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FIELDIANA: ZOOLOGY

A continuation of the
ZOOLOGICAL SERIES

of
FIELD MUSEUM OF NATURAL HISTORY

VOLUME 39

NATURAL HISTORY SURVEY
FEB 11 1971

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'*» HISTORY >\

FIELD MUSEUM OF NATURAL HISTORY
CHICAGO, U.S.A.

•° FIELDIANA • ZOOLOGY

Published by
CHICAGO NATURAL HISTORY MUSEUM

Volume 39 November 17, 1960 No. 40

Bermudan Cephalopods1

Gilbert L. Voss

The Marine Laboratory, University of Miami

The cephalopods reported upon here were collected off Bermuda
during the summer of 1948 by an expedition of Chicago Natural His-
tory Museum (CNHM) based at the Bermuda Biological Station.
The collections were made by means of the Woods Hole Oceano-
graphic Institution's research vessel Caryn. A variety of nets was
used, none of which were of the closing type. As a result, definite
statements as to the depth of capture cannot be made. The depth
fished has been calculated as one third of the amount of wire out for
those stations not listed by Grey (1955), from which the station list
was partly compiled.

A brief resume' of the Bermudan cephalopods is given for the sake
of completeness. Since the Bermudas are oceanic islands located far
from other land masses, the boundaries have been set to contain that
area within a circle whose radius is 200 miles.

I am indebted to Dr. Fritz Haas, Curator of Lower Invertebrates,
Chicago Natural History Museum, for permission to study the collec-
tions, and for providing useful data. The collection consists of 50
specimens representing 21 species, of which three are new to science.
Only three of these species have previously been recorded from
Bermuda. This study has been supported by a grant-in-aid from the
National Science Foundation (NSF-G-5853) which I gratefully
acknowledge.

FORMER RECORDS

Apparently the first record of a cephalopod from the Bermuda
Islands was given by Verrill (1880a) . He listed three species collected
on the beach by G. Brown Goode: Stenoteuthis pteropus, Sepioteuthis
sepioidea, and Loligo pealei. In his report of the Challenger expedition,
which touched at the islands, Hoyle (1886) named a new species of

1 Contribution No. 276 from The Marine Laboratory, University of Miami.
Library of Congress Catalog Card Number: 60-531^59
No. 900 419

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420 FIELDIANA: ZOOLOGY, VOLUME 39

Octopus, 0. bermudensis, from the reefs, and also listed a squid,
Onychia caribaea, from Station 30, between St. Thomas and the
Bermudas, but Peile (1926) states that this station was actually near
Bermuda and he includes it in his list. Heilprin (1888) described
Octopus chromatus from a specimen collected at Flatts, and also re-
ported that Octopus vulgaris was seen but not collected. In his mono-
graphic revision of the Oegopsida Pfeffer (1912) listed three species
from the area: Onychoteuthis banksi, Onychia caribaea and Brachio-
teuthis riisei. In the report of the material collected by the United
States survey ship Bache, Berry (1920) described a new species of
cranchiid, Teuthowenia corona, from near the islands (32° 29'N., 68°
22'W., 0-100 meters).

A paper published by Peile (1926) contains a short section on
cephalopods. He lists nine species: Spirula australis, Loligo pealei,
Sepioteuthis sepioidea, Onychia caribaea, Stenoteuthis bartrami, S.
pteropus, Octopus rugosus, O. bermudensis (= chromatus Heilprin), and
Argonauta argo var. americana Dall. To this may be added two
species described by Joubin (1931, 1935) : Octopodoteuthis danae and
Egea inermis.

Two other works must be mentioned, both by Pickford. In 1945
this author published her revision of the littoral octopods of the west-
ern Atlantic. Two species are recorded : Octopus vulgaris and O. mac-
ropus. Later, in 1950, in her work on the Vampyromorpha she listed
Vampyroteuthis infernalis from Bermudan deep waters.

For some years Mr. Louis Mowbray, Curator of the Government
Aquarium, Flatts, has sent me small collections of cephalopods from
Bermuda. In one of these collections was a number of small loliginids
which were identified as Doryteuthis plei. This is the first record of
this tropical species from the islands. I am not at all certain that the
Loligo pealei recorded by Verrill did not belong to Doryteuthis, as I
have never seen Loligo pealei among the collections in my possession.

A list of the Bermudan cephalopods follows. All of those species
marked with an asterisk are new records for the area, as nearly as can
be determined from the literature.

Subclass Coleoidea
Order Sepioidea

Family Spirulidae

1. Spirula spirula (Linnaeus)
Order Teuthoidea

Suborder Myopsida
Family Loliginidae

2. Loligo pealei Le Sueur

3. Sepioteuthis sepioidea (Blainville)

4. *Doryteuthis plei (Blainville)

•

Mat. V- . Sur/ev

VOSS: BERMUDAN CEPHALOPODS

421

Suborder Oegopsida

Family Enoploteuthidae

5. *Abraliopsis morisii (Verany)

6. *Pyroteuthis margaritifera (Ruppell)

7. *Pterygioteuthis giardi Fischer
Family Octopodoteuthidae

8. Octopodoteuthis danae Joubin
Family Onychoteuthidae

9. Onychia caribaea Le Sueur

10. Onychoteuthis banksi (Leach)
Family Bathyteuthidae

11. *Bathyteuthis abyssicola Hoyle

12. *Cienopteryx siculns (Verany)
Family Histioteuthidae

13. *Calliteuthis celetaria, n. sp.

14. *Calliteuthis sp.
Family Mastigoteuthidae

15. *Mastigoteu(his spp.
Family Ommastrephidae

16. Ommastrephes pier opus Steenstrup

17. 0. bartrami Lesueur
Family Brachioteuthidae

18. Brachioteuthis riisei Steenstrup
Family Cranchiidae

19. *Leachia cyclura Le Sueur

20. *Galileuthis armata Joubin

21. *Megalocranchia megalops (Prosch)

22. *Megalocranchia papillata, n. sp.

23. Megalocranchia corona (Berry)

24. *Carynoteuthis oceanica, n. sp.

