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ZOOLOGISCHE STATION ZU NEAPEL

,/ '*VfLLIAM I'i.

Sect ional libf^ary
DlYiSION OF MOLLUSKS

n

FAUNA and FLORA
of the BAY of NAPLES

Monograph No. 35
ADOLF NAEF

CEPHALOPODA (systematics)

Part I, Vol. I, Fascicle U ( End of Vol. / )

TRANSLATED FROM GERMAN

Published for the Smithsonian Institution
and the National Science Foundation, Washington, D.C.
by the Israel Program for Scientific Translations

ZOOLQGISCHE STATION ZU NEAPEL
Zoological Station, Naples

Fauna and Flora
of the Bay of Naples

(Fauna e Flora del Golfo di Napoli)

Monograph No. 35

CEPHALOPODA (systematics)

by

Adolf Naef

Part E Vol. I, Fascicle II (End of Vol. I)

Berlin

In Kommission bei R. Friedlander and Sohn
1923

Translated from German

Israel Program for Scientific Translations
Jerusalem 1972

TT 68-50343/2

This is a Limited Edition by Special Agreement between
THE SMITHSONIAN INSTITUTION
and the

STAZIONE ZOOLOGICA DI NAPOLI

Israel Program for Scientific Translations Ltd.
IPST Cat. No. 5110/2

Translated by A. Mercado
Edited by Prof. O. Theodor

Printed in Jerusalem by Keter Press
Binding: Wiener Bindery Ltd., Jerusalem

313 Chapter 18

GENUS ON YCHOTEUTHIS-
Lichtenstein, 1818

ONYCHOTEUTHIS BANKSI (Leach, 1817) d'Orb., 1839
DIAGNOSIS

A large yellowish brown luminous organ on ventral side of eyeball. Two
lens- shaped luminous organs in median part of mantle cavity (the anterior
organ behind the anal papilla, the posterior in the ink sac, before the renal
papillae). Gladius with a posteriorly well developed crest in its greater part
(except in the anterior part); crest visible as distinct dark line through the
dorsal skin. Rostrum short, about as long as the gladius. Carpal pad
rounded, with 6—10 suckers and the same number of knobs. Club with 10—12
hooks in each row; fifth hook (often fourth and fifth or fifth and sixth)
smaller than preceding and following hooks and displaced toward the
ventral row of hooks.

LITERAT URE

1817 Leach, Loligo banksii (p.l41).

1818 Leach, Loligo banksii (p. 411).

1818 Lichtenstein, O ny c h o te u th i s bergii (Abh. Ak. Berlin, p. 223).

1818 Lichtenstein, O n y ch o te u th i s molinae (Abh. Ak. Berlin, Plate 4) (Copied in d'Orbigny 1839,
Onychot. Plate 5, Figures 1—3).

1821 Lesueur (2), O n y k i a angulata (p. 99, Plate 9, Figure 3). (One specimen determined by Lesueur as
O. angulata in d’Orbigny (1839, Plate 4) .

1821 Lesueur, Loligo bartlingii (p. 95) (Drawing copied in d'Orbigny, 1839, Onych. Plate 3, Figures 1,

2. 2a-d).

1824 Quoy and Gaimard (2), Lolig o uncinata Vol. I, p. 410, Plate 66, Figure 7).

1830 Lesson, Onychoteuthis lessoni Fer. 1825, p. 240, Mollusques, Plate 1, Figure 3 (Drawing copied
in d'Orbigny 1839, Onychot. Plate 2, Figure 1).

1830 Lesson, Onychoteuthis fleuryi Renaud (Centurie ZooL, p. 61, Plate 17) (Figure copied in
d’Orbigny 1839, Onychot. Plate 9, Figure 1).

314 1839 Ferussac and d'Orbigny, Onychoteuthis banksii (p. 330, Onychot., Plates 1, 2, 3 to 4, 5, 6, 7, 9, 12,
Plate 12, Figures 1—9) ("O. angulata, lessoni, bartlingii, lesueurii, bergii, fleuryi”).

1849 Gray, Onychoteuthis banksii (p. 53), bartlingii (p. 54), krohnii(p. 55).

1851 Verany, Onychoteuthis krohnii (p.80, Plate 29, Figure d, e).

1851 Verany, Loligo bianconii (p. 100, Plate 35, Figures i—1).

* Onychoteuthinae: occipital folds distinct. Gladius with a rudimentary flag. Club of tentacle without
marginal rows on the hand part after the median rows have been converted into hooks (see p.3ll).

293

1852 Gould, Onychoteuthis rutilus (p. 482, Plate 50, Figure 595).

1868 Gabb, Onychoteuthis aequimanus (p. 23, Plate 2).

1881 Verrill, Onychoteuthis banksii (Plate 15, Figures 6 a, b, c; a^ , b^ , Ci).

1881 Owen,Onychoteuthis raptor (p. 146, Plate 29).

1889 Posselt, Onychoteuthis banksii (pp. 144-145).

1889 Cams, Onychoteuthis krohnii (p. 499).

1890 Norman, O nychoteuthis krohnii (p. 475).

1890 Girard, O n y cho t e u th i s banksii (p. 266).

1891 Lbnnberg, Onychoteuthis banksii (p. 37),

1896 Goodrich, Onychoteuthis banksii (p. 11).

1896 Jatta, Ancistroteuthis lichtensteinii pars (p. 103, Plate 13, Figures 19, 21).
1’896 Jatta, Te 1 e o te u th i s krohnii (p. 97, Plate 13, Figures 24 to 34, 42).

1900 Joubin, Teleoteuthis carolii.

1900 Pfeffer (2),Onychoteuthis banksii (p. 159).

1900 Pfeffer, T etronychoteuthis krohnii.

1904 Jatta, T etronychoteuthis krohnii (p. 200).

1904 Hoyle (diagn. Key), T etronychoteuthis krohnii (p. 19).

1904 Hoyle (Albatr.), T e t r o n y c h o t e u th i s krohnii (p,35).

1906 Hoyle, Tetronychoteuthis krohnii juv. (p. 160).

1907 Hoyle, T e t ro n y ch o t eu th is krohnii (p. 14).

1908 Pfeffer, Tetronychoteuthis krohnii (p. 65, Figures 71—77).

1909 Hoyle, T etronychoteuthis krohnii (p. 3).

1912 Pfeffer,0 n y cho teuthi s banksii (pp. 70— 88, Plate 3, Figures 13— 25, Plates 4— 6).
1916 Naef,0 n y ch o t eu th i s banksii (p. 15).

1921 Naef, O nychoteuthis banksii (p. 537).

1921 Grime (North Sea), Onychoteuthis banksii (p. 298).

STRUCTURE OF THE ADULT ANIMAL

I have no mature specimens of this species, but a large number of young
specimens near maturity. These specimens have completed their full
development, except that they are smaller and the genitalia are not fully
developed. Sexual differences are not recognizable. A female is shown
in Figure 153.

The mantle sac has already the typical form, it is cylindrical anteriorly,
pointed posteriorly. The gladius is visible as a dark mediodorsal line which
begins behind the middle of the fins and corresponds anteriorly to the
median stripe of the groovelike rhachis, posteriorly to the crest of the
rhachis.

The neck folds show the typical character of the subfamily, i. e. the
presence of "occipital folds" (pp. 311 and 313, Figure 153a) which occupy the
area between the anterior and posterior transverse neck folds on each side
of the upper side of the neck, forming parallel lamellae which extend
medially and slightly laterally from the typical position of the first longi-
tudinal fold. Embryology shows (p.320) that the largest (usually the outer
fold but one) of these lamellae corresponds to the first longitudinal neck
fold. The other folds are named accessory occipital folds. There are 6
(5—7) medially directed and 1 (rarely 2) laterally- oriented folds, together
usually 8 (7—9) such folds. They are delicate, crescent- shaped folds, the
inner folds not always distinct, in addition to the typical neck folds. The
315 anterior transverse fold extends on each side from the funnel pit to the
occiput, where it meets the fold of the other side in the middle at an angle

294

which opens anteriorly. The posterior fold extends from the third longi-
tudinal neck fold to the occiput almost without projecting beyond the median
accessory occipital folds, and does not reach the middle of the body. The
fourth longitudinal fold ends free posteriorly on the neck, and only the third
longitudinal fold reaches the posterior transverse fold on the prominent
area which bears the olfactory organ. The second longitudinal fold is
normal.

FIGURE 153. Onychoteuthis b a n k s i, almost mature female. Note the
form of mantle and fins; the translucent mediodorsal rhachis of the gladius;
neck and folds; funnel pit; funnel adductors; the typical lid opening; the
swimming margin of the three upper pairs of arms; the tentacle clubs with
hooks and carpal pads.

The lid is of typical form: transversely truncate posteriorly, with a
pointed sinus anteriorly. The head is narrower before the eyes, but other-
wise its outline resembles that of the Ommatostrephidae (cf. Figure 238).

295

The arms are rather long, whiplike. The third pair is longer, the first
pair shorter than the others. Formula: 3, 4, 2, 1. The distal half of the
dorsal arms has a narrow swimming margin in the middle of the outer side.
The swimming margins of the second pair are wider, they extend to the base
316 and occupy the lateral edge of the outer surface. The margins of the third
pair are still more strongly developed, and form a blunt lateral corner and
taper suddenly to a mere edge or a very narrow margin. The fourth arm

bears the typical outer margin.

FIGURE 154. Right clubs of Onychoteuthis
banksii (a) and Ancistroteuthis lich-
te nstcini (b). (a - 2X; b - V3X.) Note the
different number of suckers on the carpal pad,
the prehensile and rotating hooks. In b the
position of the lost suckers of the outer rows
is marked by crosses:

Et - terminal part; Fh - prehensile hooks;

Dh — rotating hooks; Ss — protective margin
(widening); Cp - carpal pad; So - dorsal
protective margin; x.y - displaced hooks;

Sm - swimming margin; Sf - lateral surface
of club axis.

The tentacle stalks are moderately long and project about one -third of their
length beyond the end of the arms, if extended. The clubs are shown in
Figures 153 and 154. The proximal 4(3—4) hooks of the ventral row are of
the clasping or rotating type, like those of the dorsal row; the others are
firmly attached prehensile hooks, directed anteriorly and downwards, the
third or fourth are the largest. The dorsal row consists entirely of rotating
hooks. The fifth hook (y) is very small and displaced toward the row of
large hooks, often together with the sixth (x) or fourth hook. The carpal pad
usually consists of 8 suckers and knobs (according to Pfeffer, 6—10). There
are only 3—4 rows with 4 small suckers in the distal part.

296

The mantle cavity of the adult shows the same condition as the juvenile
stage (p. 320), if it is not changed by the development of the genitalia
(Figures 155 and 157).

317 LATER POSTEMBRYONIC STAGES

Older juvenile stages which can be identified continue the series of
younger larvae described on pp. 304—309 (Figures 145—152), which cannot
be determined with certainty. They have been described in the past as
"O n y c h o t e u t h i s krohni." Already at a mantle length of 11 mm
(preserved specimens), they can be recognized by the length of the fins
(over 4 mm, actual measurements 4.3 and 4.4 mm), the gladius is visible
along the whole dorsal mid line, and the form of the cone is ventrally
recognizable as a hollow cone (cf. p.318). The head becomes thicker (in
comparison with Figure 151), the arms are longer, and the fins attain their
typical form (Figure 157); hooks develop in the ventral median line of the
clubs and swimming margins appear on the clubs and arms (Figure 152).
At a dorsal mantle length of 17.5 mm, the animal shows the very charac-
teristic form of "Teleoteuthis krohni" (Figure 156).

FIGURE 155. Onychoteuthis b a n ks i i, half- mature male and female. V2X. Mantle situs.
In the male, the terminal part of the gonoduct (Go) projects from the pocket of the branchial
base, the sperm duct (SI) and spermatophore gland (Sp) are visible behind the gill. The corres-
ponding place in the female is occupied by the widened posterior part of the oviduct gland
(Dr), posteriorly connected with the short coelomic oviduct (Od); between them the partly de-
veloped nidamental glands (Nd) in normal position and form. From the nidamental glands
extend (dotted) fascicles of cutaneous muscles toward the hind intestine. In the mature animal
(cf. Figures 207 and 225) these fascicles support the ligaments which bear the glands. The
other conditions resemble those in Figure 157, with slight differences in size and position of
the organs.

297

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298

The fins of the well-preserved specimen described above are 7.9 mm
long, i. e. about half of the mantle length, and already have "earlobes" and
lateral corners. Their posterior end is only slightly pointed and produced.
The fin span exceeds even Vs of the mantle length. The apex of the fins is
level with the posterior end of the body, which is supported by the rostrum.
Proximally, however, the fins are raised markedly because of the form of
the rostrum, which is markedly directed dorsally and laterally compressed,
and situated entirely on the dorsal side of the gladius. The fin is attached
only to the dorsal edge of the rostrum. The two fins are fused in the median
line in the area of rostrum and cone, forming a single structure. This
process advances later further anteriorly. (Such a fusion takes place also

318 in the Ommatostrephidae and Gonatidaei its anatomical consequences are
discussed on p.227.) The gladius is visible as a dark line which almost
reaches the mantle margin, but this line does not represent a median keel;
it widens slightly to the anterior end of the base of the fins, forming the
remnant of a lanceola (Figure 151 on p.310); it tapers rapidly further on
and widens again only anteriorly.

The mantle of this specimen has the usual form, it is thick and spindle-
shaped posteriorly but its anterior part already resembles the cylindrical
form of the adult and may even bulge in the middle. Preservation often
causes marked deformations, e. g. the ventral bend of the gladius, which is
observed especially in younger specimens. Careful preservation is
necessary to avoid the retraction of the head into the mantle, as this changes
the natural habitus.

The typical neck folds are already indicated. The third longitudinal fold,
with a prominent olfactory tubercle is particularly distinct. The eyes are
much larger and strictly lateral in position. The eyelid has a sinus, and a
window (dark in the figure) is already present on the ventral side of the
eyeball. The lens is strongly convex and projects markedly.

The third pair of arms has already reached the length of the second, and
the fourth pair that of the first, but the lateral arms are distinctly longer
than the dorsal and ventral arms. A rudimentary swimming margin is

319 present on the terminal part of the first arms. Such a margin is more
distinct on the greater part of the second pair of arms but is best developed
on the third pair. The fourth pair bears a typical outer margin. The
inner surfaces of the arms show almost their definitive condition, but the
number of suckers is still small. The buccal funnel is typically differenti-
ated: its long points and supports surround a markedly projecting mouth
cone, which is not visible in this drawing. The tentacle club is further
developed than in Figure 152: three or four prehensile hooks are already
strong and functional, and other hooks in the ventral median row are being
formed. The suckers of the dorsal median row are slightly larger but
show no trace of conversion, at least not under the magnifying lens. There
is a well developed protective margin in the proximal part of the club,
along the ventral edge, but not in the other parts. The swimming margin

is well developed.

Figure 157 and Figure 1 of Plate IV show a similar but larger and older
specimen with opened mantle cavity. The fins of this specimen are larger
and occupy over half of the dorsal mantle length. The posterior end of
the body shows the usual conditions (p. 302).

299

The funnel of this stage shows differentiation and enlargement of the
funnel bonds and the relative reduction of the projecting part of the funnel.

32 0 There is a strong contraction of the primary lid fold. The position of the
window (Fe) is indicated in this stage, and also the large, yellowish brown
luminous organs (Oi, see below). The neck folds are thicker; they consist
of an anterior transverse fold and the 4 typical longitudinal folds, occipital
folds are still absent. The edge surrounding the funnel pit is also distinct.
The olfactory tubercle projects at the posterior end of the second longi-
tudinal neck fold as a flat, slightly ovoid appendage.

