CLYDE F. E. ROP

RICHARD E. YOl

and GILBERT L. VC

An Illustrated

Key to the

Families of the

Order Teuthoidea

Cephalopoda)

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • 1969 NUMBER 13

SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 13

Clyde F. E. Roper, An Illustrated Key

5K?Z" to the Families of the

Order Teuthoidea

(Cephalopoda)

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Contents

Page Plate

Introduction 1

Key to the Families 3

Table (Comparison of basic characters) 5

Suborder Myopsida 4

Loliginidae 4 1A,

Pickfordiateuthidae 4 1c, D

Suborder Oegopsida 6

Gonatidae 6 2A-D

Enoploteuthidae 6 3A-C

Octopoteuthidae 7 3D

Onychoteuthidae 7 4A-C

Lepidoteuthidae 8 4D

Ctenopterygidae 8 5c

Batoteuthidae 8 6A, B

Brachioteuthidae 8 7A, B

Lycoteuthidae 9 7c, D

Histioteuthidae 9 8A, B

Bathyteuthidae 9 9A-C

Psychroteuthidae 9 8c, D

Neoteuthidae 9 10A-D

Architeuthidae 10 lie, D

Ommastrephidae 10 11A, B

Thysanoteuthidae 10 12A

Chiroteuthidae 11 13A-E

Mastigoteuthidae 11 12B, C

Joubiniteuthidae 12 6c, D

Cycloteuthidae 12 14A-E

Promachoteuthidae 12 5A, B

Grimalditeuthidae 12 15A

Cranchiidae 12 15B-D, 16A-D

Untenable Families 13

Selected Faunal Works 13

References 14

ABSTRACT

A dichotomous key to the twenty-five families of cephalopods of the order Teuthoidea

is presented. A supplementary chart of basic, external teuthoid characters is included. Representatives of each family are illustrated. The current state of systematics

within each family is briefly discussed.

Introduction

No comprehensive key to the families of the Teuthoidea has been published since Hoyle's key to the

dibranchiate Cephalopoda (1904). Hoyle's excellent

work also includes a key to the genera. Three major

keys on the suborder Oegopsida also were published

around the turn of the century. The first of these keys

was presented by Goodrich (1892) and dealt only

with the genera of oegopsids. This was followed in 1900

by Pfeffer's synopsis of the oegopsid cephalopods

which included a comprehensive key to the twelve

known families. The remaining key to appear on

oegopsids was published in Pfeffer's extensive monograph (1912) ; this was a modification of his earlier

key, but utilized external characters only. Although not

a key, Thiele's handbook (1935) includes brief diagnoses of the genera of Recent cephalopods known at

that time. This work should be used with caution, however, because some of the diagnoses are incomplete and

misleading.

Advancements in the systematics of recent cephalopods have decisively altered earlier concepts and the

former keys no longer reflect current knowledge. The

need for a modern key is particularly pressing in the

order Teuthoidea in which the number of families has

nearly doubled since the last major revision (i.e.,

Pfeffer, 1912). Furthermore, the rapid expansion in

the fields of fisheries science and biological oceanography has created a greater need by nonteuthologists

for information about cephalopods. The cephalopods

with which the programs in these fields are involved

belong primarly to the order Teuthoidea.

The present work is designed for rapid identification

of the families of the order Teuthoidea by utilization

of a dichotomous key, a supplementary chart of basic

Clyde F. E. Roper, Department of Invertebrate Zoology,

Smithsonian Institution, Washington, D.C. 20560. Richard E.

Young, University of Hawaii, Honolulu, Hawaii 96822.

Gilbert L. Voss, Institute of Marine Sciences, University of

Miami, Miami, Florida 33149.

characters, illustrations, and discussions of individual

families. We believe this is the most effective method

of presentation permitting the familial identification

of both whole and damaged specimens. This paper is

designed for use by the nonspecialist as well as the

specialist.

The dichotomous key utilizes only external characters which have been kept to a minimal number. It is

designed primarily for the identification of adult specimens; therefore, one should use caution when dealing

with larval forms. The order in which the families are

presented is not necessarily indicative of phylogenetic

relationship; some closely related groups naturally will

fall together, while others may be widely separated.

The diagrams in Figure 1 illustrate most of the morphological terms presented in the key. The ventral arms

are often modified in males (hectocotylization), and

the tips of the other arms occasionally may be modified; these modifications should not be confused with

the typical characters of the arms used in the key. Some

groups are characterized by the absence of tentacles,

but lack of tentacles may also result from injury. Injured specimens, however, invariably show stubs of the

missing tentacles.

The chart presents a brief summary of the basic

characters for each family. It should be particularly

useful if damaged specimens lack certain characters

that are utilized in the key. Representative species of

each family are illustrated, and atypical forms are

illustrated where necessary. A discussion is presented

which more fully characterizes each family, briefly

surveys the genera, and reflects the current status of

their systematics. References to recent works dealing

with the systematics of families and genera are given

where possible.

ACKNOWLEDGMENTS.—Many of the illustrations are

reproduced from recent works of the authors and were

originally drawn by Miss Constance Stolen, as was the

diagram of the basic teuthoid features; we gratefully

acknowledge her contribution. The remaining illustra1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

B.

SuckersBuccal membrane

Oegopsid eyeFunnel

Funnel-mantle fusion

Funnel locking cartilage —

Mantle locking cartilageMantle

PhotophoresFin (posteriorly concave)

Arm I

Arm I I

Dactylus

Fin (posteriorly convex)

Buccal lappet

Tail

Arm H I

Buccal suckers

Buccal connective (ventrally attaching)

Buccal connective (dorsally attaching)

Arm U

D F H

FIGURE 1.—A, A composite diagram illustrating basic teuthoid features, B, Diagram of oral

surface of brachial crown and buccal membrane. Basic types of funnel locking-cartilage:

c, -H-shaped; D, J.-shaped; E, oval with inward projecting knobs; F, simple, straight; o, subtriangular; H, oval.
NUMBER 13

tions come from published teuthological works of other from the Institute of Marine Sciences for testing and

authors. The work was partially supported by grants commenting on the key: E. S. McSweeny, G. Y. HenGB 57294 and GA 1493 from the National Science drix, E. T. LaRoe, and students of the invertebrate

Foundation. We wish to thank the following persons taxonomy class.

Key to the Families of the Order Teuthoidea

1. Eye covered by a transparent membrane (cornea) (Figure 1A) (Myopsida) 2

Eye without cornea and in open contact with seawater (Figure 1A) (Oegopsida) 3

2. Four rows of suckers on manus of tentacular clubs. Medial posterior border of fins concave

(i.e., curves posteriorly toward midline) (Figure 1A) Loliginidae (Plate 1A, B)

Two rows of suckers on manus of tentacular clubs. Medial posterior border of fins convex (i.e.,

curves anteriorly toward midline) (Figure 1A) Pickfordiateuthidae (Plate lc, D)

3. Funnel free from mantle; a funnel-mantle locking apparatus present 4 *

Funnel fused to mantle on each side; no funnel-mantle locking apparatus present (Figure 1A) .2 4

4. Funnel-mantle locking apparatus a simple, straight groove and ridge (Figure IF) ... . 5 b

Funnel-mantle locking apparatus not a simple, straight groove and ridge (Figures lc, D, E,

G, H) 18

5. Arms with hooks or with suckers in four rows on the proximal half of the ventral arms . . 6

Arms without hooks and with suckers in two rows on the proximal half of the ventral arms. 8

6. Armature of arms in two rows 7

Armature of arms in four rows Gonatidae (Plate 2A—D)

7. Tentacles present; fully developed clubs present Enoploteuthidae (Plate 3A—c)

Tentacles and clubs absent in adults although present in larvae or occasionally juveniles

(Taningia) but always with rudimentary clubs Octopoteuthidae (Plate 3D)

8. Buccal membrane connectives attach to the ventral sides of arms IV (Figure 1B) 9

Buccal membrane connectives attach to the dorsal sides of arms IV 13°

9. Hooks present on tentacular clubs (tentacles and clubs unknown in Chaunoteuthis)

Onychoteuthidae (Plate 4A-C)

Hooks lacking on tentacular clubs 10

10. Cartilaginous scales present on mantle (may be minute) (Figure 2). Tentacular clubs with

four longitudinal rows of suckers Lepidoteuthidae (Plate 4D )

Cartilaginous scales lacking. Tentacular clubs with more than four longitudinal rows of suckers

on some areas 11

11. Fins nearly as long as the mantle, supported by strong, transverse, muscular ribs. Minute

suckers present on oral surface of buccal lappets (Figure 1B) . Ctenopterygidae (Plate 5c)

Fins less than half the body length and without supporting ribs. No suckers on buccal lappets.

