RICHARD E. TOUNG A Monograph of the

LYDE F. E. ROPER ^ r 7 l r ' 1

Cephalopoda of the

North Atlantic:

The Family

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • 1969 NUMBER 5

SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 5

Richard E. Young A Monograph of the

and Clyde F. E. Roper ^ 1 1 -• r •»

Cephalopoda ot the

North Atlantic:

The Family

Cycloteuthidae

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Contribution No. 1005 from the Institute of Marine Sciences, University of Miami, and

Paper No. 64 from the Dana Oceanographical Collections

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

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ABSTRACT

A new family of oegopsid cephalopods, Cycloteuthidae, is erected as a result of the

elevation of Naef's subfamily Cycloteuthinae. Cycloteuthis sirventi Joubin, 1919, is

redescribed based on new material from the Atlantic Ocean. A new genus, Discoteuthis,

and two new species, D. discus and D. laciniosa, are described from the Atlantic Ocean.

Larvae of C. sirventi and D. laciniosa are described. The distributions of the species of

cycloteuthids in the Atlantic and Indian Oceans are presented. The relationships of

the family, genera, and species are discussed.

Among the cephalopods captured during the 1964

cruise of the University of Miami, Institute of Marine

Sciences' RV John Elliot Pillsbury to the Gulf of

Guinea, one specimen presented a particular problem

in identification even at the familial level. The specimen bore some resemblance to Joubin's poorly known

Cycloteuthis sirventi on which no additional material

has been reported since the original description of the

species in 1919. A search through the extensive cephalopod collections at the Institute of Marine Sciences

yielded a number of specimens that included not only

the new form, but also Cycloteuthis sirventi. A study

of these specimens, which are primarily from collections of IMS vessels and the RV Dana Expeditions,

revealed that the species represented by the Gulf of

Guinea specimen indeed showed relationship with

Cycloteuthis sirventi but was generically distinct from

it. After further investigation, a second new species

was discovered to be referable to the new genus, Discoteuthis. The present material of Cycloteuthis and

Discoteuthis, although not great in volume, has provided sound information to confirm that these genera

form a taxonomic unit sufficiently distinct from other

oegopsid familial groups to necessitate the erection of

a separate family, the Cycloteuthidae.

The coauthors wish to thank M. R. Clarke for the

loan of a Discovery specimen, E. Bertelsen for the Dana

material, and G. E. Maul for the Madeira specimens.

The remaining specimens were taken on the Institute

of Marine Sciences' research vessels Pillsbury and

Gerda and during Ocean Acre cruises. We are also

grateful to G. L. Voss for his support and for his comments on the manuscript. The illustrations were preRichard E. Young, Institute of Marine Sciences, University

of Miami, Miami, Florida 33149. Clyde F. E. Roper, Division

of Mollusks, Smithsonian Institution, Washington, D.C.

20560.

pared by Miss Constance Stolen, to whom we are

greatly indebted. The work was supported primarily

by research grants (GB-3808, GB-2204, GB-5729Y)

from the National Science Foundation and the National Geographic Society-University of Miami DeepSea Biology Program.

DEFINITIONS.—The following definitions are

designed for the Cycloteuthidae and do not necessarily

apply to other groups.

Mantle length (ML): The length of the mantle measured

along the dorsal midline from the free anterior margin

to the extreme posterior tip of the body. In the Cycloteuthidae, therefore, the mantle length is equal to the

pen length.

Tail: The posterior extension of the body beyond the point

where the posterior margin of the fins turns abruptly

posteriorly near the mantle.

Fin length (FL) : The length from the anterior extension of

the fin lobes to the extreme posterior tip of the body,

measured along the middorsal line of the mantle.

Historical Resume

Joubin (1919) described a new genus and species,

Cycloteuthis sirventi, and indicated that it was closely

related to Lycoteuthis diadema; he placed these two

genera in a new family, the Parateuthidae, which he

considered to be intermediate between the Enoploteuthidae and the Histioteuthidae. Since a familial

name must be based on the name of the type genus,

Joubin's name is invalid. Independently, Thiele (1921)

erected the genus Paratcuthis based on a larval

oegopsid captured in the Antarctic Ocean (he did not

assign it any oegopsid family) ; this genus, however, has

no connection with Joubin's Parateuthidae. Naef

(1922, p. 298) recognized that Joubin's family was not

a natural grouping and transferred Lycoteuthis to the

Enoploteuthidae and Cycloteuthis to the Onychoteu1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

thidae. Shortly afterward, Naef (1923, p. 312) established for Cyloteuthis a separate subfamily, the

Cycloteuthinae. He also indicated (p. 266) that

Cyloteuthis, although an onychoteuthid, shares characters with the Enoploteuthidae.

Joubin (1924) repeated his earlier description and

retained the position of Cycloteuthis in his Parateuthidae, apparently unaware of Naefs changes and

Thiele's description of Parateuthis.

Although Grimpe (1925) listed Cycloteuthis in the

Onychoteuthidae, he did not maintain Naefs subfamily and stated that the position of the genus was still

uncertain.

Thiele (1935) gave a brief diagnosis of the genus

and included it in the Onychoteuthidae, apparently

also rejecting Naefs subfamily.

Clarke (1966) resurrected Joubin's invalid family,

Parateuthidae, in which he placed four unrelated

genera, including Cycloteuthis; he gave no explanation for this unwarranted action.

Family CYCLOTEUTHIDAE Naef, 1923

PARATEUTHIDAE Joubin, 1919, p. 2 [in part; refers to

Cycloteuthis only].

CYCLOTEUTHINAE Naef, 1923, p. 312.

DIAGNOSIS.—Buccal connectives attach to ventral

borders of arms IV; arms with biserial suckers; tentacular club with tetraserial suckers; club compact, expanded with well-defined manus and dactylus, ill-defined carpus; funnel locking-cartilage subtriangular;

fins large in adults (over 70 percent of mantle length) ;

photophores present.

Genus Cycloteuthis Joubin, 1919

Cycloteuthis Joubin, 1919, p. 1.—Naef, 1922, p. 298; 1923,

pp. 266, 809.—Thiele, 1935, p. 965.

DIAGNOSIS.—Tail present; gladius only slightly

thickened; conus hollow, elongate; suckers in a transverse series on manus nearly equal in size; large photophore on ink sac; a series of small photophores on eye;

fins large but not equal to the mantle length in adults.

TYPE-SPECIES.—Cycloteuthis sirventi Joubin, 1919-

By monotypy.

Cycloteuthis sirventi Joubin, 1919

PLATES 1; 2; 7E, F, H, K ; 8c, F ; 9c—u

Cycloteuthis sirventi Joubin, 1919, p. l.figs. 1-2.—Naef, 1923,

p. 312.—Joubin, 1924, p. 53, pi. 7: figs. 3-6; pi. 8: figs.

1_11.—Grimpe, 1925, pp. 96, 103.—Thiele, 1935, p.

965.—Clarke, 1966, p. 161.

DESCRIPTION.—The redescription of Cycloteuthis

sirventi is based primarily on the male specimen with

a ML of 42 mm (Gerda 212). A clear indication is

given when other specimens are discussed. Table 1 lists

the measurements of available specimens.

The mantle is conical and tapers posteriorly to a

sharp point (Plates 1A, 2C-D) . The mantle wall is thin

and invaded by gelatinous connective tissue between the

inner and outer layers of muscle. Posteriorly the muscular portion of the mantle terminates in advance of

the posterior margin of the fins at the beginning of the

conus of the pen. In the largest specimen the mantle

width is about one-third of the mantle length. The

mantle narrows relatively with growth; smallest specimens have mantle widths about 50 percent of the

mantle length.

