucker

on each arm. The suckers disappear on the very attenuate arm tips and only the conical sucker bases re-
NUMBER 9 7 97

TABLE 26.—Measurements in mm for Vampyroteuthis infernalis Chun, 1903.

Characters

Sex

P.L

P.W

Egg Diam

Sucker D

GillL

Gill W

Filament L

Ov. G. Diam

California

Station Numbers of the VELERO

8796 10259 10259

9 <? <? cf

74 58 79 60

31 28 34 26

0.9 — —

1.5 1.2 1.7 1.4

L R L R L R L R

21 22 21 21 21 22 15 16

11 11 9 8 8 10 7 8

12 11 9 8 10 10 7 8

2 2 — — — — — —

Africa

Station Numbers of the PILLSBURY

308 308 308 308

9 9 9c ?

78 64 56 49

38 35 29 25

4.0 2.8 — —

2.6 2.2 1.7 1.7

L R L R L R L R

17 16 13 12 12 — 10 —

7 7 5 6 4— 4 —

7 7 7 7 4— 5 —

6 6 5.5 6 — — — —

main; also the lateral cirri fuse together to form a

membrane along each distal margin. The arms are

joined by an extensive web that extends approximately to the point where the lateral cirri fuse, but

not quite to the point where the suckers terminate.

The web is equally developed between all arms. Between arms I and II on either side, a slender, retractile filament extends from a pocket on the dorsal

surface of the web. The filament is frequently completely retracted within this pocket and may be difficult to locate.

There is no buccal membrane.

Numerous small photophores are scattered over the

surface of the head, mantle, fins, and arms, but are

more abundant on the ventral surfaces of the animal.

Photophores are lacking on the oral surface of the

arms and web and from the aboral surface of the

web. A single large photophore is located just posterior to each fin. An oval patch of thickly packed

small photophores is located on the dorsolateral surface of the body at the level of the mantle opening.

In females there is a flask-shape vesicle imbedded

anterior to each eye which functions as a spermatophore receptacle. Eggs are about 3 mm in diameter at maturity. Nidamental glands are lacking.

The oral face of the arms and web is jetblack. The

aboral surfaces, including the fins and mantle, are

also nearly black, but seem to have a slight purple

tint in the living specimens.

In the specimens counted, the gills each had from

28-31 filaments.

Larval specimens have a single posterior pair of

fins. With increased size an anterior pair begins to

develop that eventually surpasses the larval fin in

size. The larval fin degenerates and is represented in

adult specimens by a small pocket behind the adult

fins.

TYPE LOCALITY.—Region of the Cape Verde Islands.

LOCATION OF TYPE.—Berlin Museum, Berlin,

Germany.

DISCUSSION.—A single species, V. infernalis, is

presently recognized in the order. I compared specimens from the Gulf of Guinea which is not far from

the type locality, with specimens from California and

find that there are a number of differences. The most

apparent difference is the great discrepancy in beak

size which can be seen by the illustrations on Plate

36. In addition, the suckers are smaller in the California specimens, the gills are much larger, the females mature at a much larger size, and the eggs at

maturity seem to be smaller. Most of these features

are somewhat variable and must be used with caution. There can be, however, no doubt that the California and African specimens are significantly different. At present, these differences are interpreted as

representing geographical variation; however, a more

intensive study of the problem is warranted.

DISTRIBUTION.—Pickford (1946) has demonstrated

that Vampyroteuthis infernalis has a worldwide distribution primarily within the tropical and subtropical oceans, but with northern and southern
98 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

limits near the fortieth parallels of latitude. This

species has previously been known to occur off the

coast of southern California (Berry 1912) and has

recently been reported off the coast of Oregon

(Pearcy 1965) at a latitude of approximately 44° N.

In the present study, Vampyroteuthis was found to be

relatively abundant in all the areas sampled. The

highest frequency of capture (1.19 specimens/hour)

occurred in zone 6 and lowest (0.23 specimens/hour)

occurred in zone 1 (San Pedro Basin).

Records of Additional Pelagic Species from off

Southern California

A number of other species of pelagic cephalopods

occur in the southern California area, but were not

included in this study since material was either lacking or inadequate for proper treatment. These species

consist of the following: Dosidicus gigas, Moroteuthis

robusta, Loligo opalescens, Thelidioteuthis sp., Pterygioteuthis giardi, Argonauta pacifica, Branchioteuthis

sp., Taonius sp., Liocranchia sp., and Eledonella sp.

Dosidicus gigas commonly occurs off the western

coasts of Central and South America and Mexico.

Occasionally, it is found in the waters off California

and has been reported as far north as Monterey Bay.

In the years 1934-1937, it was particularly abundant

in local waters. Croker (1937) in recording its occurrence stated: "The plague of jumbo squid has been

worse in 1937. Set line, net and troll commercial

fishermen are still bothered by them, and in addition

sport fishermen have been harassed all spring. No

sooner does a pleasure boat start to fish than a horde

of squid appears on the scene to crowd the game fish

away and seize all the baits. When one is hooked, it

proceeds to shower boat and fishermen with ink and

water, and then delights in biting its captor with its

parrot-like beak. Several fishermen have been seriously bitten this year. Although squid fishing is hilarious sport for a few minutes, it becomes too much of

a good thing day after day." Dosidicus gigas attains

considerable size: 4/ 2 feet (Classic 1949) and a

weight of more than 30 pounds (Croker 1937) have

been recorded for specimens off California, but much

larger specimens have been reported off South America. Specimens of D. gigas can easily be distinguished

from other local ommastrephids by their attenuate

arm tips bearing many small suckers which increases

the total sucker count to well over 200 suckers per

arm. This is more than three times the sucker count

of other local ommastrephids.

Moroteuthis robusta has been captured off the west

coast of north America, along the Aleutian Islands,

and off northern Japan. Although this species has

been recorded a number of times from California,

only one record (Croker 1934) occurs as far south as

the present study area. This species attains a mantle

length of at least 7/ 2 feet (Berry 1912b:315). Although the I.K.M.T. (Isaacs-Kidd Midwater Trawl)

is perhaps not well suited for capturing specimens of

M. robusta, it is somewhat surprising that no larvae

were taken. It seems probable that southern California is at the extreme limit of its distribution. M.

robusta can easily be distinguished from Onychoteuthis borealijaponicus, the other local onychoteuthid,

by the relatively greater length of the arms, by the

larger number of hooks on the tentacular clubs

(about 36 in M. robusta and 25-27 on O. borealijaponicus) , and by the lack of photophores.

Loligo opalescens is probably the most abundant

cephalopod species occurring off the California coast.

This species is epipelagic and fast moving, and therefore difficult to capture with an I.K.M.T. Over 80

specimens, however, were taken during the present

study, but most were larval forms under 10 mm M.L.

Fields (1965) did a detailed study of the biology of

this species, and nothing could be added from the

present collection. L. opalescens is fished commercially off California, primarily at Monterey. Over the

past 17 years, the average catch has been approximately 6,500 tons per year; the maximum catch of

19,000 tons occurred in 1946 (Fields 1965).

L. opalescens is the only member of the sub-order

Myopsida occurring off California. It can be easily

separated from all other squid of this area by the

presence of a cornea covering the lens of each eye.

The several specimens of Taonius sp. taken during

the present program were too badly damaged to

treat systematically. Taonius has also been reported

from off Oregon (Pearcy 1965). This cranchiid appears very similar to Galiteuthis, but instead of the

central club suckers being modified into hooks, each

looks like a set of claws (i.e., a number of elongate

teeth on each sucker).

Berry (1912b) recorded a small, finned octopod

captured off San Diego, California as "? Stauroteuthis

sp." The identity of this form is still uncertain, since

no additional material has been taken. Judging, how-
NUMBER 9 7 99

ever, from the small size of the fins and the large size

of the eyes, it is possible that this is a juvenile specimen of Opisthoteuthis californiana Berry, 1949.

Argonauta pacifica has been reported from off

California on a number of occasions (see Berry

1912b). It is not yet certain whether this form is

specificially distinct from the Atlantic Argonauta

argo. Several larval specimens of the "paper nautilus"

are present in the VELERO collections from off Baja

California, but can only be referred to Argonauta sp.

Specimens of Argonauta can easily be recognized.

The tips of the dorsal arms in females are each provided with an extensive flaglike web which is responsible for secreting the paper-thin shell that serves

both as a broad chamber and a flotation device.

Males are dwarfed and lack the modifications of the

dorsal arms; however, the third left arm is hectocotylized, and enrolled in a sac; this feature distinguishes Argonauta from all other cephalopods.

Recntly McGowan (1967) has indicated the presence of Pterygioteuthis giardi in the southern California area. This species is very similar to P. gemmata, but can easily be separated by the presence of

biserial hooks in the middle of the arms. Apparently,

this warm-water species occurs off southern California only during certain years.