12

12. Tentacular clubs with six uniform rows of suckers. Long spike-like tail present (greater than

the fin length) Batoteuthidae (Plate 6A, B)

Tentacular clubs with four rows of suckers on distal portion, numerous rows on proximal

portion. No tail present Brachioteuthidae (Plate 7A, B)

13. Ventral surface of eye with a row of photophores. Buccal membrane with eight separate lappets.

Lycoteuthidae (Plate 7c, D )

No photophores on eyes. Buccal membrane with seven lappets or less 14

14. Surface of mantle, head, and arms covered with numerous photophores (usually large and

distinct) Histioteuthidae (Plate 8A, B)

Surface of mantle and head without photophores (arms may have a few photophores). . 15

15. Minute suckers present on oral surface of buccal lappets... . Bathyteuthidae (Plate 9A-C)

No suckers on oral surface of buccal lappets 16

a

 See footnote 1 in Table 1.

b

 See footnote 2 in Table 1.

e

 See footnote 3 in Table 1.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to the Families of the Order Teuthoidea—Continued

16. Many small to minute suckers (or suckers and knobs) at the proximal end of the manus . . 17

No cluster of small suckers at proximal end of the manus . . . Psychroteuthidae (Plate 8c, D)

17. Medial posterior borders of fins slightly convex. Carpal knobs in a single dorsal row or absent.

Small size Neoteuthidae (Plate 10A-D)

Medial posterior borders of fins concave. Carpal knobs in a cluster alternating with carpal

suckers. Attains gigantic size Architeuthidae (Plate lie, D)

18. Funnel locking-cartilage with a longitudinal and a transverse groove, J_-shaped or —(-shaped

(Figure ID or c) 19

Funnel locking-cartilage oval, triangular or oval with inward projecting knobs (Figures 1H,

O, E) 20

19. Funnel locking-cartilage with a longitudinal groove crossed by a transverse groove at its posterior

end, _L-shaped (Figure ID). Fins less than 60% of mantle length

Ommastrephidae (Plate 11A, B)

Funnel locking-cartilage with a longitudinal groove from which a shorter groove branches

medially, -(-shaped (Figure lc). Fins more than 80% of mantle length

Thysanotcuthidae (Plate 12A)

20. Funnel locking-cartilage oval with one or two knobs directed toward the center of the concavity

(Figure 1E) 21<*

Funnel locking-cartilage oval or subtriangular, without knobs (Figures lo, H) 22

21. Club with only four rows of suckers Chiroteuthidae (Plate 13A-E)

Club with many (more than 15) rows of minute suckers . . Mastigoteuthidae (Plate 12B, C)

22. Suckers on arms in four to six rows. Tail extremely long (greater than the mantle length). . .

Joubiniteuthidae (Plate 6c, D)

Suckers on arms in two rows. Tail short (less than half the mantle length) or absent.. . 23

23. Suckers on the tentacular clubs in four longitudinal rows. Mantle free dorsally

Cycloteuthidae (Plate 14A-E)

Suckers on the tentacular clubs in eight or more longitudinal rows. Mantle fused dorsally to

head Promachoteuthidae (Plate 5A, B)

24. Mantle free dorsally—articulates with head by ridge and groove

Grimalditeuthidae (Plate 15A)

Mantle fused dorsally with head Cranchiidae (Plates 15B-D and 16A-D)

d

 See footnote 4 in Table 1.

Suborder MYOPSIDA

This suborder contains two families of near-shore,

usually shallow water, squids, many of which form

the basis of commercial fisheries (see Fields, 1965).

Members of the Myopsida can be recognized by the

presence of corneal membranes covering the eyes.

LOLIGINIDAE sensu restricto Steenstrup, 1861

PLATE 1A, B

Presently eight genera are recognized in the family:

Loligo, Doryteuthis, Lolliguncula, Sepioteuthis, Loliolopsis, Loliolus, Alloteuthis, and Uroteuthis. Loligo

is the most common and speciose genus. Members of

the family are primarily distributed throughout the

warm waters of the world, although some species of

Loligo occur in colder regions.

The characteristic features of this family include a

straight, simple locking apparatus, seven buccal lappets each with a few small suckers, buccal connectives

that attach to the ventral borders of arms IV, four

rows of suckers on the manus, two rows of suckers on

the arms, and fins with concave posterior borders.

Photophores are present in some species.

PICKFORDIATEUTHIDAE Voss, 1953

PLATE 1C,D

This recently described family (Voss, 1953) contains a single species, Pickjordiateuthis pulchella,

which is known only from the tropical western Atlantic. It is a small species that has a mantle length

of about 13-17 mm at maturity. The family characteristics include the presence of only two rows of

suckers on the manus of the tentacular clubs; biserial

suckers on the arms; small, round fins with convex

posterior borders; buccal membranes with reduced

lappets and no suckers; buccal connectives that attach

to the ventral sides of arms IV; and a straight, simple

funnel locking apparatus. Photophores are absent.
TABLE 1.—Comparison of basic characters for all families of the Teuthoidea

Families

Funnel locking apparatus

^ , 2 1

Co I

i

1,8

Buccal connectives to

arms IV

Arms I-III

{excluding tips)

4

 I

Clubs

Loliginidae

Pickfordiateuthidae

Gonatidae

Enoploteuthidae

Octopoteuthidae

Onychoteuthidae

Lepidoteuthidae

Ctenopterygidae

Batoteuthidae

Brachioteuthidae

Lycoteuthidae

Histioteuthidae

Bathyteuthidae

Psychroteuthidae

Neoteuthidae

Architeuthidae

Ommastrcphidae

Thysanoteuthidae

Chiroteuthidae

Mastigoteuthidae

Joubiniteuthidae

Cycloteuthidae

Promachoteuthidae

Grimalditeuthidae

Cranchiidae

X

X

X

X

X

X

X

X

X

X

1

X

X

X'

X

X

X

X

X

X

X*

X

X

X

X

X

X

X

X

X

X

X

X

X

x»

X

X

X

7

 X 7

X

8

X

X"

X

X

X

X

X » X »

X

X »« X »

X» X »

X

X

X

X

X

X

X" X"

X

17

X

X

Xio

X

1 8

1

 See, however, Symplectoteuthis, under the discussion of the

Om mas trephidae.

2

 The classification "simple and straight" includes some

locking apparatuses that show considerable variation. For

example, in the Octopoteuthidae and the Histioteuthidae

the central groove is fairly broad and may curve slightly

and in Gonatopsis borealis (Gonatidae) the central groove

flares broadly at its posterior end. The homogeneity of

this classification becomes apparent when this type of locking-cartilage is contrasted with the highly specialized types

illustrated in Figure 1.