Sex

1 cf

1 cf

1 cf

1 cf

1 juv.

1 L '

3 L

4 L

1 L

1

1

ML, mm

42

41

27

17

15

10

8.5

4-9

8

head only

head only

Station number

Gerda 212

Acre 2-2

Dana 829

Dana 4017 VII

Dana 1172 II

Dana 1157 III

Dana 1030

Dana 830

Dana 1157 XII

Dana 1157 XII

Dana 4003 IX

List of Material

Location

23°32'N 82°37.5'W

32°26'N 63°44'W

25°O7'N 19°20'W

29°13'N 14°12'W

07°22'N 46°51'W

21°57'N 22°58'W

35°31'N 55°58'W

24°44'N 22°32'W

21°57'N 22°58'W

21°57'N 22°58'W

08°26'S 15°H'W

20

6

13

27

14

27

1

15

27

27

9

Date

I 1964

III 1968

IV 1920

III 1930

XI 1921

X 1921

XII 1913

IV 1920

X 1921

X 1921

III 1930

Estimated

depth, m

650

140

100

200

200

100

10-400

100

200

200

100

1

 L refers to larval specimens.
NUMBER 5

TABLE 1.—Measurements (in mm) of Cycloteuthis sirventi

Characters

Sex

Mantle length

Mantle width

Head length

Head width

Fin length

Fin width

Arms I

II

III

IV

Tentacle length

Club length

Gerda

212

&

42

14

22

13

34

37

26

36

37

36

70

15

Dana

829

d"

27

11

10

11

19

26

16

22

22

21

40

11

Dana

1172 II

juv.

15

7

6

6

6.5

8.5

5.5

7.5

7.5

7.5

26

4

Dana

1157 III

larva

10

4.5

3.0

4 . 2

3 . 0

5.2

2.5

5.0

5.0

4 . 5

11

3 . 2

Dana

1030

larva

8.5

4 . 0

2 . 2

4 . 0

2 . 0

5.0

1.5

3.2

3 . 0

3 . 0

7.2

2.5

Dana

1030

larva

8.5

-

2.5

3.8

2 . 0

5.0

1.8

3.6

3.6

2.8

8.0

2.2

Dana

1030

larva

8.5

4 . 0

2.5

4 . 0

1.8

4 . 2

2.0

3.5

3.0

2.0

7.0

2.0

Dana

1157

XII

larva

8.0

4.0

4.0

5.0

3.5

7.0

2.0

5.5

5.5

5.0

13.0

3.2

Dana

1157

XII

larva

-

_

-

5.0

-

-

-

-

-

-

11.0

3.2

Acre

2-2

c?

41

14

18

15

33

40

23

33

-

34

64

16

The fins are long, about 80 percent of the mantle

length, and broad, about 90 percent of the mantle

length. The fins are very muscular and elliptical, except

posteriorly where they are drawn out along the lateral

sides of the tail. The anterior fin lobes are only slightly

produced. The fins are separated by a narrow gap for

most of their length and are not fused in the dorsal

midline. The bases of the fins insert directly on the

shell sac of the gladius.

The funnel is small and reaches just past the posterior edge of the eyelids. The funnel locking-cartilage

on either side of the funnel is approximately triangular

in shape, with the lateral side nearly parallel to the body

axis and the anterior side transverse to the body axis

(Plate 7E). The groove of the locking-cartilage is

broadly V-shaped paralleling the lateral and anterior

sides. A deep, hemispherical depression lies at the

anterolateral apex of the lock and is indented into the

lateral wall. Medially the groove gradually becomes

shallow toward the inner edge of the cartilage. The

mantle component is a V-shaped ridge with a large,

swollen apical knob. The transverse limb extends ventrally and the oblique limb extends posteriorly. A distinct groove lies anterior to the transverse limb and

a more shallow depression parallels the oblique limb.

A long, narrow funnel valve is present. The dorsal

portion of the funnel organ is an inverted V-shaped

pad (Plate 7H). The limbs are very long and broad;

they bear high flaplike median ridges. The apical area

bears a separate, flaring median ridge that has an

anteriorly directed point located slightly posterior to

the apex of the organ. The ventral pads are large and

nearly oval, although slightly pointed anteriorly. Two

pairs of bridles attach to the dorsal surface of the funnel near its free anterior end.

The head is short and oval with large eyes that occupy almost the entire lateral portions of the head; the

circular eyelids have a small anterior sinus. Nuchal

folds and the nuchal crest are lacking. A small olfactory

papilla lies posterior to each eye and lateral to the

funnel. The nuchal locking-cartilage is long and narrow ; a deep groove with raised lateral ridges runs along

the midline (Plate 7F) .

The length of all arms except arms I is approximately 85 percent of the mantle length in the largest

specimens. Arms I are about 60 percent of the mantle

length. The arm formula is 2 = 3 = 4>1. Gelatinous

aboral keels are present on arms I—III and lateral keels

are present on arms IV. The arms are slightly damaged

and the total extent of the keels cannot be determined.

Low, undulating, trabeculate protective membranes

are present on arms I—III, but are lacking on the

ventral arms. The suckers are arranged biserially in

longitudinal rows along the entire lengths of all the

arms.

The sucker stalks on arms IV arise from large

gelatinous bases that extend transversely across the

arms and give the oral faces a ladder-like appearance.

The suckers on arms IV are much smaller, more closely

packed and more numerous than on the other arms.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The inner rings of the arm suckers bear numerous, low,

truncate teeth around the entire margin of the ring.

The teeth on the distal part of the rings are larger than

those on the proximal part. About 20 teeth occur on

the larger suckers, fewer on smaller suckers (Plate

9Q-T) . The outer chitinous rings are very narrow; they

bear numerous minute granulations. The number of

suckers on the arms and clubs of the largest specimen

of C. sirventi (Gerda 212, 42 mm ML) is as follows:

Arm

I

II

III

IV

Club

Left

46 +

-

66

78

117

Right

48

-

-

70

122

The tentacles are long and robust; the tentacular

stalks are heavily pigmented and lack suckers. The

tentacular club is expanded (Plate 1B) ; it bears suckers

on long, slender stalks in four longitudinal rows; suckers of the manus grade in size into the smaller suckers

of the dactylus. Each transverse row on the manus has

suckers of nearly equal size. The inner rings of the

suckers from the manus bear approximately 20 teeth;

a few of the most distal teeth are the longest and most

pointed; these grade into teeth on the proximal margin that are short and truncate (Plate 9o). The broad

outer rings from the suckers of the manus are made

up of narrow, fused radiating strips (Plate 9u).

The suckers of the dactylus grade in size in a transverse series from the largest in the ventral row to the

smallest in the dorsal row. The flattened, expanded tip

of the dactylus bears an irregular circlet of about 14

suckers with three or four additional scattered suckers.

The inner rings of the dactylus suckers have about

22-28 teeth with larger, more pointed teeth distally

(Plate 9P).

The carpal locking-apparatus consists of an irregular

grouping of 5-6 suckers and pads at the base of the

manus with an additional one or two suckers and pads

extending up the dorsal margin of the manus.

Well-developed protective membranes, supported by

broad, flat trabeculae, occur on the club. The dorsal

membrane terminates at the base of the dactylus and

the ventral membrane extends to the tip of the dactylus.

A broad, aboral keel exists on the distal half of the

manus and along the dactylus.

Seven buccal lappets occur on the buccal membrane.

The connectives of the buccal membrane attach to the

dorsal borders of arms I and II and to the ventral borders of arms III and IV. The oral surface of the membrane is slightly rugose.