Other species taken in the present study include

larval specimens of Liocranchia sp., Thelidioteuthis

sp. from zone 7, Brachioteuthis sp. from zone 6, and

one damaged specimen of Eledonella sp. from zone 3.

Checklist of Cephalopods Definitely Known to Occur in Southern Californian or Neighboring Waters

Order Sepioidea

Family Sepiolidae

Rossia pacifica Berry, 1911 t

Order Teuthoidea

Suborder Myopsida

Family Loliginidae

Loligo opalescens Berry, 1911

Suborder Oegopsida

Family Enoploteuthidae

Abraliopsis falco, new species*

Abraliopsis felis McGowan and Okutani,

1968*

Pyroteuthis addolux, new species *

Pterygioteuthis gemmata Chun, 1908 *

Pterygioteuthis giardi Fischer, 1895

Thelidioteuthis sp.

Family Ommastrephidae

Dosidicus gigas d'Orbigny, 1835

Ommastrephes bartramii (Lesueur, 1821)*

Symplectoteuthis luminosa Sasaki, 1915 *

Family Neoteuthidae

Neoteuthis sp.*

Family Histioteuthidae

Histioteuthis heteropsis (Berry, 1913)*

Histioteuthis dofleini (Pfeffer, 1912)*

Histioteuthis corona berryi Voss, 1969

Family Octopoteuthidae

Octopoteuthis deletron, new species *

Family Gonatidae

Gonatus onyx, new species *

Gonatus berryi Naef, 1923 *

Gonatus pyros, new species *

Gonatus calif or niensis, new species *

Gonatopsis borealis Sasaki, 1923 *

Family Onychoteuthidae

Onychoteuthis borealijaponicus Okada, 1927 *

Moroteuthis robusta Verrill, 1876

Family Brachioteuthidae

Brachioteuthis sp.

Family Mastigoteuthidae

Mastigoteuthis pyrodes, new species *

Family Chiroteuthidae

Chiroteuthis calyx, new species *

Valbyteuthis oligobessa, new species *

Valbyteuthis danae Joubin, 1931 *

Family Grimalditeuthidae

Grimalditeuthis bomplandii (Verany, 1837)*

Family Bathyteuthidae

Bathyteuthis berryi Roper, 1968 *

Family Cranchiidae

Cranchia scabra Leach, 1817 *

Leachia dislocata, new species *

Galiteuthis phyllura Berry, 1911 *

Galiteuthis pacifica Robson, 1948 *

Helicocranchia pfefferi Massy, 1907 *

Taonius sp.

Liocranchia sp.

Order Octopoda

Suborder Cirrata

Family Opisthoteuthidae

Opisthoteuthis californiana Berry, 1949

Suborder Incirrata

Family Bolitaendiae

Japetella heathi (Berry, 1911)*
100

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Japetella sp.*

Eledonella sp.

Family Ocythoidae

Ocythoe tuberculata Rafinesque, 1814*

Family Alloposidae

Alloposus mollis Verrill, 1880 *

Family Argonautidae

Argonauta pacifica

Family Octopodiae

Octopus bimaculatus (Verrill, 1883)

Octopus bimaculoides Pickford and McConnaughey, 1949

Octopus leioderma (Berry, 1911)

Octopus californicus (Berry, 1911)

Octopus rubescens Berry, 1953

Octopus micropyrsus Berry, 1953

Octopus dofleini (Wulker, 1910)

Octopus penicillifer Berry, 1954

Octopus pricei Berry, 1913

Order Vampyromorpha

Family Vampyroteuthidae

Vampyroteuthis infernalis Chun, 1903 *

t Berry has divided R. pacifica, a benthic sepiolid, into

two subspecies: R. p. pacifica and R. p. diegensis.

* Species described in this report.

For a complete review of the early work done

along the Pacific coast of North America, the reader

is referred to the excellent paper of Berry (1912b).

Summary

SYSTEMATICS

1. Thirty-three species of pelagic cephalopods and

the pelagic larval stage of a benthic octopus are described, and a key to their identification is presented.

2. The following 10 species are described as new

to science: Abraliopsis falco, Pyroteuthis addolux,

Octopoteuthis deletron, Gonatus onyx, Gonatus pyros,

Gonatus californiensis, Chiroteuthis calyx, Valbyteuthis oligobessa, Mastigoteuthis pyrodes, and Leachia

dislocate

3. It is demonstrated that Abraliopsis felis and

Watasenia scintillans do not belong in the same subgenus.

4. Pyroteuthis margaritifera oceanica, Pyroteuthis

margaritifera aurantiaca are considered to be invalid

subspecies.

5. Pterygroteuthis gemmata is shown to undergo

considerable geographical variation in the number

of arm hooks.

6. The photophore patterns of Ommastrephes bartramii and Symplectoteuthis luminosa are described.

7. The genus Neoteuthis is delineated and the family Alluroteuthidae is synonymized with the family

Neoteuthidae.

8. Octopoteuthis longiptera is declared a doubtful

species. It is suggested that all identifications of Octopoteuthis sicula from the Atlantic Ocean must be

held in question until the photophore pattern of

specimens from the type locality in the Mediterranean is determined. Octopodoteuthopsis is submerged for the second time into Octopoteuthis.

9. The formerly monotypic genus Gonatus is

shown to consist of at least 10 species, 4 of which live

sympatrically off California. The number of arm and

club suckers in members of the genus are shown to

become fixed at a very early stage and constitute an

important systematic character. Each species off California is shown to have a distinctive pattern at the

time when various arm and club hooks appear.

10. Larval Gonatopsis borealis are shown to possess

tentacles which are lost abruptly at 8-11 mm M.L.

Gonatopsis borealis makko is elevated to specific

status.

11. Onychoteuthis borealijaponicus is shown to be

a valid species. It is further suggested that "O.

banksii" represents a complex of species.

12. The formerly monotypic genus Valbyteuthis is

shown to consist of at least 5 species, 2 of which are

undescribed.

13. The enlarged photophore embedded in the eyelid of certain Mastigoteuthis is shown to be a widespread feature in the genus. M. grimaldii and M. iselini are declared doubtful species. M. levimana is

transferred to the genus Valbyteuthis. Mastigoteuthis

glaukopis atlantica is elevated to specific status. M.

dentata is briefly redescribed from material taken at

the type locality off Panama.

14. Pyrgopsis is shown to represent a larval stage

of Leachia.

15. Galiteuthis phyllura is considered to be a valid

species, although evidence for its separation from G.

armata is not conclusive. Taonidium pacificum is

shown to be a larval Galiteuthis and a valid species.

16. Helicocranchia is demonstrated to be a valid

and distinctive genus. The enigmatical genus Ascocranchia is shown to be a close relative of Helicocran-
NUMBER 9 7 101

chia. The identification of Helicocranchia pfefferi off

California needs confirmation through the comparison of adults from the Atlantic and Pacific oceans.

17. Two sympatric species of Japetella are shown

to occur off California, but their taxonomic separation has been incomplete.

18. The abundant octopus larvae appear to belong

predominantly to one species, but it has not been

identified.

19. It is suggested that Berry's Strauroteuthis sp.

may be the young of Opisthoteutnis californiana.

20. Species known to occur off California, but not

encountered in the present trawling program, are

briefly distinguished from those described in this

paper.

DISTRIBUTION

1. Northern zones within the area studied have

predominantly Subarctic and Transitional water

species.

2. Southern zones have predominantly tropical

species. The two most numerous of which appear to

be associated with the north Pacific central water.

3. Forty-two species of pelagic cephalopods are

now known from the waters off southern California,

of which 36 have been identified to the specific level.

Only 10 of these species appear to be cosmopolitan;

however, in only 4 of the 10 species were detailed

comparisons with Atlantic specimens possible. All 4

showed marked geographical variation.

4. All 10 apparently cosmopolitan species are associated primarily with tropical and subtropical

waters.

5. It is suggested that Histioteuthis heteropsis and

Octopoteuthis deletron are antitropical species.

6. Gonatus onyx, G. califoriensis, and G. pyros appear to occur primarily in the inshore waters over

the Continental borderland while Onychoteuthis

borealijaponicus, larval Abraliopsis felis, and Pyroteuthis addolux showed preference for more oceanic

waters.

Literature Cited

Adam, W.

1952. Ce"phalopodes. RSsultats Scientiftques Expedition

0' ceanographique Beige Atlantique Sud, 3 (3): 1—

142.

1954. Cephalopoda. 4. Clphalopodes a Pexclusion des

genres Sepia, Sepiella et Sepioteuthis. Siboga

Expeditie, 55c: 121-198.

1960. Les Cephalopodes de PInstitut Frangais d'Afrique

Noire. Bulletin de I'Institut frangaise d'Afrique

Noire, SSrie A, 22:465-511.

Akimushkin, I.I.

1963. "Cephalopods" 233 pages. Moscow. [In Russian.]

(Translation by A. Mercado. Cephalopods of the

Seas of the U.S.S.R. 223 pages. Jerusalem, 1965.)

Allan, J.