25. Egea inermis Joubin

26. *Bathothauma lyromma Chun

Order Vampyromorpha

27. Vampyroteuthis infernalis Chun
Order Octopoda

Family Octopodidae

28. Octopus vulgaris Cuvier (=0. rugosus)

29. O. macropus Risso (=0. chromatus Heilprin and 0. bermudensis

Hoyle)
Family Bolitaenidae

30. *Eledonella pygmaea Verrill

31. *Japetella diaphana Hoyle
Family Vitreledonellidae

32. *Vitreledonella richardi Joubin
Family Argonautidae

33. Argonauta argo Linnaeus (=var. americana Dall)

A few remarks on habitat seem worth while. The only bottom
dwellers are Octopus vulgaris and 0. macropus, the "rock" and "grass"
scuttles, respectively. They live in shallow water and are both found
on the mainland, vulgaris from as far north as Long Island Sound,
macropus from southern Florida and the Bahamas.

422 FIELDIANA: ZOOLOGY, VOLUME 39

All of the loliginids, Loligo pealei, Sepioteuthis sepioidea, and Dory-
teuthis plei, are coastal species, seldom being found far from land.
While L. pealei occurs as far north as Cape Cod, S. sepioidea and D.
plei are both tropical species, the former found along the mainland
only as far north as Cape Canaveral on the Florida coast and the
latter recorded as far north as Sea Island, Georgia {Pelican record).
They undoubtedly were introduced from the south.

All of the remaining species are inhabitants of the open ocean,
occurring far from land. Their occurrence at Bermuda was expected
and the list will grow materially as more net collecting is carried out
in the vicinity of the islands. A few are truly deep species, as will be
noted under the individual records, but most are commonly found in
the upper surface, either dwelling there in the young stage or migrat-
ing upward to it during the early evening.

SYSTEMATIC SECTION

Family ENOPLOTEUTHIDAE

Abraliopsis morisii (Verany, 1837)

Onychoteuthis Morisii Verany, 1837, Mem. Accad. Torino, 1: 2.

Material— One male, mantle length 26.5 mm.; CNHM no. 78299;
haul 41.

This beautifully preserved specimen has retained its rich reddish
brown color, among which the light organs on the ventral surface of
the mantle, funnel, head and arms are plainly visible. The right eye-
ball is protruding and bears five small round photophores in a row on
the ventral surface, the terminal organs about twice as large as the
inner three. The right ventral arm is hectbcotylized and the penis
contains numerous spermatophores.

This species may be easily distinguished from the other Atlantic
enoploteuthids by the large, black, swollen three-parted photophores
on the distal part of the ventral arms. It is an upper bathypelagic
species, occurring from the surface to a depth of about 500 meters.

Pyroteuthis margaritifera (Ruppell, 1844)

Enoploteuthis margaritifera Ruppell, 1844, Intorno ad alcuni cefalopodi del
mare di Messina, p. 129.

Material. — One female, mantle length 11.5 mm.; CNHM no. 78300;
haul 30.

This is another bathypelagic species, dwelling apparently in the
upper 600 meters. It is commonly distributed over the Atlantic in

VOSS: BERMUDAN CEPHALOPODS 423

temperate and warm waters. Development is incomplete in this
specimen and for full details Chun's monograph (1910) of the Val-
divia expedition, with colored plates, is unexcelled.

Pterygioteuthis giardi Fischer, 1896

Pterygioteuthis giardi Fischer, 1896, Jour. Conch., 43: 205.

Material. — One juvenile, mantle length 8.5 mm.; CNHM no. 78301;
haul B30.

Only one specimen of this common bathypelagic species was cap-
tured. The definitive number of photophores on the eyeball has not
yet developed, and the tentacular club is immature in appearance.
This species has been exquisitely illustrated by Chun (1910) in the
Valdivia monograph. It probably lives between the surface and about
500 meters, or near the limit of light penetration.

Family BATHYTEUTHIDAE

Bathyteuthis abyssicola Hoyle, 1885

Bathyteuthis abyssicola Hoyle, 1885, Rep. Sci. Res. Voy. Challenger, 1: 272.

Material.— One juvenile, mantle length 7.0 mm.; CNHM no. 78302;
haul 50.

Only one specimen of this widespread species was captured. It is
somewhat faded, only the arms retaining the deep reddish color.
The light organs, six in number, are present on the base of each of
the three upper pairs of arms. They appear as long, oval, dark
bodies with light centers.

The characteristic color, the protruding and forward-directed eyes,
the separate terminal fins, and the six light organs easily serve to
identify this interesting deep sea species. It occupies considerable
depths, probably from near the surface to 1500 meters or more.

Ctenopteryx siculus (Verany, 1851)

Sepioteuthis siculus Verany, 1851, Mem. Accad. Torino, 1: 51.

Material. — Two juveniles, mantle length 9.0 and 11.0 mm.;
CNHM no. 78303; haul 23.

This species is widely represented in deep hauls and is easily rec-
ognized by the peculiar comb-like structure of the fins. In the early
larvae the fins are small and terminal, but they begin to grow ante-
riorly and soon show strong lateral supports and transverse muscle
bundles connected by a thin, delicate membrane, the flat surface of

424 FIELDIANA: ZOOLOGY, VOLUME 39

the fins. In almost all cases the membrane is torn between the sup-
ports, leaving a curious comb or fringe. In adults the fins attain the
full length of the mantle. Although closely related to Bathyteuthis,
Ctenopteryx appears to be limited to the upper 400 meters of the sea.

Family ONYCHOTEUTHIDAE

Onychoteuthis banksi (Leach, 1817)
Loligo banksi Leach, 1817, Zool. Misc., 3: 141.

Material. — One juvenile, mantle length 6.5 mm.; CNHM no.
78304; haul 46. One juvenile, mantle length 11.5 mm.; CNHM
no. 78305 ; haul 30. One juvenile, mantle length 8.0 mm. ; CNHM no.
78306; haul 48.

This species of cephalopod, above all others, may be called cos-
mopolitan. It is known in all oceans from as far north as Hammer-
fest, Norway, to the Southern Ocean, and is found in nearly all
collections. It often "flies" onto the decks of ships at sea. Oceanic
in habitat, it is usually found from the surface to about 150 meters.
The young may easily be identified from all other squids by the sleek,
compact appearance with partially withdrawn head, nearly terminal
fins, beyond which projects the sharp, tapered, slightly curving and
transparent conus of the gladius, and the dark streak along the dorsal
mid-line of the mantle composed of the visible rib of the gladius with
a streak of closely set brown chromatophores over it.