The mantle cavity shows the conditions typical for the Oegopsida. The
anal papilla, funnel retractors, renal papillae, gills and the posterior aorta
and its branches are typical. The same applies to the translucent organs:
the branchial hearts with their appendages and coelomic pockets, the venous
appendages on the roots of the posterior pallial veins, the venous branches
and the course of the vena cava. The heart is still wide, as is characteristic
for the young forms. Also typical is the rudimentary gonoduct near the
left branchial heart and the formation of "lateral pockets" which penetrate
toward the dorsal part of the mantle between the body and the origin of the
funnel retractors. Specific characters are the narrow form of the gladius,
which is visible in the upper part of the visceral sac, and especially the
formation of the two luminous organs. These are situated asymmetrically
on the left side near the hind intestine but displaced to the middle. Their
"anlagen" are present already in the youngest stage which can be identified
asOnychoteuthis banks i, with a dorsal mantle length of about 1 1 mm,
when the hind intestine is still strictly median (see Figure 149 on p. 308).

The luminous organs appear at first dorsally as mere thickenings of the
skin. The smaller anterior organ is situated on the left side close to the
hind intestine, closely behind the anal papilla, but it is gradually displaced
to a median position, and is visible through the intestine. The larger
posterior organ is situated at the point where the hind intestine sinks
deeper or rises to the surface. Like the corresponding left organ of
Octopodoteuthis sicula (Figure 1 66), it appears as a thickening of
a thin cutaneous muscle, the now distinct musculus rectus abdominis.*

Such a symmetrical organ does not develop, but the left luminous organ
moves to the median line, pushing the hind intestine aside, and develops
rapidly (see also Figure 155). At the same time it sinks into the ink sac,
which probably serves as a reflector. This condition resembles that in
Rondeletiola and the Sepiolidae in general (q. v.), although the typical
basis is quite different (see also Plate IV, Figure 3).

Both organs appear as glassy lenses in the living animal. They are
opaque and white in alcohol- preserved material. The organs are circular.

I did not observe luminescence in freshly killed animals.

These luminous organs of the mantle cavity were already known to Hoyle
(1909). There are, in addition, 2 large, yellowish brown, padlike organs on
the ventral side of the eyeball, medially, anterior and below the windows.

321 According to their form and position, these can only be luminous organs.
They are already visible in the youngest specimens identified as this
species, i. e. at a dorsal mantle length of about 11 mm. In our specimen.

Only microscopic studies will show whether the right muscle bears a smaller rudiment of a homonomous
organ.

300

each organ has a process toward the anterior side of the eyeball which
occupies the position of the anterior organs of the Pyroteuthinae (Figure 127).

The club is almost completely developed; also the dorsal median row
of the hand part bears hooks or at least suckers with a strong, hooklike
tooth, but the large hooks of the ventral median row have already reached
their definitive form. The suckers of the marginal rows are very small,
retarded but still present; those of the carpal part form a group but not a
carpal pad. Stalked suckers are absent. There are only 1—2 small
suckers which eventally fall off between the carpal and hand part.

Further development rapidly leads to the adult stage. A young specimen
from Capri, with a dorsal mantle length of 53 mm, may be described as
follows. Habitus as in Figures 156 and 157. Coloration reddish brown
ventrally, violet brown dorsally. Fins 21 mm long. Neck with accessory
occipital folds: first typical longitudinal fold joined by a median row of
5 parallel folds extending toward the midline; there is a slightly weaker
fold laterally. There are thus 7 longitudinal folds on each side of the
occiput, the second from the outside is homologous to the typical second
longitudinal fold of the Teuthoidea.

The tentacle club of this specimen already shows the definitive character.
There are no marginal rows in the hand part; the suckers of the carpal
part (8) are situated on an oval, slightly raised, edged pad. On the right club
there is a scar at the place of the fallen- off sucker between hand and carpal
part. Protective margins are present on both sides at the base of the hand
part; the ventral edge bears a particularly distinct, rounded, wide margin
without distinct supports.

301

322 Chapter 19

GENUS CHAUNOTEUTHIS
Appellof, 1891

CHAUNOTEUTHIS MOLLIS Appellof, 1891
DIAGNOSIS

Body gelatinous. Habitus resembling that of Onyc h o t e u th i s banksi.
Gladius not visible through the dorsal skin as a dark line. Gladius with
only a short, tuberclelike rostrum; flag widest behind the middle. Sperma-
tophores attached during copulation in special longitudinal lateral grooves
on outer surface of mantle of female.

LITERATURE

Appellof (1891,pp. 3— 29, Plates 1—4),

Lonnberg (1896, pp. 603 — 612).

Pfeffer (1900, p.l60). (1912, pp. 89-91, Plates 7 and 8.)

Grimpe (1920 (VI. pp. 289 — 296, Figure 3).

For a more detailed description see Appellof and Pfeffer, 1912, and for some additions, Grimpe 1920.

STRUCTURE OF THE ADULT ANIMAL

I do not have intact mature specimens (the fragment in Leipzig does not
show anything new), and the following description is based only on younger
stages.

Chaunoteuthis occurs in the Mediterranean and Atlantic. It differs
from the closely related O n y c h o t e u th i s banksi in the swollen,
gelatinous consistence of flesh and skin. Pfeffer (1900) considered it,
therefore, as a deepwater variety of O. banksi, but this is hardly justified.
Another assumption is that the characteristic consistence of the tissues is
due to partial digestion in sharks or whales which ejected the animals.

This is also not very probable and could only be settled by. comparison of
well preserved specimens.

323 The general form closely resembles that of Onyc ho teu thi s banksi,
but head and arms are apparently shorter. The recorded differences in
the form of the mantle sac are caused by changes due to preservation and

302

contraction. The gladius resembles that of O. banski, but it is slightly
wider and has a weaker keel, a deeper cone and a shorter rostrum. The
gladius is not visible as a dark median dorsal line. Fins, neck and occipital
folds, funnel pit, lid fold, buccal funnel and coloration are the same as in
O. bank si. The arms are short (slightly more than V3 mantle length) and
differ little in length. The first pair is distinctly shorter than the others,
the fourth pair only slightly shorter (formula: 2 =3, 4, 1). Tentacles are
apparently absent (3 specimens); they have apparently been lost during
postembryonic development, leaving short stumps with pigmented scars.

Little is known of the sexual dimorphism, as the available material
consists only of 4 females. The spermatophores are attached in longitu-
dinal grooves on the right and left of the ventral side of the mantle sac.

The impression is gained that the grooves are wounds resulting from
copulation, after which the spermatophores sank into the skin (cf. Vol. Ill
on the function of the sperm mechanism). It is also possible that the
spermatophores dig themselves in, after having been placed there by the
male. This seems more reasonable because of the absence of hecto-
cotylization in the Onychoteuthidae. (We know unfortunately nothing about
the deposition of spermatophores in other species of the family.)

Luminous organs have not been found in the adult. If organs of the size
and form in O. bank si would have been present in the mantle cavity (on
the ink sac), they would not have been overlooked by Appellof. They
probably disappeared during development (see p. 325).

According to Appellof (Plate IV), the radula bears only unicuspid teeth.

POSTEMBRYONIC STAGES

Appellof (1891, p. 26) suggested that the embryo of Grenacher (1874) is
a Chaunoteuthis because it has stump- shaped tentacles. This can only
be based on ignorance of the typical stages of Oegopsida. Not the tentacles,
but the 3rd and 5th arms of the Oegopsida are inhibited and stump- shaped
(p.234). We do not know the early stages of Chaunoteuthis which could
in any case not be distinguished from those of other Onychoteuthidae
(p. 304—309). On the other hand, I place in this species an animal the posi-
tion of which was doubtful for a long time (cf. also Grimpe, 1920, p. 295).

It was caught at a depth of about 150 m in the Naples plankton on 7 January
1903. Its proportions and the degree of development of most organs closely
resemble those of the corresponding stages of Onychoteuthis. The
habitus, however, is so different that one can hardly identify it as Onycho-
324 teuthis. Mantle, fins, head and arms of the specimen have a very delicate
thin-skinned, gelatinous consistency, while the corresponding stages of
Onychoteuthis are thick-skinned to thick- fleshed (I never found any
different specimens among numerous specimens examined). Artificial
changes of the tissues can be excluded; I do not know whether the "larva"
was caught alive, but it was certainly in good condition before preservation
and not macerated or otherwise damaged. A drawing made two days after
preservation (by Merculiano, Figure 158) shows that the specimen had the
same appearance as now, and had distinct, contracted chromatophores on
arms, head and mantle which are now partly bleached, in the same arrange-
ment as in the young Onychoteuthis.

303

FIGURE 158. Young Chaunoteuthis mollis. 2x. Drawn
by Merculiano after a specimen from the Naples plankton.

The jellylike consistency of the body is evident. Note also
the folds on the ventral side of the mantle.

Except for the characteristic consistency of the tissue and the flaccid
condition of the body, which is never found in Onychoteuthis, the larva
of Chaunoteuthis is identical with larvae of the same stage of Onycho-
teuthis. This proves only that the species to which the larva belongs is
more closely related to Onychoteuthis than to Ancistroteuthis.

A larva of Ancistroteuthis of this size could be identified with certainty,
but this is not the case with much smaller specimens (cf. p. 309).

This larva has a dorsal mantle length of 12.5 mm and a fin length of
4.5 mm. A small O n y c h o t e u t h i s from Messina has the same measure-
ments and form of fins. However, the mantle of this specimen is cylindrical
anteriorly and has a typically pointed, not spindle-shaped or bulging posterior
part. The gladius is visible only anteriorly and between the fins. The
gladius is covered by the muscular mantle before the fins. This is an
important difference from Onychoteuthis (cf. Figure 156), especially
because the tissue is transparent enough to show the whole rhachis.
Otherwise, the larva closely resembles Ancistroteuthis, in which the
corresponding part of the gladius is also covered by the muscular mantle.

In Onychoteuthis of similar size, on the other hand, the gladius is
visible as a dark median line which becomes slightly narrower before the
fins, but remains distinct, and a crest appears at an early stage.

32 5 The head shown no characteristic differences from Onychoteuthis,
except the state of contraction. The head projects free anteriorly; in
Onychoteuthis of similar size it is always retracted (Figure 157),
unless the animal is preserved with great care after being anesthesized.

As in many preserved larvae of Oegopsida, the left eye projects partly from
the orbit due to withdrawal of the flaccid lid from the eyeball. This shows
an important character: the ventral side of the golden brownish eyeball
bears a padlike luminous organ, as in On y c h o t e u th i s (p.304). It is
yellowish brown and has a process toward the anterior side of the eyeball.

The organ is directed slightly toward the middle and is also visible on the
other eye through the wall of the orbit. The rounded lid margin of the right
eye shows a modification of the sinus. Both eyes have a wide open iris mar-
gin which projects all around; the lens has a blackish central part.

304

The neck fold which bears the olfactory tubercle is indictinct. The
funnel is very delicate and more weakly developed than in Onychoteuthis.
On the other hand, the arm apparatus is developed as in O. banks! of
similar size, except for the gelatinous consistency of the tissue. The arms
bear 2 rows of very small suckers. The greater part of the tentacle clubs
is occupied by 4 rows of still smaller suckers and with 2 rows at the base.
The dorsal and ventral arms are distinctly shorter than the lateral arms.
Swimming margins are already visible on the third pair. The tentacles are
1.5 times as long as the third arms. The buccal funnel has 7 distinct points
and the typical bonds.

Opening of the mantle cavity shows almost the same picture as in
O. banks! of similar size. All organs have a typical arrangement (inter-
mediate between Figures 149 and 157). A rounded, lens- shaped brownish
luminous organ is already well developed in the median part of the ink sac.

As in similar stages of Onychoteuthis, examination with a lens shows
no trace of the anterior luminous organ near the anus. The posterior
organ has not yet pushed the hind intestine aside, as in slightly older stages
of Onychoteuthis, but is situated below the hind intestine (cf. p. 319).

The intestine is still more or less median.

The jaws already show the typical characters of the Oegopsida (Plate XVII,
Figure 5). They appear at an early postembryonic stage and are identical
with those of the young Onychoteuthis.

Our larva thus differs from stages of Onychoteuthis as follows:

1) tissues gelatinous; 2) middle of gladius surrounded by the muscular
mantle; 3) funnel small.

305

326 Chapter 20

GENUS ANCISTROTEUTHIS
Gray, 1849

ANCISTROTEUTHIS L I C H T E N S T E IN I
(d'Orb., 1839) Gray, 1849

DIAGNOSIS

Luminous organs absent in the mantle cavity or on the eyeball.

Median ridge of gladius not visible through dorsal skin and forming a distinct
crest only posteriorly. Rostrum particularly long (V? of length of gladius),
flat-oval in cross section, chitinous-cartilaginous. Carpal pad elliptical,
with 8—12 suckers and an equal number of knobs. Clubs with 10—12 hooks
in each row.

LITERATURE

1839 D’Orbigny, Onychoteuthis 1 i ch t e n s t e in i i (p. 334, Plate 8, Figures 8— 12; Plate 14, Figures 1—2).
1849 Gray, A n c is t ro te u th is li chte nste inii (p. 55).

1851 Verany, On y c ho te u t his 1 i chte ns te in i i (p. 75, Plate 29).

1869 Targioni-Tozzetti, A n c is t r o t e u t h i s 1 i c h t e n s t e in i i (p. 50).

1889 Cams, A n c is t r o t e u t h is 1 ichtenste inii (p.449).

1896 jatta, A n c i s t r o t e u th i s lichtensteinii part (Figure 13, p. 11; Figures 61, 62, p. 28; Plate 13,
Figures 13—23 mainly of Onychoteuthis banksi).

1900 Joubin, A n c i s t ro t e u th is lichtensteinii (p. 62).

1900 Pfeffer, A ncistroteuthis lichtensteinii (pp. 149—160).

1904 Hoyle, A n c i s t r o t e u th i s lichtensteinii (p. 18).

1904 jatta, A n c i s t r o t e u th i s lichtensteinii (p. 200).

1912 Pfeffer, A ncistroteuthis lichtensteinii (pp. 92—98).

1916 Naef (Syst.), A n c i s t ro t e u th i s lichtensteinii (p. 15).

1921 Naef (Syst.), A ncistroteuthis lichtensteinii (p. 537).

STRUCTURE OF THE ADULT ANIMAL

A well preserved adult specimen is illustrated by Pfeffer (1912, Plate IX,
Figure 1; Plate X, Figure 1). All my specimens are deformed and badly
preserved and need reconstruction for any idea of the natural relationships .
to be obtained. I have therefore not given a drawing of the whole animal.
The specimens resemble O. banksi so closely that Jatta (1896) confused
the two species. The main figures of Jatta (14 and 19, Plate XIII) certainly

306

327 represent Onychoteuthis and probably also the other drawings, but an
exact determination is not always possible because of the inadequate
drawings.

Ancistroteuthis is in general more slender than Onychoteuthis.
The arms are longer and its color is lighter (orange-yellow, wine red to
reddish brown) and is easily distinguished from Onychoteuthis, because
the dark longitudinal line on the dorsal side (crest of gladius) is absent, as
the gladius is completely surrounded by the mantle. A number of additional
differences are given below.

The gladius (Pfeffer, 1912, Plate X) consists mainly of a wide rhachis;
the flag is quite rudimentary and present only in the posterior part; it
passes into the spoon- shaped, also reduced cone, the weak development of
which is already evident in the young stages (p. 328, Figure 159; cf.