1945. Planktonic Cephalopod Larvae from the Eastern

Australian Coast. Record of the Australian Museum, 21:317-350.

Berry, S.S.

1909. Diagnoses of New Cephalopods from the Hawaiian

Islands. Proceedings of the United States National

Museum, 37:407-419.

1911. Preliminary Notices of Some New Pacific Cephalopods. Proceedings of the United States National

Museum, 40:589-592.

1912a. Some Necessary Changes in Cephalopod Nomenclature. Science, 36 (932): 643-646.

1912b. A Review of the Cephalopods of Western North

America. Bulletin of the Bureau of Fisheries, 1910,

30 (761): 267-336, 25 plates.

1912c. On a Cephalopod New to California, with a Note

on Another Species. Collected Reports of the

Laguna Marine Laboratory, 1:83—87.

1913. Notes on Some West American Cephalopods. Proceedings of the Academy of Natural Sciences of

Philadelphia, 913:72-77.

1914. The Cephalopoda of the Hawaiian Islands. Washington; A Bulletin of the Bureau of Fisheries,

32:255-362.

1916. Cephalopoda of the Kermadec Islands. Proceedings of the Academy of Natural Sciences of Philadelphia, 68:45-66.

1920. Preliminary Diagnoses of New Cephalopods from

the Western Atlantic. Proceedings of the United

States National Museum, 58 (2335):293-300.

1949. A New Opisthoteuthis from the Eastern Pacific.

Leaflets in Malacology, 1(6): 23-26.

1953. Preliminary Diagnoses of Six West American Species of Octopus. Leaflets in Malacology, 1(10) :

51-58.

1954. Octopus Penicillifer, New Species. Leaflets in

Malacology, 1(11) :66.

1955. On Recent Californian Occurrences of the Rare

Octopod Ocythoe. California Fish and Game, 41:

177-181.

1963. A "Doratopsis" Larva of the Squid Family Chiroteuthidae in Californian Waters. California Fish

and Game, 49:128-139.

Boone, L.

1933. Scientific Results of Cruises of the Yachts "Eagle"

and "Ara," 1921-1928. Mollusca. Bulletin of the

Vanderbilt Marine Museum, 4:165-210.
102

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Brinton, E.

1962. The Distribution of Pacific Euphausiids. Bulletin

of the Scripps Institute of Oceanography, 8(2) :

51-270.

Burgess, L.A.

1966. A Study of the Morphology and Biology of

Octopus hummelincki, Adam, 1936 (Mollusca:

Cephalopoda). Bulletin of Marine Science, 16(4):

762-813.

Chun, C.

1903. Aus den Tiefen des Weltmeeres. Jena:Gustav

Fischer.

1908. Ueber Cephalopoden der Deutschen Tiefsee-Expedition. Zoologischer Anzeiger, 33:86-89.

1910. Die Cephalopoden, I: Oegopsida. Wissenshaftliche

Eregebnisse der Deutschen Tiefsee-Expedition,

Valdivia, 18(1): 1-410; atlas of 61 plates.

1911. Cirrothauma, ein blinder Cephalopod, PromotionsReuntictions Programm der Philosophischen Fakultat der Universitat Leipzig, 1910, 11:1-21.

1913. Cephalopoda from the "Michael Sars" North

Atlantic Deep-Sea Expedition, 1910. Report on

the Scientific Results of the "Michael Sars" North

Atlantic Deep Sea Expedition, 1910, 3:1-21.

Clark, F.N., and J.B. Phillips

1936. Commercial Use of the Jumbo Squid, Dosidicus

gigas. California Fish and Game, 22(2): 143-144.

Clarke, M.R.

1962. The Identification of Cephalopod "Beaks" and

the Relationship Between Beak Size and Total

Body Weight. Bulletin of the British Museum

(Natural History), Zoology, 8(10):419-480.

1965. Large Light Organs on the Dorsal Surfaces of the

Squids Ommastrephes pteropus, "Symplectoteuthis

oualaniensis" and "Dosidicus gigas." Proceedings

of the Malacological Society of London, 36:319—

321.

1966. A Review of the Systematics and Ecology of

Oceanic Squids. Advances in Marine Biology, 4:

91-300.

Classic, R.F.

1949. Squid. Fish Bulletin, California Fish and Game

Commission, (74): 172-174.

Croker, R.S.

1934. Giant Squid Taken at Laguna Beach. California

Fish and Game, 20:297.

1937. Further Notes on the Jumbo Squid Dosidicus

gigas. California Fish and Game, 23:246-247.

Degner, E.

1925. Cephalopoda. Report on the Danish Oceanographic Expeditions 1908-10 to the Mediterranean

and Adjacent Seas, 2:1-94.

Ekman, S.

1953. Zoogeography of the Sea. London: Sedgwick and

Jackson, Limited.

Emery, K.O.

1960. Sea off Southern California. New York:John

Wiley and Sons, Inc.

Ferussac, A. de

1835. Note sur des cephalopodes nouveaux. Annales des

Sciences Naturelles, series 2, 3:339-390.

Ferussac, A. de, and A. d'Orbigny

1835-48 Histoire naturelle ginerale et particuliire des

cephalopodes acetabuliferes vivants et fossiles.

Paris, 2 volumes, text and atlas. [These volumes

appeared in numerous separate sections during

this period.]

Fields, W.G.

1965. The Structure, Development, Food Relations, Reproduction, and Life History of the Squid Loligo

opalescens Berry. California Department of Fish

and Game, Fish Bulletin, 131:1-108.

Fields, W.G., and V.A. Gauley

In press. Preliminary Description of an Unusual Gonatid

Squid (Cephalopoda: Oegopsida) from the North

Pacific. Journal of the Fisheries Research Board

of Canada.

Fischer, H.

1896. Note preliminaire sur le Pterygioteuthis Giardi,

cephalopode nouveau recueilli dans le cours de

l'Expedition scientifique du Talisman (1883).

Journal de Conchyliologie, 43:205-211.

Fischer, H., and L. Joubin

1906. Notes sur les cephalopodes captures au cours des

expedition du "Travailleur" et du "Talisman."

Bulletin du Musium d'historie naturelle, 12:202—

205.

Furtado, A.

1887. Sur une nouvelle espece de Cephalopode appartenant au genre Ommatostrephes-. Memorias da R.

Academia das Sciencias de Lisboa.

Goodrich, E.S.

1896. Report on a Collection of Cephalopoda from the

Calcutta Museum. London, Transactions of the

Linnean Society, series 2, 7:1—24.

Gray, J.E.

1849. Catalogue of the Mollusca in the Collection of

the British Museum: I, Cephalopoda Antepedia.

164 pages. London.

Grimpe, G.

1922. Systematische iibersicht der europaischen Cephalopoden. Sitzungsberichte der Naturforschenden

Gesellschaft zu Leipzig, 45-48:36-52.

1925. Zur Kenntnis der Cephalopoden-fauna der

Nordsee. Wissenshaftliche Meeresuntersuchungen

der Kommission, Kiel, 16(3): 1-124.

Hoyle, W.E.

1885. Diagnosis of New Species of Cephalopoda Collected During the Cruise of HMS Challenger, II.

The Decapoda. Annals and Magazine of Natural

History, series 5, 16:181-203.

1886. Report on the Cephalopoda Collected by H.M.S.

Challenger During the Years 1873-76. Report of

the Voyage of the Challenger, Zoology, 16(44):

1-246, 33 plates.

1904. Reports on the Cephalopoda. Bulletin of the Museum of Comparative Zoology, 43(1): 1-71.
NUMBER 9 7 103

Issel, R.

1908. Molluschi. 1. Cefalopodi planctonici raccolte

planctoniche fallo della R. Nave "Liguria." Pubblicazioni dell'Istituto di studi superiori pratici e

di perfezionamento in Firenze, 1:201-243.

Jatta, G.

1896. I cefalopodi viventi nel Golfo di Napoli (Sistematica). Fauna und Flora Neapel, Monograph 23,

268 pages.

Joubin, L.

1894. Note preliminaire sur les Cephalopodes provenants

des Campagnes du Yacht, L'Hirondelle. Memoires

de la Soctite" Zoologique de France, 7:211-216.

1895. Cephalopodes. Resultats des campagnes scientifiques accomplies par le Prince Albert I, 9:3-63.

1896. Observations sur divers cephalopodes. 1. Abraliopsis Pfefferi (nov. gen. et spec). Bulletin de la

Sociite scientifique et medicate de I'Quest, 5:19-

35.

1898a. Observations sur divers Cephalopodes. Quatrieme

note: Grimalditeuthis Richardi. Bulletin de la

Sociite Zoologique de France, 23:101-113.

1898b. Note sur une nouvelle farnille de cephalopodes.

Annales des sciences naturelles, series 8, 6:279-

292.

1899. Liste des cephalopodes recueillis pendant les derniers campagnes de la "Princesse-Alice" (1895-

1897). Bulletin de la Sociiti Zoologique de France,

24:62-74.