The beaks are illustrated on Plate 8F. The rostrum

of the upper beak is long and sharply pointed; the

palatine lamella is broad. The lower rostrum of the

mandible is short and moderately blunt. No lateral

ridge occurs across the gular lamella. The beaks, especially the rostrae, are deeply pigmented.

The radula has seven rows of teeth in a transverse

series (Plate 7K.) The rhachidian tooth has a long,

sharply pointed median cusp and two short, pointed

lateral cusps. The first lateral tooth has a moderately

long median cusp and two irregularly shaped lateral

cusps. The second lateral tooth has a broad base and a

single long, pointed cusp. The third lateral tooth has

an extremely long, narrow, pointed cusp. Marginal

plaques are present.

A large oval photophore is embedded on the ventral

surface of the ink sac immediately to the left of the

intestine (Plate lc). Photophores also exist on the eye.

Joubin described these photophores around the eye as

small, yellow organs but did not give further details.

Although the eyes were damaged, photophores could

be distinguished on the specimen from off Bermuda

(Acre 2-2). A row of about 20 small round to oval

irregularly-spaced photophores with a pale golden

sheen rims the edge of the iris (Figure 1). A thick

golden band occurs on the ventrolateral surface of the

FIGURE 1.—Pattern of photophores on the eye of Cycloteuthis

sirventi (Acre 2-2; partially reconstructed).
NUMBER 5

eye and diminishes dorsally. No discrete organs exist

along the band, but the entire thickened part, at least,

appears to be a luminescent organ. In addition, an inverted U-shaped band of golden tissue lies embedded

on the ventral surface of the liver, dorsal to the anal

section of the intestine. This organ, too, appears to be

a photophore.

The rhachis of the gladius is long and slender; it

extends nearly to the posterior tip of the gladius. The

vane arises near the anterior end of the gladius, but it

remains narrow for nearly half the length of the gladius

then broadens suddenly. The edges of the vane fold

ventrally in the posterior third of the gladius and fuse

to form a long, hollow conus (Plate 8c).

While the largest specimens available are males (41,

42 mm ML), they are not sufficiently mature to have

developed spermatophores. In fact, the entire reproductive system at this stage is very weakly developed.

Although Joubin's original and subsequent descriptions of C. sirventi were relatively lengthy, they did

not include an abundance of pertinent, specific characters. The specific features that he did mention, however, do not differ significantly from those described

above.

LARVAE (Plate 2A-B) .—The larvae are elongate and

bear small terminal fins. The posterior tail that is long

in the adult is only a small bump between the posterior margins of the fins in the larvae. The visceral

light organ is barely visible and lies ventral to the

intestine instead of at the side as in the adult. The

photophores on the eyes are not present in larvae.

The locking apparatus is similar to that of the adult.

The eyes protrude noticeably, but they are not stalked.

The arms are short and in the order of 2 = 3^4>1 .

The tentacular clubs are miniature versions of the

adult clubs. The larval pen is similar to that of the

adult, except that it is not drawn out posteriorly into

a long conus. The vane widens at the level of the fins,

and posteriorly the margins of the vane turn ventrally

to form a very short conus.

LOCATION OF HOLOTYPE.—Oceanographic Museum,

Monaco.

TYPE-LOCALITY.—Approximately 800 km west of

Madeira at 30°45'N, 25°47'W, 0-500 m.

DISTRIBUTION.—Throughout Atlantic warm waters:

Western Atlantic off Brazil, Straits of Florida, off Bermuda; Eastern Atlantic off Northwest Africa, off St.

Peter and Paul Rocks (Figure 2). Specimens have

been captured at depths between 100 m and 650 m.

Discoteuthis, new genus

DIAGNOSIS.—Tail lacking; gladius greatly thickened; conus absent or only slightly developed; medial

suckers on manus greatly enlarged; external photophores present, visceral photophores lacking; fins very

large and round, equal to the mantle length in adults.

TYPE-SPECIES.—Discoteuthis discus, new species. By

original designation.

Discoteuthis discus, new species

PLATES 3; 4; 7A, B, G, I ; 8A, D ; 9A-G

DESCRIPTION.—The mantle is nearly conical but

tapers posteriorly to a slightly rounded point (Plate

3A). The mantle is widest (about 50 percent of the

mantle length) at its free anterior end. The anterior

margin of the mantle exhibits no pronounced projections or lobes. The mantle wall is relatively thick, but

it is not heavily muscled. A central layer of gelatinous

connective tissue lies between thin outer and inner

sheets of strong, circular muscles. The mantle and

head are covered with moderately thick semigelatinous

tissue. Table 2 lists measurements of specimens.

Sex

HOLOTYPE

d"

ML, mm

53

OTHER SPECIMENS:

9

9

cf

9?

juv.

45

25

24

24

16

Station number

Pillsbury 36

Dana—no data

Dana 1284 I

Pillsbury 473

Dana 1178 II

Dana 1172 II

list of Material

Location

03°50'N O2°37'W

14°38'N 61°16'W

13°08'N 61°55'W

10°24'N 54°38"W

07°22'N46°51'W

29

27

1

19

14

Date

V 1964

X 1921

VIII 1966

XI 1921

XI 1921

Estimated

depth, m

750

300-400

450

200

200
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The fins are huge and broadly oval in outline (Plates

3A, 4C). They extend nearly the entire length of the

mantle (98 percent of the mantle length in the holotype) and are very broad. Their width is about 40

percent greater than the length of the mantle. For

most of their length the fins do not fuse in the dorsal

midline but are separated by a narrow gap. The bases

of the fins insert directly on the shell sac of the thick,

sturdy gladius. The attached median borders of the

fins diverge anteriorly forming a distinct V. The anterior fin lobes are short and free. The posterior margin

of the fins near the midline is almost straight and has

no pronounced posterior projection. The fins are very

heavily muscled.

The funnel is short and broad; it extends anteriorly

to the level of the posterior border of the eyelids. The

musculature of the funnel is similar to that of the

mantle. The anterior end of the funnel is free; the

remaining portions of the funnel at the collar appear to

be bound to the head by a continuous integumentary

sheath, but the condition of the specimen makes this

uncertain. The funnel locking-cartilage is approximately subtriangular and is encircled by a broad,

nearly membranous flange (Plate 7A). Its longest axis

lies at a diagonal to the longitudinal axis of the body.

The sulcus has a broad, shallow depression anteriorly;

it becomes shallower in the midportion and then

deepens to a hemispherical depression posterolaterally.

The mantle component consists of a diagonally-set,

rounded, cartilaginous ridge that has a prominent

nose-shaped bulge at the posterolateral end. The bulge

articulates with the deep depression on the funnel

component. A moderately deep furrow extends along

the anterior side of the ridge and curves around the

ends; this conforms to the prominent anterior edge of

the funnel component.

The dorsal member of the funnel organ is a large,

swollen, inverted V-shaped pad (Plate 7G ). A large,

thumblike papilla protrudes from its apex and a low,

indistinct ridge extends partway along each limb. The

ventral members are large, elongate, rounded pads.

The funnel valve is a small, thin, semicircular flap.

The head is broad and short; it bears huge eyes that

occupy nearly the entire lateral region of the head.

Each circular eyelid has a small, anterior sinus. The

funnel groove is indistinct (or nonexistent). A small

olfactory papilla protrudes from the posterolateral

surface of the head. No nuchal ridges or folds appear

to have been present on the slightly damaged holotype

or on the other specimens.

The nuchal locking-cartilage is oval, and it has a

distinct ridge on each side of the deep median sulcus.

The mantle component has a complementary arrangement with a median ridge and lateral grooves (Plate

7B).

The arms are large (60-85 percent of the mantle

length) and robust, but they are not heavily muscled;

most of the arm-tips are missing from the holotype.