Figure 156). The ventral margin of the conical cone of Onychoteuthis
is apparently lost. On the other hand, the rostrum is particularly long,
more than V? of the length of the gladius. It is also slightly more massive
than in Ony c h o t e u th i s, but less firm and more cartilaginous. The
characteristically pointed form of the mantle sac is shown in Figure 162,

The fins are pointed posteriorly and longer than wide, in contrast to the
young animal (Figure 160). They are widest slightly before the anterior
third (cf. Pfeffer, 1912, p. 93).

The neck folds show the typical conditions of the family (p. 302) and
subfamily (p. 311). There are about 10 occipital folds on each side, those
directed toward the middle become progressively smaller and less distinct.
The occipital folds are situated between two sharp transverse folds. The
funnel pit is more sharply delimited than in Onychoteuthis.

The arm apparatus is strong and very long. The arms reach ^4 of the
mantle length. The second arms are the longest, but are only slightly
longer than the third pair, and the V arms are only slightly shorter than
the third pair. The D arms are the shortest (formula: 2, 3, 4, 1). The
swimming and protective margins show no special characters. The tentacles
are also very long. My largest specimen measures 32 cm, including the
arms; the tentacles project by the same length beyond the end of the arms
and are unusually strong. The fully developed club (Figure 154) differs
from that of the young stage described below (Figure 161) as follows:

1. The hand part bears completely developed hooks, and the suckers finally
disappear at the beginning and end of the median rows of the hand part.

2. The marginal rows are lost; the suckers have fallen off. 3. The carpal
suckers and knobs are concentrated in a longitudinal oval carpal pad, which
still shows an arrangement in rows and contains 9—12 alternating suckers
and knobs. The pad is slightly raised and surrounded by a swollen edge
the lateral parts of which have developed from the primary protective mar-
gins. The number of hooks varies; I found 21, 22, 23, 23, 22 hooks on the
right club of 5 specimens and 21, 22, 21, 22, 22 on the left clubs.

The radula shows the typical characters of the Teuthoidea. As in most
(all?) Oegopsida, there are only traces of the marginal rows, and the teeth
on each side of the median rows have a distinct lateral cusp, which is weakly
developed in Onychoteuthis (Plate XV, Figures 2 and 3).

328 The jaws are also slightly more differentiated than in Onychoteuthis
(Plate XVII, Figures 4 and 5). The biting process of the upper jaw is very
long and slender; the outer and inner plates meet behind it in an edge which
forms a blunt angle.

307

The lower jaw has an accessory tooth directed inwards and slightly
covered by the margin in addition to the typical structures (note the position
of the arrow). The inner plate has an accessory strengthening ridge close
to the connection with the outer plate.

The mantle cavity shows the same picture as inOnychoteuthis,
except the characters caused directly by the absence of luminous organs
(cf. Figures 155, 157). The hind intestine is straight.

POSTEMBRYONIC DEVELOPMENT

329

Among the young stages of Onychoteuthidae with a dorsal mantle length
of less than 1 1 mm, there are certainly specimens of Ancistroteuthis

which could be identified by the absence
of "anlagen" of luminous organs. The
following young specimen certainly belongs
to A n c i s t r o t e u th i s (Figure 159).

This specimen has a dorsal mantle
length of 16 mm. Extended and including
the tentacles, it is about 3 cm long. It
differs from a stage of O. bank si of
similar size (Figure 156) in the short fins,
the absence of a dark dorsal median line
and the absence of luminous organs on
the eyes. The gladius is surrounded by
the muscular mantle in its greater part,
at least in the anterior half. Between
the anterior end of the bases of the fins
it is still visible through the skin as a
dark stripe which narrows rapidly
anteriorly and more gradually posteriorly.
The fins are connected in the posterior
quarter of the base and cover the gladius.
Examination of the ventral side shows a
special character of the gladius: the cone
is shortened to a minute spoon, while in
Onychoteuthis of equal size it still has
a deep- conical end part. The fins are about
5.6 mm long, i. e. slightly over Vs of the
dorsal mantle length; their span is Vs of
the dorsal mantle length. The fins are trans -
verse-rhomboidal, with distinct "earlobes."
Also characteristic is the distribution of the chromatophores, most of
which are preserved on the dorsal side. They include a median row of large
brownish red spots in the area where the muscular mantle covers the
rhachis. In stages of Onychoteuthis, there are alternating chromato- '
phores on each side of the median dark line in the same area as in
Ancistroteuthis and homologous to them. The mantle margin is
typical.

FIGURE 159. Young form of Ancistro-
teuthis lichtensteini. 2x. Col-
lected in the deep plankton of Naples in
1909. Compare with the specimen of
O ny c h o t e u th i s in Figure 156. Note
particularly the form and size of the fins,
the mediodorsal chromatophores, the
barely visible gladius behind the chro-
matophores, the weak development of
the cone and the proportions of the arms.

308

The third longitudinal neck fold is already distinct, with a prominent olfac-
tory tubercle posteriorly; traces of the second longitudinal and anterior
transverse folds are also present. The funnel shows the typical structure
of the Onychoteuthidae, but the funnel cartilage is particularly long and
projects anteriorly beyond the funnel pockets.

The head has the typically rounded form of young Onychoteuthidae. The
eyes have large, convex lenses, and the lid margin has a rounded sinus.
Luminous organs are apparently absent on the ventral side of the eyeball.

The tentacle club resembles that of similar stages of Onychoteuthis
(Figure 152). Four rows of suckers occupy its entire length; the stalked
suckers are already lost. The knobs and pad of the carpal part are still
absent, and the suckers of this zone pass without boundary into those of the
hand part. Five suckers of the ventral median row of the hand part are
already transformed into hooks; the suckers, proximal and distal to them,
show some beginning of conversion, e. g. a more or less distinct tooth.

The other suckers are normal, slightly ovoid, those of the short distal part
very small, the last only "anlagen" and not functional. The hooks are still
short but typical in principle, with a wide base attached to the surface. A
protective margin is present on the ventral edge of the hand surface, and
the dorsal margin is absent or only indicated, especially in the carpal area.
The swimming margin extends over % of the club, proximally reaching
the base of the hand part; it tapers suddenly to a mere edge just before the
proximal end, as in O n y c h o t e u th i s (Figure 152). Both protective mar-
gins end as parallel edges near the proximal part of the stalk.

The arms show the same relationships as in an Onychoteuthis of
the same size (Figure 156). Both lateral arms are much longer than the
dorsal and ventral arms; the first pair is slightly longer than the fourth,
the second slightly longer than the third (formula: 2, 3, 1, 4). Swimming
margins are already recognizable: they are indicated in the distal third
of the first pair, as a narrow outer edge along the whole second pair, and
as a distinct margin in the distal part of the third pair, which tapers
proximally to a narrow edge. All arms bear 2 rows of suckers which are
particularly dense on the ventral arms and smaller than the others. Pro-
tective margins are distinct everywhere, but not widened and without
differentiated supports. The function of the supports is taken over by the
pillars of the suckers, which are fused with the margins.

330 The buccal funnel has the 7 typical supports, points and bonds. The

mantle cavity of this specimen, a young male, shows the typical relationships,
the only differences from those in Figure 157 being the absence of luminous
organs and the unchanged position of the hind intestine, because of the
absence of the posterior luminous organ.

The specimen still shows traces of a bright, iridescent metallic sheen,
greenish on the mantle, dark bluish on the head.

An older male is shown in Figure 162. Measurements: length (without
tentacles) 53 mm, dorsal mantle length 30 mm, fin length 15 mm. The
general habitus is that of a stage of Onychoteuthis of similar size,
but the tentacles are longer and the fins slightly shorter. There is almost
no trace of the dark dorsal line, the rhachis being completely surrounded
by the muscular mantle. The rostrum, 3.6 mm long, occupies V4 of the
length of the fins and is visible on the ventral side of the apex (Figure 157).
Also visible is the very low margin of the cone which is spoon- shaped at a

309

point which is still deep and conical in
such stages of Onychoteuthis. The
fins are transverse-rhomboidal, only half
as long as the mantle sac in the dorsal
midline, with "earlobes" and a slightly
pointed posterior end. Their span is
4.8 mm. The mantle sac is typical,
widened anteriorly, with blunt dorsal and
sharp ventral marginal corners.

The neck folds are further developed.

The fourth longitudinal fold is distinct, the
third is strong and bears posteriorly the
olfactory tubercle, which is directed
outward. The lateral second longitudinal
fold is weaker. Still less developed are
two dorsally adjacent folds, of which the
larger lateral fold is, in my opinion, the
first normal longitudinal fold, the other
belongs to the secondary "occipital folds."
However, the median fold may represent
the first normal longitudinal fold, and the
other would be an intercalated fold which
is also present in Onychoteuthis. At
any rate, the whole system of occipital
folds is related to the first longtitudinal
neck fold which it replaces in a more de-
veloped form (see p. 314). Transverse
folds are also indicated, but only the
anterior is distinct. The fourth longitudinal
neck fold is adjacent to the marginal edges of the funnel pit, which converge
anteriorly almost at a right angle.

There is no trace of luminous organs on the eyeballs. The greater part
of the lens is covered by the contracted lid fold which has a distinct sinus.

The whole arm apparatus is relatively stronger and longer. The third
pair is almost as long as the second; the fourth pair is much longer than
the first, but still shorter than the others (formula: 2 = 3, 4, 1). Swimming
margins are distinct in the terminal part of the first arm and on the outer
edge of the second arm, but are widened in the middle of the third arm,
where they taper distally and proximally form only a thin edge.

Marked further development is evident in the tentacle clubs (right club
shown in Figure 161 in greater detail). The 4 primary longitudinal folds are
still present along the whole club, but with numerous modifications. The
carpal part bears 2 oblique rows of 4; one of these rows is incomplete and
then follows a further row of 4 in which only the 2 dorsal suckers belong to
the carpal part. The third sucker of this row (marked with an x on
Figure 161a, enlarged in Figure 16 If) shows a transition to a hook and
belongs to the ventral row of hooks. The rows of 4 carpal suckers alternate
with rows of knobs, of which only one is complete. The knobs correspond
to the suckers of the other club, on which there are 1, 3, 4, 3, 1 suckers in
332 the successive rows; the last 2 rows are complete in themselves, but only
the outer, dorsal suckers belong to the carpal part. The number of knobs

FIGURE 160. An older stage of A nc is -
troteuthis lichtensteini.male,
caught in Naples on 23 May 1913. 2x.
Compare with the preceding stage, and
note particularly the formation of ten-
tacles, arms, neck folds and fins. The
gladius is not visible through the dorsal
222 skin, as it is completely surrounded by
the muscular mantle.

310

is accordingly 2, 4, 4, 2, the last of them indistinct. There are thus 12
suckers and 12 knobs in the carpal part, which is not yet transformed into
a carpal pad. Then follows the hand part, which consists of 2 median rows
which consist mainly of hooks, and 2 marginal rows of suckers, with a
number of slightly enlarged suckers in the distal part of the ventral row.
Three suckers are already lost in the distal half of the dorsal marginal
row, symmetrically on both clubs. The position of these suckers is marked
with small crosses in Figure 161a. The dorsal median row begins on the
right club with a very small booklet (situated obliquely before x) and ends
in an intermediate form between sucker and hook, shown in Figure 161a (y)
and 161 e. There are thus 11 elements, of which 10 are true hooks, in the
dorsal median line. The ventral median line begins with x, a small sucker
with a hooklike tooth (slightly larger than in Figure 161e) and with 10 other
hooks, of which the sixth is the largest.

(331)

FIGURE 161. Right tentacle club of the stage in
Figure 160. 6x. With highly magnified hooks
and hooklike suckers. The small crosses in the
main figure (a) mark the position of lost suckers
of the dorsal marginal row: b) large prehensile
hook of the lower side; c) the same, lateral;
d) rotating clasping hook of the dorsal median
row; e) recently formed, undeveloped clasping
hook of the left hand; f) sucker marked with an
X in the main figure, highly magnified, showing
transition to a hook;

Kp — hood; Hk -hook; St - stalk of hook;

Hr — horny ring of original sucker; Rz - grasp-
ing ring of same; Rd - marginal ring of same;

Ba - basal part of (modified) horny ring; Ei -
typical invagination of horny ring; Fo - process
of this; Tr - pillar of sucker or hook; y — distal
hooklike sucker (transitional stage, like e);

Sm — swimming margin; Ds — dorsal protective
margin; Ve — ventral protective margin; z —
carpal knob.

On the left club, each hand row has 11 elements, the first and last of which
are transitional stages as in Figure 161e, which shows the terminal element
of the dorsal row. Each of these tows bears 9 well developed hooks, although
some of them show still some marked juvenile characteristics. If this
specimen were adult, it would probably bear 23 and 22 hooks on the right
and left clubs, respectively. The hooks of the dorsal row are typical clasping
hooks, those of the ventral row are prehensile and are constantly rounded
anteriorly and downward, but the 3 proximal hooks are again of the clasping
type, as far as their character is already distinct. The distal part of the
club bears only 3 rows of 4 small suckers, followed by 3, then 2 single
suckers (on the left side 4X4, then 3); it is evident that the first row
passes completely (on the left side) or partly (on the right side — the

311

2—3 upper suckers, as y in Figure 161a) into the hand part. These suckers
are larger and less flattened than the others. Some of them probably
develop into hooks.

This distal part resembles a typical end part (p.229), and it could be
assumed that a typical distal part of Onychoteuthidae has been included into
the area bearing hooks which we consider here as the hand part.

A typical swimming margin reaches to the base of the hand part along
about 74 of the club, leaving the apex free. The protective margins begin
before the carpal part as thin ridges continuing the edges bordering the

inner side of the stalk. The dorsal mar-
gin rises slightly at the base of the hand
part and tapers further on. to a low,
almost blunt edge which forms the distal
boundary of the inner surface of the club.
The ventral protective margin widens
from the middle of the carpal part onward
to a thin, smooth- edged skin fold without
distinct supports. In the distal half, this
margin tapers gradually, and becomes not
only narrower, but also stronger and is
connected with the pillars of the suckers
so that these are apparently situated on
the base of the margin. The protective
margins of the arms are low edges with
barely visible supports which cause a
slight dentition corresponding to the
suckers.

The mantle cavity resembles that of
a young Onychoteuthis (Figure 157),
except that luminous organs are absent
and the gonoduct is further developed and
has a prominent proximal part.

The proportions and habitus of this
young stage of Ancistroteuthis change
markedly during further development.

The arms become relatively longer, the
tentacles very long and strong (thick when
contracted). The changes are most
marked in the posterior region because of
the growth of the apex and the enlargement
of the fins. This growth is associated with
the formation of a long, strong rostrum at
the end of the gladius, but also with the
relative narrowing of the posterior part of the mantle sac (as in Alloteu-
this. Figure 104). At the same time, the fins grow, forming a terminal
point and finally occupying almost Vs of the dorsal mantle length. Figure 162
shows the posterior part of one of the largest specimens.

There is no marked change in the typical neck folds. However, further
"occipital folds" appear medially from the longitudinal folds described, to
10 on each side.

Mature specimens are not known, but this species certainly grows to a
greater size than Onychoteuthis.

(333)

FIGURE 162. Body of an adult A nc ist ro
teuthis. V2 X. Note the larger size and dif-
ferent form of the fins and the form of the
mantle sac. The narrowing behind the
middle of the mantle sac is not always so
distinct and the transition to the tail- like
posterior end may be more gradual, accord-
ing to the state of contraction (cf, Pfeffer,
1912, p. 92).