1900. Cephalopodes Provenant des Campagnes de la

Princesse-Alice (1891-1897). Risultats des campagnes scientifiques accompliis par le Albert I,

17:1-135.

1913. Etudes preliminaires sur les cephalopodes recueillis

au cours des croisieres de S.A.S. le Prince de

Monaco. 3. Mastigoteuthis magna nov. sp. Bulletin

de I'Institut oceanographique de Monaco, (275):

1-11.

1924. Contribution a Petude des Cephalopodes de PAtlantique Nord. 4e series. Resultats des campagnes

scientifiques du Prince de Monaco, 67:1-113, 16

plates.

1931. Notes Preliminaires sur les Cephalopodes des

Croisieres du Dana (1921-1922). 3e Partie. Annales de I'Institut Oceanographique, new series,

10(7): 167-211.

1933. Notes Preliminaires sur les Cephalopodes des

Croisieres du Dana (1921-1922). 4e Partie. Annales de I'Institut Oceanographique, new series,

13(1):1-49.

Lamarck, J.B.

1798. Bulletin de la Sociite Philomathique de Paris.

2:129.

Lavenberg, R.J., and A.W. Ebeling

1967. Distribution of Midwater Fishes Among Deepwater Basins of the Southern California Shelf.

Proceedings of the Symposium on the Biology of

the California Islands, Santa Barbara, California.

Botanic Garden, pages 185-201.

Leach, W.E.

1817. Synopsis of the Orders, Families, and Genera of

the Class Cephalopoda. Zoological Miscellany,

3:137-141.

Lesson, R.P.

1830. Mollusques. Voyage dans la Coquille, 2:239-246.

Lesueur, C.A.

1821. Descriptions of Several New Species of Cuttlefish.

Journal Academy Natural Sciences Philadelphia,

2:86-101.

Lichtenstein, K.M.H.

1818. Onychoteuthis, sepien mit krallen. Isis, 1818:

1591-1592.

Lonnberg, E.

1896. Notes on Some Rare Cephalopods. Ofversigt K.

Vetenskaps-Akademiens Forhandlingar. Stockholm,

8:603-612.

1899. On the Cephalopods Collected During the Swedish Arctic Expedition 1898 Under the Direction

of Professor A.G. Nathorst. Ofversigt af K. Vetenskapsakademiens forhandlingar. 1898. Pages 791—

792.

Lynn, R.J.

1967. Seasonal Variation of Temperature and Salinity

at 10 Meters in the California Current. California

Cooperative Oceanic Fisheries Investigations Reports, 11:157-174.

MacDonald, R., and WJ. Clench

1934. Descriptions of a New Genus and Two New

Species of Squids from the North Atlantic. Occasional Papers of Boston Society of Natural History, 8:145-152.

Massy, A.L.

1907. Preliminary Notice of New and Remarkable

Cephalopods from the South-west Coast of Ireland. Annals and Magazine of Natural History,

series 7, 20:377-384.

1909. The Cephalopoda Dibranchiata of the Coasts of

Ireland. Scientific Investigations of the Fisheries

Branch. Department of Agriculture for Ireland,

1907,: 1-39.

1928. The Cephalopoda of the Irish Coast. Proceedings

of the Royal Irish Academy, series B, 38:25-37.

McGowan, J.A.

1967. Distributional Atlas of Pelagic Molluscs in the

California Current Region. California Cooperative

Oceanic Fisheries Investigations Atlas no. 6, 218

pages.

1968. The Thecosomata and Gymnosomata of California. The Veliger. Supplement to 3:103.

McGowan, J.A., and T. Okutani

1968. A New Species of Enoploteuthid Squid, Abraliopsis (Watasenia) felis, from the California Current.

The Veliger, 11(1): 72-79.

Middendorf, A.T.

1849. Beitrage su einer Malacozoologie Rossica. Zapiski

Imperatorskoi akademu nauk, series 6, 6:67—275,

330-596.
104

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Moller, H.P.C.

1842. Index Molluscorum Groenlandiae. Naturhistorisk

Tidsskrift. Copenhagen, 4:76-97.

Muus, B.J.

1956. Development and Distribution of a North Atlantic

Pelagic Squid. Family Cranchiidae. Meddelelser

fra Kommissionen for Damnarks Fiskeri og Havundersegelser, 1:3-13.

1962. Cephalopoda. Meddelelser om Granland, 81:4-21.

Naef, A.

1921. Das System der Dibranchiaten Cephalopoden und

die Mediterranean Arten derselben. Mitteilungen

aus der Zoologischen Station Zu Neapel, 22:527-

542.

1923. Die Cephalopoden: Systematik. Fauna und Flora

Neapel, Monograph 35, l(part 1, number 2):ixiv, 149-863.

Nesis, K.N.

1971. The Family Gonatidae—Abundant Squids of the

North Pacific (their Distribution, Ecology, System and Phylogeny). Fourth Meeting of the Investigation of Molluscs. Academy of Sciences.

USSR Zoological Institute, 60-63. (In Russian.)

Okada, Y.K.

1927. Cephalopodes japonais des collections du museum.

Bulletin du Museum d'histoire naturelle, Paris.

Pages 93-98.

Okiyama, M.

1969. A New Species of Gonatopsis from the Japan Sea,

with the Record of a Specimen Referable to

Gonatopsis sp. Okutani, 1967 (Cephalopoda:

Oegopsida, Gonatidae). Publications of the Seto

Marine Biological Laboratory, 17(1): 19-32.

Okutani, T.

1966. Studies on Early Life History of Decapodan Mollusca-II. Planktonic Larvae of Decapodan Cephalopods from the Northern North Pacific in Summer

Seasons During 1952-1959. Bulletin of Tokai Regional Fisheries Research Laboratory, 45:61-79.

1967. Preliminary Catalogue of Decapodan Mollusca

from Japanese Waters. Bulletin of Tokai Regional

Fisheries Research Laboratory, (50): 1—16.

Okutani, T., and J.A. McGowan

1969. Systematics, Distribution, and Abundance of the

Epiplanktonic Squid (Cephalopoda, Decapoda)

Larvae of the California Current April, 1954-

March 1957. Bulletin of the Scripps Institution of

Oceanography, 14:1-90.

Okutani, T. and T. Nemoto

1964. Squids as the Food of Sperm Whales in the Bering

Sea and Alaskan Gulf. Scientific Reports of the

Whales Research Institute, (18): 111-122.

Orbigny, A.D.

1845. Mollusques vivants et fossiles ou description de

toutes les espices de coquilles et de mollusques

classees suivant leur distribution geologique et

geographique. 605 pages. Paris.

Parker, R.H.

1964. Zoogeography and Ecology of some Macroinvertebrates, Particularly Mollusks, in the Gulf of California and the Continental Slope off Mexico.

Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening 126:1-178.

Pearcy, W. G.

1965. Species Composition and Distribution of Pelagic

Cephalopods from the Pacific Ocean off Oregon.

Pacific Science, 19(2) :261-266.

Pearcy, W. G., and G. L. Voss

1963. A New Species of Gonatid Squid from the Northeastern Pacific. Proceedings of the Biological Society of Washington, 76:105-112.

Pfeffer, G.

1884. Die Cephalopoden des Hamburger Naturhistorischen Museums. Abhandlungen aus dem Gebiet

der Naturwissenschaften hrsg. vom Naturwissenschaftlichen Verin in Hamburg, 8:63-90.

1900. Synopsis der Oegopsiden Cephalopoden. Mitteilunegn Naturhistorischen Museum in Hamburg

(2 Beiheft Z. Jahrbach der Hamburgischen Wissenschaftlichen Anstalten), 17(2): 147-198.

1912. Die Cephalopoden der Plankton-Expedition.

Ergebnisse der Plankton-Expedition der Humboldt-Stiftung, 2:i-xxi, 1-815; atlas of 48 plates.

Phillips, J. B.

1961. Two Unusual Cephalopods Taken Near Monterey.

California Fish and Game, 47(4): 416-417.

1966. Additional Records of Flapjack Devilfish from

California Waters. California Fish and Game,

52(2):124.

Pickford, G.E.

1946. Vampyroteuthis infernalis Chun, an Archaic Dibranchiate Cephalopod. I. Natural History and

Distribution. Dana-Report No. 29, 40 pages.

1949. Vampyroteuthis infernalis Chun, an Archaic Dibranchiate Cephalopod. II. External Anatomy.

Dana-Report No. 32, 132 pages.

1964. Octopus dofleini (Wiilker). Bulletin of the Bingham Oceanographic Collection, 12(4): 1-66.

Pickford, G.E., and B.H. McConnaughey

1949. The Octopus bimaculatus Problem: A Study in

Sibling Species. Bulletin of the Bingham Oceanographic Collection, 12(4): 1-66.

Prosch, V.

1849. Nogle nye Cephalopoder. Kongelige Danske Videnskabernes Selskabs Skrifter, series 5, 1:53-72.

Rafinesque, C.S.