The arms appear to be approximately subequal. The

outer integument has been rubbed off the arms and

no traces of aboral keels remain. Low, fleshy protective

TABLE 2.—Measurements {in mm) of Discoteuthis discus

Characters

Sex

Mantle length

Mantle width

Head length

Head width

Fin length

Fin width

Arms I

II

III

IV

Tentacle length

Club length

Pillsbury 36

d"

53

27

27

27

52

74

34 +

44 +

33 +

36 +

132

26

Dana

9

45

25

26

28

44

65

-

-

-

-

117

25

Dana 1284 1

9

25

16

-

_

23

43

_

_

-

-

68

12

Pillsbury 473

&

24

12

10

13

22

44

_

_

_

_

_

-

Dana 1178 11

9?

24

14

12

14

20

30

_

18

_

_

61

11

Dana 1172 11

juv.

16

8

5

7

10

16

_

-

-

-

36

6
NUMBER 5

membranes, supported by fleshy trabeculae, occur on

all of the arms.

The biserially arranged suckers on the arms are

large, globular, and relatively few in number. The

suckers are small at the base of each arm but rapidly

increase in size and attain their maximum dimensions

along the middle one-third of the arm. Suckers on arms

IV do not increase greatly in size and are only about

half the size of the largest suckers on the other arms.

The suckers have narrow stalks which are set on large,

robust, conical bases. The inner chitinous rings on the

basal suckers are smooth. Distally, however, they have

about 7-9 low, broad, rounded teeth on the distal

borders (Plate 9C-F). In young specimens the teeth

tend to be slightly more pointed.

The numbers of suckers on the arms of the holotype

are 43 on right I, 38 on left II, and 46 on right IV.

Tips are missing from the remaining arms, so the

number of suckers that occur along the proximal

portions of the arms for a distance equal to 50 percent

of the mantle length are recorded:

Arm

I

II

III

IV

Lejt

25

21

-

31

Right

25

22

21

27

There are 81 suckers on the right tentacular club

and 80 on the left.

The tentacles are very long—about two and a half

times the length of the mantle. The stalks are moderately robust and muscular; they bear no suckers. The

clubs are expanded and flattened (Plate 3B). The

ill-defined carpal area of the club bears 2 or 3 small,

irregularly arranged suckers. A few shallow, round

depressions, apparently containing flattened knobs,

alternate with the suckers and correspond in position

to the location of the suckers on the opposite club. The

carpal suckers bear small, irregular teeth around their

entire circumference. The carpal area blends distally

with 2-4 small to slightly larger suckers with more

distinct teeth that are truncate and slightly larger on

the distal margin of the sucker ring. It is uncertain

whether the smaller of these are true carpal suckers;

however, corresponding pads could not be found.

The manus bears four rows of suckers. The 6-9

suckers in the two medial rows on the manus are

greatly enlarged; they are closely packed and are set

in deep, thin-walled pockets on short, thick stalks.

Approximately 32 very low, flattened to slightly

rounded teeth line the entire circumference of the inner ring of each of these suckers (Plate 9A) ; the outer

rings are very narrow and smooth. Distally, the large

suckers give way to one or two suckers of intermediate

size with more elongate, rounded to truncate teeth on

the distal margin of the ring. The suckers of the two

marginal rows are small and widely spaced; they are

attached close to the marginal edges of the club by

long, slender stalks. Two proximal suckers of the dorsal

marginal row have very short stalks and therefore are

closely attached to the club. These two suckers closely

resemble the carpal suckers in size and appearance.

The stalked marginal suckers bear long, slender,

pointed to truncate teeth on the distal margin on the

inner ring, but shorter more irregular teeth on the

proximal margin; the total number of teeth is about 36.

Four longitudinal rows of small suckers set on long

stalks extend the length of the dactylus and terminate

just short of the tip. In a transverse series, the suckers

grade in size from the largest at the ventral end to the

smallest at the dorsal end. The inner rings of the

dactylus suckers are each armed with numerous (about

34) sharply pointed teeth; those of the distal half are

much longer than those of the proximal half. The

dactylus suckers and those of the marginal rows of

the manus each have broad outer rings which have

several rows of minute bumps; the bumps of the inner

row are longer and look like minute teeth. An irregular circle of about 11 or 12 small suckers lies on

the flaplike extreme tip of the dactylus.

Protective membranes are present along the dorsal

and ventral borders of the manus. The marginal

suckers arise from the trabeculae that support these

membranes. The dorsal protective membrane disappears along the base of the dactylus while the ventral

membrane continues along the dactylus to the tip of

the club as a low membrane with flattened trabeculae.

A low keel extends along the distal quarter of the

tentacular stalk. A broad swimming keel originates at

about the midportion of the aboral surface of the club

and extends along the dorsal margin of the dactylus.

It terminates abruptly at the base of the flaplike tip

of the dactylus.

The buccal mass is large and protruding. The seven

strong, muscular supports of the buccal membrane

have connectives that attach to the dorsal surfaces of

arms I and II and to the ventral surfaces of arms III

and IV. The seven buccal lappets are small and triangular. The buccal membrane is fleshy and rugose.

328-974 O - 69 - 2
SMITHSONIA N CONTRIBUTIONS TO ZOOLOGY

100' 90" 80' 70" 60" 5 0' W 30' 20 ' 10* 0* ID* 20*

500 mime

Cycloteuthis sirvent

Discoteuthis discus

Discoteuthis laciniosa

Holotype

90" 70* 20* 20"

FIGURE 2.—Geographical distribution of Cycloteuthis sirventi, Discoteuthis discus, and

Discoteuthis laciniosa.

The upper and lower mandibles are illustrated on

Plate 8D. The upper beak has a long, curved, pointed

rostrum and a semicircular palatine lamella. The rostrum of the lower jaw is triangular in silhouette. The

gular lamella is large and has a moderate lateral ridge

across its surface. The rostrae are darkly pigmented.

The dorsal and lateral region of the palatine lamella

of the upper jaw is pigmented.

The radula consists of seven rows of teeth in a transverse series (Plate 7i). The rhachidian tooth has a

broad, pointed median cusp and two small, triangular

lateral cusps. The first lateral tooth has a short, sharp

primary cusp and a small, pointed lateral cusp. The

second lateral tooth has a broad base that tapers into

a single pointed cusp. The third lateral tooth has a

single, very long, narrow, curved, sharply pointed

cusp. Small marginal plaques are present.

At least four light organs occur around the margin

of the eyelid, but this area is damaged so the exact

number and arrangement of photophores is uncertain.

A dark, round photophore is located on the ventral

surface of the posterior tip of the mantle. Reddishbrown chromatophores occur on the mantle, head, fins,

and arms where the skin is not damaged.

The gladius has a total length of 47 mm in the holotype, of which the anterior 16-18 percent consists

almost exclusively of the rhachis except for a thin,

narrow anterior flap 1 mm in length (Plate 8A). The

rhachis is widest at the point where the vane originates

(3 mm wide); it then gradually tapers posteriorly and

disappears 3-4 mm from the terminal end of the

gladius; therefore, the posterior portion of the gladius

consists entirely of the vane. The vane is long and

broad. Soon after its origin near the anterior end of

the gladius, the vane widens rapidly, and for the

greater part of its length the lateral edges lie nearly

parallel. Posteriorly the vane forms a rounded, spoonshaped end (not a conus) with the terminal edge

folded slightly ventrally and anteriorly to form a narrow, delicate lip.

The most striking feature of the pen is its extreme

thickness. Almost the entire concavity ventral to the
NUMBER 5

vane is occupied by a solid amber-colored substance.