312

334 Chapter 21

FAMILY OCTOPODOTEUTHIDAE
Berry, 1912

(For "Veranyidae" Chun, 1910)

DIAGNOSIS

Suckers on arms biserial, their greater part converted into hooks. Ten-
tacle clubs with strongly reduced number of suckers already in youth;
tentacles later lost. Buccal funnel with only 6 points, supports and bonds.
Fins firmly fused in the middle, covering finally the greater part of the
mantle sac.

This family was first named by Chun (1910) "Veranyidae" (Veranya
Krohn = Octopodoteuthis Riippell) as he recognized its independence
from the Enoploteuthidae. Pfeffer (1912) placed it again in the Enoplo-
teuthidae as the subfamily Octopodoteuthinae. Berry (1912) corrected the
terminology to its present form. In addition to Octopodoteuthis, the
family includes only two little known species, Octopodoteuthopsis
megaptera (Verrill, 1885) Pfeffer, 1912 and Cucioteuthis molinae
(d'Orb., 1839) Pfeffer, 1912. I, therefore, give no further description of
the family.

GENUS OCTOPODOTEUTHIS
Riippell, 1844

OCTOPODOTEUTHIS SICULA Ruppell, 1844
DIAGNOSIS

Body short, compact. Skin swollen, gelatinous. Arms swollen apically,
club-shaped. Clubs of young forms with only 7—8 suckers in 2 rows. Arm
formula: 2, 3, 1, 4.

335 LITERATURE

1844 Ruppell, Octopodoteuthis sicula (p. 135).

1845 Krohn, Octopodoteuthis sicula (p.47, Plate 5, Figures A— F),
1847 Krohn, Veranya sicula (p.39).

1849 Gray, Octopodoteuthis sicula (p. 51).

313

1851 Verany, Ve ra ny a sicul a (p. 86, Plate 28) and also Loligo meneghinii (p. 105, Plate 35,
Figure c, dd).

1884 Pfeffer, Octopodoteuthis sicula (p.28).

1886 Hoyle, Ve r a n y a sicula (p. 6, Figures 12 — 23).

1889 Weiss, Verany a si c ul a (p. 87, Plate 8, Figures 1—3).

1896 Jatta, Veranya sicula (p. 92, Plate 7, Figure 14; Plate 13, Figures 1—12),

1899 Ficalbi, Veranya sicula (p. 83).

1900 Pfeffer, Octopodoteuthis sicula (p. 166).

1907 Massy, Octopodoteuthis sicula (p. 381).

1909 Massy, Octopodoteuthis sicula (p. 28).

1910 Chun, Octopodoteuthis sicula (pp. 139— 147) , Plate XVII (species?).

1912 Pfeffer, Octopodoteuthis (pp. 213— 222, Plate 19, Figures 1—16).

1916 Naef ( Syst.), Octopodoteuthis sicula (p. 15).

1921 Naef (Syst.), Octopodoteuthis sicula (p. 537).

Adult specimens of this species are not known. The oldest juvenile
stages appear like adults and we describe them in connection with the onto -
genetic development.

POSTEMBRYONIC DEVELOPMENT

Lo Bianco (1909, p.657) was of the opinion that the spawn the embryos of
which are shown in Plate Vlll (Vol. II) belong to Octopodoteuthis. This
is definitely incorrect; compare the volume dealing with embryology and
note especially the rudimentary tentacle of Octopodoteuthis in com-
parison with that of the Oegopsid x, which already bears numerous suckers
at hatching (Figure 67).

Chun (1910, Plate XVII, Figures 11, 12) described very young postembry-
onic stages of Octopodoteuthidae. These forms certainly belong to this
family, but their species cannot be determined. They obviously belong to
Octopodoteuthis, as shown by the structure of the tentacles: "The
tentacle club has 5 suckers. The 2 small proximal suckers belong to the
carpal part; the 2 following suckers, which are much larger, and the small
distal third sucker belong to the later hand part of the club (but see p. 339),
which is already bent obliquely from the tentacle stalk" (Chun, p. 146). The
third and fourth arms are still only "anlagen" without suckers; the first
and second pair bear one small sucker each, as is typical for the youngest
larvae of Oegopsida. The fins are still very small, separated, subterminal.
The dorsal mantle length is 1.2 and 1.7 mm.

Similar young stages are illustrated by Pfeffer (1912, Plate XIX, Figures
14 and 15, 11 and 12). The fins of these forms are attached to a wide
lanceola (Figure 131).

Figure 163 shows a more advanced stage which will be described below
(See also Pfeffer, 1912, Plate XIX, Figures 5-8; Chun, 1910, Plate XVII,
Figures 5 and 6). The general appearance of all these forms is essentially
336 as in Figure 163, but the arms are less developed and terminal swellings
are present only on the DL arms. Arms and tentacles are still very short.
The youngest specimen is shown in Figure 163. This is a strongly shrunken,
balsam preparation, so that its size does not correspond to its stage of

314

development. The dorsal mantle length of the live animal would have been
about 4 mm, not slightly over 3 mm as now. Before mounting, the animal
had the typical thick, gelatinous skin of a young Octopodoteuthis. The
gladius is not well visible. But the relationship between gladius, mantle
and fins certainly resembles that shown in Figure 131, as proved by the
conditions in later stages. The fins are already large, rounded lobes, still
widely separated and attached to the outer surface of the gladius.

FIGURE 163. The youngest available stage of Oc topodoteu this sicula
from the deep plankton of Naples (old material). 4x. The dotted line along
the outline of the mantle indicates the thickness of the markedly swollen
skin. Note also the form of the fins, the prominent eyes, the length of the
arms, the club-shaped terminal swellings of the arms, and the form of the
tentacle clubs.

The mantle margin shows the 3 typical corners which are still little
developed. The eyes are very prominent but rather small and project
because of the form of the inner parts. The lid margin is rounded.

The head is characteristically constricted before the eyes so that the
arms are situated on a short, outer support which would be even thinner if
the constriction would not be made indistinct by the thick skin. The ventral
arms are shorter than the dorsal arms, and the laterodorsal longer than the
lateroventral arms, and they never reach their length; formula: 2, 3, 1, 4.

All arms bear 2 rows of hooklike suckers and terminal luminous organs
situated in club-shaped swellings. (I cannot determine the state of develop-
ment of the suckers.)

The tentacles are apparently complete (cf. the following stage).

Already at this stage, the larva described by Chun (1910, Plate XVII,
Figures 3, 4, 9) from the Agulhas Current differs from the Mediterranean
specimens. The stage of development is similar but the arm apparatus is
markedly less developed. There are much fewer suckers and terminal
swellings are absent, even on the second pair, which is the longest. On the
other hand, the tentacles are relatively strong. These are obviously two
different species and I propose the name Octopodoteuthis indica
nov. spec, for this larva. It is significant that this larva already has a
buccal funnel consisting of 6 parts at a dorsal mantle length of 3.8 mm.

337 The oldest stage described by Chun (Plate XVII, Figures 1, 2, 7, 8)
probably belongs to a third species. Although it is larger and more
developed, it shows no evidence of conversion of suckers into hooks. I
name this form Octopodoteuthis persica nov. spec. O. persica
is certainly distinct from O. sicula, of which another, much larger specimen
will be described below. The young O. persica has a mantle length of
only 4.7 mm, compared with 6.5 mm of the specimen of O. sicula described
below. The fins of the specimen of O. persica, however, are much larger,
occupying Vs of the mantle length and their span is more than 1.5 times that

* This resembles the Onychoteuthidae, which Octopodoteuthis also resembles in other aspects. A median
bond of the two primary ventral pillars presupposes a buccal funnel of 6 parts (but cf. also Plate III, Figure 2).

315

of the mantle length. On the other hand, the arm apparatus of O. persica
is even less developed than that of the stage described above (Figure 163):
the third and fourth arms are still short, the third arms are shorter than
the first, and terminal swellings are only present on the second pair. All
these differences are so marked that they cannot belong to the same species.

A stage younger than the stage described below but belonging to the same
species has been described by Chun (1910, Plate XIX, Figures 5, 6, 10).

o

FIGURE 164. Young Oc topodoteuthis sicula from the deep
plankton of Naples. 4x. Note the characteristic habitus: the thick,
gelatinous skin of head and mantle (marked with the dotted inner
line): the general form and median fusion of the fins: the part of
the gladius not covered by the muscular mantle, but only by skin:
the thick, swollen eye stalks: the presence of hooks and terminal
swellings on the arms: the characteristic clubs, the suckers of which
are shown in the small figure according to size and position.

I obtained a live young stage of O. sicula from the deep plankton of
Naples on 19 March 1910. This was a perfectly transparent specimen with
orange red chromatophores which rapidly widened and contracted. Gills,
branchial hearts, heart, aortae, ink sac, statocysts, optic ganglia, the optic
section of the eye and other details were easily recognizable during
cocainization.

The preserved specimen is shown in Figure 164. The gladius is appar-
ently Loli go-like, in the form of a wide stalked leaf, probably with a flat,
spoon- shaped cone at the posterior end. Only the anterior part of the
338 gladius is visible, as the posterior part is covered by the fins; at any rate,
the surrounding mantle is still little advanced. The fins are rounded, very
large (the form shown in Pfeffer, Plate XIX, Figures 3 and 4 is certainly an

316

artifact) and still without distinct basal "earlobes" or lateral corners.

Their bases are contiguous on the gladius and begin to become fused. The
posterior margin of the fins projects distinctly beyond the apex, forming a
median incision. There is a weak indication of an apex so that the posterior
margin of the fins is not directly attached to the posterior curvature of the
mantle in the middle either. The posterior apex becomes later stronger.

The three corners of the mantle margin are only indicated, the margin is
transverse, with slight dorsal and ventral indentations. The funnel is rela-
tively weak. The appearance of the animal suggests a planktonic, floating
form rather than a strong swimmer.

The head is characteristic and the relatively small eyes are situated on
stalks. The eyes appear shorter and larger than they actually are by the

swollen skin. The resulting form strikingly
resembles the head of the middle embryonic
stages of typical Decapoda (cf. Vol. 11, Plate IV).
The lid fold is normal and the olfactory organ
is also a flat, oval papilla in the typical position.
A flat, differently colored tubercle is already
distinct on the ventral side of the eyeball of
these stages. The tubercle is sometimes
yellowish brown, like the corresponding luminous
organ of Onychoteuthis, with which it is
certainly homologous.

The ventral arms have remained the shortest.
The dorsal arms are slightly longer and the
lateroventral arms are still longer. The
laterodorsal arms are distinctly the longest
(formula: 2, 3, 1, 4). This condition is definitive,
but the differences are less distinct in the
largest specimens.

The gelatinous skin is markedly raised on
the outer side of the second and third arms, and
indicates swimming margins which later re-
semble the form in the Onychoteuthidae, but
they remain gelatinous, swollen and indistinctly
defined.

All 8 arms have club-shaped terminal swell-
ings which taper to a blunt apex. These swell-
ings are undoubtedly luminous organs, analogous
to those of Abraliopsis. Suckers are absent
on the terminal swellings. The arms bear 2
inner rows of prehensile organs, partly suckers,
partly hooks and transitions between them. There are suckers with round
rings at the base, followed by characteristic, elongated, hooklike suckers
with a hooklike tooth which becomes increasingly marked toward the middle
of the arms, where they form short hooks. This is reversed toward the
apex, which bears barely distinguishable "anlagen" of suckers. The suckers
of the ventral arms are smaller and not so far advanced in the middle part
that they could be considered as true hooks. The protective margins are
only low edges with tubercles corresponding to the supports.

FIGURE 165. Embryo of Sepia
officinalis. Note the surpris-
ing resemblance to Figure 164,
especially in the form of the head,
cheek and eye regions. Further
explanations in Vol. II:

1—4 — arms; 5 — tentacles; 6 —
yolk sac; 7 — eye; 8 — iris; 9 —
posterior border of "head cover"
(see Vol. II); 10 - occipital bond;

339 II _ cheek tubercle; 12 - funnel
pocket; 13 — shell; 14 — first
septum; 15 - Hoyle’s organ; 16 -
apex of body; 17 - fin.

317

The tentacles are delicate, barely as long as the third pair of arms and
the small clubs are characteristic. It is impossible to distinguish between
hand and carpal parts because, comparatively speaking, the whole armature
of suckers corresponds to the first stalked suckers of other larvae of
Oegopsida (p. 235), which usually appear singly and are arranged in 2 rows.
The formation of further components is generally retarded in Octopodo-
teuthis, so that we have a purely larval structure before us. The club
normally bears 7—8 suckers (7 on the left side of our specimen) of which
the first 2 are very small, the next 3 much larger, and the distal 3 again
decreasing in size. The surface of the club which bears them is slightly
widened and pointed, with apparently protective margins on the distinct
lateral edges.

The stalk is thick at the base and tapers gradually. Only its inner side
has a denser, muscular structure which probably corresponds to the mus-
cular axis of the tentacle stalk. The other mass consists of gelatinous
tissue which probably belongs to the skin, and there is the question whether
such swollen structures can still be forcefully projected. There are
distinct superficial circular fibers the contraction of which possibly pro-
duces a moderate, not too marked lengthening of the stalk or contributes
to the lengthening caused by the muscular axis. At any rate, the stalk has
a small efficiency, and the tentacles are apparently ephemeral structures.
The larvae probably feed on easily obtainable small plankton, weak enough
to be caught by this imperfect apparatus. The tentacles apparently remain
functional only until the arms attain their definitive form and armature of
hooks. They are thus true larval organs, since they are later lost. This
takes place when the dorsal mantle length is about 11 mm as both tentacles
are present in smaller animals, while only little older specimens have only
small basal stumps."*'

340 The oldest specimen that still had well preserved tentacles had a dorsal
mantle length of 10.3 mm and fins 6.8 mm long. The fins have thus grown
not only absolutely but also relatively. Their form is also different: small
"earlobes" begin to develop, and the posterior indentation has disappeared.
The two fins together form a transverse oval the width of which is two-thirds of
the mantle length and which is distinctly longer than the dorsal mantle length
(11.5 mm). The posterior apex of the body is already distinct and resembles
in principle that of the young Abraliopsis (Figure 141). The swollen
skin, however,. encloses the apex completely, so that it is recognizable, only
because of its transparency. In later stages, it is barely recognizable.

The arm apparatus resembles that of the preceding stage, except that each
tentacle bears 8 normal suckers and the transformation of suckers into
hooks is more advanced. The arms are stronger, longer and the constric-
tion before the eyes is less marked.

" Verany (1851, Plate XXVIII) illustrates a specimen which is perhaps slightly larger (magnification not
given) and still has tentacles, without further details. Loligo meneghiniiof this author (Plate XXXV)
is a younger specimen of this species and certainly not a Calliteuthis (q.v.). This is proved by the
length of the arms, the form of the fins, the biserial pattern on the clubs and its general habitus. Verany
(p.98) describes the body as fleshy ("charnu"), but this does not say much. The method of preservation
(alcohol) can obliterate the gelatinous character. This may be one of the numerous minor errors of obser-
vation or an inaccuracy of description, which matters little in view of the exact morphological data
and drawings. Another error is apparently that he states that there are more than 8 suckers on the club
(Plate XXXV, Figure e).