1814. Precis des decouvertes somiologiques ou zoologiques

et botanique. Palermo.

Rathke, H.

1835. Perothis, ein neues Genus der Cephalopoden.

Memoires presentes a academie Impiriale des

Sciences de Saint-Petersbourg, 2:1-43.

Reid, J.L., Jr., G.I. Roden, and G.J. Wyllie

1958. Studies of the California Current System. California Cooperative Oceanic Fisheries Investigations Report 1, pages 28-56.

Rice, D.A.
NUMBER 9 7 105

1963. Progress Report on Biological Studies of the

Larger Cetacea in the Waters off California.

Norsk Hvalfangsttid. 52:181-187.

Robson, G.C.

1924. On the Cephalopoda Obtained in South African

Waters by Dr. J. D. F. Gilchrist in 1920-21.

Proceedings of the Zoological Society of London.

Pages 589-686.

1932. A Monograph of the Recent Cephalopoda. Part

II. The Octopoda. British Museum. 359 pages.

1948. The Cephalopoda Decapoda of the Arcturus

Oceanographic Expedition, 1925. Zoologica,

33(3):115-132.

Rochebrune, A.T.de

1884. £tude monographique de la famille des Loligopsidae. Bulletin de la Societe Philomathique de

Paris, series 7, 8:7-28.

Roper, C.F.E.

1963. Observations on Bioluminescence in Ommastrephes

pteropus (Steenstrup, 1855), with Notes on its

Occurrence in the Family Ommastrephidae (MolIusca: Cephalopoda). Bulletin of Marine Science

of the Gulf and Caribbean, 13(2):343-353.

1968. Preliminary Descriptions of Two New Species of

the Bathypelagic Squid Bathyteuthis (Cephalopoda: Oegopsida). Proceedings of the Biological

Society of Washington, 81:161-172.

Roper, C.F.E., and R.E. Young

1967. A Review of the Valbyteuthidae and an Evaluation of its Relationship with the Chiroteuthidae

(Cephalopoda, Oegopsida). Proceedings of the

United States National Museum, 123(3612): 1-9.

Roper, C.F.E., R.E. Young, and G.L. Voss

1969. An Illustrated Key to the Families of the Order

Teuthoidea (Cephalopoda). Smithsonian Contributions to Zoology, 13:1-32.

Ruppell, E.

1844. Intorno ad alcuni cefalopodi del mare di Messina.

Giornale del Gablnetto di Messina, 26:1-7.

Sasaki, M.

1915. On Three Interesting New Oegopsids from the

Bay of Sagami. Journal of the College of Agriculture, Tohoku Imperial University, 6(6): 131-150.

1916. Notes on Oegopsid Cephalopods Found in Japan.

Annotationes Zoological Japonesis, 11:89-120.

1920. Report on Cephalopods Collected During 1906 by

the United States Bureau of Fisheries Steamer

"Albatross" in the Northwestern Pacific. Proceedings of the United States National Museum, 57:

163-203.

1923. On a New Eight-armed Squid from Hokkaido,

Gonatopsis borealis n.sp. Annotationes Zoological

Japonesis, 10:203-207.

1929. A Monograph of the Dibranchiate Cephalopods

of the Japanese and Adjacent Waters. Journal of

the College of Agriculture, Hokkaido Imperial

University, 20 (supplementary number): 1-357.

Shepard, F.P., and K.O. Emery

1941. Submarine Topography off the California Coast.

Geological Society of America, Special Papers,

31:1-171.

Steenstrup, J.

1855. Referat of Beskrivelse af et Par Kjaeber af en

kolossal Blacksprutte. Oversigt Danske Videnskabernes Selskab Forhandlinger, 1855:199-200. (English translation: A. Volse, J. Knudsen, and W.J.

Rees, 1962. The Cephalopod Papers of Japetus

Steenstrup. Copenhagen: Danish Science Press.)

1857. Oplysning om en ny art of Blacksprutter, Dosidicus Eschrichtii. Oversigt Danske Videnskabernes

Selskab Forhandlinger, 1857:11-14. (English

translation: A. Volse, J. Knudsen, and W.J. Rees.

1962. The Cephalopod Papers of Japetus Steenstrup. Copenhagen: Danish Science Press.)

1880. Orientering i de Ommatostrephagtige Blacksprutters indbyrdes Forhold. Oversigt Danske Videnskabernes Selskab Forhandlinger, 1880-81:73-110.

(English translation: A. Volse, J. Knudsen, and

W.J. Rees. 1962. The Cephalopod Papers of

Japetus Streenstrup. Copenhagen:Danish Science

Press.)

Thiele, J.

1912. Die Cephalopoden der Deutschen Siidpolar Expedition. Dentsche Siidpolar Expedition (1901-

1903) 16, Zoologie, 8:433-466.

Thiele, J. (and C. Chun)

1915. Die Cephalopoden. II Teil: Myopsida Octopoda.

Wissenschaftliche Ergebnisse der Deutschen

Tiefsee-Expedition auf dem Dampfer "Valdivia"

1898-1899, 18:403-552. Jena.

Thore, S.

1949. Investigations on the "Dana" Octopoda. Part I.

Bolitaenidae, Amphitretidae, Vitrelendonellidae,

and Alloposidae. Dana-Report 33:1-85.

Van Hyning, J.M., and A.R. Magill

1964. Occurrence of the Giant Squid (Moroteuthis

robusta) off Oregon. Fish Commission Research

Briefs, (Portland, Oregon) 10:67-68.

Ve"rany, J.B.

1837. Memoire sur six nouvelles especes des cephalopodes touvees dans le M£diterrane£ a Nice.

Memorie della Accademic Scienze dell'Instituto

di Torino, series 2, 1:91-98.

1851. Mollusques mSditerranee observes, decrits, figure's,

et chromolithographies d'apres le vivant. Premiere

partie. Ceph.alopod.es de la Mediterranee. Genes,

132 pages.

Verrill, A.E.

1876. Note on Gigantic Cephalopods—A Correction.

American Journal of Science, series 3, 14:236-

237.

1881a. The Cephalopods of the Northeastern Coast of

America, Part II. The Smaller Cephalopods, Including the Squids and Octopi, with Other Allied

Forms. Transactions of the Connecticut Academy

of Arts and Sciences, 5:259-446.

1881b. Report on the Cephalopods, and on Some Addi-
106

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

tional Species Dredged by the U.S. Fish Commission Steamer "Fish Hawk," During the Season of 1962.

1880. Harvard College, Bulletin of the Museum of

Comparative Zoology, 8(5): 99-116.

1882. Report on the Cephalopods of the Northeastern

Coast of America. Report of the United States 1963.

Commissioner of Fisheries, 1879:1-240.

1883. Reports on the Results of Dredging. . . . Report 1967.

24. Supplementary Report on the "Blake" Cephalopods. Harvard College, Bulletin of the Museum of Comparative Zoology, 11 (5): 105-114. Voss, N.

1885. Third Catalogue of Mollusca Recently Added 1969.

to the Fauna of the New England Coast and the

Adjacent Parts of the Atlantic. Transactions of

the Connecticut Academy of Arts and Sciences, Watase,

6:395-452. 1906.

Voss, G.L.

1956. A Review of the Cephalopods of the Gulf of Mex- Wiilker,

ico. Bulletin of Marine Science of the Gulf and 1910.

Caribbean, 6(2):85-178.

1960. Bermudan Cephalopods. Fieldiana. Zoology, 39

(40):419-446.

Ascocranchia joubini, a New Genus and Species of

Cranchiid Squid from the North Atlantic. Bulletin

de I'Institut oceanographique de Monaco, (1242):

1-6.

Cephalopods of the Philippine Islands. United

States National Museum Bulletin, 234:1-180.

Some Bathypelagic Cephalopods from South African Waters. Annals of the South African Museum,

50(5):61-88.

A Monograph of the Cephalopoda of the North

Atlantic. The Family Histioteuthidae. Bulletin of

Marine Science, 19(4) :713-867.

S.

On Luminous Cephalopods. Zoological Society,

Zoological Magazine, Tokyo, 18:195-196.

G.

Uber japanische Cephalopoden. Beitrage zur

Kenntnis der Systematik und Anatomie der Dibranchiaten. Miinchen, 71 pages.

Appendix 1

After the present manuscript was completed, a

paper by Okutani and McGowan (1969) on the systematics and distribution of larval squid from the

California current was published. Fourteen species of

larval squid are described and information is presented on spawning and distribution of some of these.

The authors attempted to identify the larvae by tracing successive developmental stages to the identifiable adult (page 7). This is a difficult task under the

best circumstances; however, it becomes doubly difficult when the systematics of the adults is in an unstable state as it was when these authors undertook

their project. Laboring under these handicaps, it is

inevitable that errors will be made; however, as long

as the following emendations are considered, little

information will be lost from their paper because of

systematic difficulties.