Family HISTIOTEUTHIDAE

Calliteuthis celetaria, new species. Figure 73.

A single specimen of this species is available to me. It differs so
strikingly from other known species of the genus Calliteuthis that it
is here referred to a new species.

The mantle (fig. 73, a) is thick, firm, and more elongate than in most other
species of the genus. Its greatest width is at the anterior margin; the sides are
somewhat parallel. The posterior end tapers to a blunt point. The anterior mar-
gin is slightly produced dorsally and excavated ventrally below the funnel, with
sharp lateral angles.

The fins are large, longer than half the mantle length, with their greatest width
in the posterior third. The notch between the fins is deep, starting at the mantle
tip, but the edges of the lobes extend well beyond the mantle.

The funnel is small and strong, without a groove and with two dorsal supports.
The funnel-mantle locking apparatus is typically calliteuthid, the mantle member
crescent-shaped, heavier posteriorly. The funnel organ (fig. 73, 6) is normal. The

Fig. 73. Calliteuthis celetaria, new species, a, Ventral view of holotype; man-
tle length 39.0 mm. b, Funnel organ, c, Nuchal folds and olfactory crest, d, Left
tentacular club, e, Horny ring of large tentacular sucker. /, Horny ring of postero-
ventral tentacular sucker, g, Light organs around right eye. h, Light organs
around left eye.

425

426 FIELDIANA: ZOOLOGY, VOLUME 39

dorsal member is A-shaped with long arms and either a pointed apex or a short
papilla (damaged). It has no ridges or flaps and is fleshy but not puffy. The ven-
tral pads are long ovals.

The head is large, slightly wider than the mantle and flattened dorsally. The
left eye is about twice the size of the right eye, with a correspondingly larger eyelid
opening. There appears to be no sinus on either eyelid. There are two nuchal
folds (fig. 73, c) on each side of the head. The dorsal fold is smoothly curved down-
ward, terminating in a simple point. The ventral fold is also the olfactory crest and
turns smoothly upward, terminating posteriorly in a raised bar-like olfactory organ.

The arms are long and stout, in the order 3.2.1.4, with arms III and II nearly
equal in length. Arms I and II were slightly damaged so that it could not be deter-
mined whether there were swimming keels on them or not. On arm III the keel
originates just above the base of the arm, broadens to a high ridge at about the
middle of the arm, and then decreases to a low ridge. The skin of the ends of the
arms was damaged, so it is not known whether the keel extended to the arm tip or
not. The arm suckers are biserial, larger than in most other species and nearly
equal in size from the base to near the tip, where they diminish in size over a few
rows and then abruptly become minute. The horny rings of the suckers are smooth
except on arm IV, where a few have several very low round teeth on the distal mar-
gin. The suckers of arm IV are about one-third the size of the suckers of the
other arms.

The tentacles are short and stout, with thick, round stalks and only slightly
expanded clubs (fig. 73, d). There is no cleft on the aboral surface of the club.
The stalks are slightly flattened on the oral surface with a fine groove in the mid-
line. The carpal suckers originate about one club length down the stalk and extend
as a single line of suckers and pads from the ventral side across the mid-line to the
club and then in a straight line to the dorsal edge of the club. There are about
10 rows of subequal suckers on the club with no real disparity in size either upon
the hand or between those of the hand and the attenuated distal portion. The
median hand suckers (fig. 73, e) have about 12 or 13 sharp triangular teeth on the
distal margin. The proximal margin is smooth. The suckers are bordered by a
wide chitinous denticulate collar; on the suckers of the proximal ventral border
(fig. 73, /) this tends to become broadened into a plate, giving the suckers of this
region a triangular appearance. There is a low protective membrane on each side
of the club and an aboral swimming membrane on the distal half of the club.

The buccal membrane bears 7 lappets and 7 supports. The web between the
arms is vestigial.

The photophores are of two types, large and small. On the ventral surface of
the mantle the large light organs are numerous, extending over three-fourths of the
ventral mantle length. Posterior to the large light organs are a few rows of small
photophores. The dorsal surface of the mantle bears numerous small light organs
over all the surface except near the posterior end. The dorsal surface of the head
bears a few small light organs but the ventral surface bears numerous large photo-
phores with a row of four light organs in the median line excluding the organs of
the posterior row. The right eyelid (fig. 73, g) bears a circlet of 17 light organs
closely and regularly arranged around the border with a single organ at the mid-
point on each side and separated from the series. The left eyelid (fig. 73, h) has a
series of 7 large organs in an arc on the anterior border and about 4 or 5 small light
organs in two series separated from the others.

VOSS: BERMUDAN CEPHALOPODS 427

There are light organs on all of the arms. On arms I, II and III there is a
ventral row of large light organs from the base to near the tip. In addition, ob-
scured by the damaged skin, there is a dorsal row of small light organs, few in
number, near the bases of the arms. On arm IV there are three rows of large
organs. The ventral row of light organs, consisting of about 10 photophores, ex-
tends over about three-fourths of the arm length. The middle row consists of
about 10 organs, then a break, and then a group of about 5 light organs diminishing
in size to the tip. The dorsal row of light organs consists of 7 photophores, which
extend over about half the length of the arm.

The color in alcohol is vinous red with a gold sheen underlying the skin.

Holotype. — Chicago Natural History Museum no. 78308. A
female, mantle length 39.0 mm.; from haul 37, 32° 10' N., 34° 45' W.,
730-820 meters, 0405-0838 hours, 35-foot otter trawl, August 5, 1948.

Table 1. — Measurements of Holotype of Calliteuthis celetaria, n. sp.

mm. mm.

Total length 96.3 Arm 1 34.8

Mantle length 39.0 Arm II 36.0

Mantle width 19.9 Arm III 36.2

Head length 19.3 Arm IV 32.8

Head width 22.0 Tentacle length 80.0

Fin length 21.0 Club length 10.2

Fin width 28.0 Sucker diameter (arm II) . . . 2.0

Discussion. — This new species of Calliteuthis, distinct from any
of the other known species, is known only from the type from Ber-
muda. It would be desirable to give diagnostic characters at this
time for separation of this species from those previously known, but
this must wait until the revision of the genus has been completed.
This revision has permitted the clear identification of all of the species
of Calliteuthis of the Atlantic Ocean based upon distinct and in the
main non-variable characters. I must thank my wife, Nancy A. Voss,
who is presently engaged in revising the genus, for pointing out the
distinctness of the present species and for preparing the detailed
illustrations and preliminary descriptions.

