RICHARD E. TO

and CLYDEF. E. R'

A Monograph of the

Cephalopoda of the

W North Atlantic:

The Family

Joubiniteuthidae i

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • 1969 NUMBER 15

SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 15

Richard E. Young A Monograph of the

and Clyde F. E. Roper Cephalopod a Q f

North Atlantic:

The Family

Joubiniteuthidae

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1969
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ABSTRACT

Joubiniteuthis portieri (Joubin, 1912) is redescribed on the basis of new material

from the Atlantic Ocean. Valdemaria danae Joubin, 1931, is synonymized with

Joubiniteuthis portieri, and the peculiar hectocotylus previously described for V. danae

is shown to be an artifact. The validity of the family Joubiniteuthidae Naef is confirmed. The Joubiniteuthidae is thought to be most closely related to the Mastigoteuthidae and Promachoteuthidae.

Introduction

Joubin (1912) established Chiroteuthis portieri, which

he characterized only by the great length of the

arms. Later, the species was more completely described

by Joubin (1916). Only a single mutilated specimen

was available, and in spite of its peculiar characters,

Joubin felt that it should be placed in Chiroteuthis

within the Chiroteuthinae. Berry (1920), however,

considered Joubin's species distinct enough to merit

the establishment of a separate genus, Joubiniteuthis.

Shortly afterward, Naef (1922, p. 299) created the

family Joubiniteuthidae without giving reasons other

than his belief that Joubiniteuthis belonged in no existing family. Joubin (1924) repeated his earlier description of the species and maintained its position in

Chiroteuthis, apparently unaware of the changes introduced by Berry and Naef. Grimpe (1925) listed the

Joubiniteuthidae between the Chiroteuthidae and

Grimalditeuthidae.

In a preliminary report on the cephalopods collected

by the Dana, Joubin (1931) described Valdemaria

danae, a new genus and species, which he placed in the

Mastigoteuthinae. The peculiar hectocotylus of a second specimen of V. danae was described by Joubin in

1933. Thiele (1935, p. 977) gave a brief diagnosis of

the Joubiniteuthidae, which he listed after the Grimalditeuthidae and before the Cranchiidae; he also

briefly characterized Valdemaria, which he retained in

the Mastigoteuthinae.

Recently Clarke (1966) listed Chiroteuthis portieri

in the Chiroteuthinae, apparently unaware of the nomenclatural changes that have occurred. He also listed

Valdemaria danae under the Chiroteuthinae but gave

Richard E. Young, University of Hawaii, Honolulu, Hawaii

96822. Clyde F. E. Roper, Department of Invertebrate Zoology, Smithsonian Institution, Washington, D.C. 20560.

no reason for this change from the Mastigoteuthinae.

The collections of cephalopods at the Institute of

Marine Sciences, University of Miami, contain material that conforms to the descriptions of Joubin's two

species. Joubin's male specimen of V. danae is also in

these collections. The study of this material has shown

that Valdemaria danae is a synonym of Joubiniteuthis

portieri and that the Joubiniteuthidae is a valid family.

The coauthors wish to acknowledge E. Bertelsen of

the Carlesberg Foundation for the loan of the R/V

Dana material, H. Bullis for the R/V Silver Bay material, and G. E. Maul for the loan of the specimen

from Madiera. We also would like to thank G. L. Voss

for his support and his comments on the manuscript.

The illustrations have been prepared by Miss Constance Stolen to whom we express our gratitude. The

work was supported in part by research grants from

the National Science Foundation, GB-57294, and the

National Geographic Society-University of Miami

Deep-Sea Biology Program. This article continues the

series of papers which was initiated by G. L. Voss

(1962) on the families of North Atlantic Cephalopoda.

Family JOUBINITEUTHIDAE Naef, 1922

JOUBINITEUTHIDAE Naef, 1922, p. 299; 1923, p. 795.—

Grimpe, 1925, pp. 97, 103.—Thiele, 1935, p. 977.—Young

and Roper, 1968, p. 197, et seq.

DIAGNOSIS.—Buccal connectives attach to ventral

borders of arms IV; funnel locking-cartilage oval without tragus or antitragus; conus elongate with ventral

fusion; tail elongate, filiform, longer than mantle

length; arms I—III extremely long with six longitudinal rows of suckers; ventral arms much shorter and

with four longitudinal rows of suckers; tentacular clubs

laterally compressed with 8-12 suckers in a transverse

row; no photophores.