1. Specimens identified as Octopodoteuthopsis sp. almost certainly belong to Octopoteuthis deletron.

2. Onychoteuthis banksii should read Onychoteuthis

borealijaponicus. Their illustration (figure 9c) shows

a tentacular club with 26 hooks, a feature distinctive

of this latter species. Their records of this species are

shown to extend far south to the tip of Baja California. The most southernly records possible belong to a

member of the "O. banksii" complex. This is somewhat suggested by the shape of the individual in figure 9h (no locality data are given for this figure);

however, even if this proves to be the case, it will not

significantly effect the conclusions reached by Okutani

and McGowan.

3. The larvae identified as Gonatus fabricii probably

belong to the most dominant species in the genus off

California, Gonatus onyx, and this is attested to by

their figure of a club belonging to this species (page

28, figure 10a). Larvae of G. pyros and G. californiensis are extremely close to those of G. onyx and

since these species are quite common, I suspect some

of their specimens belong to these species also. The

authors mention that 10 larvae had more slender and

opaque mantles. I suspect that these are larvae of

Gonatopsis borealis and not Berryteuthis magister as

is suggested. Okutani and McGowan state that this

latter animal is known from the California coast. I

have not been able to find this record, but its occasional occurrence here would not be surprising since

it has been reported from Oregon. Even if the species

gets as far south as southern California, I doubt that

it breeds here as no larvae or juveniles have been

taken in the University of Southern California's

trawling program.

4. Juveniles identified as Meleagroteuthis heteropsis

consist of two species: Histioteuthis heteropsis (figure

12c) and H. dofleini (figure I2d).

5. The specific identification of their Ctenopteryx

must be held in question. The genus is in a systematic
NUMBER 9 7 107

jumble at present.

6. Larvae identified as Chiroteuthis veranyi almost

certainly belong to C. calyx.

7. Species identified as Pyrogopsis pacificus belong to

Leachia dislocata as evidenced by figure 166 which

shows the displaced tubercle characteristic of the

species.

8. Species identified as Teuthowenia megalops almost

certainly belong to Galiteuthis phyllura.

These authors add two species to those now known

from southern Galifornian and adjacent waters.

These are Liguriella sp. from 28° 17' N, 118°58' W

and 29° 12' N, 115°38' W, and Ctenopteryx sp. from

27°33'Nandll5°53'W.

VELERO

Station

Number

7157

7221

7279

7280

7343

7387

7389

7393

7394

7409

7410

7411

7412

7414

7415

8018

8022

8024

8025

8026

Stations

Date

30 September 1960

9 December 1960

2 February 1961

2 February 1961

7 April 1961

15 August 1961

15 August 1961

16 August 1961

16 August 1961

6 September 1961

6 September 1961

6 September 1961

6 September 1961

7 September 1961

7 September 1961

19 July 1962

20 July 1962

7 August 1962

7 August 1962

8 August 1962

Cited

Midpoint

Latitude

and

Longitude

33°36'N

118°28'W

33°32'N

118°24'W

33°32'N

118°23'W

33°29'N

118°20'W

33°33'N

118°24'W

33°31'N

118°18'W

33°29'N

118°20'W

33°30'N

118°22'W

33°33'N

118°22'W

33°36'N

118°24'W

33°30'N

118°18'W

33°29'N

118°19'W

33°37'N

118°24'W

33°28'N

118°19'W

33°32'N

118°26'W

33°18'N

118°40'W

33°19'N

118°42'W

33°21'N

118°46'W

33°16'N

118°38'W

33°22'N

118°43'W

Appendix 2

Wire Out

(meters)

1845

1385

2000

925

2300

1075

1690

770

2300

1690

2000

2300

2300

1690

2000

3090

3090

1385

1690

1385

VELERO

Station

Number

8027

8028

8030

8031

8111

8112

8114

8116

8117

8120

8121

8123

8237

8238

8243

8292

8697

8716

8795

8796

Stations

Date

8 August 1962

8 August 1962

9 August 1962

9 August 1962

21 August 1962

22 August 1962

22 August 1962

23 August 1962

23 August 1962

5 September 1962

5 September 1962

5 September 1962

22 October 1962

25 October 1962

26 October 1962

8 November 1962

23 May 1963

7 June 1963

18 July 1963

July 1963

Cited

Midpoint

Latitude

and

Longitude

33°22'N

118°43'W

33°15'N

118°32'W

33°09'N

118°23'W

33°10'N

118°22'W

33°21'N

118°45'W

33°21'N

118°44'W

33°15'N

118°31'W

33°19'N

118°44'W

33°19'N

118°45'W

33°2O'N

118°43'W

33°12'N

118°32'W

33°15'N

118°35'W

33°15'N

118°25'W

33°23'N

118°48'W

33°2O'N

118°45'W

33°22'N

118°45'W

33°22'N

118°46'W

33°15'N

118°37'W

33°21'N

118°46'W

33°22'N

118°44'W

Wire Out

(meters)

1690

2000

1690

1385

2300

2300

2000

1385

2000

3090

770

770

2300

3380

1540

770

3695

925

2460

1845
108 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

VELERO

Station

Number

8878

8881

8934

9056

9249

9349?

9602

9603

9605

9661

9890

9899

9901

9905

9960

10259

10260

10265

10399

10401

Stations Cited

Date

20 August 1963

21 August 1963

18 September 1963

14 November 1963

25 January 1964

16 April 1964

17 April 1964

17 April 1964

14 May 1964

5 August 1964

7 August 1964

7 August 1964

8 August 1964

September 1964

13 January 1965

13 January 1965

14 January 1965

7 March 1965

8 March 1965

(continued)

Midpoint

Latitude

and

Longitude

33°01'N

119° 04' W

33°03'N

119°00'W

33°11'N

118°28'W

33°18'N

118°42'W

33°19'N

118°38'W

Catalina Basin ?

33°17'N

118°40'W

33°22'N

118°54'W

33°20'N

118°37'W

32°59'N

119°02'W

32°48'N

120°56' W

29°58'N

119°12'W

29°33'N

118°59'W

29°19'N

119O21'W

Catalina Basin

33°19'N

118o4r\V

33°18'N

118°42'W

33°17'N

118°40'W

33°04'N

119°08'W

32°59'N

119°01'W

Wire Out

(meters)

3090

925

1540

3380

1540

1540

1075

770

3090

310

1845

310

2460

1845

2125

1075

3370

1385

1845

VELERO

Station

Number

10402

10673

10677

10679

10838

10840

10841

10844

10897

10906

10973

10976

10985

11022?

11097

11100

11101

11181

11183

11225

Stations Cited

Date

8 March 1965

27 August 1965

28 August 1965

28 August 1965

24 November 1965

25 November 1965

25 November 1965

26 November 1965

5 January 1966

6 January 1966

16 February 1966

17 February 1966

18 February 1966

14 April 1966

14 April 1966

14 April 1966

2 August 1966

2 August 1966

30 August 1966

(continued)

Midpoint

Latitude

and

Longitude

33°08'N

119°13'W

29°37'N

118°33'W

29°05' N

118°06'W

28°54'N

118°08'W

31°14'N

118°45'W

30°23'N

118°29'W

30°00' N

118°26'W

28°54'N

117°49'W

31°42'N

118°30'W

29°12'N

118°09'W

32°45'N

120°31'W

32°27'N

12O°27'W

31°18'N

117°22'W

No data

32°33'N

118°04'W

32°35'N

118°06'W

32°29'W

118°01'W

29°40'N

118°48'W

29°36'N

118°35'W

30°18'N

118°53'W

Wire Out

(meters)

925

Surface

1230

93

1230

4000

3090

4000

1230

3090

3090

4000

3090

1385

40

4000

4000

3090

3090
PLATES 1-38

NUMBER 9 7 111

PLATE 1.—A, Abraliopsis falco, ventral view, male, M.L. 37 mm; B, same specimen, left side of

head; c, Abraliopsis felis, ventral view, male, M.L. 41 mm; D, same specimen, left side of head.
112

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

M

PLATE 2.—A-G, Abraliopsis falco: A, tentacular club, M.L.

approximately 40 mm; B. arms IV, mature male; c, Largest

hook from arm III; D, distal sucker, arm III; E, largest club

hook; F, dactylus sucker (C,D,F maintain relative sizes, E has

half enlargement of others, all M.L. approximately 40 mm);

G, ventral view of eye. H-N, Abraliopsis felis: H, tentacular

club, M.L. 40 mm; i, arms IV, M.L. 41 mm; j , largest hook

from arms III; K, distal sucker, arm III; L, largest club

hook; M, dactylus sucker (j, K, M maintain relative sizes,

L has half enlargement of others, all M.L. 40 mm); N, ventral view of eye.
NUMBER 9 7 113

PLATE 3.—A, Pterygioteuthis gemmata, ventral view, M.L. 23 mm; B, Pyroteuthis addolux,

ventral view, holotype, M.L. 30 mm.
114

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 4.—A-H, Pterygioteuthis gemmata: A, tentacular club, M.L. 28 mm; B, medial view,

hectocotylus, M.L. 27 mm; c, largest hook, arm III; D, ventral view of eye; E, chitinous plate

from hectocotylus, M.L. 24 mm; F. chitinous plate from hectocotylus, M.L. 27 mm; G, inner

ring from largest sucker, arm III; H, inner ring, largest sucker, manus (c, o, H maintain relative sizes, M.L. 23 mm), i—<j, Pyroteuthis addolux: i, tentacular club, holotype; j , hectocotylus, holotype; K, spermatophore, holotype; L, enlargement of tip of ejaculatory apparatus,

holotype; M, largest club hook, tentacular club; N, inner ring of largest club sucker; o, largest

hook, arm III; p, inner ring, second proximal sucker, arm IV, (M-P maintains relative size, M.L.