8

 This character is difficult to detect in some histioteuthids

because of the development of secondary connectives.

4

 In some mastigoteuthids the knobs are difficult to distinguish. See also Mastigoteuthis magna under discussion of the

Mastigoteuthidae.

8

 The arm characters used here refer only to the dorsal

three pairs of arms; ventral arms fiequently arc modified.

The characters of the extreme tips of the arms are also

excluded from this chart because the tips occasionally are

modified by an increase in the number of sucker rows.

• Males of Berryteuthis anonychus lack hooks. Females of this

species have hooks near the bases of arms I-III.

7

 Ctenopteryx has sucker rows that increase distally to four or

sometimes six rows.

332-752 O—«G 2

8

 The rows of suckers on the arms of Bathyteuthis are irregularly

arranged, beginning proximally with one or two rows but

increasing to three or four rows.

* Only one species, Mtsonychoteuthis hamiltoni, has hooks on

the arms.

10

 Two genera, Pholidoteuthis and Tetronychoteuthis, have four

rows of suckers on the clubs; Lepidoteuthis lacks tentacles.

11

 Most, but not all, species of Histioteulhis have more than

four rows of suckers on the club.

12

 The species of Psychroteuthis have from four to seven irregular

rows of suckers on the club.

IJ

 Young members of the Neoteuthidae have four rows of

suckers on the clubs, however, with growth numerous rows

of suckers appear at the proximal end of the manus and on

the tentacular stalk.

14

 Architeuthids have four rows of suckers on the manus and

dactylus but more than four rows in the carpal locking region.

15

 Illex has eight rows of suckers on the dactylus. Other

members of the family have four rows of suckers on the manus

and dactylus.

16

 Berryteuthis has clubs with only suckers and no hooks;

Gonatus has some hooks; Gonatopsis lacks tentacles.

17

 One genus, Pterygioteuthis, has only suckers on the clubs.

18

 Several genera, e.g., Galiteuthis and Mesonychoteuthis, have

some hooks on the clubs.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Suborder OEGOPSIDA

Species of this suborder generally inhabit the open

sea from the surface to great depths. The group is

very diverse in species and exhibits a wide variety of

forms.

The Oegopsida is distinguished from the Myopsida

primarily by the absence of a corneal membrane over

the eye. The oegopsid eye is directly bathed in sea

water. The muscular integument of the head encircles

the eye, forming an oval eyelid capable of a sphincterlike contraction. A contracted eyelid in a preserved

specimen may give the false impression that the eye

is covered by a cornea. If one is careful, however, there

should be no difficulty in distinguishing the two suborders of squids.

GONATIDAE Hoyle, 1886

PLATE 2A-D

This family is characterized by tetraserial armature

on the arms. Arm tips, however, in one species, Gonatopsis octopedatus, are modified and have up to twelve

rows of minute suckers. The two medial rows of arms

I—III consist of hooks in all species, except Berryteuthis anonychus, which has only suckers, except at the

bases of some arms in females. All members of the

family have a simple, straight funnel locking-cartilage

which may flare slightly at the posterior end; buccal

connectives that attach to the ventral borders of arms

IV; numerous irregular rows of suckers on the clubs

with additional hooks in some forms (i.e., Gonatus

species). Only one species in the family has photophores {Gonatus, new species) ; these are oval patches

located on the ventral surfaces of the eyes.

Three described genera, Gonatus, Berryteuthis, and

Gonatopsis, are known. Gonatus is characterized by at

least a single large hook in the center of the tentacular

club and only five longitudinal rows of radular teeth.

Berryteuthis is characterized by the absence of hooks

on the tentacular club and by seven longitudinal rows

of radular teeth. This genus has previously been considered a synonym of Gonatus or at best a subgenus.

However, with the recent discovery of a number of

new species of Gonatus, the generic differences are now

more clearly defined and the evidence indicates that

Berryteuthis should be considered a valid genus

(Young, in manuscript). Gonatopsis is characterized

by the complete loss of tentacles, except in the early

larval forms (Young, in press).

Species in this family are cold water forms and are

among the most abundant squids in higher latitudes.

ENOPLOTEUTHIDAE Pfeffer, 1900

PLATE 3A-C

All members of this large family possess a straight

locking apparatus; biserial armature (except occasionally at the arm tips) with at least some hooks on the

arms; tetraserial armature on the clubs (marginal

suckers of the manus may be lost with growth in some

species) ; photophores; eight buccal lappets and buccal

connectives that attach dorsally to the ventral arms

{Enoploteuthis dubia Adam, 1960, is unique in having connectives that attach dorsally to all arms). All

species, except those of the genus Pterygioteuthis,

possess hooks on the tentacular clubs.

The Enoploteuthidae is divided into three longestablished subfamilies: Enoploteuthinae, Pyroteuthinae, and Ancistrochirinae. The Enoploteuthinae is

characterized by the absence of nidamental glands,

presence of numerous, small photophores over the

suface of the mantle, head and arms, but not on the

tentacles or viscera, and fins with concave posterior

borders. This is the most speciose subfamily and contains the following genera: Enoploteuthis, Abralia, Abraliopsis, and Watasenia. The Pyroteuthinae has nidamental glands, no photophores on the surface of

the mantle, head or arms, but large photophores embedded in the tentacles and others present on the

viscera, and fins with convex posterior borders. Two

genera, Pyroteuthis and Pterygioteuthis, are included

in this subfamily. The Ancistrochirinae has nidamental

glands, a few large scattered photophores on the surface of the mantle, head, and tentacles, but not on the

arms or viscera, and huge fins with convex posterior

borders. Two genera, Thelidioteuthis and Ancistrochirus, are known, but there is a possibility that the first

is a synonym of the latter. Occasionally Abralia, Abraliopsis, and Watasenia have been split from the Enoploteuthinae as a separate subfamily, the Abraliinae.

This division, however, cannot be maintained on the

basis of present knowledge.

Enoploteuthids are primarily inhabitants of the

tropical and subtropical oceans of the world. Some

species of Pterygioteuthis, Abraliopsis, and Watasenia,

however, inhabit temperate waters as well. (Recent

works include Roper, 1966, and Voss, in manuscript.)
NUMBER 13

OCTOPOTEUTHIDAE Berry, 1912

PLATE 3D

The family is characterized by biserial hooks on the

arms (usually replaced by small biserial suckers near

the arm tips); a lack of tentacles; buccal connectives

that attach to the ventral borders of arms IV; a

simple, straight, slightly broad funnel locking-cartilage; very large fins; light organs at the tips of at

least some of the arms.

The family presently consists of two genera, Octopoteuthis and Taningia. Tenacles are present in larval forms of all species but are lost very early in

Octopoteuthis, while they remain for some time in

Taningia as small rudimentary filaments which bear

a few distal suckers. Octopoteuthis bears small spindle-shaped photophores at the tips of all eight arms,

but Taningia bears very large photophores only at

the tips of the second arms. The genus Octopodoteuthopsis PfefTer, 1912, was synonymized with Octopoteuthis by Robson (1948). Recent workers have

retained the genus Octopodoteuthopsis; however, this

genus does not differ significantly from Octopoteuthis

and the genus cannot be maintained (Young, in press).

Considerable confusion concerning the correct name

of this family and its type genus has existed from the

time of Riippell's original description of Octopoteuthis

sicula to the present day. Therefore, it is necessary to

review the nomenclatural history of the family in detail.

Riippell's description of Octopoteuthis sicula was

published in March 1844. The new generic name was

mentioned three times, always spelled Octopoteuthis.

Riippell stated that the new genus was called Octopoteuthis because it united the characters of Octopus,

Loligo, and Enoploteuthis. Regardless of the soundness

of such a relationship, Riippell's intentions are clear—

he used the stem of Octopus in forming the name.