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Genus Joubiniteuthis Berry, 1920

Chiroteuthis Joubin, 1912, p. 396.

Joubiniteuthis Berry, 1920, p. 152.—Naef, 1921, p. 147; 1922,

p. 299.—Thiele, 1935, p. 977.—Young and Roper, 1968,

p. 201.

Valdemaria Joubin, 1931, p. 176; 1933, p. 44.

TYPE-SPECIES.—Joubiniteuthis portieri (Joubin,

1912). [By monotypy.]

Joubiniteuthis portieri (Joubin, 1912)

Chiroteuthis portieri Joubin, 1912, p. 396; 1916, p. 1; 1924,

p. 80.—Clarke, 1966, p. 203.

Joubiniteuthis portieri, Berry, 1920, p. 152.—Naef, 1922, p.

299; 1923, pp. 237, 810.—Grimpe, 1925, pp. 97, 103.—

Thiele, 1935, p. 977.

Valdemaria danae Joubin, 1931, p. 176; 1933, p. 44.—Clarke,

1966, p. 206.

Sex

M

M i

M

?

F

juv.

ML, mm

105

89

74

94

70

34

List

Sta. no.

Pillsbury 344

Dana 1185 XIII

Dana 3978 VI

Silver Bay 3729

Silver Bay 3738

Aphanopus ca'bn stomach,

of Material

Location

09°26'N

17°41'N

30°15'S

29°24'N

29°58'N

Madeira

77°36'W

60°58'W

13°15'E

80°08'W

80°10'W

10

27

13

3

5

4

Date

VI

XI

XI

II

II

I

1966

1921

1930

1962

1962

1960

Estimated

depth, m

800-900

2,000

2,500

350

330

-

1

 Joubin's specimen (1933).

DESCRIPTION.—The mantle is long and slender (Figures 1A, D) ; its walls are heavily invaded with gelatinous tissue. The muscular portion of the mantle terminates in the region of the anterior one-third of the

fins. The small fins are thick and gelatinous medially

but are thin around the free edges. They are slightly

longer than broad and are oval in outline; no free

lobes occur (see measurements in Table 1). Posteriorly

the fins terminate abruptly. The extremely long, slender tail is slightly longer than the length of the mantle.

The needle-like tail tapers gradually to a sharp point;

for its full length it is primarily comprised of the

elongate conus of the gladius which is covered only by

soft, thickened integument.

The gladius (Figure 1B) is an extremely long, thin,

tapering rod. The rhachis is free for nearly one-third

the length of the gladius. The vane is weakly developed ; it folds ventrally from its point of origin, and

the edges fuse in the posterior one-third of the gladius

to form a long conus.

The funnel extends to the level of the middle of the

eyes. Only the short, narrow exhalent tip is free, and

the remainder of the funnel is embedded in the soft

tissue of the head. The funnel is thick-walled and has a

semigelatinous consistency. Two pairs of muscular

bridles connect the anterior portion of the funnel to

the head. The funnel locking-cartilage is small and

oval; it consists of a deep, oval concavity that deepens

posteriorly and undercuts the posterior edge of the

cartilage. A membranous flange encircles the cartilage.

Both tragus and antitragus are lacking. The bulbous

mantle component protrudes conspicuously from the

inner surface of the mantle (Figure 2D). The dorsal

member of the funnel organ is an inverted U-shaped

pad; the limbs are broad and rounded. The surface

of the organ has convolutions. A low, median ridge on

the organ extends anteriorly as a short, spatulate, apical papilla. The ventral members are small, oval pads

(Figure 2F) . A funnel valve is present.

The head is elongate and cylindrical; it bears

moderate-sized eyes. The finger-like olfactory papillae

arise at the back of the head just lateral to the anterior

portion of the funnel.

The arms are extraordinarily long, slender, and

whiplike. The bases of the arms are broad and robust.

The arms taper gradually and terminate in minute,

blunt, or conical tips. Arms I—III are nearly subequal

and are two to four times longer than the length of the

mantle (exclusive of the tail). The fourth arms are

relatively short and slender, approximately 80 percent

as long as the mantle.