The name celetaria is derived from the Greek KeXeraptos (graceful)
and is descriptive of the beautiful little animal here described.

Calliteuthis sp.

Material. — One juvenile, mantle length 15.0 mm.; CNHM no.
78307; haul 55.

A second specimen of Calliteuthis was found in the collections.
It is a juvenile, but sufficient characters are present to distinguish it
from the preceding species. The mantle is cone-shaped, with light
organs on the anterior two-thirds of the ventral mantle surface con-

428 FIELDIANA: ZOOLOGY, VOLUME 39

sisting of about 3 transverse rows of large organs and 3 rows of small
ones. There are a few small ones scattered dorsally.

The head is normal, with the left eye twice as large as the right.
There are 17 light organs around the right eye and there is a sinus.
The funnel is small and contains a typically shaped calliteuthid fun-
nel organ, but the dorsal member bears a fleshy ridge on each limb
which ends posteriorly in a flap. The olfactory organ is a simple
papilla. There are no nuchal folds.

The arms are small and subequal, and bear suckers with a number
of low rounded teeth on the distal borders. The tentacular club has
no aboral cleft. The swimming keel is more than half the club length.
The suckers are in about 6 or 7 rows.

At the present time the identity of this specimen must be left
in doubt.

Table 2. — Measurements of a Juvenile Calliteuthis from Bermuda

mm. mm.

Total length 49.5 Arm 1 22.0

Mantle length 15.0 Arm II 24.1

Head length 9.8 Arm III 25.0

Head width 13.0 Arm IV 21.2

Fin length Tentacle length 36.5

Fin width 12.0 Club length 5.0

Family MASTIGOTEUTHIDAE

Mastigoteuthis spp.

Material— One!, mantle length 25.5 mm.; CNHM no. 78309;
haul 29. One head with arms; CNHM no. 78310; haul 36.

The two specimens listed above both belong to the genus Masti-
goteuthis, a peculiar group of deep-dwelling, soft-bodied squids, some
of which attain a rather large size. Because of the condition of the
specimens, definite identification was not attempted. The complete
specimen, lacking only the tentacles, bears no light organs on the
body but has two pairs of large light organs on the eyes, one postero-
ventral and the other anterior. The end of the body is drawn out
into a narrow point beyond the large fins, which occupy about four-
fifths of the mantle.

The mutilated specimen consists of only the head, 'arms and
neck, but much of the skin is intact and this bears numerous large,
circular photophores on the head and arms.

VOSS: BERMUDAN CEPHALOPODS 429

Family BRACHIOTEUTHIDAE

Bra chioteu this sp.

Material— One?, mantle length 19.0 mm.; CNHM no. 78311;
haul 22.

A single specimen, tentatively identified as a Brachioteuthis, is in
the collections. It is quite small and badly flexed, and the tentacles
are missing. Despite this, the size of the fins and their shape, and
the type of locking mechanism point toward this genus. Pfeffer has
recorded Brachioteuthis riisei from Bermudan waters and the present
specimen may belong to this species.

Family CRANCHIIDAE

Subfamily CRANCHIINAE

Leachia cyclura Le Sueur, 1821

Leachia cyclura Le Sueur, 1821, Jour. Acad. Nat. Sci. Philadelphia, 2: 90.

Material. — One female, gravid, mantle length 45.0 mm.; CNHM
no. 78312; haul 36. One female, gravid, mantle length 55.0 mm.;
CNHM no. 78313; haul 24.

As is typical of this rare species, both specimens lack tentacles,
the basal section only remaining as stumps between arms III and IV.
The mantle bears no pigmentation, but the head region is strongly
colored with reddish brown. The head is small but bears large eyes
whose openings could not be determined in either specimen. The
eyeballs bear four large light organs on the ventral periphery with
another smaller organ halfway between these and the pupil. There

Table 3. — Measurements of Two Specimens of Leachia cyclura Le Sueur

mm. mm.

Total length 67.0 81.0

Mantle length 45.0 55.0

Head width 9.0 10.0

Fin length 11.0 13.0

Fin width 22.0 21.0

Arm 1 4.0 4.5

Arm II 9.5 9.0

Arm III 15.0 15.0

Arm IV 7.5 7.0

is a small, erect olfactory papilla on the ventro-posterior surface of
the covering of the eye. The fins are large, terminal, circular, or
nearly so, in outline. Both specimens are females with small com-

430 FIELDIANA: ZOOLOGY, VOLUME 39

pact ovaries bearing numerous rather large undeveloped eggs. Most
noticeable within the mantle cavity are four large, pigmented bodies,
split at the anterior end, lying rather loosely, two lateral, two ven-
tral, on the visceral mass. They appear to be nidamental glands, but
are of enormous size in comparison to the remaining viscera, and are
united to the mass by only a small section of the posterior end of each.
Little is known of the bathymetric range of this species but it is pre-
sumed to live between 700 to 1000 meters in depth.

Subfamily TAONIINAE

Galiteuthis armata Joubin, 1898

Galiteuthis armata Joubin, 1898, Ann. Sci. Nat. Zool., 6: 279.

Material. — One, sex indet., mantle length 82.0 mm.; CNHM no.
78314; haul 56. One juvenile, mantle length 58.0 mm.; CNHM
no. 78315; haul 22.

The two specimens are in rather poor shape, the mantle greatly
contracted and distorted, and no valid measurements can be obtained.
This species is only infrequently reported in the literature and much
needs to be learned of its developmental history, especially of the
adults and maximum size attained. It apparently lives in the upper
1000 meters. In the older stages it has large hooks on the tentacular
clubs. Probably its most characteristic feature is the unusually long
and finely drawn-out tail, which becomes threadlike toward the
extremity.

Megalocranchia papillata, new species. Figure 74.