Krohn (1845) described the species in more detail

and discussed the generic relationships. The generic

name was used only twice, in the title and in the final

sentence. Both times it was spelled Octopodoteuthis.

Krohn gave no reason for the emended spelling; he

simply stated that, following Riippell's suggestion, he

called the genus Octopodoteuthis. This is an "unjustified emendation" according to the International

Code of Zoological Nomenclature (Article 33). The

original spelling was correct, and Krohn's name is a

junior objective synonym.

Krohn added to the confusion futher in 1847 when

he changed the name of "Octopodoteuthis" to Verania

because he discovered that the young of O. sicula have

tiny tentacles—thus have 10 appendages—so the original name no longer applied to the condition of the

species. A change of this sort, of course, is untenable

(Article 18); Verania is merely a junior synonym of

Octopoteuthis.

Chun (1910) accepted the name Octopodoteuthis

and created it for a new family. Chun, however, based

the name of the family, the Veranyidae, on the genus

Veranya which he admitted was a junior synonym.

Therefore, although Chun was the first to have a concept of the family, his name is inadmissible because it

was originally based on a generic name that was not

currently valid. Berry (1912) correctly pointed out

that Chun's Veranyidae is untenable because Veranya

is an "exact synonym of Octopoteuthis." He concluded

that Octopoteuthidae (or Octopodoteuthidae if

Krohn's emendation could be accepted) must replace

Veranyidae.

Pfeffer's monograph came out in 1912 also; he

called the group Octopodoteuthinae, based on the incorrectly emended generic name. This would require

subsequent emendation to Octopoteuthinae and elevation to Octopoteuthidae. We have been unable to determine the exact date of publication of Pfeffer's monograph and, therefore, must consider it as published on

the last day of the year. The authorship of the family

name must then be attributed to Berry.

Adam (1952) correctly used Octopoteuthis and gave

a good synonymy.

ONYCHOTEUTHIDAE Gray, 1849

PLATE 4A-C

The family is characterized by tetraserial armature

generally present on the tentacular clubs (the tentacles of Chaunoteuthis are unknown), of which the

two median rows consist of hooks and the marginal

rows of suckers (in some forms the marginal rows are

absent; straight simple locking apparatus; two rows of

suckers on the arms; buccal connectives that attach to

the ventral sides of arms IV. Most members of the

family are strong, active swimmers.

Five genera are currently recogized: Onychoteuthis,

Onykia, Moroteuthis, Ancistroteuthis, and Chaunoteuthis. The generic boundaries, however, are not well

defined and the family is in need of revision.

Onychoteuthis is the most widely distributed genus

and the only genus of the onychoteuthids known to possess photophores. Onykia contains the smallest forms
8

in the family and has a number of poorly defined species in tropical seas. Moroteuthis is the giant of the

family; the largest recorded specimen had a total length

greater than four meters (Verrill, 1876). Chaunoteuthis differs from the other genera by having a gelatinous consistency.

LEPIDOTEUTHIDAE Pfeffer, 1912

PLATE 4D

Formerly only a single species, Lepidoteuthis grimaldii, was included in this family. We, however, include

two additional genera, Pholidoteuthis and Tetronychoteuthis, although a detailed analysis of the three genera

is necessary to confirm their relationships. The lack of

sufficient material has resulted in the uncertain position of these genera. Pholidoteuthis was previously

placed in its own family, the Pholidoteuthidae; Tetronychoteuthis had been included in the Onychoteuthidae, although its placement there is clearly

unwarranted.

The family Lepidoteuthidae is characterized by the

presence of distinct "scales" on the mantle (Figure 2),

buccal connectives that attach to the ventral borders of

arms IV, straight, simple funnel locking-cartilage, biserial suckers on the arms, and tetraserial suckers on the

tentacular clubs, except in Lepidoteuthis, which lacks

tentacles in the adults. Photophores are absent.

Species of Lepidoteuthis and Pholidoteuthis are

known to attain large size: 97 cm mantle length in

Lepidoteuthis (Clarke and Maul, 1962), and 78 cm

mantle length in Pholidoteuthis (Voss, unpublished).

FIGURE 2.—Cartilaginous scales on the surface of the

mantle in the Lepidoteuthidae: A, Tetronychoteuthis; B,

Pholidoteuthis.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The most characteristic feature of this family is the

existence of distinct cartilaginous "scales" that occur

only on the surfaces of the mantle. The posterior end

of the mantle, however, is devoid of "scales." The scaleless area on the dorsal side extends nearly to the anterior margin of the fins, but on the ventral side it occurs only on the posterior two-thirds of the area covered by the fins.

CTENOPTERYGIDAE Grimpe, 1922

PLATE 5C

The only genus in this family, Ctenopteryx, is readily

distinguished by its long, ribbed fins; suckers on the

lappets of the buccal membrane; eight to fourteen

longitudinal rows of small suckers on the tentacular

clubs; four to six rows of suckers on the distal half of

arms I—III; buccal connectives that attach to the ventral borders of arms IV; straight, simple, locking-cartilage; photophores in some species. A number of species

exist; however, due to the confused state of the systematics within the genus, all forms have long been

lumped under the name C. sicula.

BATOTEUTHIDAE Young and Roper, 1968

PLATE 6A, B

This family has recently been discovered in Antarctic waters. The arms bear two rows of suckers, and

the tentacles bear very elongate clubs with six longitudinal rows of suckers. The buccal connectives attach to the ventral borders of arms IV; the funnel

locking-cartilage has a simple, straight groove; the

posterior end of the body is drawn out into a long,

sharp tail. The single species, Batoteuthis scolops, is

known only from Antarctic waters.

The title page of the journal (Antarctic Research

Series) in which the description of the family first

appeared inadvertently listed the publication date as

1967. The actual date of publication was May 1968.

BRACHIOTEUTHIDAE Pfeffer, 1908

PLATE 7A, B

This family is monotypic and characterized primarily by the presence of numerous rows of small

suckers on the proximal portion of the tentacular clubs,

straight, simple funnel locking-cartilage, ventral attachment of the buccal connectives to arms IV, two

rows of suckers on the arms, and long slender bodies
NUMBER 1 3

with relatively short terminal fins. Some species of

Braehioteuthis have peculiar larvae with characteristic, long, slender necks. The family is badly in need

of a revision.

LYCOTEUTHIDAE Pfeffer, 1908

PLATE 7C, D

The family is distinguished by biserial suckers on

the arms, tetraserial suckers on the tentacular clubs,

straight, simple funnel locking-cartilage, buccal membrane with eight lappets and supports, buccal connectives that attach to the dorsal borders of arms IV, and

photophores at least on the viscera and the ventral

surfaces of eyes.

Two subfamilies are known. The Lampadioteuthinae represented by a single species, Lampadioteuthis

megaleia, is distinguished by the peculiarities of its pen

and by the presence of only four light organs on the

eye. The Lycoteuthinae has a characteristic pen and

five photophores on the ventral surface of the eye.

This subfamily has four genera: Lycoteuthis, Selenoteuthis, Nematolampas, and Oregoniateuthis. The family recently was monographed by Voss (1962).

HISTIOTEUTHIDAE Verrill, 1881

PLATE 8A, B

This monotypic family is easily distinguished by

large, anteriorly directed photophores over the surface

of the mantle, head and arms; large head with the

left eye considerably larger than the right; six or

seven buccal lappets; buccal connectives that attach

to the dorsal border of arms IV; a straight or slightly

curved and slightly broad, simple, funnel lockingcartilage; suckers on the tentacular clubs arranged in

four, or more commonly, greater than four irregular

rows; suckers on the arms biserial.