45 mm); Q, ventral view of eye. R. Pyroteuthis margaritifera, second hook proximal to pad on

ventral row of hectocotylus, Gulf of Mexico specimen, M.L. 28 mm; s. Pyroteuthis addolux,

second hook proximal to pad on ventral row of hectocotylus, holotype, M.L. 30 mm.
NUMBER 9 7 115
116

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 5.—A, Symplectoteuthis luminosa, ventral view, M.L. 136 mm; B, Ommastrephes

bartramii, ventral view, M.L. 211 mm.
NUMBER 9 7 117

PLATE 6.—A, Symplectoteuthis luminosa, photophore pattern, ventral view, M.L. 163 mm; B,

Same, dorsal view; c, Ommastrephes bartramii photophore pattern, ventral view, M.L. 215 mm.
118

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 7.—A-I, Symplectoteuthis luminosa, M.L. 136 mm; A, tentacular club; B, inner ring of

large sucker from 6th row of arm III; c, inner ring of basal sucker of arm III; D, inner ring of

sucker from 18th row of arm III; E, inner ring of sucker from middle of dactylus, dorsal row;

F, enlargement of E; G, inner ring of largest sucker, manus, marginal row; H, inner ring of

largest sucker, manus, medial row; B-E, G-H maintain relative sizes); i, right funnel lockingcartilage showing area of fusion, J-R, Ommastrephes bartramii from off Guadalupe Island,

Pacific Ocean, M.L. 303 mm: j , right funnel locking-cartilage; K, inner ring of sucker from

middle of dactylus, ventral row; L, large medial sucker of manus showing fleshy, marginal papillae; M, tentacular club; N, inner ring of basal sucker, arm III; o, inner ring of 5th row sucker,

arm III; p. inner ring of 14th row sucker, arm III; Q, inner ring of largest sucker, marginal

row of club (K, N-Q maintain relative sizes); R, inner ring of largest sucker, medial row of club,

enlargement half that of other suckers; s, Ommastrephes bartramii from off Florida, M.L. 302

mm, inner ring of largest sucker, marginal row of club; T, same specimen, inner ring of largest

sucker, medial row of club (enlargement of s and T is the same as Q and R, respectively).
NUMBER 9 7 119
120

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 8.—Neoteuthis sp., M.L. 83 mm: A, ventral view; B, tentacular club; c, dorsal view.
NUMBER 9 7 121

PLATE 9.—A, Histioteuthis heteropsis, ventral view, photophore pattern only approximate, M.L.

35 mm; B, Histioteuthis dofleini, ventral view, M.L. 150 mm.
122

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

v * *

M

PLATE 10.—A, Histioteuthis dofleini, tentacular club, M.L.

150 mm; B, same, large club sucker, M.L. 150 mm; c, same,

inner ring of sucker from arm III, 6th row, M.L. 14 mm;

D, Histioteuthis heteropsis, tentacular club, M.L. 35 mm;

E, same, inner ring of large club sucker, M.L. 65 mm; F,

same, inner sucker ring, arm III, 11th row; o, same, inner

sucker ring, arm IV, eleventh row (E-G maintain relative

sizes), H-M, Octopoteuthis deletron; H, tip of arm III,

M.L. 167 mm; i, ventral view of mantle and fins, M.L. 167

mm; j , inner ring from tip of arm III, fifth row from hooks,

M.L. 167 mm; K, large hooks, arm III, M.L. 20 mm?; L,

large hook, arm III, M.L. 109 mm; M, large hook, arm III,

M.L. 167 mm.
NUMBER 9 7 123

i

,

PLATE 11.—A, Octopoteuthis deletron, holotype, M.L. 109 mm; B, same showing

photophore pattern.
124

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 12.—A-D, Octopoteuthis deletron showing metamorphosis, relative sizes maintained, M.L. 31, 32, 34, 39 mm, respectively; E, Octopoteuthis A, large arm hook. M.L. 49 mm; F, Octopoteuthis A, ventral view of mantle and fins. M.L. 49 mm;

G, Octopoteuthis neilseni, ventral view of mantle and fins, M.L. 36 mm; H, Octopoteuthis danae, ventral view of mantle and

fine, M.L. 53 mm; i, Octopoteuthis megaptera, ventral view of mantle and fins, M.L. 38 mm.
NUMBER 9 7 125

PLATE 13.—A, Gonatus onyx, ventral view, P.L. 69 mm; B,

P.L. 39 mm.

Gonatus pyros, ventral view,
126

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 14.—A, Gonatus onyx, tentacle, P.L. 69 mm; B, Gonatus pyros, tentacle, P.L. 35 mm?;

Gonatus onyx, P.L. 69 mm relative sizes maintained except D: C, inner ring of large marginal

sucker, arm III; D, large hook, arm III, enlargement half that of suckers; E, inner ring of large

medial row sucker, arm IV; F, inner ring of large marginal sucker, arm IV; o, inner ring of

largest sucker of ventral marginal zone of club; H, inner ring of sucker from middle of dactylus,

ventral row; i, inner ring of sucker taken one-fifth of tentacle length from base of club, J-Q,

Gonatus pyros, P.L. 39 mm, relative sizes maintained except for L: J, inner ring of arm III

sucker tilted to show frequent bicuspid nature of teeth; K, inner ring of large marginal sucker,

arm III; L, large hook, arm III, enlargement half that of suckers; M, inner ring of large medial

row sucker, arm IV; N, inner ring of large marginal sucker arm IV; o, inner ring of largest

sucker from ventral marginal zone on club; p, inner ring of sucker from middle of dactylus,

ventral row; Q, inner ring of sucker taken one-fifth of tentacle length from base of club.
NUMBER 9 7 127
128 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 15.—A, Gonatus berryi, ventral view, P.L. 119 mm;

B, Gonatus californiensis, ventral view, holotype, P.L. 112

nun.
129

PLATE 16.—A, Gonatus berryi, tentacle, P.L. 119 mm; B, Gonatus californiensis, tentacle, holotype, P.L. 112 mm. c-i. Gonatus berryi, relative sizes maintained except for D: C, inner ring of large marginal sucker, arm III; D, large hook, arm III,

enlargement half that of suckers; E, inner ring of large medial row sucker, ventral arm; F, inner ring of large marginal

sucker, arm IV; G, inner ring of largest sucker of ventral marginal zone club; H, inner ring of sucker from middle of dactylus, dorsal row; i, inner ring of sucker taken one-fifth of tentacle length from base of club, j-p , Gonatus californiensis,

holotype, relative sizes maintained except for K: J, inner ring of large marginal sucker, arm I: K, large hook, arm I,

enlargement half that of suckers; L, inner ring of large medial row sucker, arm IV; M, inner ring of large marginal

sucker, arm IV; N, inner ring of largest sucker of ventral marginal zone of club; o, inner ring of sucker from middle of

dactylus, ventral row; p, inner ring of sucker taken one-fifth of tentacle length from base of club.
130 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 17.—A-C, club hooks Gonatus berryi, PL. 119 m m (relative sizes maintained):

A, central hook, enlargement half that of other hooks; B, distal hook; c, first proximal hook.

D-F, Gonatus californiensis, P.L. 112 mm, club hook (relative sizes maintained): D, central

hook; E, distal hook; F, first proximal hook, c-h Gonatus pyros, P.L- 39 rnm, club hooks

(relative sizes maintained): G, central hook; H, distal hook; i, proximal hook, j , Gonatus

onyx, P.L. 69 mm?, central hook of club; K, Gonatus Pyros, P.L. 37 mm, ventral view of

eye showing photophore; L, Gonatus berryi, larva, P-L. 10 mm.
NUMBER 9 7 131
132 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

B

PLATE 18.—A, Onychoteuthis borealijaponicus, ventral view,

M.L. 50 mm; B, Gonatopsis borealis, ventral view, M.L.