The mantle (fig. 74, a) is slender and tubular, perhaps widest near the middle,
and tapers to a blunt point posteriorly. It is united to the head dorsally and on
each side of the funnel. The mantle wall is thicker and more muscular than is
usual in the cranchiids and the gladius does not show through the dorsal wall except
for the anterior end which forms a small rhombic area near the edge of the mantle.
There is a short dark streak between the anterior bases of the fins. The surface
of the mantle is thickly covered with small sharp papillae which are more prom-
inent anteriorly and on the sides and ventral surface of the funnel. They are
accompanied by traces of reddish brown chromatophores.

The fins (fig. 74, b) are very short and broad (1.4 X 7.0 mm.). They project
beyond the end of the mantle and are united to the sides of the gladius point but
fused behind it. The fins are irregular in contour.

The funnel (fig. 74, c) is large and projects far beyond the eyes and reaches
to about the basal fourth of the arms. The dorsal member is trifoliate, with a
slender pointed papilla in each lobe (fig. 74, d). The ventral pads are long and
slender and only slightly curved. There is no valve.

VOSS: BERMUDAN CEPHALOPODS

431

Fig. 74. Megalocranchia papillata, new species, a, Dorsal view of holotype;
mantle length 40.0 mm. b, Ventral view of fins, showing attachment, c, Funnel.
d, Dorsal member of funnel organ, e, Ninth ventral sucker of second left arm.
/, Oral view of left third arm. g, Semi-diagrammatic drawing of spermatophore;
length 7.4 mm. h, Details of horn and mid-organ of spermatophore; length 1.1 mm.

The head is small and somewhat injured. Only the left eye is present. The
eye is of moderate size and in the single specimen has no light organ that can be
distinguished as such.

The arms are short and rather stout. Left I is broken off at the middle and
left II and III are missing entirely. The approximate arm order is 1.3.2.4. All
arms except IV are keeled on their basal half. They are somewhat compressed
with biserial suckers which are bordered by protective membranes. The suckers

432 FIELDIANA: ZOOLOGY, VOLUME 39

of arms I, II and IV are of moderate size, rather closely set and bordered all around
their circumference by minute, closely set sharp teeth (fig. 74, e). The suckers of
III are moderate in size basally but become large very quickly; they have smooth
rings. At about the middle of the arm the suckers become much smaller for a pair
or two but beyond this they are missing except for a single large sucker near the
tip of the arm (fig. 74,/). The change in suckers on III probably represents hecto-
cotylization and the apparent enlargement of the suckers near the tip may be
comparable to that shown by Joubin (1933) for Teuthowenia (= Megalocranchia)
megalops, although the suckers show no elongation.

Table 4. — Measurements of Holotype of Megalocranchia papillata, n. sp.

mm. mm.

Mantle length 40.0 Diameter of arm sucker 0.8

Mantle width 10.5 Arm 1 12.5

Head width 10.0± Arm II 11.3

Fin length 1.4 Arm III 11.5

Fin width 7.0 Arm IV 9.0

Eye diameter 3.5

Both tentacles are missing; their bases show between arms III and IV and
appear symmetrical.

The examination of the viscera showed no points of special interest. However,
upon examination of the spermatophores it was found that the structures differed
markedly from those so far described. A complete spermatophore is shown in
figure 74, g, and the details of the horn are shown in figure 74, h. In one of these
spermatophores, mounted in Canada balsam, the horn is not tightly coiled but is
telescoped or accordioned in such a way that it appears jointed. The total length
of the spermatophore is 7.4 mm., width 0.4 mm., length of reservoir 4.3 mm.,
middle section 2.0 mm., mid organ 0.15 mm., length of horn 0.95 mm.

Holotype. — Chicago Natural History Museum no. 78316, one male,
in alcohol, mantle length 40.0 mm., from Caryn haul 60, 32° 05' N.,
64° 38' W., 1500 meters wire out in 3270 meters, 0220-0433 hours,
12-foot ring net, August 28, 1948.

Discussion. — This unique individual differs so greatly from the
other cranchiids presently known that it has been deemed necessary
to describe it as a new species. Little is known concerning the struc-
ture of the spermatophores in the cranchiids, and as these are of sig-
nificant taxonomic importance it seems advisable to have descriptions
of spermatophores tied to indisputably recognizable specimens.

The systematic position of this species is also doubtful. Appar-
ently it is closely related to Helicocranchia pfefferi Massy, 1907, and
Helicocranchia beebei Robson, 1948. Grimpe (1922) considered that
Massy's species should be placed in the genus Teuthowenia Chun,
1910. By association H. beebei should also be placed in this genus.
Muus (1956) has shown that Teuthowenia megalops Prosch, 1849, is
a developmental stage of Desmoteuthis hyperborea (Steenstrup) and

VOSS: BERMUDAN CEPHALOPODS 433

that the first has priority, thus changing the name to Desmoteuthis
megalops, preferring Desmoteuthis to Teuthowenia. As pointed out
by Muus, however, Desmoteuthis is a synonym of Taonius. In fact,
it is an absolute synonym and therefore is not available for use in
this case. Berry (1912) has reviewed this situation and pointed out
that in all essential detail^ Megalocranchia Pfeffer, 1884, is the same
as Desmoteuthis auct. and is therefore the valid name. Later he
(1916) proposed the name Verrilliteuthis for reasons that do not now
seem valid. Muus (1956) in his discussion of the genera Taonidium
and Megalocranchia states that Megalocranchia is identical with Des-
moteuthis. If this is true, one wonders why he did not then use
Megalocranchia as the valid generic name.

To return to the present species and its affinities, Muus placed
Helicocranchia pfefferi Massy, 1907, in synonymy with his Desmo-
teuthis megalops. He gives no reason for this action. If one exam-
ines both his figure 4c of a specimen 30 mm. in mantle length and
Massy's (1909) plate 3, figure 1, of a specimen with a mantle length
of 39 mm., one sees that there is no comparison in fin shape and that
Massy's larger specimen has a fin shape comparable to that of much
smaller specimens of Desmoteuthis (actually Megalocranchia). It
seems probable that the genus Helicocranchia is a synonym of his
Desmoteuthis but I fail to see the basis for placing pfefferi in the
synonymy of megalops. Perhaps there are other characters which
Muus did not elucidate. At present it seems preferable to retain
Helicocranchia as a subgenus of Megalocranchia.