The family is being monographed by N. Voss (in

press).

BATHYTEUTHIDAE Pfeffer, 1900

PLATE 9A-C

The family is distinguished by the presence of a

straight funnel locking-cartilage, buccal connectives

that attach to the dorsal borders of arms IV, suckers

on the buccal lappets, small clubs with many rows of

minute suckers, round or paddle-like subterminal fins

(i.e., posterior margins convex), suckers on the arms

arranged in irregular rows (two rows proximally increasing to four distally), and a deep maroon color.

A single photophore is embedded at the base of each

arm I—III. This is easily apparent in young specimens

but may be obscure in adults.

The family contains a single genus, Bathyteuthis,

and three species. The most common species, B. abyssicola, is a deep sea form that has been recorded from

all major oceans and is most abundant between 1000

m and 2500 m; B. abyssicola is one of the most numerous cephalopods in the Antarctic waters. The systematics and distribution of the family have been recently studied (Roper, 1969).

PSYCHROTEUTHIDAE Thiele, 1921

PLATE 8C, D

The monotypic family is based on Psychroteuthis

glacialis described from incomplete specimens from the

stomachs of penguins and seals. Because of the inadequate description, the status of this family has been

considered doubtful. Recently additional material has

become available that confirms the validity of the

Psychroteuthidae (Young and Roper, in manuscript).

The family is characterized by buccal connectives

that attach to the dorsal borders of arms IV, straight,

simple funnel locking-cartilage, absence of photophores

on the mantle and head, a tentacular club with four

to seven rows of suckers on the manus and dactylus,

and biserial suckers on the arms.

Two species, one undescribed, are presently known

from Antarctic waters.

NEOTEUTHIDAE Naef, 1921a

PLATE 10A-D

The family is characterized by biserial suckers on the

arms, four rows of suckers on the distal part of the

manus and dactylus, and numerous small suckers on

the proximal end of the manus, buccal connectives that

attach to the dorsal borders of arms IV, straight, simple

funnel locking-cartilage, anterior fin lobes lacking and

posterior fin lobes free, the absence of photophores.

Two genera, Alluroteuthis Odhner, 1923, and Neoteuthis Naef, 1921, are known. Odhner (1923) suggested that Parateuthis tunicata Thiele, 1921, might

be the larva of the monotypic Alluroteuthis antarcticus.

Although these species appear superficially to be quite

different, new material of both species presently under

study indicates that they probably are synonymous and
10

that the older genus, Parateuthis, will, therefore, be the

valid name. Before this change can be made, however,

the problem will require further investigation.

The validity of Neoteuthis has previously been considered doubtful because of the very inadequate early

description. Recent findings, however, have confirmed

the validity of the genus and have demonstrated its

close affinity with Alluroteuthis. This information has

necessitated the submergence of the family Alluroteuthidae into the synonymy of Neoteuthidae (Young, in

manuscript).

ARCHITEUTHIDAE Pfeffer, 1900

PLATE 1lc, D

This family includes the largest of all cephalopods.

Specimens occasionally are found moribund at the

surface of the ocean, but most records are from strandings or whale stomachs.

The Architeuthidae is characterized by tetraserial

armature on the tentacular clubs with large suckers in

the medial rows of the manus and small suckers in the

marginal rows; a distinct cluster of numerous small

suckers and knobs at the proximal end of the manus;

two longitudinal rows of alternating suckers and pads

on the tentacular stalks; a straight simple funnel locking-cartilage; buccal connectives that attach to the

dorsal border of arms IV; the absence of photophores.

Many species have been named in the sole genus of

the family, Architeuthis, but they are so inadequately

described and poorly understood that the systematics

of the group is thoroughly confused. Total lengths of

nearly 60 feet and mantle lengths of nearly 17 feet (not

a complete specimen) have been recorded (summary

in Clarke, 1966).

Juveniles of species from both the Atlantic and Pacific Oceans have recently been discovered (Roper and

Young, in manuscript).

OMMASTREPHIDAE Steenstrup, 1857

PLATE 11A, B

The family is characterized by an inverted T-shaped

funnel locking-cartilage, biserial suckers on the arms,

tetraserial suckers on the tentacular clubs (except I Ilex

which has eight rows of suckers on the dactylus), buccal connectives that attach to the dorsal borders of arms

IV, photophores in some genera, and a muscular bridge

anterior to the funnel locking-cartilage which passes

from the funnel to the ventral surface of the head.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The inverted T-shaped funnel locking-cartilage

easily distinguishes this family from all others, even in

the youngest stages. One genus, Symplectoteuthis, has

the funnel and mantle cartilages fused at a single

point, but they are otherwise typical.

Three subfamilies are presently recognized: Ommastrephinae, Todarodinae, and Illicinae. The Ommastrephinae is distinguished by the presence of a

central pocket (foveola) and several side pockets in

the funnel groove, and by the presence of photophores

which are often deeply buried in the tissue of the mantle, head and arms. Five genera are present: Ommastrephes, Symplectoteuthis, Dosidicus, Hyaloteuthis,

and Ornithoteuthis. Species of these genera are strong,

powerful swimmers that are commonly found in surface waters at night. Dosidicus, the giant of the family which inhabits the Peru Current, is reported to attain a total length of about four meters (Duncan,

1941).

The Todarodinae has a foveola, but lacks side pockets in the funnel groove, and lacks photophores. Two

genera are known: Todarodes and Nototodarus. Todarodes pacificus from Japan is the basis of one of the

largest cephalopod fisheries in the world. Nototodarus

is known only from the Pacific Ocean.

The Illicinae lacks both central and side pockets in

the funnel groove and lacks photophores. The two

genera, Illex and Todaropsis, are known only from

the Atlantic Ocean. Illex illecebrosus from Newfoundland is the basis of one of the largest squid fisheries

in the Atlantic.

All species of the family in which the developmental

stages are known pass through the "rhynchoteuthis"

larval stage recognized by the fusion of the tentacles

to form a trunklike proboscis. The systematics of the

family is unstable and is presently being revised by

Roper and by Clarke.

THYSANOTEUTHIDAE Keferstein, 1866

PLATE 12A

This family is characterized by a funnel lockingcartilage with a long, narrow longitudinal groove and a

short, broad transverse groove ( —\-shaped, Figure lc);

long broad fins in the shape of a rhombus that extend

nearly the full length of the mantle; nuchal-mantle lock

with two distinct knobs that fit into opposing pits;

extremely long, cirrate trabeculae on the arms; buccal connectives that attach to the ventral borders of

arms IV; two rows of suckers on the arms and four
NUMBER 13 11

rows of suckers on the manus; the absence of

photophores.

Two genera, Thysanoteuthis and Cirrobrachium,

are currently included in the family. Cirrobrachium

is known only from a few larval forms and a single,

larger fragmentary specimen; it is distinguished by

long, slender, free cirri (trabeculae) on the arms. So

little is known about the characters of the two nominal

species of Cirrobrachium that the genus is in question.

The monotypic species, Thysanoteuthis rhombus, attains a large size of at least 80 cm mantle length and

19 kg weight (Nishimura, 1966) and is a very powerful

swimmer.

CHIROTEUTHIDAE Gray, 1849

PLATE 1 3A-E

This family is often divided into two subfamilies:

the Chiroteuthinae and the Mastigoteuthinae. We

consider these groups to be closely related, but we

believe that they are distinct enough to each merit

familial ranking.