318

In a slightly larger specimen, of a dorsal mantle length of 11.6 mm, only
short basal stumps are left of the tentacles. The fins have grown to just
over Vs of fhe mantle length, and the whole development has progressed in
the direction indicated. The posterior end is already deformed, as in the
later stages: during preservation, contraction of the ventral part of the
mantle probably caused the end of the mantle sac and apex to become curved
downward together with the posterior end of the fins, so that the median

part of the mantle and fins appear markedly
shortened, blunt, so to say, drawn in. This
has been observed in all older specimens
and is shown in Figure 167, together with a
reconstruction of the conditions assumed
for the live animal. A further complication
is that the swollen skin encloses not only
the apex but also the end of the mantle sac
so that the posterior end appears atypically
blunt and the form of the fins appears wrong.
The structure of the fins adds to this wrong
impression. In Figure 167 a curve open (Ht)
posteriorly delimits a membranous postero-
median area which does not contain typical
fin muscles and is transparent in prepara-
tions (cf. p. 153). This modified area in-
cludes also the folded over terminal part of
the fin, which becomes even less distinct.

In addition, a wide stripe at the posterior
margin of the fin (Fr) is also membranous,
thin, always curved down or folded back.

This stereotypic behavior of the preserved
fin suggests that such a "deformation" of the
posterior end may take place temporarily
also during life. The animal is not a strong
swimmer, according to its plump form; its
gelatinous consistency, the thin muscular
mantle and small funnel suggest that this
is an almost planktonic form, feeding on
plankton, particularly in the young stages.

Figure 1 66 shows the situs of the mantle cavity of a half-grown specimen. The
short, very wide funnel bonds, and the very small mantle bonds are recognizable
(the funnel has a typical funnel valve and funnel gland). The mantle organs show
no special characters. The whole complex is short and wide, like the body. The
mantle cavity does not reach into the posterior part of the mantle sac.

It is not certain from the condition of the material whether the posterior
aorta is like in the Enoploteuthidae (p.264) or shows the typical characters
of the Oegopsida (the available specimen from Jatta's collection is strongly
damaged). The strong musculus rectus abdominis passes over the ink sac
on each side, as in young Onychoteuthidae (p. 320). It is thicker where it
touches the ink sac, and the thickening has the form of a lens pressed into
the sac. It is not certain whether this thickening develops from muscular
tissue; Chun (1910, p. 141) found that it is situated above a different tissue
and considered it as a luminous organ in development. At any rate.

FIGURE 166. Open mantle cavity of
Oc t o p o d o t e u t hi s sicula. 2x.
( Slightly reconstructed.) Note the
form of the funnel cartilage, the two
light-colored lens-shaped luminous
organs inside the ink sac (1), the
large musculus rectus abdominis (2)
passing over the ink sac, the form
of the posterior end (reconstruction
shown by a dotted line). Also shown
3^ are gills, branchial hearts, vena cava
(3), venous branches (4), heart (5),
posterior pallial veins (6), etc.

319

luminous organs are present in a corresponding position in the Chiroteu-
thidae (Plate IV, Figure 3) and Cranchiidae (C o r y n o mm a, Chun 1910,

Plate 60, Figure 13), and also on the left side in the Onychoteuthidae
(Onychoteuthis, p. 320; Chaunoteuthis, p. 32 5), so that homologous
structures may be expected also in this case.

In an older but still immature specimen (Figure 167), the fins are already
complete but also similarly deformed. They have "earlobes" and lateral cor-
ners, and occupy now over % of the mantle length. A dotted dorsomedian line
indicates the suture at which the musculature of the two fins is fused. The
gladius is still visible anteriorly through the skin as it is not yet surrounded
342 by the muscular mantle. The eyes are very prominent, but this is partly

obliterated by the swollen skin and the now much stronger arms. (On swim-
ming and protective margins and terminal swellings, see pp. 338—339.) All
except the very small terminal suckers are now converted into hooks.

Massy (1907, 1909) described a large (mature?) specimen with a dorsal
mantle length of 10.1 cm, from the Atlantic Ocean. It closely resembles
the specimen described above. My largest specimen, which is severely
damaged, has a mantle length of 3.6 cm.

FIGURE 167. Half-grown O c to p o d o t e u th is sicula from Naples. 2x. The dotted lines indicate the
gelatinous, swollen skin of the arms (protective margins, Ss) and mantle sac (Mm); the membranous margi-
nal parts of the fins (Fr),and the anterior part of the gladius (Gl), which is not surrounded by musculature.
Note the funnel adductors and olfactory papillae (Ro), which are visible through the mantle margin.

At - common base of arms, now internal (cf. Figure 164); Ek - terminal club (luminous organ) of an arm;
Fa — anterior margin of muscular part of fin; Ht — membranous posterior end of fin.

320

343 Chapter 22

FAMILY HISTIOTEUTHIDAE
Verrill, 1881

DIAGNOSIS

Habitus Octopus -like. Ventral side of head, arms and mantle with a
regular pattern of oblong, distinct luminous organs. Fins rounded, projecting
posteriorly beyond the calyxlike mantle sac and apparently connected
ventrally by a thin, fleshy, frenulumlike posterior apex. Left eye enlarged,
with well-developed luminous organs some distance from the lid margin and
with rudimentary organs nearer to the lid margin, in a posterior-upper
direction. Right lid margin with a dense ring of 16—18 large organs. Hand
part of tentacle with 7 rows of suckers at an early stage. Adhesion
apparatus of tentacle stalk and carpal part consisting of a loose row (or
double row) of suckers and knobs ending in a larger sucker or knob on the
carpal part. The latter elements followed by a row of 5 — 7 suckers and
knobs which reaches the dorsal marginal row of the hand part.

The Histioteuthidae are one of the most characteristic and peculiar
families of the Oegopsida. They show no close relationship to the other
families. The characteristic habitus of the Histioteuthidae develops at an
early stage, and only the youngest forms could be confused with the Ptery-
gioteuthinae (Figures 173 and 131).

TYPICAL STRUCTURE OF THE ADULT ANIMAL

The characteristic habitus of the family (Figure 168) is caused mainly
by the form of the fins, the apex of the body, the form of the mantle, the large
head, the asymmetrical eyes, the formation of the club and the very charac-
teristic luminous organs.

The gladius is Loligo -like, with an indistinct or very small, spoon-
shaped cone; it is only partly surrounded by the mantle. Not only the
median ridge of the rhachis but also a large part of the flag is visible as
a lanceola through the mantle, before the fins and between their anterior
part (Figure 181 ).

344 The fins are rounded, nearly circular, and form typical "earlobes”
anteriorly; their curvature projects posteriorly behind their base and the
apex, forming a median indentation in the middle of the posterior margin.
The fins are attached at the outer side of the gladius, which they partly

321

cover (Figures 168, 17 7). Their musculature, however, is fused in the middle
only in a small part, i. e. slightly behind the middle of their length from
where they diverge anteriorly and posteriorly (Figure 180).

FIGURE 168. Young Calliteuthis reversa Verr. from
Messina (Kolliker). 0.5 x. From the Stuttgart Museum of
Natural History. The typical habitus of the family is shown.
Note the Octopus -like habitus; the form of the fins; the
rounded head with reduced neck folds; outer and inner
funnel adductors; asymmetrical eyes, of which the left is
apparently larger; the characteristic club, which is thinner
in the middle and bears transverse ridges (5), w'hile the
dorsal margin contains a muscular cord (6). c) a single
luminous organ, slightly enlarged; d) part of the left
tentacle stalk.

1,2 - proximal normal adhesive suckers; 3 - sucker
situated on the ventral side of the tentacle stalk, which
keeps the resting tentacle atjiached to the V arm; 4 — edge;

e) cross section through proximal part of tentacle stalk;

f) cross section of distal part of tentacle stalk; g) cross
section of hand part of club.

The fins are thus not directly connected posteriorly but by a characteristic
thin, long apex (Pfeffer's "frenulum"). The mantle sac also does not extend
posteriorly to the median indentation of the fin, this place is occupied by the
apex, which has no connection with the muscular mantle.

The mantle sac is calyxlike, always widest anteriorly, with a weak, rounded
dorsal process at the anterior margin; and a distinct indentation of the
funnel, with indistinct corners which become very blunt in older animals.

The anterior transverse fold of the neck is a weak edge which reaches
the boundary of the flat funnel indentation (Figure 168). The second and
third longitudinal edge are slightly indicated. The olfactory tubercle is
usually isolated, as a large wart on the lateral-posterior side of the head.

The asymmetry of the eyes is very peculiar. My material permits nc
exact measurements, but the left eyeball and its lens always appear larger
than the right; in any case, the structure of the eyelid and its surroundings
is certainly different, and also the physiological condition, which is expressed
345 in the constant asymmetrical contraction in preserved specimens. The lid
margin of the left eye is usually flaccid and widened, that of the right eye
more strongly contracted. On the asymmetrical arrangement of the luminous
organs see p. 349. The sinus at the anterior margin of the lid is indistinct,
very shallow, and disappears if the lid is widened (Figure 171).

The arms are long but of different length; both lateral pairs are longer
than the first and fourth arms. The outer surfaces of all arms are flat,
with sharp edges which are connected by small membranes at the base, the

5110/2

322

basal membrane between the ventral arms is the smallest. The protective
membrane between the third and fourth arms is slightly better developed
than the others, and a "lateral margin" is indicated only at the base of the
fourth pair. The three upper pairs bear distally membranous swimming
margins in the middle of the outer side, between the still distinct outer
edges. Particularly strong is the margin of the third pair (Figure 185).

It occupies about the distal half of the arm, is very narrow in the distal
quarter but widens to a finlike lobe in the subapical quarter. The swimming
margin of the first arm is narrow and occupies only about one quarter of
the arm; that of the second arm is even shorter. The inner side of the
arms is bordered by distinct edges which bear well developed protective
margins, but without distinct supports. The protective margins reach the
base of the arm; between the three upper pairs of arms they are connected
at the base, forming inner membranes between two adjacent arms, in addition
to the outer membranes. The corresponding bond of the buccal funnel is
attached dorsally and dorsolaterally to the base of the inner membranes.

The arms bear two rows of suckers without teeth, which are more densely
arranged distally. The buccal funnel consists of 7 parts; the third and
fourth bond are attached on the inner side of the third and fourth arms
without passing into the protective margins.

The suckers are relatively large in early youth (Figure 174), but small
or very small in the older stages (Figure 168) resembling those of the
Sepiolidae, almost spherical. Their margin is characteristically differen-
tiated in the largest specimens: the chitinous adhesive ring (p. 98) is absent;
instead, the marginal ring extends inward to the suction chamber, forming
broad adhesive surface with glandular epithelium (Chun, 1910, p. 156) which
perhaps contains poison glands.

The tentacles are delicate in relation to the arms and bear much smaller
suckers. The tentacle stalk is thicker at the base, flattened, bandlike
between the third and fourth arms. The stalk becomes thinner and more
rounded distally. The inner surface of the tentacle is bordered on both
sides with edges which extend along its greater part and pass into the
protective margins of the club. Only the distal part of the outer side has an
edge which may bear a weak margin. The club is widened by the strong
protective margins, especially in the proximal part (Figure 168) and bears
a typical swimming margin in the distal half.

346 A loose, distally denser row of small suckers and knobs occupies the
greater part of the tentacle stalk. In the middle of the stalk, each two
knobs are followed by two suckers (Figure 169), the suckers of one tentacle
being situated exactly opposite the knobs of the other (p. 228). This row
ends a small distance before the club. Near it, but slightly more dorsally,
begins a new row which represents the carpal part and ends at the beginning
of the widening of the club in a particularly large sucker or knob which
already belongs to the hand part, which is not distinctly delimited. Sur-
prisingly, it bears 7 rows* of suckers. The median row consists of 1 or

2 small suckers, followed by about 6 larger suckers. The beginning of the

3 dorsal rows is also markedly different: it belongs to the adhesion
apparatus of 3 (2—4) suckers and the same number of knobs, arranged in

* It is possible that 8 rows were originally present, one of which was lost, but this does not agree with the
ontogenetic data (see pp. 350,355).

323

FIGURE 169. Tentacles of a young Calliteuthis reversa. 6X. The right
tentacle is shown on the left, the left on the right. About half of the stalk is shown.
Note particularly the structure and extension of the adhesion apparatus.

Vi(v) - last sucker (knob) of the stalk rows; x.x^ - first element of the carpal rows;
y,yi — last, large element of the carpal row; z,Zi — last suckers and knobs of the
adhesion apparatus of the hand part.

324

a simple or zigzag row directed upward; these elements are adjacent and
alternate with the enlarged sucker or knob at the end of the carpal row.

The beginning of the curve belongs to the third row, its end is situated in
the dorsal marginal row. The distal part of the club is not sharply delimited
from the hand part. It begins with an oblique, transverse row of 7 suckers,
followed by rows of 6, 5 and finally with the normal 4 suckers. The club
is slightly curved at the end. A distinct terminal part is apparently absent.

The funnel is weak but normal. The funnel bond closely resembles that
of the typical Enoploteuthidae and Onychoteuthidae (Plates III, IV), at least in
young forms; it apparently becomes later wider (Pfeffer, Plate 20, Figure 8;

347 Plate 23, Figure 4; Plate 23, Figures 1, 2), and of a more oblong-oval form
which is narrower anteriorly.

FIGURE 170. Luminous organ of the young Histioteuthis in Figure 185. 20 x.
a) upper view; b) median section (diagrammatic). The median section shows the
luminous body which consists of 2 parts, its anterior part (Lo) invaginated into the
posterior part (K). The posterior part is surrounded by a pigmented sheath (Ph.Ph^)
which extends to the window (Fe). Above the organ are chromatophores (Ch),
before and behind it differentiated parts of the argentea (Sp.Spp. Jr = argentea.

A chromatophore formed like an 8 (Me) covers with its anterior part the anterior
part of the luminous organ, the posterior part of the organ is covered by the posterior
part of the chromatophore, if it is expanded.

H2 — posterior margin of posterior mirror; — space above the latter; G^ — space
above anterior mirror; Ep - epidermis. About the more detailed structure see
Joubin,1893; Chun, 1910; Pfeffer, 1912; and Vol. Ill of this work.

The mantle cavity shows the typical relationships of the Oegopsida, but
the proportions are changed by the form of the mantle sac (Plate 2, Figure 3).
Luminous organs are absent. Chun (l910, Plate 20, Figure 7) observed a
noteworthy character: Calliteuthis has a functional gonoduct on both
sides, also in the male. This condition is undoubtedly typical for the

325

Decapoda and is general in the primary state of "anlage" (p. 154), but is a
unique phenomenon in mature male Teuthoidea and connected with the
bilateral hectocotylization of the distal parts of the dorsal arms (Chun,

1910, pp. 167 — 168). It is not certain whether this is also the case in other
Histioteuthidae. Examination with a lens did not reveal any distinct "anlage"
of the gonoduct in my young male (Plate III, Figure 3).

The nidamental glands of Calliteuthis develop in the typical position, i.e.
at the entrance of the posterior pallial veins into the renal sac. They
probably grow later rapidly anteriorly; in Histioteuthis, at any rate, they
are present as slightly developed "anlagen" (specimen in Figure 185)
situated further anteriorly. In later stages, their position resembles that in
other Oegopsida (Figure 207), as shown by a large specimen in the Naples
Museum of Natural History. The nidamental glands of this specimen also
closely resemble the projecting parts of the oviduct glands in form and
structure.

The luminous organs of the Histioteuthidae are particularly characteristic.
They are situated on the surface of the mantle on head and arms. Their
characteristics are in their structure, size and especially in their distri-
bution. They are relatively sparse on the upper side, where they are also
348 smaller and incompletely developed. On the lower side, however, they form
a dense cover of pearl-like structures which give a peculiar appearance to the
animal.