Megalocranchia (Helicocranchia) pfefferi has a smooth mantle, fins
which are very narrowly pedunculate but fan-shaped distally and no
teeth on the arm sucker rings. M. (H.) beebei Robson has non-
pedunculate fins which are individually semicircles with scalloped
margins, and the sucker rings are unknown. In M. (H.) papillata
the fins are very slender and non-pedunculate, the arm suckers are
finely toothed, there is no light organ on the eye (there is a light
organ present in pfefferi but not in beebei) and the mantle and fun-
nel are strongly papillated.

Megalocranchia megalops (Prosch, 1849)

Owenia megalops Prosch, 1849, K. Dansk. Vid. Selsk. Skr., ser. 2, 1 : 64.

Material. — One juvenile, mantle length 26.0 mm.; CNHM no.
78317; haul 27. One juvenile, mantle length 18.0 mm.; CNHM
no. 78318; haul 48. One juvenile, mantle length 31.0 mm.; CNHM no.
78319; haul 47. Four juveniles, mantle length 32.0, 22.0, 13.0 and

434 FIELDIANA: ZOOLOGY, VOLUME 39

12.0 mm. ; CNHM no. 78320; haul 22. One juvenile, mantle length?,
CNHM no. 78321; haul 26. One juvenile, mantle length 19.0 mm.;
CNHM no. 78322; haul 58. One juvenile, mantle length 12.0 mm.;
CNHM no. 78338; haul 38. One juvenile, mantle length 24.0 mm.;
CNHM no. 78323; haul 63. One juvenile, mantle length 17.0 mm.;
CNHM no. 78324; haul 52. Three juveniles, mantle length 17.0,
15.0 and 13.0 mm.; CNHM no. 78325; haul 56.

The fifteen specimens listed above seem referable to Megalocran-
chia megalops on the basis of Muus' 1956 paper. While some of them
still have terminal fins at a fairly large size, this is not incompatible
with our knowledge of larval development. Also in some specimens
around 9.0-20.0 mm. in mantle length there is a decided rostrum
on the ventral margin of the eyeballs covering a halfmoon-shaped
light organ. In the larger specimens the rostrum was lost. Many
of the specimens were more slender than is normal, but correspond
well with Joubin's (1933) figure of the male of Teuthowenia (Desmo-
teuthis) megalops. Evidently Muus overlooked this description, for
he states (1956, p. 12) : "Since none of the adult specimens examined
to date have been males, nothing is known on a possible sexual di-
morphism."

The species is easily recognized by the lack of tubercle rows at
the mantle fusion, absence of a funnel valve, presence of enlarged
suckers on arms II and III and semicircular light organs on the eye-
balls. The present specimens range from about 300 to 1200 meters
in depth.

The generic name of this species has been in a state of confusion
for many years. For a review of the biology, morphology and taxo-
nomic status one should consult Muus (1956) and the discussion
section of the preceding species.

Carynoteuthis, new genus

A taoniid with sessile eyes which bear two large meandering light organs; fun-
nel with a small valve, the dorsal funnel organ with two large triangular flaps; arm
suckers minutely toothed; tentacular suckers toothed; light organs present on
the ink sac.

Type species. — Carynoteuthis oceanica.
Carynoteuthis oceanica, new species. Figure 75.

Megalocranchia abyssicola Joubin, 1924, Result. Camp. Sci. Monaco, 67: 96.

The mantle (fig. 75, a) forms a slender cone with straight sides, converging to a
point posteriorly. It is widest at the anterior margin and is united by a fusion in
the nuchal region and on each side of the funnel.

VOSS: BERMUDAN CEPHALOPODS

435

Fig. 75. Carynoteuthis oceanica, new genus and species, a, Dorsal view of
holotype; mantle length 76.0 mm. b, Funnel organ, c, Right eyeball, ventral
view, showing light organs, d, Sucker from third left arm. e, Sucker from hand
portion of tentacular club, inner row. /, Details of ventral surface of the liver of
the holotype, showing arrangement of light organs.

The /ins are small, terminal, slender, attached to the posterior end of the gladius
and projecting beyond the mantle end for over half their length. They have nearly
straight posterior edges and form a long triangle.

The funnel is large and projects to about the level of the eyes. It has a U-shaped
dorsal funnel organ (fig. 75, b) with a large, thin, triangular lappet on each side.
The ventral pads are large and nearly oval. There is a broad, nearly straight-edged
valve just anterior to the margin of the dorsal funnel organ and nearly covered
over by it.

The head bears enormous round black eyes which are sessile and about as wide
as the mantle. The eyes bear two light organs. Their arrangement is shown in fig-

436 FIELDIANA: ZOOLOGY, VOLUME 39

ure 75, c. In their shape and arrangement they do not agree with the light organ
structure as given by Pfeffer for the Taoniinae, although they are probably derived
from the basic double halfmoon type. They do not correspond to any as yet de-
scribed. There is an olfactory organ below and posterior to each eye. It is borne
on a long slender stalk and is cup-shaped, somewhat resembling that illustrated by
Goodrich for Megalocranchia abyssicola. The buccal membrane is seven-pointed
and has seven connectives.

The arms are in the order 3.4.2.1 and nearly round in cross section. The suckers
are biserial with minute round teeth on the horny rings (fig. 75, d). The suckers are
somewhat smaller toward the base of the arms but they increase in size, becoming
largest about two thirds of the length of the arm; near the tip they become smaller
and then are abruptly minute. The suckers are bordered on both sides by a pro-
tective membrane consisting of large fleshy trabeculae joined by a rather delicate web.

The tentacles are slender, round in cross section and flattened on the oral sur-
face. The clubs are moderately long, little expanded and bordered on each side by
a trabeculate membrane. A narrow keel borders the distal half of the club on the
dorsal side. There is no distinct carpal cluster but the four rows of suckers con-
tinue down the stalk as minute suckers for a distance about equal to the length of
the club. The hand suckers (fig. 75, e) are borne on short stalks, and are cupped
and bordered by a broad papillate band. The apertures are armed with about
18-20 sharp teeth, large and separate on the distal half, close together and minute
proximally.