The family Chiroteuthidae, although distinctive, is

rather difficult to characterize because of secondary

modifications in certain genera (e.g., Valbyteuthis,

Chiropsis). The funnel locking-cartilage distinguishes

the Chiroteuthidae from all families except the Mastigoteuthidae. The locking apparatus is oval and bears

one or two distinct knobs that project toward the center

of the cavity. When two knobs are present, one occurs

on the medial side and the other on the posterior side

of the lock. The arms bear biserial suckers, except occasionally at the tips; the tentacles have tetraserial

suckers; the buccal connectives attach to the ventral

borders of arms IV; photophores are usually present.

Three genera are presently recognized: Chiroteuthis, Chiropsis, and Valbyteuthis. Several other genera

have been described, but their status is uncertain.

Chirosoma regnardi Joubin, 1912, was named in a

brief note on a collection of cephalopods from the eastern North Atlantic. An inadequate preliminary diagnosis was given, based on a single character (rows of

suckers), but a description of the species never appeared. Joubin (1924) discussed the species of the

family Chiroteuthidae (as Chiroteuthinae) but did

not mention his own species Chirosoma regnardi. Apparently he no longer recognized the genus or species

as distinct taxa. Clearly, this form must be considered

a genus and species dubia. Berry (1920) described a

new genus, Chiroteuthoides, based on a larval specimen from the western Atlantic. The affinity of this

taxon with the Chiroteuthidae is doubtful. Bigelowia

MacDonald and Clench (1934) possesses the typical characteristics of Chiroteuthis and clearly is a synonym of this genus, although, until now, it has never

formally been synonymized.

Most chiroteuthids belong in the genus Chiroteuthis

and possess very elongate tentacles, distinctive clubs

with suckers arranged in four rows on long, slender

stalks, greatly enlarged ventral arms, and abundant

photophores. Chiroteuthis larvae are noted for their

large size and exotic appearance.

Chiropsis has a single species, C. mega, known only

from the holotype. It is similar in appearance to

Chiroteuthis except for the short, broad, blunt ventral

arms.

Valbyteuthis differs considerably from Chiroteuthis

and Chiropsis; a recent study, however, has revealed

a striking similarity between the species of Valbyteuthis and certain larval forms ("doratopsis" type) of

Chiroteuthis (Roper and Young, 1967). Members of

the genus are easily recognized by the lack of a funnel

valve and by the presence of an oval funnel lockingcartilage bearing only a posterior knob.

MASTIGOTEUTHIDAE Verrill, 1881

PLATE 12B, C

The family Mastigoteuthidae is characterized by

an oval funnel locking-cartilage with inward projecting knobs, as in chiroteuthids, but generally the

posterior knob is weakly developed and occasionally the

medial knob is also poorly developed giving the locking-cartilage an almost oval appearance. The long,

whiplike tentacles that bear many hundreds of minute,

often microscopic, suckers in many rows (more than

15) are also very characteristic of the family. Suckers

are arranged biserially on the arms; the buccal connectives attach to the ventral borders of arms IV; the

fins generally are large; the ventral arms are commonly greatly enlarged. Often numerous, small photophores are embedded in the integument of the mantle,

fins, head, and arms. The funnel locking-cartilage of

Mastigoteuthis magna is somewhat peculiar in having

lobes that project toward the concavity from both the

medial and the lateral sides, giving the groove of the

lock a flask-shaped appearance. The Mastigoteuthidae

is closely related to the Promachoteuthidae, but, among

other characters, the Promachoteuthidae is easily distinguished by the dorsal fusion of the mantle to the
12

head, while the Mastigoteuthidae has a free mantle

margin.

Two genera, Mastigoteuthis and Echinoteuthis, are

included in the family. Echinoteuthis, represented

only E. danae Joubin, 1933, has not been reported

since its original description. It is distinguished by

numerous, small, cartilaginous tubercles over the surface of the mantle, head, funnel, and arms. Fifteen

nominal species of Mastigoteuthis are known, several

of which have been described from single multilated

specimens. The species of Mastigoteuthis are common

inhabitants of the deep sea.

JOUBINITEUTHIDAE Naef, 1922

PLATE 6C, D

This family is very distinctive and is easily recognized by the extremely elongate, slender arms which

have six rows of suckers on each arm I—III and four

rows of suckers on each arm IV. The tentacular clubs

are elongate, compressed, and bear eight to ten longitudinal rows of small suckers; the funnel lockingcartilage is oval; the buccal connectives attach to the

ventral borders of arms IV; an elongate, thread-like

tail that is longer than the length of the mantle is

present; photophores are absent.

A single species, Joubiniteuthis portieri, is known.

A recent review of the family has shown that Valdemaria danae, a species previously included in the Mastigoteuthidae, is synonymous with Joubiniteuthis portieri (Young and Roper, 1969b).

CTOLOTEUTHIDAE Naef, 1923

PLATE 14A-E

The Cycloteuthidae is characterized by buccal connectives that attach to the ventral borders of arms IV;

arms with biserial suckers; a tentacular club with tetraserial suckers; a compact, expanded club with a welldefined manus and dactylus, but an ill-defined carpus;

a subtriangular funnel locking-cartilage; fins (including tail when present) greater than 70% of the mantle

length in adults; the presence of photophores.

Naef (1923) established the Cycloteuthinae as a

subfamily of the Onychoteuthidae based on a previously described genus, Cycloteuthis Joubin, 1919,

that contains a single species, C. sirventi. Recently

Young and Roper (1969a) elevated the Cycloteuthinae

to familial status and included in it another genus,

Discoteuthis.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

C. sirventi has a prominent tail and four rows of

equal-sized suckers on the manus of the tentacular

club. Species of Discoteuthis lack a tail and have

greatly enlarged medial suckers on the manus of the

tentacular club.

PROMACHOTEUTHIDAE Naef, 1912

PLATE 5A, B

The Promachoteuthidae is known from only a few

juvenile specimens; however, its very distinctive features make it easy to recognize. The mantle is fused

dorsally to the head in the nuchal region; the eyes are

greatly reduced; the funnel locking-cartilage is oval;

the tentacular club has numerous longitudinal rows

(more than eight) of minute suckers; the buccal connectives attach to the ventral borders of arms IV;

the arms have biserial suckers; photophores are absent.

The eye openings are greatly reduced in size corresponding to the small eyes and are difficult to detect.

In the past, the misinterpretation of this condition resulted in the placement of the Promachoteuthidae in

the Myopsida. Recently the family has been removed

from the Myopsida and placed in the Oegopsida

(Roper and Young, 1968—inadvertently listed as

1967 in journal). Promachoteuthis, the only genus in

the family, contains two species, P. megaptera from the

Pacific near Japan and an unnamed species from the

waters off southern Chile.

GRIMALDITEUTHIDAE Pfeffer, 1900

PLATE 15A

The Grimalditeuthidae is easily recognized by the

fusion of the funnel and mantle locking-cartilages

while the dorsal mantle-nuchal locking apparatus is

free. Tentacles are lacking; the arms have biserial suckers; buccal connectives attach to the ventral borders

of arms IV. An extremely long tail bears an accessory

fin; however, this structure is delicate and may be

broken off during capture. Photophores may be present at the tips of the arms.

A single genus, Grimalditeuthis, is known. Two nominal species have been described but are probably

synonymous.

CRANCHIIDAE Prosch, 1849

PLATES 15B-D; and 16A-D

The Cranchiidae contains a great diversity of species

which exhibit a wide variety of basic characters. One
NUMBER 13

prominent character, however, easily distinguishes all

members of the group: the mantle is fused to the head

in the nuchal region and to the funnel at its two posterolateral corners. Buccal connectives attach to the

ventral borders of arms IV; the armature of the clubs

generally is in four longitudinal rows; the arms generally have biserial suckers; photophores are present.

The numerous cranchiid genera are divided into

two subfamilies, the Cranchiinae and the Taoniinae.