A well-developed web joins the bases of arms I—III:

it is not present between arms IV and it is very low between arms III and IV. Protective membranes are

very low and weakly developed on all the arms. Near

the bases of the arms the membranes are apparently
NUMBER 15

i

FIGURE 1.—Joubiniteuthis portieri [Dana 1185 XIII, ML 89 mm): A, Ventral aspect.

B, Gladius, entire, and cross sections, c, Tentacular club, D, Dorsal aspect.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 2.—Joubiniteuthis portieri (Dana 1185 XIII, ML 89 mm): A, Oral surface of anomalous right arm IV. B, Aboral surface of anomalous right arm IV. c, Nuchal locking cartilage.

D, Funnel and mantle locking-cartilages, E, Radula. F, Funnel organ and funnel valve, o, Upper

and lower beaks.
NUMBER 15

%i

"a

FIGURE 3.—Joubiniteuthis portieri (Dana 1185 XIII, ML 89 mm): A, Right arm III. B, Left

arm IV. Suckers from arm I: c, from fifth row; G, taken 4 cm from arm base; K, taken 6.5 cm

from arm base. Suckers from arm II: D, from sixth row; H, taken 4 cm from arm base; L, taken

6.5 cm from arm base. Suckers from arm III: E, from fifth row; i, taken 4 cm from arm base;

M, taken 6.5 cm from arm base. Suckers from arm IV: F, from fifth row; j , taken 4 cm from

arm base; N, taken 6.5 cm from arm base, o, Sucker from tentacular club, p, Sucker from

arm III taken 6.5 cm from arm base showing outer ring.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

entirely absent, but minute bumplike trabeculae are

present.

Suckers are biserially arranged at the bases of arms

I—III, but they increase to four, then to six, often irregularly arranged rows that appear to extend to the

tips of the arms (Figure 3A). The closely packed, irregular arrangement of suckers near the arm tips, however, makes it difficult to determine the exact number

of rows. The suckers on arms IV originate in two rows,

then quickly increase to four rows which remain

distinct nearly to the tips of the arms (Figure 3B) . The

arm suckers are small and extremely numerous. The

diameter of the suckers on all arms decreases as the

number of rows increases. The medial suckers are one

and one-half to two times the diameter of the marginal

suckers along most of each arm length. The basal

suckers that lie in two rows are large with small

apertures; the most proximal few suckers have smooth

sucker rings. The next suckers bear small pointed

teeth; farther distally the teeth are larger and number

between seven to nine on the distal two-thirds of the

ring. At the point where the number of sucker rows

reaches six, the suckers decrease in size and bear six or

seven pointed to truncate teeth on the distal margin

(Figures 3C-N, P) . The suckers again increase in size

toward the middle of the arms.

The tentacles are long and threadlike; the stalks are

naked. The club is elongate (about half as long as the

mantle length exclusive of the tail) and laterally compressed. The oral face is rounded and no division into

dactylus, manus, and carpus occurs. The club is covered with numerous, minute, closely packed suckers

that arise proximally in one or two rows, then increase

to 5-6 transverse rows; 8-12 rows span the width of

the club in its distal two-thirds. The tip of the club appears to be the growing area where new suckers are

formed. The distal fifth or sixth of the club is bordered

by a distinct protective membrane with no apparent

trabeculae. Protective membranes are lacking elsewhere on the club. The suckers have small apertures

and smooth rings (Figure 3c).

The buccal membrane has seven lappets. The indistinct buccal connectives are embedded in gelatinous

tissue and attach to the dorsal sides of arms I and II

and to the ventral sides of arms III and IV. The distal

area of the oral face of the buccal membrane bears

numerous long, finger-like papillae dorsally that decrease in number and size ventrally. Papillae are lacking on the proximal region of the membrane, particuFIOURE 4.—Spermatophore from mature male

(Pillsbury 344).
NUMBER 15

TABLE 1.—List of measurements of Joubiniteuthis portieri (Joubin 1912)

Station no.

Sex

Mantle length

Tail length

Mantle width

Head length

Head width

Fin length

Fin width

ARMS I

II

III

IV

Tentacle length

Club length

Pillsbury 344

R

200

240

240

70

-

105

155

-

30

-

-

-

L

200

220

220

70

-

Dana

R

172

163

175

1185 XIII

d

1

89

122

20

22

17

27

26

47 +

157

40

L

165

163

183

64

-

Dana 3978 VI

R

174

200

187

53

-

d

1

74

112

19

21

13

23

22

L

190

220

187

64

-

Silvei

R

220

240

230

78

-

Bay 3729

?