The mantle was opened and the viscera examined (fig. 75,/). The arrangement
of the liver, ink sac and what appear to be four large lenses over a diffused light
organ is unusual. The liver is large and fat, oval in outline. On the ventral surface
lies a larger rectangular ink sac, traversed ventrally by the rectum, which extends
downward posteriorly and then turns along the sac anteriorly, terminating at the
anterior edge where the ink sac duct unites with it just posterior to the anus. On
each of the four corners of the ink sac is located a large, clear, hollow sphere, easily
ruptured, lying over and united with an iridescent layer. There appear to be two
of these, one on each side, forming a somewhat saddle-shaped organ. I have re-
marked nothing like this in the cephalopods which have come to my notice.

Holotype. — Chicago Natural History Museum no. 78326. A fe-
male, mantle length 76.0 mm., from Caryn haul 61, 32° 08' N.,
64° 33' W., 3500 meters of wire out, in 1700 fathoms, 1135 hours,
Blake trawl, August 30, 1948.

Paratype. — One female, mantle length 44.0 mm.; CNHM no.
78327; Caryn haul 63, 32° 12' N., 64° 36' W., 3000 meters of wire
out, in 1300 fathoms, 12-foot ring net, 1243 hours, September 2, 1948.

Discussion. — It is with trepidation that I have erected a new genus
to accommodate this species, inasmuch as there is a possibility that
it may be the adult of Chun's Corynomma speculator. His smallest
specimen of 11.0 mm. mantle length was figured, the larger one of
32.0 mm. mantle length was not, and was only incompletely described.
We do not know what changes were brought about by growth. My
44.0 mm. specimen is very different from his, and is comparable only

VOSS: BERMUDAN CEPHALOPODS 437

in the presence of a light organ on the liver. It differs in other aspects:
the sessile eyes (an adult character?), the differently shaped funnel
organ, the presence of a valve, and others. These alone prevent its
being placed in Corynomma.

Table 5. — Measurements of Type and Paratype of Carynoteidhis oceanica, n. sp.

Holotype Paratype

mm. mm.

Mantle length 76.0 44.0

Mantle width 23.0 11.0

Head width 21.0 11.0

Fin length 20.5 11.0

Fin width 14.0 8.5

Arm I, length 13.0 6.5

Arm II, length 16.5 6.5

Arm III, length 21.5 11.0

Arm IV, length 19.0 10.2

Eye diameter 12.0 7.0

Diameter of arm sucker 1.2 0.5

Diameter of club sucker 0.6 0.4

Tentacle length 32.0 23.0

Club length 10.0 7.0

In 1924 Joubin described a specimen from off the Azores which
he called Megalocranchia abyssicola Goodrich, 1896. His description
and illustrations conform exactly to the present species. Joubin dis-
misses the differences between the Azores specimen and Goodrich's
Laccadive specimen by stating that Goodrich's description and illus-
trations were not accurate. However, he did not examine the only
specimens, which are in the Calcutta Museum. I have carefully com-
pared my specimens with Goodrich's description and illustration and
I am convinced that Joubin's assumption was erroneous. In addi-
tion, I requested the curator of the mollusk collections in the Cal-
cutta Museum to re-examine the type, but I was informed that it
was in such poor condition that nothing of value could be determined
from it. Under these circumstances I feel justified in my original
decision that M . abyssicola and Carynoteuthis oceanica are quite dis-
tinct and that Joubin was in error.

From Desmoteuthis and its complex, Teuthowenia and Megalo-
cranchia, it may be separated by its possession of internal light organs
and a funnel valve and the shape of the funnel organ. One species in
Megalocranchia is excepted, M. abyssicola Goodrich, 1896, which has
a funnel valve. This may possibly belong to Carynoteuthis. It is in-
teresting to note that Pfeffer, Chun, and Thiele all state that the
funnel valve is lacking in the Cranchiidae, despite Goodrich's state-

438 FIELDIANA: ZOOLOGY, VOLUME 39

ment and illustration and the description by Joubin (1895) of a valve
in Taonius richardi. The present specimens should dispel this error.
The name Carynoteuthis is derived from the R/V Caryn and the
Greek revrrjis or squid. The specific name oceanica refers to its
habitat.

Bathothauma lyromma Chun, 1906

Bathothauma lyromma Chun, 1906, Zool. Anz., 31: 86.

Material. — One female, mantle length 80.0 mm.; CNHM no.
78328; haul 45. One female, mantle length 60.0 mm.; CNHM no.
78329; haul 59. One juvenile, mantle length 36.0 mm.; CNHM
no. 78330; haul 39. Three juveniles, mantle length 29.0, 26.0 and
10.0 mm.; CNHM no. 78331; haul 22.

A nice series of specimens from 10.0 to 80.0 mm. in mantle length
of this poorly known and weird species is available. I have com-
pared these specimens with a male of 114.0 mm. mantle length from
the Philippines and find no essential differences. In the specimens
at hand there is a large yellowish chromatophore between the bases
of the eyestalks on the dorsal surface of the head stalk, and a row of
similar spots along the ventral mid-line of the stalk, which extends
outward along the ventral surface of the eyeball. The fins are some-
what rectangular in outline and widely separated from one another,
the blunt, round end of the mantle projecting somewhat beyond.
The mantle of these creatures may aptly be described as resembling
deflated sausage skins, both in shape and consistency.

The availability of such a wide range of sizes lends itself to a sur-
vey of the changes in proportions due to growth; the following meas-
urements are given.

Table 6. — Measurements of Six Specimens of Bathothauma lyromma Chun

(In millimeters)

Mantle length 10.0 26.0 29.0 36.0 60.0 80.0

Mantle width .... .... .... .... ....