The pen, which is usually paper-thin in other oegopsids,

therefore is approximately 2 mm thick. The vane has

the normal thin structure only along its edges; the

rhachis is typical (see cross sections). Interestingly,

when the pen is submerged in water, the thickened portion becomes nearly invisible and the pen assumes the

appearance of a typical gladius (Plate 8A) .

SUBADULT STAGES.—The amount of material available for this study is limited, but the specimens represent a wide range of sizes. Some aspects of the changes

that occur during growth are apparent on the specimens at hand. The smallest specimen that can be positively identified as Discoteuthis discus has a mantle

length of 16 mm {Dana 1172 II).

The major allometric changes during growth are

associated with the length of the fins. At 16 mm ML

the fins occupy about 60 percent of the length of the

mantle. Around 24-25 mm ML the fins increase proportionally to 80-90 percent of the mantle length. By

45 mm ML and larger the fin length is 98 percent or

more of the mantle length (Plate 4A-C). The gladius

strongly reflects the allometric increase in the size of

the fins by the relative lengthening of the vane during

ontogeny (Plate 4D) . The basal musculature of the fins

attaches to the shell sac which tightly envelops the

gladius. Therefore, the allometric growth of the pen

and fins is directly related.

LOCATION OF HOLOTYPE.—The holotype of Discoteuthis discus has been deposited in the United States

National Museum and is designated as catalog number 576214.

TYPE-LOCALITY.—Approximately 100 km SSW of

Cape Three Points, Ghana, West Africa, in the Gulf

of Guinea at 03°50'N, O2°37'W; Pillsbury Sta. 36;

29 May 1964; 750 m.

DISTRIBUTION.—Atlantic warm water: Gulf of

Guinea; off northeast coast of South America: and eastern Caribbean Sea (see Figure 2). Captured in open

nets at depths between 200 and 750 m.

ETYMOLOGY.—The generic name, Discoteuthis, is

derived from the combination of the Greek "diskos,"

a flat, circular plate or disk, and "teuthis," squid; it

refers to the huge circular fins. The specific name of the

type-species, discus, emphasizes this feature of the fins.

Discoteuthis laciniosa, new species

PLATES 5; 6A-E; 7C, D, J; 8B, E; 9H-N

DESCRIPTION.—The mantle is conical and the muscular portion terminates slightly anterior to the posterior margins of the fins. The mantle width is nearly

one-half the mantle length (see measurements in Table

3). The mantle musculature is weak and covered externally by a thick layer of gelatinous connective tissue.

A few (four or more) small, truncate, integumentary

flaps occur along the ventral and lateral borders of the

anterior margin of the mantle. In the holotype four

flaps occur ventrally between the photophores and

some flaps may occur lateral to the photophores, although this is uncertain due to the slightly damaged

condition of the mantle margin. The mantle flaps are

more pronounced in the smaller specimens (Plate 6A,

B ).

Sex ML, mm

HOLOTYPE :

p 70

OTHER SPECIMENS:

p

?

<?

p

juv.

L

3 L

56

39

35

20

14

7.5

5-9

Station number

Acre 3-3

Dana 1157 I

Discovery 4840

Gerda-TOTO

Dana 1161 IV

Dana 4009 III

Dana 1341 II

Dana 4009 II

List of Material

Location Date

33°04'N 64°37'W 4 VII 1968

Estimated

depth, m

1000

21°

30°

23°

14°

24°

33°

24°

57'N 22°58'W

02'N 22°55"VV

'52'N 76°46'W

42'N 28°04'\V

36.5'N 17°27'W

15'N 68°20'W

36.5'N 17°27'\V

27

5

20

5

18

14

18

X

IV

II

IX

III

V

III

1921

1962

1961

1921

1930

1922

1930

300-400

600

750

100

100

200

200

5 additional specimens in poor condition, 55-82 mm ML. taken as stomach contents of

the fish, Aphanopus carbo, off Madeira.
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 3.—Measurements (in mm) of Discoteuthis laciniosa

Characters

Sex

Mantle length

Mantle width

Head length

Head width

Fin length

Fin width

Arms I

II

III

IV

Tentacle length

Club length

Acre

3-3

?

70

30

31

40

71

95

41 +

63

62

51 +

84

20

Dana

11571

•>

56

26

29

27

59

88

31

51

48

38

80 +

19

Discovery

4840

?

39

12

11

10

38

51

-

-

28

28

68

10

Gerda

TO TO

d"

35

12

11

-

32

52

19

20

29

27

52

10

Dana

1161IV

•?

20

-

-

7

19

26

-

-

-

-

24

5

Dana

4009 III

juv.

14

5

-

-

12

18

-

-

-

-

-

—

Dana

4009 II

larva

9

3

3

4

7.5

12

2 .5

4 . 5

4

3.5

-

Dana

4009 11

larva

5.5

2.5

2 .5

3.0

5.0

8.0

2.0

2.5

2.5

2.5

9.0 +

Dana

4009 II

larva

5.0

2 . 5

2.0

2 .5

2.0

6.5

1.5

2 . 5

2 .5

2.2

-

Dana

1341 II

larva

7.5

-

-

-

7.5

12

-

-

-

-

-

? Dana

3996 VII

9

33

11

14

12

33

44

21

30

31

28

47

9

The round fins are very large and in adults slightly

exceed the length of the mantle (Plates 5A; 6A-C).

The width of the fins is one and one-third to one and

one-half of the mantle length. The fins are extremely

muscular, but dorsally, at least in the midportion, they

are overlain by a layer of gelatinous connective tissue.

The fins, for most of their length, do not fuse in the

dorsal midline but are separated by a narrow gap. The

bases of the fins insert directly on the shell sac of the

thick sturdy gladius. The anterior lobes are free; each

medial border forms a deep notch with the surface of

the mantle. The notch is more pronounced in the holotype than in the illustrated paratype (Plate 6c). The

lateral edges of the fins are semicircular in outline;

the posterior margins are nearly straight. No tail exists.

The medium-sized funnel extends nearly to a level

with the center of the lenses of the eyes; it lies in a

shallow groove on the ventral surface of the head. A

small funnel valve is present. The dorsal pad of the

funnel organ has an inverted V-shape with long limbs.

A low median ridge appears to be present on the

anterior half of each limb. A small papilla is present

at the anterior apex. The ventral pads are very long and

narrow. The funnel locking-cartilage is subtriangular

in shape and has a deep, V-shaped groove (Plate 7c).

A deep, oval depression lies at the anterolateral angle

of the cartilage. The groove becomes more shallow

medially and posteriorly, and remains well defined in

most specimens, although in the holotype it is somewhat less distinct than in the illustration of a paratype

(Plate 7c). A distinct rounded ridge overhangs the

groove along the anterior margin of the cartilage. The

mantle locking-cartilage consists of a transverse,

broadly V-shaped ridge and groove. The ventral end

of the groove is deep and pronounced; the groove disappears at the dorsal end. The strong, transverse (ventral) part of the ridge tapers diagonally from the bulb

that locks into the deep depression on the funnel cartilage. The posterior sides of both parts of the ridge taper

into the inner surface of the mantle.

The head is large, broad and oval; the eyes are

enormous and occupy almost the entire lateral portion

of the head. A small sinus is present in the anterior

margin of each circular eyelid. An olfactory crest is not

apparent. A narrow, fingerlike olfactory papilla occurs

on the posterior ventrolateral angle of the head. The

sensory epithelium is not restricted to the tip of the

papilla but covers most of the surface of this structure.

The nuchal cartilage is shield-shaped with a broad

anterior end. A broad central ridge that extends the

length of the cartilage bears a narrow median groove

(Plate 7D).