(349)

FIGURE 171. Lateral view of a young Calliteuthis reversa. Natural size.
Note the asymmetrical arrangement of luminous organs on the lid margins (dotted).
The lower figure (left side) shows also the outer protective membranes betvv'een
the arms.

Most luminous organs have the same structure, and only a few of them
are markedly different. The main part of the organ consists of an ovoid
luminous body, which is invaginated into the skin; it consists of an anterior
part (Lo) and a posterior part (K). The anterior part is invaginated into
the posterior part, which is enveloped by a pigment sheath (Ph, Phi) with a
round window (Fe) opening posteriorly and upward. The pigment sheath
corresponds to a chromatophore: it also develops from a chromatophore,
and the window can probably be widened and constricted in life. Light can
be emitted in two directions from the luminous body:* posteriorly through

* This may be connected with the emission of two colors of light, observed by Verany (1851, p. 119) in a
living Histioteuthis from Nice. Verany speaks of a bright sapphire blue and an even more beautiful
topaz yellow. Histioteuthidae have been caught sometimes among other nektonic Oegopsida on dark
autumn nights near Capri, using light to attract them. I could, unfortunately, not obtain a good specimen;
my knowledge is based on the vague reports of fishermen of radiant "totari," covered with pearls.

326

the window, through the transparent tissue (or fluid) which fills the cavity
Gi, or anteriorly, through the similar cavity G. The bottom of this cavity
is covered with a strongly shining part of the argentea or iridocyte layer
of the skin, which acts as a mirror (Sp) and in well-preserved specimens
apparently emits light, because of the strong reflection of incident light.

An argentea also covers the bottom of the cavity Gi, producing a weaker
sheen. According to Pfeffer, framelike thickenings are present around the
mirror and window of older specimens (l912, pp. 247—248).

The whole organ is surrounded and covered with regularly arranged
chromatophores. One of these usually has the form of a figure 8 and its
anterior part is situated on the free surface of the anterior part (Li), the
posterior part covers the window (Fe). The extended chromatophore
probably covers the window completely; some of the neighboring chroma-
tophores also partly cover the anterior part. Several larger and smaller
extended chromatophores cover the mirror (Sp), preventing the emission
of light if it becomes a danger to the animal. This takes place also in other
Cephalopoda, Not all luminous organs attain the typical development. Some
remain more or less rudimentary, especially on the upper side, but there
may be rudimentary organs also among the large, well developed luminous
organs on the lower side of head, mantle and arms. Other luminous organs
lack an anterior mirror while the window area is particularly well
developed. In H i s t i ot e u th i s, these organs form large, blackish brown
349 spots on the dorsal side of head and mantle (Figure 185), behind the left
eye and on the ventral side (Figure 177), The organs apparently have two
functions, emitting light posteriorly and anteriorly with different auxiliary
apparatus. One of these functions may be reduced in favor of the other.

The luminous organs have a very characteristic distribution on the
surface of the body, i. e. on the outer side of the arms, on the head and
mantle sac. The luminous organs on the arms form regular longitudinal
rows; they are most numerous on the ventral arms, where there are two
crossing systems of short, oblique transverse rows. A similar pattern is
present on the ventral side of the head, which forms a continuation of the
ventral arms in this respect. Near the margin of the ventral side of the
mantle there is a row of small organs; then follow a number of curved
rows, almost parallel to the former, their organs arranged in oblique,
also crossing rows. On the dorsal side of head and mantle, the number of
luminous organs is much smaller and their distribution is similar but less
regular.

Of particular interest is the arrangement of luminous organs in the eye
region, which shows the above-mentioned asymmetry.

The margin of the right eyelid (Figure 171) bears a dense row of
16—18 rather large organs which form a continuous border. Large, typical
luminous organs are also present on the adjacent ventral parts of the head.
The margin of the eyelid often appears bare at first, but it obviously differs
from that of the other eye. Closer examination of the anterior and lower
side shows a number of larger, normal organs some distance from the
margin; nearer to the margin there are very small, easily overlooked
rudimentarly organs. There are quite rudimentary organs on an almost
bare area behind the left eye.

327

The jaws of Histioteuthidae are typical for the Oegopsida, without any
special characters (Plate XVII, Figure 6). The biting processes are
moderately strong; the anterior edge behind them has an irregularly
jagged part. The radula is also typical for the Oegopsida (Plate XV,
Figure 1 ).

350 POSTEMBRYONIC DEVELOPMENT

The youngest Histioteuthidae'^" known to me are shown in Figures 173
and 182; their typical habitus is better shown in Figure 173. The character

of the family is already recognizable by the
presence of 7 rows of small suckers on the
tentacle clubs, the length of the two dorsal
arm pairs, with their at first relatively large
suckers, the form of the fins and of the
posterior end of the body, with its narrow
apex. These larvae are most closely
related to those of the Enoploteuthinae
(Figures 173 and 131 ), but the chromato-
phores on the mantle have a different distri-
bution. The third pair of arms is already
markedly shorter than the first and second
pairs, the fourth pair even shorter
(formula: 2,1; 3,4). (Cf. also Pfeffer,
Plate 20, Figures 2, 3, pp. 262—264).

Luminous organs appear on the ventral
side of the head in stages about twice as
large as the above. The characteristic
habitus of the family is also recognizable in
other characters (Figures 174, 175). The
fins and the growing posterior end of the
body are particularly characteristic.

Luminous organs appear at first on the
ventral side of the head in the Histioteu-
thidae (Figure 175). The "anlagen" are at
first of a simple type; they are oblong, white, opaque, subcutaneous growths
(epithelial invaginations?) which form tubercles on the epidermis after
preservation. On the posterior part of this growth, there is always a
chromatophore from which the pigment sheath later develops. Older stages
(Figure 177) show the condition shown in Figure 172. Windows are still
absent and light can only be emitted anteriorly. In 3 organs near the left
eye (see p. 349 ), however, the anterior part of the typical structures is already
retarded, and the large chromatophore covering the posterior part already
has a typical window. This confirms the view that the pigment sheath
develops from this chromatophore (p.348).

FIGURE 172. Luminous organs of the
young Calliteuthis reversa in
Figure 177. 40 x. a) longitudinal
section; b) upper view; c) one of the
heteromorphous luminous organs on the
ventral side of the head behind the left
eye.

Lk - luminous body; Jr - argentea;

Rf- shining sheath; Sp — mirror; Ch —
chromatophores; Hch^ — chromatophore
of sheath. In c), the window is visible in
the chromatophore of the sheath, while
the anterior part.e.g. the mirror (not
marked) is less developed. Li - surface
of lens.

Young stages of Histioteuthidae (Calliteuthis) are described by Tilesius (Krusenstern) Plate 38,
Figures 32-33; Ferrusac and d’Orbigny, 1835 (Squid, Plate 1, Figures 2-4; Loligopsis tilesii from
Japan); Hoyle, 1907 (Antarct. Figure).

328

351 VARIATION OF THE TYPE OF HISTIOTEUTHIDAE

I place the following four genera in the Histioteuthidae: Calliteuthis,

M e 1 e ag r o t e u t h i s, H i s t i o p s i s and Hi st i ot euth i s. These can be
distinguished as follows:

1. Ventral side of body with numerous, uniformly distributed luminous
organs of typical structure (Figure 170). Ventral arms with about

8 longitudinal rows of luminous organs at the base. Protective mar-
gins forming very weak membranes between the bases of the 3 upper

pairs of arms. Horny rings of suckers with teeth

Meleagroteuthis Pfeffer, 1900

2. Ventral side of body with differently developed, rudimentary and typical
luminous organs. Ventral arms with 3 rows of well developed luminous
organs at the base, with 2 longitudinal rows apically. The 3 upper arm
pairs with one row of typical, large organs along the outer (ventral)
edge and irregularly distributed smaller organs inside (dorsal to)
them

a) Buccal funnel consisting of 7 parts, protective membrane as in 1
Calliteuthis V err ill, 1880

b) Buccal funnel consisting of 7 parts; membrane as above, but more

strongly developed (Figure 181) Histiopsis Hoyle, 1889

c) Buccal funnel consisting of 6 parts; membrane well developed also
between ventral and later oventral arms . . Histioteuthis d'Orb., 1839

These 4 genera apparently form a series of transitions in the sense of the
introduction (p. 17). Division into subfamilies is therefore useless, as it
would only confuse the natural sequence.

329

352 Chapter 23

GENUS CALLITEUTHIS

Verrill, 1880

DIAGNOSIS

Large luminous organs forming only 3 rows on base of ventral arms.
Buccal funnel consisting of 7 parts. Protective margin only moderately
developed, also on the 3 dorsal arm pairs, and connected only at the base
of the arms into a rudimentary membrane.

In addition to C. r e v e r s a, the genus contains a number of closely related
species placed by Pfeffer (1912, pp. 279— 290) in the genus Stigmatoteu-
this; these resemble C.reversa in all important characters.

CALLITEUTHIS REVERSA Verrill, 1880
DIAGNOSIS

Suckers without teeth. Large suckers of tentacles with accessory
chitinous structures (?). Margin of eyelid with 18 luminous organs, the
organ situated in the middle of the upper margin markedly smaller than the
adjacent organs. Fins occupying less than half of mantle length.

LITERATURE

Contrary to what Pfeffer (1912)* states, the following are the original records of this species (cf. footnote
on p. 339):

1880 Verrill (U. S.), C alliteuthis reversa (p. 393; first reliable record).

1880 Verrill (Proc. N.H.), C alii ten this reversa (p. 262).

1881 Verrill (Conn. Ac.), Calliteuthis reversa (Plate 46, Fig. 1, p. 295).

1881 Verrill (Hawk), C a 1 1 i t . reversa (p. 112, Plate 7, Fig. 1). .

353 1881 Verrill (Rep. for 1879), Calliteuthis reversa (p. 327, [117] , Plate 22, Fig. 1).

Pfeffer (1912) considers this species as identical with "Loligo meneghinii" Verany (1851, Plate 35,
Figures c— e). I accepted this view at first (Naef, 1916, p. 16) although it is quite untenable. Verany’s
good figure shows a young Octopodoteuthis sicula. This is proved by the general habitus, the
different length of the median and lateral arms, the form of the fins (see above, p. 337, Figure 164) and the
rudimentary tentacle clubs. No other known juvenile Teuthoidea have biserial clubs in a so far advanced
stage (cf. Idiosepius). Figure c in Verany, which shows this structure, is apparently not very exact, but
the author would certainly not have failed to recognize the relationships in the young Calliteuthis
(Figure 176).

330

1899 Joubin (list), Calliteuthis reversa (p. 72).

1900 Joubin, C a 1 1 i t e u th i s reversa (p. 96),

1900 Joubin, Histioteuthis r ii p p e 1 1 i i (p. 98; echantillon A).

1900 Pfeffer, Calliteuthis reversa (p. 170) pars.

1903/4 Jatta (Lo Bianco), Histiopsis atlantica (Plate 8, Fig, 22, p. 172).

1904 Hoyle?, Calliteuthis reversa (p. 42).

1907 Massy, Histioteuthis bonelliana (p, 381) pars.

1909 Massy, Histioteuthis bonelliana (p. 29) pars.

1909 Russel, Calliteuthis reversa (p. 446).

1910 Chun, Histioteuthis juv. (p, 176, Plate 18, Fig. 6, 7, Plate 19, Fig. 1—4).

1910 Chun, C a lli t eu this reversa (p. 174, Plate 18, Fig .2-4, Plate 19, Fig. 5, Plate 20, Fig. 3, 4,6,
Fig.11,13,14).

1912 Pfeffer, Calliteuthis meneghinii (Plate 20, 21, 22, Fig. 13-19).

1916 Naef (Syst.), C a 1 li te u this meneghini (P.15).

1921 Naef (Syst.), C a 11 i t e u th i s reversa (p. 538).

STRUCTURE OF THE ADULT ANIMAL

I could not obtain large Calliteuthis near Naples and the following
description is based mainly on VerrilL Chun, and Pfeffer, after examination
of Chun's original specimens.

The general habitus is typical for the family, also in most details. A
striking difference from Stigmatoteuthis is the marked reduction of
the luminous organs on the whole surface of the body (see p. 351 ). The fins
are relatively short, occupying one-third to two-fifths of the dorsal mantle
length. The posterior apex is weak so that the fins project only slightly
beyond the mantle sac. Only the anterior transverse neck fold is stated to
be distinct; the only remnant of the third longitudinal fold is the elliptical,
knoblike olfactory tubercle.* The funnel and buccal skin show no special
characters. Arm formula: 3, 2; 1=4. Swimming and protective margins
are typical.

The arm suckers are widely separated proximally, very dense in the
distal part. They are largest on the third and second arms, smaller on the
first and smallest on the fourth arms. The suckers have mainly smooth
margins, only the terminal suckers have 8 — 10 crenelated teeth with narrow
intervals and with well developed adhesive rings. The area of the adhesive
rings on the proximal suckers is soft, curved outward and glandular. The
tentacles are characteristic for the family: the stalks are pressed in
between the third and fourth arms at the base, they are bandlike, with fine
longitudinal striation, flat, narrow inner surface and rounded outer surface
(Figure 168e). The inner side is turned ventrally in the distal part (f), the
outer side bears a margin. The hand part is markedly widened, the muscular
axis is apparently divided, and the gap covered with a membrane with
muscular cross bridges (Figure 168a: 5, 6, g). The narrow distal part bears
a typical swimming margin, the hand part protective margins which continue
on the stalk as edges anteriorly and posteriorly.

According to Pfeffer (l912, p. 255), the suckers of the hand part bear
"large, chitinous, accessory thickenings, especially in the dorsal part which

* But compare Figure 168a and b, also p. 344.

331

354 completely deform the suckers."'’' The distribution of these suckers is as
described above for the family (p.346). Hectocotylization has not been
observed in this species. It probably resembles the condition in C. hoy lei
(Chun, 1910, p. 167, Text Plate l) (=Stigmatoteuthis chuni Pfeffer,

1912, p. 286), i. e. it affects the two dorsal arms, especially their distal parts,
where the elongated sucker pads enclose a transversely ridged groove,
like a palissade, apparently for the transfer of the spermatophores (cf. the
Sepiolinae). The jaws and radula probably resemble those of Histio-
teuthis (q. V. ).

About the gladius of the same species, see Chun (1910, p. 157, Figure 23a, b);
it is markedly L o 1 i g o-like, with a completely flattened cone.

POSTEMBRYONIC DEVELOPMENT

The youngest known stage (Figure 173) is from the plankton in Naples.
It was probably 5 mm long in life; it has now a dorsal mantle length of
2 mm. The habitus of the family is already distinct, much better than in
specimens of similar size of H i s t i o t e u t h i s (Figure 182).

FIGURE 173. Youngest known larva
of Calliteuthis reversa from
the Naples plankton. 10 x, a) pre-
paration with spread fins, deformed,
b) reconstruction of normal relation-
ships. Note the form of fins, pos-
terior end of body, cone, m.antle,
head, arms, and ventral corner of
cone; eyelids widely opened,
showing eyeball and lens. Cf.
Figures 131, 167.

FIGURE 174. Young stage of Calliteu-
this reversa. 6x. This specimen was
collected, together with that in the next
figure, at a depth of 250 m near the "Amon-
tatura" on 23 April 1912. The arms are
slightly damaged, but the chromatophores
are well preserved and of characteristic
distribution. The luminous organs on the
ventral side of the head are still indistinct
and rudimentary; their arrangement is as
in the following specimen (Figure 175).

The dorsal side shows the wide, Lo 1 i g o-like gladius, covered only by skin.
The spoon-shaped cone with its median projecting corner is visible ventrally.