The Granchiinae is characterized by cartilaginous strips

bearing tubercles on the ventral surface of the mantle

that originate at the funnel-mantle fusions and extend

posteriorly, the lateral fusion of the funnel to the ventral surface of the head in the adult stage, and one or

more rows of small, round photophores on the eye.

This subfamily contains the following genera: Cranchia, Liocranchia, Leachia, and Drechselia. Pyrgopsis

generally has been regarded as a separate genus; however, a complete series of the developmental stages of a

new species of Leachia has shown that young specimens

possess the generic characters of Pyrgopsis and that this

genus must fall into the synonymy of Leachia (Young,

in press). The genus Egea was previously placed in

this subfamily. One of us (G.L.V.) has recently examined the type and found that there are no cartilaginous

strips on the mantle contrary to what Joubin described

and that the genus clearly belongs in the Taoniinae.

Liguriella has also been placed in the Cranchiinae but

it is known only from larval specimens and its true affinities are uncertain. Clarke (1966) erroneously

placed Crystalloteuthis, a member of the Taoniinae,

in this subfamily.

The Taoniinae is characterized by the absence of

cartilaginous strips that extend posteriorly from the

funnel-mantle fusions, by the presence of a funnel that

is free laterally, and by the presence of one to three

generally crescent-shaped photophores on the eyes.

This subfamily contains numerous and diverse species

that are arranged in many ill-defined genera. In addition, many of the genera are based on larval forms

(e.g., Fusocranchia, Teuthowenia, and Taonidium);

therefore, a listing of the nominal genera would serve

no useful purpose.

Mesonychoteuthis, represented only by M. hamiltoni, has large hooks on both the tentacles and arms;

it attains a total length of at least 3J4 meters (Clarke,

1966). A related genus, Galiteuthis, has large hooks

only on the tentacular clubs. Taonius has a few greatly

enlarged teeth, closely approximating hooks, that arise

333-752 O—69 3

13

from the sucker rings on the tentacular clubs. The systematic position of Ascocranchia has been uncertain

because of the atypical eye photophores in the only

species. However, the close similarity of the peculiar

attachment of the paddle-like fins and the large funnel

with those of Helicocranchia indicates that Ascocranchia belongs to the Taoniinae.

Untenable Families

PARATEUTHIDAE JOUBIN, 1919.—Clarke (1966) reestablished the family Parateuthidae, and it is therefore necessary to make some comments concerning it.

The family Parateuthidae was originally erected by

Joubin (1919) to include the genera Lycoteuthis and

Cycloteuthis. Not only is the familial name invalid

since it is not based on a type genus, but also these

genera do not form a natural group. The family Parateuthidae has been rejected by teuthologists since it

was first established. Thiele (1921) described the genus

Parateuthis based on a larval oegopsid captured in the

Antarctic Ocean; this genus has no connection whatsoever with Joubin's Parateuthidae. Because of its uncertain position, Parateuthis has never been placed in

any family, although Odhner (1923) thought it might

be a larval Alluroteuthis. Larvel specimens of Alluroteuthis in the collections of the I.M.S. indicate that

this, indeed, probably is the case (see page 9).

The family Parateuthidae is invalid and the four

genera that Clarke includes in it (Cycloteuthis, Psychroteuthis, Parateuthis, and Alluroteuthis) do not

form a natural group (except the last two as noted

above).

ALLUROTEUTHIDAE ODHNER, 1923.—The Alluroteuthidae has been synonymized with the Neoteuthidae. See page 9 for discussion.

PHOLIDOTEUTHIDAE ADAM, 1950.—The Pholidoteuthidae has been synonymized with the Lepidoteuthidae.

See page 8 for discussion.

VALBYTEUTHIDAE JOUBIN, 1931.—The Valbyteuthidae has been synonymized with the Chiroteuthidae.

See page 11 for discussion.

VERANYIDAE CHUN, 1910.—The Veranyidae was

incorrectly established and has been replaced by Octopoteuthidae. See page 7 for discussion.

Selected Faunal Works

Systematic works of broad general application include Chun (1910), Pfeffer (1912), and Naef (1923).
14

The following brief list is presented as a preliminary

guide to the literature dealing with the teuthoid fauna

of local areas:

North Sea—Grimpe, 1925; Jaeckel, 1958

Northwestern Atlantic—Verrill, 1882

Mediterranean—Mangold-Wirz, 1963

Gulf of Mexico—Voss, 1956

West Africa—Adam, 1952

South Africa—Voss, 1967

Indian Ocean—Adam, 1939

Australia—Berry, 1918

New Zealand—Dell, 1952

Philippines—Voss, 1963

Japan—Sasaki, 1929

Hawaii—Berry, 1914

California—Young, in press

References

(Including the works in which the familial names were first

introduced.)

Adam, W.

1939. The Cephalopoda in the Indian Museum, Calcutta.

Records of the Indian Museum, 41(1): 61-110.

1950. Un Cephalopode Nouveau: Pholidoteuthis boschmai gen. et sp. nov. Koninklijke Nederlandse

Akademie van Wetenschappen, 53(10): 1-8.

1952. Cephalopodes. Resultats Scientifiques Expedition

Oceanographique Beige Atlantique Sud, 3(3): 1—

142.

1960. Cephalopoda from the Gulf of Aqaba. Number

16 in "Contributions to the Knowledge of the Red

Sea." Bulletin of the Sea Fisheries Research Station,

Haifa, 26:1-27.

Berry, S. S.

1912. Some Necessary Changes in Cephalopod Nomenclature. Science, 36(932): 643-646.

1914. The Cephalopoda of the Hawaiian Islands. Bulletin

of the Bureau of Fisheries, Washington, 32:255-

362.

1918. Report on the Cephalopoda Obtained by F.I.S.

Endeavour in the Great Australian Bight and

Other Southern Australian Localities. Biological

Results of the Fishing Experiments by Endeavour,

1909-1914, 4(5):203-298.

1920. Preliminary Diagnoses of New Cephalopods from

the Western Atlantic. Proceedings of the United

States National Museum, 58(2335): 293-300.

Chun, C.

1910. Die Cephalopoden, I: Oegopsida. Wissenshaftliche

Ergebnisse der Deutschen Tiefsee-Expedition, Valdivia, 18( 1): 1-410; atlas of 61 plates.

Clarke, M. R.

1966. A Review of the Systematics and Ecology of

Oceanic Squids. Advances in Marine Biology,

4:91-300.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

1967. A Deep-Sea Squid, Taningia danae Joubin, 1931.

Symposium of the Zoological Society of London,

19:127-143.

Clarke, M. R., and G. E. Maul

1962. A Description of the "Scaled" Squid Lepidoteuthis grimaldi Joubin, 1895. Proceedings of the

Zoological Society of London, 139(1) :97—118.

Dell, R. K.

1952. The Recent Cephalopoda of New Zealand. Dominion Museum Bulletin, 16:1-157.

Duncan, D. D.

1941. Fighting Giants of the Humboldt. National Geographic Magazine, 79:373-400.

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.

Goodrich, E. S.

1892. Note on a Large Squid (Ommastrephes pteropus

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of the United Kingdom, 2:314-321.

Gray, J. E.

1849. Catalogue of the Mollusc a in the Collection of the

British Museum: I, Cephalopoda Antepedia. 164

pages. London.

Grimpe, G.

1922. Systematische Ubersicht der europaischen Cephalopoden. Sitzungsberichte der Naturforschenden

Gesellshaft 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.

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. A Diagnostic Key to the Genera of Recent Dibranchiate Cephalopoda. Memoirs and Proceedings of

the Manchester Literary and Philosophical Society,

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Jaeckel, S. G. A., Jr.