94

101

16

29

17

26

24

L

215

-

-

75

-

—

Silver Bay 3738

9

70

83

12

22

10

19

15

R L

-

175

195

-

-

R

86

135

115

31

90

20

Madeira

juvenile

34

51

6

12

5

11

11

L

100

114

-

31

-

larly ventrally where only small crenulations occur.

The lips are thick and bear many low grooves and folds

but lack papillae.

The beaks are illustrated on Figure 2G. The upper

beak has a slender, sharply pointed rostrum and a long,

triangular hood; the insertion plate of the palatine lamella is long. The rostrum of the lower beak is blunt

and triangular; no ridge occurs across the gular

lamella.

The radula consists of seven rows of teeth in a transverse series (Figure 2E). The rhachidian tooth has a

broad, blunt median cusp and two small, pointed lateral cusps. The first and second lateral teeth have

broad, blunt primary cusps that are only slightly smaller

than the primary cusp of the rhachidian tooth; small,

sharp lateral cusps occur on the first lateral teeth. The

third lateral teeth are long, slender, and pointed. Rectangular marginal plaques are present.

Photophores are absent.

The males lack hectocotylization. In the mature

male the penis is very large and reaches about the level

of the anterior end of the funnel. A spermatophore is

illustrated in Figure 4.

The surfaces of the mantle, fins, funnel, head, and

arms bear small brown or purplish-brown chromatophores. The oral surface of the brachial crown, and

particularly the buccal region, are pigmented with a

dull purple color.

LOCATION OF TYPE.—Oceanographic Museum,

Monaco? (not verified).

TYPE-LOCALITY.—29°03'N, 16°08'30"W, in the region of the Salvage Islands.

DISTRIBUTION.—The geographic distribution of all

known records of /. portieri is plotted in Figure 5. All

specimens were captured in open nets. The type of /.

portieri was taken at 0-3500 m; the type of V. danae

came from an estimated depth of 500 m; the present

specimens came from estimated depths that range from

330 m to 2500 m.

Discussion

COMPARISON OF Joubiniteuthis portieri AND Valdemaria danae.—Joubin's lengthy descriptions of Joubiniteuthis portieri (1916, 1924) and Valdemaria danae

(1931, 1933) give few characters that can be used to

separate these forms, and it is difficult to understand

why he placed the two species in separate families (his

subfamilies). The apparent differences, which Joubin

may have considered important, can be easily reconciled. Two distinctive features of Valdemaria, the very

elongate tail and the peculiar tentacles, were missing

in the badly damaged specimen of Joubiniteuthis portieri. (The original illustrations of V. danae and /. portieri are reproduced in Figure 6). Although Joubin

described a funnel valve in V. danae, he was unable

to detect one in /. portieri, also undoubtedly due to its

deteriorated condition. The lengths of arms I—III differ

considerably in the two descriptions. The type of /.

portieri has arms three to four times longer than the
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

MO" 90' 80* 70" 60' SO- SO* 20" 10 0- » " 20* 30"

_ 500 m line

XT Zf 2ff JO*

FIGURE 5.—Geographical distribution of Joubiniteuthis portieri based on all known records.

length of the mantle, while the type of V. danae has

arms about two and one-half times the mantle length.

The lengths of arms I—III in our specimens show variation from two to nearly four times greater than the

length of the mantle, depending on the condition of

the specimen. The differences in the lengths of the arms

of Joubin's specimens clearly are of no significance. The

only remaining difference between the two nominal

species concerns the number of rows of suckers on

arms IV. V. danae is described as having only two rows

of suckers on each of these arms with additional, irregularly interspersed suckers (Joubin, 1931). In the subsequent description of V. danae (1933), however, Joubin stated that all of the arms except the hectocotylus

have four to six rows. Our examination of the same

specimen {Dana 1185 XIII) shows the ventral arms

to have a maximum of four rows of suckers. Joubin

described six rows of suckers in the midportion of the

ventral arms of /. portieri. Specimens that are not in

perfect condition, however, generally have suckers that

are very irregularly arranged along the twisted, contracted, serpentine arms, and it is difficult to determine

the exact number of rows. Poor condition of the specimen also probably accounts for the discrepancy in the

small number of rows of suckers reported at the tips

of the arms in /. portieri.