240 mm.
NUMBER 9 7 133

M

PLATE 19.—A-G, Onychoteuthis borealijaponicus, M.L. 81 mm; A, tentacular club; B, ninth hook from proximal end of club

in ventral row from club in A; C-F, largest suckers from arms I-IV respectively (B-F retain relative sizes) ; G, visceral photophores. H, Onychoteuthis "banksii" from off Florida, visceral photophores, M.L. 80 mm. I-P, Gonatopsis borealis:

i, larva, M.L. 9 mm; j , Tentacle from larva, M.L. 7 mm; K-P, Gonatopsis borealis, M.L. 200 mm, relative sizes retained

except in M: K, inner ring of largest sucker, arm IV, median rowjL, inner ring of largest sucker, arm IV, marginal row;

M, largest hook, arm III, enlargement half that of suckers, N, inner ring of sucker, arm III tip, medial row; o, inner ring

of largest sucker, arm III, marginal row; p, inner ring of sucker, arm III tip, marginal row.
134

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 20.—Chiroteuthis calyx: A, advanced doratopsis stage, ventral view, M.L. 49 mm?;

B, ventral view, subadult, M.L. 54 mm.
NUMBER 9 7 135

PLATE 21.—A, Chiroteuthis calyx, oral view of tentacular club, M.L. 60 mm; B, same specimen,

aboral view of tentacular club; c, Chiroteuthis veranyi from Mediterranean Sea (DANA 4030

XVIII), M.L. 61 mm, aboral view of tentacular club.
136 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 22.—Chiroteuthis calyx except L and M which are C. veranyi. Inner sucker rings in A-I

maintain relative sizes except in E, M.L. 60 mm?, A, arm III, second row; B, arm III, thirteenth row; c, arm III, eighteenth row; D, arm III, fourteenth row; E, D enlarged four times;

F, arm I, sixth row; c, arm II, tenth row; H, arm IV, first row; i, arm IV, twelfth sucker, j ,

inner ring of large club sucker; K, largest arm III sucker from advanced doratopsis larva, M.L.

56 mm; L, C. veranyi from Mediterranean Sea (DANA 4139 IV), M.L. 68 mm, inner sucker

ring, twelfth sucker of arm IV; M, same, inner sucker rings, thirteenth row of arm III;

N, ventral view of eye showing photophores, holotype; o, left funnel locking-cartilage, M.L.

60 mm; Q, medial and marginal tentacular suckers and stalks, holotype.
NUMBER 9 7 137
138

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 23.—A, Valbyteuthis oligobessa, mature female, holotype; B, Valbyteuthis danae, ventral

view, redrawn from Roper and Young (1967), M.L. 34 mm; c, Valbyteuthis oligobessa, juvenile, M.L. 23 mm.
NUMBER 9 7 139
140 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 24.—A-G. Valbyteuthis danae redrawn from Roper and Young (1967); A, tentacular

club, M.L. 47 mm; B-E, largest suckers from arms I-IV, respectively, M.L. 47 mm; F, largest

club sucker M.L., 47 mm, (B-F maintain relative sizes); G, mantle and funnel locking-cartilages, M.L. 47 mm. H-N, Valbyteuthis oligobessa, V-10976, M.L. 27 mm; H-K, largest

suckers from arms I-IV, respectively; L, largest club sucker (H-L maintain relative sizes);

M, mantle and funnel locking-cartilages; N, tentacular club; o, Grimalditeuthis bomplandii,

large sucker and base from arm III, M.L. 89 mm; p, same specimen, inner ring of large sucker

from arm III; Q, Valbyteuthis oligobessa, section of distal portion of outer ring of arm sucker,

M.L. 34 mm; R, same specimen, scale from proximal portion of outer ring of arm sucker

(slightly diagrammatic).
NUMBER 9 7 141
142

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 25.—Mastigoteuthis pyrodes, holotype, M.L. 110 mm.
NUMBER 9 7 143
. . . SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

144

PLATE 26.—A-H, Bathyteuthis berryi: A, right arm I, holotype; B, tentacular club, M.L. 19 mm;

C-F, suckers from base of arms I-IV, respectively, relative sizes maintained, holotype; o, inner

ring of buccal sucker from dorsal lappet, holotype; H, club sucker, M.L. 19 mm. I-Q, Mastigoteuthis pyrodes: I-L, largest suckers from arms I-IV respectively, relative sizes maintained, holotype; M, inner ring of club sucker, holotype; N, club sucker enlarged three times relative to arm

suckers, holotype; o, left funnel locking-cartilage, holotype; p, base of tentacular club of holotype; Q, aboral view of tentacular club of holotype; R, Mastigoteuthis dentata from off Cape

Mala, Panama (PILLSBURY 510), large sucker from arm III; s, same specimen, club sucker;

T, same specimen, inner ring of club sucker.
NUMBER 9 7
146

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 27.—A, Bathyteuthis berryi, ventral view, holotype; B

Grimalditeuthis bomplandii, ventral view, M.L. 89 mm.
NUMBER 9 7 147

PLATE 28.—A, Heliocranchia pfefferi, ventral view, M.L. 41 mm; B, Cranchia scabra, ventral

view, pattern and shape of tubercules diagrammatic, M.L. 107 mm.
148

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGV

PLATE 29.—A-I, Heliochanchia pfefferi, male, P.L. 39 mm: A, tentacular club; B, left arm I;

c, left arm II; D, largest club sucker; E, inner ring of same sucker; F-I, largest suckers from

arms I-IV, respectively, relative sizes maintained, J-Q, Cranchia scabra, male, M.L. 107 mm:

j , tentacular club; K, hectocotylized right arm IV; L, largest sucker from tentacular club; M,

enlarged sucker from tip of hectoctylus; N-Q, largest suckers from arms I-IV, respectively.

L-Q maintain relative sizes.
NUMBER 9 7 149

ope

If.0 §
150

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 30.—A, Galiteuthis pacifica, ventral view, P.L. 42

mm; B, Lechia dislocate, VELERO 8028, ventral view, P.L.

110 mm.
NUMBER 9 7 151

•

PLATE 31.—Lechia dislocate: A, Larva, "pyrgosis stage,"

P.L. 27 mm; B, mature female, P.L. 145 mm, tips of arms

III added from another specimen.
152

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 32.—Lechia dislocata: A, Immature hectocotylus of holotype; B, cartilagenous band

and tubercules, left side, P.L. 136 mm: c, right tentacular club of holotype; D, largest sucker

from arm III, female, P.L. 136 mm; E-H, largest suckers from arms I-IV, respectively, relative

sizes maintained, holotype; i, enlarged sucker from tip of hectoctylus, P.L. 137 mm; j , marginal sucker from manus, holotype, enlargement double that of arm suckers; K, medial sucker

from manus, holotype, same enlargement as arm suckers, L—s, development of eye photophores

showing lateral and ventral views of eye at each stage: L, P.L. 16 mm; M, P.L. 22 mm; N,

P.L. 25 mm; o, P.L. 35 mm; r, P.L. 44 mm; Q, P.L. 64 mm; R, P.L. 125 mm, holotype; s,

P.L. 143 mm.
NUMBER 9 7 153
154

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 33.—Galiteuthis phyllura, ventral views:

A, M.L. 79 mm; B, M.L. 33 mm; c, M.L.

170 mm.
NUMBER 9 7 155

-

.

PLATE 34.—A, Galiteuthis phyllura, tentacular club, P.L. 79 mm; B, same, tentacular club, P.L. 359 mm; c, Galiteuthis

pacifica, tentacular club, P.L. 36 mm; D, G. phyllura, larva, P.L. 13 mm; E-H, G. phyllura, P.L. 211 mm, largest

suckers from arms I-IV, respectively, relative sizes maintained; i, same specimen; inner ring of largest dactyl sucker,

enlarged four times relative to arm suckers; j , G. pacifica, largest dactyl sucker, P.L. 36 mm; K, G. pacifica, largest

sucker of arms III, P.L. 65 mm; L, G. phyllura, cartilage at funnel-mantle fusion, P.L. 173 mm; M, G. pacifica, cartilage

at funnel-mantle fusion, P.L. 65 mm.
156

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 35.—A, Vampyroteuthis infernalts, dorsal view, P.L. 50 mm; B, Japetella heathi,

lateral view.
NUMBER 9 7 157

•

PLATE 36.—A, Japetella sp., V-9661, lateral view, M.L. 72 mm; B, same specimen, radula; c, same specimen, viscera; D,

Japetella heathi, viscera, M.L. 75 mm; E, Vampyroteuthis infernalis, upper beak from Atlantic Ocean specimen, P.L. 78

mm; F, same, Californian specimen, P.L. 74 mm; G, same, Atlantic Ocean specimen, P.L. 64 mm; H, Gonatopsis borealis,

juvenile, M.L. 35 mm.
158

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 37.—Ocythoe tuberculata: A, radula, M.L. 80 mm; B, immature male, M.L. 14 mm;

c, immature female, M.L. 80 mm.
NUMBER 9 7 159

•

W

PLATE 38.—A, Octopus sp. pelagic juvenile. M.L. 19 mm; B, Alloposus mollis, female, M.L. 115 mm.

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