Although the arms are difficult to measure, they

appear to be in the order of 2 = 3^4>1 . The longest

arms are approximately 90 percent of the mantle

length and the shortest (the dorsal pair) are about 60

percent of the mantle length. Gelatinous aboral keels

are present on arms I—III; those on arms III are the

largest and those on arms I, the smallest. Lateral keels

are present on arms IV. The medial ventral margin of
NUMBER 5 11

each arm IV is sharply angled forming a narrow keel.

The arms are very attenuate. Protective membranes

occur on all arms; they are supported by the conical

bases of the sucker stalks and not by typical independent trabeculae. The arms bear two rows of large

suckers with broad apertures. The largest suckers of

the ventral arms are considerably smaller than the

largest suckers on the other arms. The distal half to

two-thirds of each of the large inner sucker rings bears

10-12 low, scalloped teeth; the proximal portion of

each ring is smooth. The number of teeth decreases on

the rings of smaller suckers (Plate 9J-M) . Outer

chitinous rings are extremely narrow and poorly developed; they are covered with minute granules. The

number of suckers on the clubs and on the proximal

portions of the arms for a distance equal to 50 percent

of the mantle length of a paratype (Dana 1157 I, 56

mm ML) is given below. The total number of suckers

on each arm is enclosed in parentheses.

Arm

I

II

III

IV

Club (total)

Left

39 (42-broken)

26 (80)

25 (90)

36 (60)

67

Right

37 (57)

28 (38-broken)

27 (84)

38 (66)

56 +

The tentacles are robust and relatively short—between one and two times the length of the mantle;

the tentacular stalks are naked. The clubs are expanded and bear four transverse rows of suckers on

the manus (Plates 5B; 6D). NO distinct carpal cluster

occurs, but a series of 4 or 5 minute suckers occurs

along the dorsal margin of the manus and carpal

region. These suckers are about one-tenth the diameter

of the largest manal suckers and about one-half the

diameter of the suckers in the true marginal series of

the manus. The carpal suckers bear small truncate

teeth, only slightly differentiated in size, around the

entire margins of the inner rings. The proximal three

suckers in this series alternate with three smooth, round

depressions that are not typical carpal knobs. The

distal two suckers in this series do not alternate with

depressions and may occur adjacent to small manal

suckers of the marginal series. The distalmost "carpal"

sucker may be absent.

The manus bears four rows of suckers. Six or seven

of the suckers in the two medial rows are enormously

enlarged and have very broad apertures. The outer

chitinous rings on the large suckers are very narrow

and are made up of numerous minute, curved rods;

no granulation exists. The dentition on the inner rings

is very weak; it consists of numerous (50-60) minute,

irregular, low, scalloped teeth around the entire

margin (Plate 9 H ). The suckers of the two medial

rows on the manus decrease rapidly in size proximally;

these small proximal suckers of the manus have long,

curved, pointed teeth distally and shorter, truncate

teeth proximally on the inner rings. The suckers on

both marginal manal rows have rings that bear

numerous, small, pointed teeth around their entire

margins. The teeth are larger, more pointed and

curved on the distal margin of the inner ring. The

outer chitinous rings are well developed with radial

striations around the outer margin. The proximal two

suckers in the dorsal marginal series of the manus

may be absent or may occur adjacent to the distal

carpal suckers.

The suckers of the dactylus decrease in size from the

ventral to the dorsal row. The suckers of the dorsal row

diminish rapidly in size and disappear about halfway

out the dactylus. Distal to this point, the suckers of the

dorsomedial row decrease abruptly in size but continue

to the tip of the dactylus. The suckers of the two ventral

rows maintain a relatively large size up to the distal

two transverse rows where they decrease rapidly to

nearly equal the size of the remaining distal suckers

of the dorsomedial row. Eleven to fifteen small suckers

are arranged in a circlet or patch around the margin

of the slightly expanded, flattened tip of the dactylus.

Protective membranes are present; strong muscular

rods connected with membranous integument serve as

trabeculae and as support for the small suckers of the

marginal rows on the manus. The ventral membrane

extends to the end of the dactylus, but the dorsal membrane terminates about halfway out the dactylus. A

short, broad swimming keel occurs on the aboral surface of the dactylus .

The buccal membrane bears seven lappets and supports; the connectives of the buccal membrane are

attached to the dorsal borders of arms I and II and to

the ventral borders of arms III and IV. The buccal

membrane is fleshy and rugose.

The beaks are illustrated on Plate 7E. The rostrum

of the upper mandible is slightly curved and bluntly

pointed; the lateral sides of the palatine lamella are

very broad and are rounded in outline. The rostrum

of the lower jaw is triangular in outline and has a

sharp, projecting tip. The gular lamella is small and
12

has a slight lateral ridge. The rostrae are darkly pigmented. The pigment on the palatine lamella of the

upper beak is concentrated only along the dorsolateral

surface.

The radula possesses seven rows of teeth in a transverse series (Plate 7j). The rhachidian tooth has a relatively narrow, pointed median cusp and two, small,

pointed lateral cusps. The second lateral tooth has a

broad base, a broad, triangular primary cusp, and a

small, sharp lateral cusp. The cusp of the second lateral tooth has a broad base that tapers to a sharp point.

The cusp of the third lateral tooth is extremely long

and narrow; it is slightly curved and sharply pointed.

Small marginal plaques are present.

A small photophore is embedded near the edge of

the membrane that passes between the third and

fourth arms on each side, and two small photophores

lie ventrolaterally near the free margin of the mantle

(Plate 5A) . No photophores are present on the surface

of the eye. No other photophores can be detected on the

holotype or on other specimens, but all the material

is damaged to varying degrees.

The gladius of a paratype {Dana 1157 I) is about

55 mm in length (Plate 8B). The free anterior end of

the rhachis is U-shaped in cross section, short, and

broad (about 7 mm long and 3.5 mm wide) with a

small terminal flap. The rhachis tapers posteriorly from

the point where the vane arises and terminates short of

the end of the gladius. The deep U-shape of the rhachis

provides the gladius with a high dorsal ridge. The vane

is long and broad. The lateral edges of the vane begin

to turn ventrally in the posterior third of the gladius;

the edges meet and fuse near the tip of the gladius to

form a simple conus. Just posterior to the free portion

of the rhachis the gladius becomes very thick dorsoventrally for its entire length; only the sides of the

vane remain thin. The thickening consists of a nearly

unpigmented, hardened substance (see cross sections on

Plate 8B).

LARVAE (Plate 6E).—Four larval specimens which

appear to belong to Discotenthis laciniosa are available; they range from 5-9 mm in mantle length. The

mantle is short and very broad in the smallest larvae,

but it becomes proportionately narrower with growth.

The huge, broad, round fins are about 50 percent of

the mantle length in the larva of 5 mm ML but rapidly

enlarge to 80-90 percent as long as the mantle by 9 mm

ML. From 10-15 mm ML the fins have reached their

full proportional development and are slightly longer

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

than the ML. The width of the fins is about 133-160

percent of the ML in width. A series of 6-8 papillalike flaps projects from the ventral two-thirds of the

mantle margin.

The eyes protrude from the sides of the head but

are not stalked; they are elliptical in an anterodorsal

to posteroventral direction. The long axis of the eye

from the specimen 9 mm in ML is 1.7 mm and the

short axis is 1.1 mm.

The funnel is long and narrow; it extends anteriorly

to the base of the oral crown. The funnel lockingcartilages are diagonally set, subtriangular-shaped

pits (very similar to those of the adult) that lock with

corresponding bumps of the mantle cartilages. The

funnel organ is a large V-shaped gland with a small

papilla at the apex and a low ridge extending along

each limb.