We conclude, on the basis of the well-preserved

material available and of the type descriptions, that

Valdemaria danae is synonymous with Joubiniteuthis

portieri. We have been unable to examine the types.

Joubin (1933) described a peculiar S-shaped hectocotylus in Valdemaria danae. We have examined this

specimen {Dana 1185 XIII) and an illustration of the

modified arm appears on Figures 2A, B. The arm is
NUMBER 15

FIGURE 6.—A, Valdemaria danae (reproduced from Joubin, 1931). B, Chiroteuthis portieri

(reproduced from Joubin, 1916).

unusual, and at first we, too, thought that it could

represent a form of hectocotylization. When we examined two other male specimens, one of which was

sexually mature, and found that no such modification

existed, it became apparent that Joubin's specimen

had an anomalous arm. Probably the arm was injured while the specimen was living and had begun

to heal prior to the capture of the squid.

FAMILIAL RELATIONSHIPS.—The peculiarities of

Joubiniteuthis clearly confirm that the inclusion of this

genus in a separate family is fully warranted. Although

the family is distinctive, some evidence for its relationship with other families can be found. Because the

buccal connectives attach to the ventral borders of

arms IV, the Joubiniteuthidae is aligned with the

following families that have the same type of attachment: Onychoteuthidae, Gonatidae, Cycloteuthidae,

Thysanoteuthidae, Ctenopterygidae, Octopoteuthidae,

Brachioteuthidae, Lepidoteuthidae, Batoteuthidae,

Chiroteuthidae, Mastigoteuthidae, Promachoteuthidae, Grimalditeuthidae, and Cranchiidae.

The presence of a conus formed by the ventral fusion of the lateral edges of the vane occurs in the

Joubiniteuthidae, Cycloteuthidae, Brachioteuthidae,

Lepidoteuthidae, Batoteuthidae, Mastigoteuthidae,

Chiroteuthidae, Grimalditeuthidae, and Cranchiidae.

(The Promachoteuthidae, although related to these

families, apparently lacks a gladius [Roper and Young,

1968].) The oval funnel locking-cartilage more closely

relates the Joubiniteuthidae to the Chiroteuthidae,

Mastigoteuthidae, and Promachoteuthidae than to any

of the other families. The numerous small suckers in

many rows on the tentacular clubs are similar to the

condition in the Mastigoteuthidae and Promachoteuthidae. The Joubiniteuthidae probably is most closely

related to these two families.
10

Literature Cited

Berry, S. S.

1920. Light Production in Cephalopods, I: An Introductory Survey. Biological Bulletin, 38(3): 141—

169.

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 Cephalopoden-fauna der Nordsee.

Wissenshajtliche Meeresuntersuchungen, Kiel, 16

(3):1-214.

Joubin, L.

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

le Prince de Monaco. Compte Rendu de I'Academie

des Sciences, Paris, 154:395-397.

1916. Etudes preliminaires sur les Cephalopodes recueillis

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

Monaco, 4"; Note: Chiroteuthis portieri. Bulletin

Institut Oceanographique, Monaco, 317:1-10.

1924. Contribution a l'etude des C6phalopodes de

1'Atlantique Nord, 4e series. Resultats des Campagnes Scientifiques du Prince de Monaco, 67: 1-

113, 16 plates.

1931. Notes prelirninaires sur les Cephalopodes des croiSMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

sieres du "Dana" (1921-1922), 3e Partie.

Annales de I'Institut Oceanographique, new series,

10 (7): 167-211.

1933. Notes prelirninaires sur les Cephalopodes des croisieres du "Dana" (1921-1922), 4C

 Partie. Annales

de I'Institut Oceanographique (n.s.), 13(1) : 1-49 .

Naef, A.

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

Fischer.

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

1968. The Promachoteuthidae (Cephalopoda: Oegopsida), I: A Reevaluation of Its Systematic Position

Based on New Material From Antarctic and Adjacent Waters. Antarctic Research Series, 11:

203-204.

Thiele, J.

1935. Handbuch der systematischen Weichtierkunde.

2(3): 779-1022. Jena: Gustav Fischer.

Voss, G. L.

1962. A Monograph of the Cephalopoda of the North

Atlantic, I: The Family Lycoteuthidae. Bulletin

of Marine Science of the Gulf and Caribbean,

12(2): 264-305.

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.

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In the bibliography (usually called "Literature Cited"), spell out book, journal, and article

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