I cannot confirm this, but it is possible that Pfeffer's material was treated with chromic acid, which causes
a wartlike swelling of the horny rings. Formol is much better for general purposes.

332

The cone bears posteriorly the fleshy apex connected laterally with the
posterior margin of the fins. The fins are very large and closely resemble
those of the adult. They are transversely elliptical, and project posteriorly
beyond the apex, forming a median indentation. The muscular mass of the
fins is apparently concentrated in a median transverse area (dotted) and
the margin is delicate and transparent. This is similar in the young
Ctenopteryx (Figure 118) but more distinct.

The 3 typical corners of the mantle margin are already distinct, funnel
and head are typically larval.

355 The second arms are the longest, the first pair only slightly shorter.

The third pair is only slightly more than half as long as the second, and the
relationship between the fourth and third arms is similar; the fourth pair
is thus the shortest. The two upper pairs of arms bear larger suckers than
the third arms; the smallest suckers are present on the fourth pair. The
tentacles are about as long as the second pair of arms. Their greater part
is covered with suckers, which form 7 rows in the widest part. The buccal
funnel consists normally of 7 parts.

It is not certain whether the specimen illustrated by Pfeffer (1912,

Plate 20, Figures 2 and 3) belongs to the same stage and species. According
to the small fins (cf. Figure 182 ), it may belong to Histioteuthis, but
the fins could be shrunken. The structure of the buccal funnel would be
decisive, which consists of 6 parts in Histioteuthis.

(356)

FIGURE 175. Young stage of Calliteuthis re versa, caught at depth of 250 m
near the "Amontatura" on 23 April 1912. 6x. Note the fins, mantle, posterior apex
of body, funnel, funnel cartilage, olfactory tubercle, arm apparatus. The gladius is
visible dorsally through the skin. Also noteworthy is the arrangement of the
"anlagen" of the first luminous organs (Lo) on the ventral side of the head.

333

The two larvae in Figures 174 and 175 certainly belong to
Calliteuthis. Their dorsal mantle length is 4 and 4.2 mm, so that they
may be considered to belong to the same stage. The habitus of the family
is already recognizable in the form of the fins, posterior end of the body,
mantle and head. The gladius is well visible dorsally and is also later
(Figures 177, 180) not surrounded more completely by the mantle. The
fins are fused in the middle posteriorly, partly covering the flag. The
anterior part of the fins diverges. The part of the fins connected with the
tail-like end of the body is transparent, membranous and clearly delimited
(dotted line in Figure 175). Each fin is nearly circular in outline. The
funnel has become smaller, head and eyes much larger. The arm apparatus
shows the same relationships in principle, although the ventral arms are
markedly longer. The club bears 7 rows in the distal part. More important
is the appearance of the first luminous organs as whitish tubercles at the
base of the ventral arms and on the adjacent part of the head. Their
distribution does not change later (Figure 175).

The tentacle club shown in Figure 176 belongs to a stage resembling
the next stage.

(35Y)

FIGURE 176. Distal part of right tentacle of a young C a 1 1 i te u th i s re versa
(collected near Naples on 11 March 1905; collection of the Zoological Station).
Note the arrangement of the suckers on the distal, hand, carpal and stalk parts.
Knobs are present on the stalk part but not yet on the carpal and hand parts. The
proximal suckers of the two dorsal rows on the base of the hand part are rearranged
in a single row which continues from the enlarged first sucker of the third row and
forms the hand part of the "adhesion apparatus." Otherwise, the suckers of the hand
part form 7 regular rows, of which the median row contains larger proximal suckers.
16 X.

334

35 6 This stage is important for the understanding of the later differentiation,

although it gives no information on the development of the characteristic
pattern of 7 rows on the tentacle club. I assumed at first that this condition
developed from a pattern of 8 rows by the loss of one row, but this is not the
case, the 7 rows develop directly. This is recognizable from the condition
in the earlier stages and the gradual transition to the normal quadriserial
pattern on the apex of the club.

This stage, of which I have 2 well preserved specimens, shows almost
all the typical characters of the external morphology of Histioteuthidae
except those connected with sexual maturity. Figure 9 of Plate XIX shows
this specimen, a female, in natural colors in swimming position (drawn after
the live animal).

(357)

c

FIGUEIE 177. Young Calliteuthis re versa, female, collected on 1 March 1913
in the deep-sea plankton of the Bay of Naples. Note the typical characters shown in
the preceding figures, especially the arrangement of luminous organs (dotted), the
protective membrane connections at the arm bases, the funnel adductors and the ol-
factory tubercle.

The dorsal side of the live animal shows its systematic position by the
form and proportions of fins, mantle and head. The arms appear longer than

335

in preserved specimens, the whole animal slightly more slender. The
general coloration is bright orange red which is characteristic for young
Oegopsida. The chromatophores show all shades between ochre yellow,
yellowish brown, vermilion, carmine and brownish red. The dark brown
violet spots probably belong to the luminous organs (Figure 177), although
their position in the color plate is not shown exactly. (The arms of the
live animal were kept close together; the fins made quick rhythmical beats
and undulate movements. Another color drawing of the freshly fixed animal
shows that the ventral side is lighter, more yellowish brown; the chroma-

357 tophores of the luminous organs are brown-violet, the mirror surfaces
(Sp in Figure 170) still with a bright silvery sheen.)

Figure 177 shows the preserved animal. The chromatophores are not
shown in the drawing (only the size and position of the luminous organs are
marked). The luminous organs are shown better in Figure 3 of Plate III,
which is drawn from a male of similar size.

The fins have become longer and more semicircular. The posterior
indentation persists, and "earlobes" are visible in the anterior part of their
base. Mantle and shell are typical,as before. The head, however, is wider
because of the enlargement of the eyes. The olfactory tubercles are now

358 situated in their definitive position and form large warts in the posterior part
of the head, so that they project sharply in dorsal and ventral view, as in
older specimens. The eyelids have a uniformly rounded margin.

FIGURE 178. Outer view of right tentacle club
of the young Calliteuthis reversa in
Figure 177 (12 x). 1) the continuation of the
stalk (St) in the axis of the club (Ax); 2) the
widening of the club into a "hand" by the large
protective margins (Ds,Vs) into which pillars
of the marginal suckers project (Figure 44 on
p. 118), so that only the outer part of the margin
has the structure of a simple protective margin;
3) the formation of a typical swimming margin
(Ss) in the distal half of the outer side of the
club.

FIGURE 179. Inner view of the two clubs
of the same young Calliteuthis. 12 x.
Note the arrangement of the suckers in the
distal part of the club; the transition into
the hand part; the alternation of suckers
and knobs in the proximal part of the dor-
sal margin of the hand part and on the
carpal and stask part.

x and Xi - main knob and main sucker of
the adhesion apparatus; y.y^ — last knob
or sucker of hand part of adhesion apparatus;
z, Zi — first knob and sucker of carpal part.

336

The arms are of about equal length, the lateral pairs only slightly longer
than the dorsal and ventral arms. The contraction does not permit exact
measurements. All arms already have the typical square cross section
and there are small connecting protective membranes at the base. The
upper 3 pairs already bear delicate swimming margins which occupy
approximately the distal half on the third pair and less on the other two
pairs. They are slightly widened on the third pair and there are low, narrow
ridges of skin between the two outer edges of the arm.

The suckers have become markedly more numerous, the protective
margins more distinct, and the whole arm apparatus essentially resembles
that of the adult. However, the connection by protective membranes between
the basal parts of the protective margin is still absent, and the tentacle club
shows still some juvenile characters.

(359)

FIGURE 180. Young Calliteuthis
re versa from the Adriatic, natural size.
Figures 169 and 171 represent the same
animal. Note the general appearance,
the distribution of luminous organs, the
form of fins and posterior end, and the
development of the arms.

(360)

FIGURE 181. Histiopsis atlantica Hoyle. Reconstructed
after Hoyle (1886), natural size. Note the resemblance to
Figure 180, except for the large membrane betv,’een the three
upper arm pairs. The animal differs from Calliteuthis in
this well developed membrane, from Histioteuthis in the
restriction of this membrane to the 3 upper pairs of arms, and
in the buccal funnel with 7 parts (cf. Figure 184).

337

Figure 3 of Plate III shows a male of similar size, collected near Naples
in 1912. The only difference between the two specimens is the absence of
359 the "anlagen" of the nidamental glands (in the specimen shown in Figure 177
they are typically situated at the exit of the posterior pallial vein from the
renal sac). Note on Plate III the formation of the luminous organs, the
combined effect of which may be imagined from the drawing, also the small
funnel cartilage, which is still quite undifferentiated, oegopsidlike, as in the
Enoploteuthidae (Plate IV, Figure 2) and the arrangement of the organs in
the mantle cavity.

The large luminous organs already have their definitive position, though
their number will increase by growth of the smaller organs. The posterior
end of the mantle sac still bears only chromatophores and small luminous
tubercles in the adjacent areas. The number of small luminous organs
certainly increases later. The degree of differentiation is shown in
Figure 172.

I obtained an older specimen from the Adriatic.* Unfortunately, this
animal is not quite intact — in particular, the protective margins of the
arms are worn. Figure 180 shows this specimen, slightly stylized.

The habitus of this specimen resembles even more closely that of the
adult in the marked change of the proportions of fins, mantle, head and arms.
The fins are already relatively smaller than in the preceding stage, so that
they had probably reached maximal development; Pfeffer (l912, p. 228,

Plate 21, Figure l) notes that the fins decrease further in size later on.

3 60 An inner protective membrane is absent between the protective margins
of the upper arms; there is a median connection between the opposite inner
margins only at the base of the dorsal arms. The dorsal margin of the
second arm is connected with the bond of the second buccal point, but not
with the ventral margin of the first arm. An inner membranous connection
is also absent between the second and third arms.

The suckers of the tentacles (Figure 169) in the median row have become
larger; the suckers and knobs of the adhesion apparatus of the hand part
are uniserial, not forming a zigzag row. There are no other important
differences; the condition is typical. The "accessory chitinous formations"
on the large sucker rings (p. 353) develop only later.

I found a Calliteuthis of about twice the above size (Figure 168) in
the Museum of Natural History in Stuttgart. This specimen is part of
Kolliker's material from Messina (l854). It is distinctly juvenile, less than
Va of the linear porportions of the specimen described by Joubin (l900). The
general habitus of this specimen apparently closely resembles that of Joubin,
so that a 7 -fold enlargement of Figure 168 would give quite an exact picture
of the adult. This specimen on the whole resembles that shown in
Figure 180; the swimming margins have to be added according to Figure 185,
especially on the LV arms. They are absent in this specimen. On the other
hand, the D arms are distinctly longer than the ventral pair, probably as a
361 preliminary to hectocotylization, and the fins are further reduced. The well-
preserved tentacles show the characteristic features of the genus and
family perhaps better than the other specimens. There is an interesting
detail: the bandlike tentacle stalk (Figure 168d) bears a number of small

* This specimen was sent to me for determination by Dr. Leidenfrost, It was collected by the "Najade"
expeditions and belongs to the National Museum in Budapest.

338

I

ventral suckers at the base which can adhere only to the ventral arm and
are distant from the other part of the adhesion apparatus on the median
side of the stalk (l, 2).

Closely related to Calliteuthis is a form which was frequently
considered (also by Pfeffer, 1912) as a young Histioteuthis (but see
Naef, 1916, p. 18). It is a close relative of both genera which is inter-
mediate in a number of important characters but is more closely related to
Calliteuthis. This is the stage described by Hoyle (188 6) as Histiop-
sis atlantica. Figure 181 is an attempt to reconstruct this specimen
from Hoyle's data and my morphological criteria, when I considered it still
as a stage of the individual development of Histioteuthis. However,
this specimen will not be described here as it is only important for under-
standing the following forms. The reader is referred to the description
and figure of Hoyle.

339

362 Chapter 24

GENUS HISTIOTEUTHIS
d'Orbigny, 1839

HISTIOTEUTHIS BONELLIANA
(Ferussac, 1835) d'Orb., 1839

DIAGNOSIS

Large luminous organs typical in structure, regularly but sparsely
distributed on the ventral surface. Smaller rudimentary luminous organs
situated among the larger organs, which form only 3 rows on the base of the
ventral arms. Buccal funnel consisting of 6 parts from early youth
(Figure 184) due to fusion of the ventral points. Not only the 3 upper arm
pairs connected by a large inner membrane formed by fusion of the pro-
tective margins, but also by a single median suture originating ventrally
on the buccal funnel and connecting the ventral protective margins of the
third arm with the inner margins of the ventral arms. Suckers of arms
and tentacles with toothed rings.

LITERATURE

1835 Ferussac (Mag. Z.), Cranchia bonelliana (Plate 66).

1839 D'Orbigny, H i s ti o te u t h is bonelliana (p. 327, Cranchies Plate 2).

1840 Verany, Cranchia bonelliana (Fig. 20).

1844 Verany, Histioteuthis riippellii (Congresso Milano).

1846 Verany, Histioteuthis riippellii (Plate 3a, p.28) and H. bonellii (p. 29)").

1849 Gray, Histioteuthis riippellii (p. 45).

1849 Gray, Histioteuthis bonelliana (p. 45).

1851 Verany, Histioteuthis riippellii (p. 117, Plates 20, 21).

1855 (1845) D’Orbigny, Histioteuthis bonelliana (p. 380, Plate 25).

1879 Verrill, (Am. J.), Histioteuthis collinsii (p. 241).

1880/81 Verrill, Histioteuthis collinsii (pp. 195, 234, 300, 404, 434, Plate 22, Plate 27,

Figures 3— 5), Plate 37, Figure 5).

1889 Carus, Histioteuthis riippellii (p. 451).

1889 Weiss, Histioteuthis riippellii (p. 83, Plate 10, Figures 8— 12).

1896 Jatta, Histioteuthis riippellii (p. 115).

1899 Joubin, Histioteuthis riippellii (p. 73).

1900 Joubin, Histioteuthis riippellii (p. 98) parts (see C a Hite u this).

1900 Pfeffer, Histioteuthis benelliana (p. 170).

1906 Fischer and Joubin, Histioteuthis riippellii (p. 205).

" H. b o n el li i is stated to differ from C r a n ch i a bo n e 1 1 i a na (p.29) as follows; "regolarita delle braccia
e membrane." "cupole cerulee, color risplendente." The suckers are described as pale, whitish and smaller.

340

1906 Chun, Histioteu this ruppellii (p. 743).

1907 Massy, Histioteuthis bonelliana (p. 381).

1908 Pfeffer, Histioteuthis bonelliana (p. 77, Figure 86).

1909 Massy, Histioteuthis bonelliana (p. 29).

1910 Chun, Histioteuthis ruppellii (pp. 147— 169, 176; 168, Plate 21).
1912 Pfeffer, Histioteuthis bonelliana (p. 297, Plates 23— 25).

1916 Naef (Syst.), Histioteuthis bonelliana (pp. 15,18).

1921 Naef (Syst.), Histioteuthis bonelliana (p. 538).

363 STRUCTURE OF THE ADULT ANIMAL

I made only a cursory examination of two large specimens of this
species in the museums in Naples and Vienna. The Naples specimen is a
female with large nidamental glands, as in other Oegopsida (Figures 207 and
222). The Vienna specimen is particularly large, about 70 cm long. After
examination of these two specimens and the original specimen of Chun
(1910, Plate 21), I can only confirm the descriptions of Chun and Pfeffer
(1912, p. 292, Plates 23—25). This species is a plump, sluggish deepwater
animal of peculiar octopodlike habitus and magnificent appearance. Fresh
specimens probably have also a strong metallic sheen, which makes the
pearl-like luminous organs (Figure 185) even more dazzling (p. 348) and can
hardly be reproduced in a drawing. Compare also Figure 185, which rep-
resents an older young stage.