The arms are weak; the arm formula is 2^3>4>1 .

No complete tentacles or clubs remain on the

specimens.

In the larvae of this genus the large fins are very

widely separated from one another by the broad pen.

This distinctive feature allows recognition of the larvae

at a glance.

LOCATION OF HOLOTYPE.—The holotype of Discoteuthis laciniosa has been deposited in the United

States National Museum and is designated as catalog

number 576215.

TYPE-LOCALITY.—Approximately 40 miles NNE of

Bermuda at 33°04'N, 64°37'W; USNS Gilliss, Acre 3 -

3; 4 July 1968; 1000 m, est.

DISTRIBUTION.—North Atlantic warm waters: off

West Africa; Madeira; Cape Verde Islands; in the

Tongue of the Ocean, Bahamas; and off Bermuda. See

Figure 2. Captured in open nets at depths between

100 and 1000 m.

ETYMOLOGY.—The specific name, laciniosa, is

derived from the Latin word "laciniosus," full of flaps

or fringed; it refers to the small flaps that fringe the

ventral half to two-thirds of the free margin of the

mantle.

Comparison of Discoteuthis discus and D. laciniosa

The two species differ distinctly in several characters. The most apparent differences concern the

distribution of photophores. Discoteuthis discus has

four light organs embedded in the tissue surrounding

each eye and a single, prominent round photophore

located ventrally near the posterior apex of the mantle.
NUMBER 5 13

Discoteuthis laciniosa has two small, round photophores on the ventral surface of the mantle near the

anterior margin and a small, round photophore on

each side near the edge of the membrane that connects the bases of arms III and IV (Plates 3A; 5A).

The total complement of photophores in each species

may not have been detected since our specimens were

partially damaged, but, based or present knowledge,

no apparent overlap exists in photophore patterns.

The dorsal row of suckers is lacking on the distal

half of the dactylus in D. laciniosa, but it is complete

in D. discus (Plates 3B; 5B; 6D) . The enlarged suckers

on the manus are globular in D. discus and flattened in

D. laciniosa. The percent of the maximum height

of the chitinous ring in relation to the diameter is 53-

66 percent in D. discus and 33-38 percent in D.

laciniosa (Plate 9G, N ).

The funnel locking-cartilage of D. laciniosa has a

distinct V-shaped groove with a strong apical depression; the posterior limb of the groove is lacking in

D. discus (Plate 7A, C).

Small flaplike papillae project from the free border

of the mantle in D. laciniosa; these are most prominent in small specimens. The papillae are absent in

D. discus (Plates 3A; 4A, B; 5A; 6A, B).

The length of the fins approximates the length of

the mantle at 10-15 mm ML in D. laciniosa, but this

does not occur until a minimum of 25-30 mm ML in

D. discus (Plates 4B ; 6A) .

The gladius of D. laciniosa has a small, distinct

conus, but no conus occurs in D. discus. This difference in pens is unusual for such closely related species

(Plate 7A, B).

More suckers appear to occur on the arms of D.

laciniosa than on D. discus, but further material is

needed to substantiate this condition.

ADDITIONAL MATERIAL.—Our material includes several specimens which do not clearly belong to the

species of Discoteuthis described here, but they are

in such poor condition that we are not able to characterize them sufficiently. Most of these specimens

were taken within the distributional ranges of the

known species. Additional material is needed in order

to delineate more fully the species of the Cycloteuthidae in the central Atlantic region.

One mutilated specimen of Discoteuthis about 35

mm in mantle length is available from the Indian

Ocean (07°26'N, 64°40'E). Although specific identification is impossible, this specimen extends the known

range of the Cycloteuthidae into the Indian Ocean.

A larval specimen, in good condition except for the

loss of the tentacles, with a mantle length of 9 mm

(Dana 1335 XV, 28°02'N, 62°26'W) does not appear

to be referable to either D. discus or D. laciniosa, so

it is described here briefly.

The mantle is long and narrow; it tapers to a blunt

tip posteriorly (Plate 6F) . Several large integumentary

flaps project from the mantle margin. The large, round,

separate fins are about 72 percent of the ML and about

as broad as the mantle is long.

The head is broad, relatively short. The eyes are

huge and occupy the entire sides of the head. They are

teardrop-shaped with a large rostrum directed posteroventrally. The long axis of the eye is 2.6 mm and the

short axis is 1.6 mm. The lenses are very large.

The funnel extends anteriorly to a level with the

center of the eyes. The locking apparatus is subtriangular and generally similar to other cycloteuthids.

The arm formula is 2 — 3>4 = 1. Tentacles are missing but the stumps that remain are robust.

Comparison of Cycloteuthis and Discoteuthis

No difficulty should arise in separating these two

distinctive genera. The generic diagnoses list the pertinent differences that are briefly summarized here.

Cycloteuthis: tail present, gladius thin, conus present,

suckers on manus equal-sized, photophores on eyes and

viscera, fins moderate—do not attain 100 percent of

ML. Discoteuthis: tail absent, gladius thick, conus

minute or absent, suckers on manus unequal-sized,

photophores in integument, fins huge—attain 100 percent of ML.

The fundamental similarities that unite these genera

have been listed in the diagnosis of the family. Some

features, however, require further comment. Within

oegopsid families the gladius is generally a homogeneous structure and is often diagnostic of the family

(e.g., Ommastrephidae, Onychoteuthidae), but the

differences between the gladii of Discoteuthis discus

and Cycloteuthis sirventi are pronounced. Cycloteuthis

sirventi has a long, well-developed conus, while D.

discus completely lacks a conus. Discoteuthis laciniosa

has a small conus that is somewhat intermediate, but

it seems to be a different type than that of C. sirventi.

Although these differences in the pen are difficult to

rationalize in the absence of a comprehensive study on

teuthoid gladii, and although they may cast some doubt
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

on the close relationship of the genera, other characters

are sufficiently similar at present to require the inclusion of the two genera in the same family.

Familial Relationships

The attachment of the buccal connectives to either

the dorsal or ventral border of arms IV recently has

been shown to separate the families of the Oegopsida

into two major subdivisions (Young and Roper, 1968)

as follows:

Dorsal attachment Ventral attachment

Ommastrephidae Cycloteuthidae

Bathyteuthidae Onychoteuthidae

Lycoteuthidae Gonatidae

Enoploteuthidae Thysanoteuthidae

Histioteuthidae Ctenopterygidae

Psychroteuthidae Octopoteuthidae

Architeuthidae Brachioteuthidae

Neoteuthidae Lepidoteuthidae

Batoteuthidae

Chiroteuthidae

M as tigoteu thidae

Promachoteuthidae

Joubiniteu thidae

Grimalditeu thidae

Cranchiidae

As the list shows, the Cycloteuthidae is aligned by

this fundamental character with the majority of the

oegopsid families. In the group with ventral attachments to arms IV, the gladius exhibits three basic

modifications. The most common type has a narrow,

generally elongate conus formed by a ventral folding

of the vane with a fusion of the edges in the ventral

midline. This type of gladius is characteristic of the

Brachioteuthidae, Lepidoteuthidae, Batoteuthidae,

Chiroteuthidae, Mastigoteuthidae, Joubiniteuthidae,

Grimalditeuthidae, and Cranchiidae (secondary modifications occur in some genera). The Promachoteuthidae, although aligned with this group, apparently lacks

a pen. The second type of gladius, found in the

Onychoteuthidae and Gonatidae, has a small, short

conus without evidence of a ventral fusion. The remaining families, Thysanoteuthidae, Ctenopterygidae,

and Octopoteuthidae, lack a conus. The Cycloteuthidae

has representatives with two of these types of gladii:

(1) C. sirventi has a conus with a ventral fusion; (2)

D. discus lacks a conus. Discoteuthis laciniosa has a

small conus, but it does not seem to conform to either

of the two basic types found in the family. Present

knowledge does not enable us to establish definitely

which type of gladius is primitive in the family. On the

basis of the pen, the Onychoteuthidae and Gonatidae

can be excluded from close relationship with the Cycloteuthidae. In determining relationships with the

other families, however, the pen is of little value.