1958. Cephalopoden. Die Tierwelt der Nord und Ostsee,

37(9 b,):479-723.

Joubin, L.

1900. Cephalopodes Provement des Campagnes de la

Princesse-Alice (1891-1897). Risultats des Campagnes Scientifiques du Prince de Monaco, 17:1—

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1912. Sur les Cephalopodes Captures en 1911 par S.A.S.

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NUMBER 13

1924. Contribution a l'etude des Cephalopodes de l'Atlantique Nord. 4e Series. Re suit at s des Campagnes

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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

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13(1):1-49.

Keferstein, W.

1866. Cephalopoda. In Bronn, Die Klassen und Ordnungen Tierreichs, 3(2): 1307-1464.

MacDonald, R., and W. J. 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.

Mangold-Wire, K.

1963. Biologie des Cephalopodes Benthiques et Nectoniques de la Mer Catalane. Vie et Milieu, Supplement, 13:1-285.

Naef, A.

1912. Teuthologische Notizen. I. Die Familien der

Myopsiden. Zoologischer Anzeiger, 39(7): 241-244.

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542.

1921b. Die Cephalopoden. Fauna und Flora Neapel,

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1922. Die Fossilen Tintenfische. 322 pages. Jena: Gustav Fischer.

1923. Die Cephalopoden: Systematik. Fauna und Flora

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1900. Synopsis der oegopsiden Cephalopoden. Mitteilungen Naturhistorischen Museum in Hamburg (2

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15

1908. Die Cephalopoden. Nordisches Plankton, 2(pt. 9,

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Roper, C. F. E.

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Redescription of the Type Species, E. leptura

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Roper, C. F. E., and R. E. Young

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(Cephalopoda: Oegopsida). Proceedings of the

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16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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

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1881b. Report on the Cephalopods, and on Some Additional Species Dredged by the U.S. Fish Commission Steamer "Fish Hawk," During the Season

of 1880. Bulletin of the Museum of Comparative

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Coast of America. Report of the United States

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Voss, G. L.

1953. A New Family, Genus and Species of Myopsid

Squid from the Florida Keys. Bulletin of Marine

Science of the Gulf and Caribbean, 2(4): 602-609.

1956. A Review of the Cephalopods of the Gulf of Mexico. Bulletin of Marine Science of the Gulf and

Caribbean, 6(2): 85-178.

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Atlantic, I: The Family Lycoteuthidae. Bulletin

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Voss, N. A.

MS. A Monograph of the Cephalopoda of the North

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Atlantic. The Family Cycloteuthidae (Suborder

Oegopsida). Smithsonian Contributions to Zoology,

number 5:1-24.

1969b. A Monograph of the Cephalopoda of the North

Atlantic. The Family Joubiniteuthidae (Suborder

Oegopsida). Smithsonian Contributions to Zoology,

number 15:1-10.

MS. Rediscovery of the Family Psychroteuthidae from

Antarctic Waters.
NUMBER 13 17

PLATE 1.—Loliginidae: A, Loligo, ventral view; B, Loligo, tentacular club. Pickfordiateuthidae:

c, Pickfordiateuthis, dorsal view (from Voss, 1953); D, Pickfordiateuthis, tentacular club (from

Voss, 1953).
18

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

iooooooc

PLATE 2.—Gonatidae: A, Gonatus, ventral view (from Young, MS); B, Gonatus, tentacular club

(from Young, MS); c, Berryteuthis, tentacular club (from Pearcy and Voss, 1963); D, Gonatopsis, ventral view (from Young, MS).
NUMBER 13 19

PLATE 3.—Enoploteuthidae: A, Abraliopsis, ventral view (from Young, MS); B, Abraliopsis,

tentacular club (from Young, MS); c, Pyroteuthis, ventral view (from Young, MS). Octopoteuthidae: D, OctopotetUhis, ventral view.
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 4.—Onychoteuthidae: A, Onychoteuthis, ventral view (from Pfeffer, 1912); B, Onychoteuthis, tentacular club (from Young, MS); c, Chaunoteuthis, ventral view (from Pfeffer, 1912).

Lepidoteuthidae: D, Pholidoteuthis, dorsal view (from Voss, 1956).
NUMBER 13 21

PLATE 5.—Promachoteuthidae: A, Promachoteuthis, ventral view (from Roper and Young,

1968); B, Promachoteuthis, tentacular club (from Roper and Young, 1968). Ctenopterygidae:

c, Ctenopteryx, ventral view (from Joubin, 1900).
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

V

Ctt

PLATE 6.—Batoteuthidae: A, Batoteuthis, ventral view (from Young and Roper, 1968); B,

Batoteuthis, tentacular club (from Young and Roper, 1968). Joubiniteuthidae: c, Joubiniteuthis,

tentacular club (from Young and Roper, 1969b); D, Joubiniteuthis, dorsal view (from Young and

Roper, 1969b).
NUMBER 13 23

PLATE 7.—Brachioteuthidae: A, Brachioteuthis, ventral view; B, Brachioteuthis, tentacular club.

Lycoteuthidae: c, Lycoteuthis, ventral view; D, Lycoteuthis, tentacular club. (All from

Chun, 1910).
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 8.—Histioteuthidae: A, Histioteuthis, ventral view (from Young, MS); B, Histioteuthis,

tentacular club (from Young, MS). Psychroteuthidae: c, Psychroteuthis, ventral view (from

Young and Roper, MS); D, Psychroteuthis, tentacular club (from Young and Roper, MS).
NUMBER 13 25

PLATE 9.—Bathyteuthidae: Bathyteuthis: A, Dorsal view; B, oral view or arms and buccal

membrane; c, tentacular club. (All from Roper, 1969.)
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 10.—Neoteuthidae: A, Alluroteuthis, ventral view; B, Alluroteuthis, tentacular club; c,

Neoteuthis, dorsal view (from Young, MS); D, Neoteuthis, tentacular club (from Young, MS).
NUMBER 13 27

PLATE 11.— Onunastrephidae: A, Ommastrephes, ventral view (from Pfeffer, 1912); B,

Symplectoteuthis, tentacular club (from Young, MS). Architeuthidae: c, Architeuthis, dorsal

view (redrawn from Verrill, 1882); D, Architeuthis, tentacular club (from Verrill, 1882).
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 12.—Thysanoteuthidae: A, Thysanoteuthis, dorsal view. Mastigoteuthidae: B, Mastigoteuthis, ventral view (from Chun, 1910); c, Mastigoteuthis, ventral view (from Young, MS).
NUMBER 13 29

*.„ m

f>

PLATE 13.—Chiroteuthidae: A, Chiroteuthis, ventral view (from Young, MS); B, Chiroteuthis,

tentacular club (from Young, MS); c, Valbyteuthis, ventral view (from Roper and Young,

1967); D, Valbyteuthis, tentacular club (from Roper and Young, 1967); E, ValbyteuthUj ventral

view (from Young, MS).
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 14.—Cycloteuthidae: A, Cycloteuthis, tentacular club; B, Cycloteuthis, visceral photophore; c, Cycloteuthis, ventral view; D, Discoteuthis, tentacular club; E, Discoteuthis, ventral

view. (All from Young and Roper, 1969a.)
NUMBER 13 31

PLATE 15.—Grimalditeuthidae: A, Grimalditeuthis, ventral view (from Joubin, 1900). Cranchiidae: B, Helicocranchia, ventral view (from Young, MS); c, Galiteuthis, tentacular club

(from Young, MS); D, Bathothauma, dorsal view (from Chun, 1910).
32

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 16.—Cranchiidae: A, Cranchia, ventral view; B, Leachia, ventral view; c, Leachia,

tentacular club; D, Galiteuthis, ventral view. (All from Young, MS.)
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