Head width 12.5 30.0 21.0 33.0 39.0 52.0

Fin length 5.0 6.0 7.0 9.5 10.5

Fin width 5.0 5.5 5.0 9.0 10.0

ArmI 0.8 2.4 2.5 2.0 6.5 8.0

Arm II 0.8 3.0 3.3 2.3 8.5 10.0

Armlll 0.8 3.1 3.6 2.3 9.5 11.0

Arm IV 0.8 3.2 3.0 2.8 8.5 10.0

Tentacle length 14.0 41.0 58.0 36.0 61.0 62.0

Club length

Head stalk length 8.0 18.0 13.0 16.5 18.0 21.5

Eye stalk length 6.0 13.0 11.5 13.0 17.5 22.0

VOSS: BERMUDAN CEPHALOPODS 439

From the data presented in the table it can be seen that in the
younger specimens the mantle is rather short and wide (mantle width
could not be measured because of the flattened condition of the man-
tle and extreme wrinkling), with long slender head stalks, long ten-
tacles and short arms. Growth of the head stalk is very slow in
comparison to mantle growth so that the general appearance in the
adult is of a long tubular body, short stalk and only comparatively
long tentacles. Arm growth does not seem to be at the same rate as
the mantle, for under 36 mm. the arms are very short, but between
36 and 60 mm. the arms greatly increase in length, becoming long
and slender. Only fin growth seems to remain at the same rate
throughout. The depth range of this species is not known since all
records are from open nets, but one may conjecture that it occurs at
1000 meters and above.

OCTOPODA
Family OCTOPODIDAE
POctopus vulgaris Cuvier, 1797

Octopus vulgaris Cuvier, 1797, Tabl. Elem. l'Hist. Nat. Anim., p. 380.

Material. — Two juveniles, mantle length 7.0 and 8.0 mm. ; haul 37.

In the same haul with some specimens of Japetella diaphana were
two small octopods. While not in good shape, part of the head pig-
mentation was present. Conceivably, these two specimens entered
the net in the upper layers. Of the two species of octopus known to
occur in Bermuda, 0. macropus and 0. vulgaris, both spawn small eggs
and have planktonic larvae. The former has a long, slender, pointed
mantle, while the latter has a round, sac-like mantle. The present
specimens correspond to 0. vulgaris and are tentatively placed in
this species.

Family BOLITAENIDAE

Eledonella pygmaea Verrill, 1884

Eledonella pygmaea Verrill, 1884, Trans. Connecticut Acad. Sci., 6, (1), p. 145.

Material. — Two females, mantle length 38.0 and 31.5 mm.;
CNHM no. 78332; haul 57. One male, mantle length 25.0 mm.;
CNHM no. 78333; haul 30.

Thore's (1949) masterful treatment of this species leaves little to
be said of interest concerning these specimens. The females are
richly covered with dark brown chromatophores, the head and web

440 FIELDIANA: ZOOLOGY, VOLUME 39

region being darker than the body. The male, conversely, is rather
light and transparent, with little pigmentation. The eyes are small,
the suckers are rather large and are set one or two diameters apart
and rather well embedded in the tissue. The male has the third right
arm modified into a hectocotylus, the distal portion bearing four im-
mense suckers, the largest 3.2 mm. in diameter, the second 3.0 mm.,
the third 2.0 mm., and the terminal one 1.0 mm. Thore states that
there are 6-8 small suckers but I found only one. As Thore first re-
ported from the Dana material, there is a distinct ligulus on the distal
portion. In the present specimen, it varies somewhat from the shape
shown by him and the ligula is more of a typical octopodid spoon
shape. It is possible that the discrepancies noted in the shape of the
ligula and the reduced number of small suckers beyond the enlarged
three is a function of growth and represents immaturity. Thore has
shown that this species is truly bathypelagic, the young occurring
just beneath the discontinuity layer from 100 to 200 meters, and that
no specimens over 35.0 mm. in mantle length occur in less than
333 meters; they go down to 2500 meters, with an average distance
from the bottom of 3038 meters and never less than 700 meters.

Japetella diaphana Hoyle, 1885

Japetella diaphana Hoyle, 1885, Rep. Sci. Res. Voy. Challenger, 1: 232.

Material. — One female, mantle length 14.0 mm.; CNHM no.
78334; haul 37.

This species has also been revised by Thore (l.c,) . It is a common
and widely distributed species, easily recognized by the close-set urn-
shaped suckers and rather large laterally directed eyes. The present
specimen is evidently a juvenile and measurements could not be taken.
Thore has given a detailed account of his calculations concerning the
depths at which this species lives. According to him, the young are
found at about 200 meters, just below the discontinuity layer, from
whence they migrate downward, adults over 40.0 mm. living below
333 meters and concentrated at 1750 and 2500 meters.

Family VITRELEDONELLIDAE

Vitreledonella richardi Joubin, 1918

Vitreledonetta richardi Joubin, 1918, Bull. Inst. Oceanogr. Monaco, no. 340, p. 1.

Material. — One male, mantle length 34.0 mm.; CNHM no. 78335;
haul B16.

VOSS: BERMUDAN CEPHALOPODS 441

This species has been so well studied and discussed by Thore that
nothing new can be added here. The animal is in rather poor con-
dition and very flaccid, so that accurate measurements are impos-
sible, but excellent data have been given by Thore. The species may
easily be distinguished by its clear gelatinous appearance, the small,
rectangular eyes, and long, slender, pointed liver. The left third arm
is hectocotylized on the distal portion by a marked reduction in size
of the suckers and dense crowding, terminating on the tip by a distal
spoon-shaped ligula. According to Thore the young of this species
hatch out at a depth below 1000 meters but grow to near adulthood
just below the discontinuity layer and thence move back downward,
living out their life between 1000 meters and 1750 meters.

Family ARGONAUTIDAE
Argonaut a argo Linnaeus, 1758

Argonauta argo Linnaeus, 1758, Syst. Nat., ed. 10, p. 708.

Material. — One female with shell, ventral mantle length 18.5 mm. ;
CNHM no. 78336; haul 22. Two females, ventral mantle length 7.0
and 6.5 mm.; CNHM no. 78337; haul 48.

The two small specimens are very immature, the larger one bear-
ing a shell or egg case. This is a very common species with a wide
range. The vertical range is not known but certainly it is restricted
to the upper layers, rather than at the depth the net was fishing. It
has been well figured by Chun (1910) from the living animal.

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444 FIELDIANA: ZOOLOGY, VOLUME 39

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Chun, Carl

1906. System der Cranchien. Zool. Anz., 31: 82-86.

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Cuvier, G.

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Goodrich, Edwin S.

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Grey, Marion

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Grimpe, Georg

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Heilprin, A.

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Joubin, Louis

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VOSS: BERMUDAN CEPHALOPODS 445

1931. Notes preliminaires sur les cephalopodes des croisieres du "Dana" (1921-
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446

FIELDIANA: ZOOLOGY, VOLUME 39

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