The straight, simple funnel locking-cartilage seems to

be primitive in the Oegopsida. Independent modifications of this type of locking-cartilage have arisen that

are stable traits at least at the familial level. Of the

FIGURE 3.—Diagrammatic cross section of body showing relationships between structure of gladii

and attachment of fins: A, Discoteuthis; B, Octopoteuthis.
NUMBER 5 15

families with ventral attachments of the buccal connectives to arms IV, an ear-shaped funnel lockingcartilage is present, with a few exceptions, in the

Chiroteuthidae and Mastigoteuthidae; and oval lock

occurs in the Promachoteuthidae and Joubiniteuthidae; a fused lock occurs in the Grimalditeuthidae and

Cranchiidae; a lazy-T lock (i.e., a straight longitudinal

groove with a medial groove projecting from its midregion) exists in the Thysanoteuthidae. The remaining

families have a straight, simple cartilage, except the

Cycloteuthidae, which has a subtriangular lock with a

deep depression at the anterolateral corner. This distinctive locking apparatus separates the Cycloteuthidae from all other families and gives no clear indication

of its familial relationships.

Specialized tentacular clubs may indicate relationship between families (e.g., Promachoteuthidae, Joubiniteuthidae, and Mastigoteuthidae), but the generalized structure of the clubs of cycloteuthids offers few

clues to indicate relationships.

Superficially, Discoteuthis resembles Octopoteuthis

in the shape and size of the fins. The attachment of the

fins, however, is fundamentally different in the two

genera. The fins of Discoteuthis are separate and insert

directly onto the shell sac, whereas the fins of Octopoteuthis are fused and continuous across the dorsal midline, thereby passing over the shell sac (Figure 3). This

basic difference is also reflected in the structure of the

pens. The pen of Discoteuthis is a very thick, rigid

supporting structure, while that of Octopoteuthis is

very thin and fragile. The possession of hooks on the

arms and the lack of tentacles in Octopoteuthis, among

other features, further substantiate the remoteness of

the relationship between these two groups.

At our present state of knowledge, it is difficult to

determine the relationships of the Cycloteuthidae, other

than to align it with the families that possess the ventral

attachment of the buccal connective to arms IV.

Within this group, the family must be assumed to occupy an isolated position.

Literature Cited

Clarke, M. R.

1966. A Review of the Systematics and Ecology of

Oceanic Squids. Advances in Marine Biology,

4:91-300.

Grimpe, G.

1925. Zur Kenntnis der Cephalopodenfauna der Nordsee.

Wissenshaftliche Meeresuntersuchungen {Helgoland), 16(3):1-124.

Joubin, L.

1919. Etudes preliminaries sur les Cephalopodes recuilles

au cours des croisieres de S.A.S. le Prince de

Monaco. 7e. Note: Cyclotheuthis sirventi nov. gen.

et sp. Bulletin Institut Oceanographique, Monaco,

351:1-7, 2 figures.

1924. Contribution a l'etude des Cephalopodes de l'Atlantique Nord. 4e serie. Resultats des Campagnes

Scientifiques du Prince de Monaco, 67:1-113, 16

plates.

Naef, A.

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

Fischer.

1923. Die Cephalopoden: Systematik. Fauna u Flora

Neapel, Monograph 35, l(pt. 1, no. 2): i-xiv+

149-863, 473 figures.

Thiele, J.

1921. Die Cephalopoden der Deutschen Sudpolar-Expedition. Deutschen Sudpolar Expedition (1901-

1903), 16, Zoology, 8:433-466.

1935. Handbuch de Systematischen Weichtierkunde, 2

(3):779-1154.

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

1968. The Batoteuthidae, a New Family of Squid (Cephalopoda: Oegopsida) from Antarctic Waters.

Antarctic Research Series, 11:185-202.
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 1.—Cycloteuthis sirventi, Gerda Sta. 212, 42 mm ML: A, ventral view; B, right tentacular

club; c, photophore on ink sac.
NUMBER 5 17

PLATE 2.—Cycloteuthis sirventi: A, Margrethe Sta. 1030, 8.5 mm ML, ventral view; B, Dana

Sta. 1172 II, 15 mm ML, ventral view; c, Dana Sta. 829, 27 mm ML, ventral view; D, Gerda Sta.

212, 42 mm ML, dorsal view.
18

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 3.—Discoteuthis discus, holotype, PUlsbury Sta. 36, 53 mm ML: A, ventral view; B, right

tentacular club.
NUMBER 5 19

PLATE 4.—Discoteuthis discus: A, Dana Sta. 1172 II, 16 mm ML, ventral view; B, Dana Sta.

1178 II, 24 mm ML, ventral view; c, Pillsbury Sta. 36, 53 mm ML, dorsal view of holotype;

D, gladii of A-C respectively.
20

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 5.—Discoteuthis laciniosa, paratype, Dana Sta. 1157 I, 56 mm ML: A, ventral view;

B, left tentacular club.
NUMBER 5 21

PLATE 6.—Discoteuthis laciniosa: A, Dana Sta. 4009 III, 14 mm ML, ventral view (eyes

reconstructed) ; B, Dana Sta. 1161 IV, 20 mm ML, ventral view; c, Dana Sta. 1157 I, 56 mm ML,

dorsal view of holotype; D, Dana Sta. 1161 IV, 20 mm ML, left tentacular club; E, Dana Sta.

4009 II, 5 mm ML, ventral view of larva. Discoteuthis species: F, Dana Sta. 1335 XV, 9 mm

ML, ventral view of larva.
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 7.—Discoteuthis discus: A, Dana Sta. unknown, 45 mm ML, funnel-mantle locking apparatus; B, holotype, Pillsbury Sta. 36, 53 mm ML, nuchal locking apparatus; G, holotype, funnel

organ and funnel valve; i, holotype, radula. Discoteuthis laciniosa: c, funnel-mantle locking

apparatus; D, paratype, Dana Sta. 1157 I, 56 mm ML, nuchal locking apparatus; j , paratype,

radula. Cycloteuthis sirventi, Gerda Sta. 212, 42 mm ML: E, funnel-mantle locking apparatus;

F, nuchal locking apparatus; H, funnel organ and funnel valve; K, radula.
NUMBER 5 23

PLATE 8.—Gladii (ventral views and cross sections) and upper and lower beaks: A, D, Discoteuthis

discus, holotype, Pillsbury Sta. 36, 53 mm ML; B, E, D. laciniosa, paratype, Dana Sta. 1157 I,

56 mm ML; c, F, Cycloteuthis sirventi, Gerda Sta. 212, 42 mm ML.
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 9.—Discoteuthis discus, Dana Sta. unknown, 45 mm ML, suckers: A, G, from manus of

tentacular club; B, from dactylus; C-F, from arms I—IV respectively. Discoteuthis laciniosa,

paratype. Dana 1157 I, 56 mm ML, suckers: H, N, from manus; i, from dactylus; J-M, from

arms I-IV respectively. Cycloteuthis sirventi, Gerda Sta. 212, 42 mm ML, suckers: o, from

manus; p, from dactylus; Q-T, from arms I-IV respectively; u, from manus showing striated outer

chitinous ring.

U. S. GOVERNMENT PRINTING OFFICE : 1969 O - 328-974






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