The hand part of the tentacle club shows some special characters. Not
only the suckers of the middle row but also the dorsally adjacent suckers
are markedly enlarged; the two following suckers are smaller and incom-
plete proximally. The 3 ventral rows are also proximally reduced to 2 rows
which contain supernumerary suckers. These adhesive suckers and knobs
show the relationship as shown in Figure 169. Teeth are absent, while they
number to 45 elsewhere. Adhesive rings are not distinct anywhere. The
distal part is quadriserial, but its very narrow end bears 2—4 rows.

According to Chun (1910, p. 167, Plate 1), hectocotylization affects the ends
of the 2 dorsal arms. The stalks of the suckers are lengthened into pinlike
structures with a more or less quadriserial arrangement and border a
groove, like a pallisade.

The gladius is distinctly Loli go-like (cf. d'Orbigny, 1845, Plate 25,
Figure 10; Tryon, 1879, Plate 71, Figure 277).

POSTEMBRYONIC DEVELOPMENT

Young stages of Histioteuthis have often been mentioned in the
literature. These forms are stated to resemble Calliteuthis and to
have a normal, buccal funnel with 7 parts. Most of these young forms
(Chun, 1910, Plate 18, Figures 6—7; Plate 19, Figures 1—4) undoubtedly
belong to C a 1 1 i t e ut hi s (q.v.). On the other hand, a larger form,
Histiopsis atlantica Hoyle (p. 360) is certainly a valid species and

* Named Loligopsis reinhardti

341

does not belong to H i s t i o t e u t h i s. The young H i s t i o t e u t h i s has a
buccal funnel with 6 parts at a mantle length of only a few millimeters,
long before the protective membrane develops (Figure 184; Plate III,

Figure 2). The young Histioteuthis differs also in other characters
from other related species (Figure 183).

The youngest stage examined (Figure 182) is a typical small larva of
Oegopsida the systematic position of which is proved by the resemblance

364 to the next stage. The habitus of this larva differs markedly from that of
the youngest Calliteuthis (Figure 173) and the typical character of the
family is even less recognizable.

The gladius is situated on the surface of the mantle, with a cone at its
posterior end. The fins are situated dorsally on the gladius, still separated

in the middle . Unlike in Calliteuthis, the
fins are at first very small and project
markedly beyond the posterior end; they form
a quadrant, attached at the corner (cf. also
Figure 183). There is a small apex between
the fins which is probably formed mostly by
the gladius and less by the fleshy part. As
in the next stage, the mantle is more cylindri-
cal than calyx- or cone-shaped and shows the
typical formation of the anterior margin. The
funnel apparatus and head show no special
characters. The funnel bond is still relatively
long and formed as in the Enoploteuthidae.

The eyes protrude slightly.

The arms are much shorter than in the
youngest stage of Calliteuthis, probably
due to preservation (the animal was kept in
formol). The arms show the normal propor-
tions (formula; 2, 1,3,4). The tentacle club
bears several row s of suckers, the oblique distal transverse rows consist
of 7 suckers. The buccal funnel is developed only as a low edge and consists
apparently of 6 parts, but this is not certain since the ventral supports and
points are not yet recognizable.

More important and better known is the next stage (Figure 183). This
young specimen does not show the habitus of the family as distinctly as the
young Calliteuthis (cf. Figure 173, p. 354), but the typical "anlagen" of
the luminous organs prove that it belongs to this species. The luminous
organs are already numerous (cf. Figure 175), but the animal is still re-
tarded in other respects. The development of this genus differs markedly
from that of Calliteuthis. The fins are still separated, but contiguous
in the posterior part of the base. They still resemble those of the preceding
stage in form. Strangely enough, there is no trace of the apex shown in
Figure 182 which persists in all juvenile stages of Calliteuthis. The

365 cone has probably become indistinct during growth, and the fleshy apex has
been taken up by the median connection between the fins. The mantle sac
is cylindrical-sac-shaped, with a typical margin. The funnel part also does
not show any special characters, although there is already a slight
widening of the funnel bonds. The eyes are already large and distinctly
lateral, but still without distinct asymmetry. The third pair of arms is

FIGURE 182. Youngest larva of H i s t i o -
teuthis bonelliana. lOx. From the
Naples plankton, September 1912.

342

already longer than the first pair and almost as long as the second; the
fourth arms are distinctly the shortest (2, 1= 3, 4). The tentacles show the
tjrpical condition of young Histioteuthidae. A loose row of suckers and
knobs occupies about the distal half of the stalk; the knobs are still in-
distinct, A second row with a few suckers begins at the base of the club,
slightly dorsal to the first row. The hand part is at first biserial and
passes rapidly into a pattern of 4 or more rows, without recognizable
regularity, until there are 7 distinct rows. In the distal part the number of
rows decreases again until there are only 4 rows near the apex. The
development of the club is comparable to that in Figure 176, except that a
swimming margin is still absent.

\\ b /f c A

FIGURE 183. Young stage of Histioteuthis bonel-
liana. ^VaX. The specimen was caught at a depth of
200 m in the Bay of Nisida on 20 February 1913. Note
the general outline, especially the form of fins, mantle
and head, arms and tentacles, distribution of chroma-
tophores (hatched) and luminous organs (dotted).

Of particular interest are the buccal funnel (Figure 184; Plate III,

Figure 2) and the adjacent mouth area. There are 6 distinct buccal points,
supports and bonds, which is caused by the fusion of the two ventral supports.
Otherwise, the mouth area is normal. The fusion cannot be attributed to
an approximation of the two small structures, but is connected with a trans-
formation of the whole ventral part of the mouth area, particularly the
366 premature approximation of the bases of the tentacles, the formation of the
tentacle pockets and the displacement of the fusing parts of the buccal

343

funnel dor sally, i. e. toward the mouth cone. These changes would have
in any case caused an approximation of the ventral supports, as shown by
a comparison of the normal formation of these parts (Figures 79 and 113;
see also p. 180). For a better understanding of the whole change, the early
stages of the buccal funnel have to be remembered, when the two ventral
points are still indistinct (Figure 88b) and the bonds (cf. Vol. II, Plate XXI)
which draw the corners apart and away from the mouth cone are not yet
present (Figure 113). This apparently does not take place in Histio-
t e u t h i s, and the fusion of the two ventral buccal pillars removes the
obstacle. The fusion takes place as with the dorsal pillars. This process
also prevents a phenomenon observed in other Teuthoidea (Figure 113), i. e.
that the buccal funnel is drawn down beyond the point (Figure 184) where
the ventral protective margins of the third arms can be united in the middle,
which takes place later. At any rate, the double bond (Ls) of the single
ventral buccal point is still visible. As in the Enoploteuthidae and in con-
trast to the Onychoteuthidae, this bond passes at first more on the outer
than on the inner side of the two ventral arms (cf. pp.241 and 303). Two
further pairs of projecting ridges extend from the base of the buccal pillar:
the first pair are the "ligaments" (Lb) of the tentacles, the second (not
marked) originates on the superficial adductors of the third arm. The
ventral protective margins of the third arms later reach the median line at
X, where the bonds of the ventral buccal pillar form the basis for the fusion
of the median protective margins of the ventral arms. This explains the
formation of the very characteristic and until now puzzling parts of the

FIGURE 184. Mouth area and buccal funnel of a young Histioteuthis
(Figure 183). 24x). Compare with Plate III and Figures 79 and 113:

1 — 4 — buccal pillars; v — connecting membranes; Bt^ — Bts — buccal
pockets; A1 — outer lip; II — inner lip; Hei-4 — bonds; Ch — chroma-
tophores; Sn - suckers; Sh - membranes of arms; As - outer margin of
ventral arm; Tr — funnel; Ls — ridges radiating from the ventral bonds;
Te — base of tentacle (cut ofO; Lb — ligament of tentacle; Cp — head;

X — point at the base of the ventral buccal pillar, at which the ventral
protective margins of the third arms and the ventral arms become united.

344

protective membrane in Histioteuthis, although the development will
only be understood after further young specimens have been obtained. The

367 structure of the similar apparatus in Pyroteuthis (Figure 12 8) should be
remembered in which there is also a median connection of the ventral
protective margins of the third pair of arms, and also a connection of the
margins between each two adjacent arm bases of the 3 upper pairs. In
Pyroteuthis, however, the buccal funnel has quite a different function:
the points provide a direct connection between the 3 upper arm pairs, so
that the funnel is included in the formation of the membrane. The ventral
margins of the third pair, however, are contiguous their whole length
under the perforated attachments of the buccal funnel. There is thus no
closer relationship.

Protective margins are not yet developed on the arms of our specimen,
but their "anlagen" are already present in some places as edges. The
extreme apex of the arms is bare; an area of about the length of 10 of the
small distal suckers shows no trace of "anlagen" of suckers (cf. Octopo-
d o t e u th i s. Figure 167a).

Further stages of Histioteuthis are not known to me'-' and I, therefore,
describe below only a young but fully developed (except for the genitalia)
animal from Messina.

The habitus is even more Octopus-like than in other Histioteuthidae,
because of the large membrane. As typical for the family, the gladius is
only partly surrounded by the muscular mantle. The fins occupy slightly
more than half of the dorsal mantle length and otherwise closely resemble
those of the young Calliteuthis (Figure 177). They form together a
transverse oval with typical anterior "earlobes" and a narrow, deep, median
posterior incision. Surprisingly, a distinct apex is absent here as well
(Figure 180), and it seems that the fins simply become fused in the middle
and project slightly beyond the mantle sac at this point. In fact, this struc-
ture is present anatomically, perhaps contracted, and only does not project
outside. The mantle sac is calyx- shaped, rounded posteriorly. The mantle
margin forms a narrow funnel indentation with blunt corners corresponding
to the very small funnel. The mantle margin forms a wide, rounded medio-
dorsal convexity, as in Calliteuthis (Figure 180).

The anterior transverse neck fold is distinct and reaches ventrally to
the border of the very shallow funnel pit without a sharp corner. Only the
wartlike olfactory tubercle remains of the third longitudinal fold; and the
second longitudinal fold is indicated dorsal to it. I cannot give exact data
on the asymmetry of the eyes, as they are damaged.

The arms are very strong, with typical edges, and with small outer
membranous connections which form shallow pockets at the base. The two
lateral pairs are longer than the ventral arms; the dorsal arms are the
shortest (formula: 3, 2, 4, 1). The 3 upper pairs have swimming margins.

368 These are stronger on the third arms, where they occupy the distal half,
forming at first a wide lobe which tapers toward the apex. On the first and
second pairs the margins are thin ridges of membrane occupying the distal
third and fifth of the arms, respectively. All arms are connected by a large
inner membrane which is formed by a modification of all protective margins

* The youngest specimen recently described by Grimpe and Hoffmann (1921) already shows all the character-
istics of the following stage, and has therefore misled the authors in their views on the early development
of H istioteuthis.

345

except the narrow, edgelike outer margins of the ventral arms. The mar-
gins are connected in the intervals between the bases of the 3 upper arm
pairs, and the muscular membrane extends over the greater part of the
arms, passing distally into the normal protective margins. The ventral

FIGURE 185. Young H i s ti o t e u th i s bonelliana from Messina , natural size,
size. The drawing is a reconstruction of a deformed specimen. The empty eyes
are probably incorrectly shown here to be symmetrical (cf. Figure 168). The clubs
are missing. Note the form of mantle, fins, small funnel, large head, the large
membrane, the swimming margins, the terminal organs of the arms, and the
distribution of the luminous organs.

margins of the third arm and those of the fourth arm (Figure 184) are
connected in the middle below the ventral buccal pillar (Figure 185a) and
form a common suture which projects toward the end of the arms. The
membrane thus connects about half of the ventral arms and about Vs of the

346

lateroventral arms. As in the Octopoda, the suckers of the 3 upper arm
pairs are directed inwardj those of the ventral arms are situated on the
outside^ toward the stalk of the tentacle, and their function is doubtful. All
369 suckers of the arms are very small, dentate, as in the Sepiolidae; the

largest sucker is barely 1 mm long. The whole membrane is apparently
a trap for small planktonic organisms, or a device to hold the prey caught
by the arms and tentacles until it is immobilized by poison or bite, as in
the Octopoda. The membrane can be closed on the prey like a sac. The
tentacle stalks are situated outside the membrane owing to the character-
istic connection between the third arms. The tentacles would only interfere
inside the membrane.

The protective margins are very low at the end. They are mere edges
at the apex and there are no suckers in the last 1—2 mm of the arm. The
last centimeter of the outer side of the 3 upper arm pairs is occupied by a
characteristic structure which is certainly a luminous organ.''' This is a
black, oblong formation which tapers apicallyi it is situated in the dorsal
skin and the narrow swimming margin passes above it. The young stages
described above show no trace of such an organ, at least on external exami-
nation. The buccal funnel, consisting of 6 parts, is situated inside, at the
base of the membrane, not affected by the surrounding membrane.

The tentacles have been lost. According to Pfeffer's description, they are
almost exactly as in Calliteuthis, but their suckers have a uniformly
dentate margin.

Opening of the mantle cavity shows mainly the same relationships as in
Calliteuthis (Plate III, Figure 3). The male gonoduct of H i s t i o t e u -
this, however, is developed only on the left side.

The luminous organs are in general as in Calliteuthis (Figures 180,
168) in their distribution, but there is one difference: several organs on
the dorsal side, mainly a pair on the head, are only large, dark, almost black
spots without a mirror or a distinct window. Several ventral organs behind
the left eye are similar. The lid margin of the right eye bears 17 organs,
one of them slightly separated.

The jaws are shown in Plate XVII, Figure 6, the radula in Plate XV,

Figure 1 .

The coloration of the live animal probably resembles that of Calli-
teuthis (Plate XIX): i. e. orange red, with a strong metallic sheen on
head, arms and mantle. The specimen preserved in alcohol is mainly wine
red to reddish brown, with a violet tone in the middle of the dorsal side.

The luminous organs are violet brown, with a yellowish, metallic mirror;
membrane and buccal funnel are dark violet brown, the mantle, head and
especially the arms have a silvery sheen with greenish, reddish, blue- violet
and yellowish iridescence.

* This was recently confirmed by Grimpe and Hoffmann (1921).

347

370 Chapter 25

FAMILY BRACHIOTEUTHIDAE

Pfeffer, 1908

GENUS BRACHIOTEUTHIS Verrill, 1881
DIAGNOSIS

Small, slender Oegopsida. Gladius consisting only of the rhachis in its
greater part. A narrow cone flag present in the region of the fins and
forming posteriorly a conical cone. Club with numerous proximal rows of
minute suckers and only 4 distal rows of slightly larger suckers. Area of
neck characteristically elongated in the young forms.

The genus contains only a few species of uncertain status which closely
resemble the species described below. The typical relationships will there-
fore not be described as they would be identical for the most part with the
description of the species.

Pfeffer (1912) distinguishes 2 subgenera. B r a c h i o t e u t h i s (s. str.)
differs in the longer fins, the presence of a funnel pit, the thicker head and
the dentition of the suckers from the other subgenus, Tracheloteuthis
Steenstr., with the single species:

BRACHIOTEUTHIS (TRACHELOTEUTHIS)

RUSE I (Steenstrup, 1882) Pfeffer, 1912

a. DIAGNOSIS

Fins less than half of the mantle length also in the adult. Funnel pit
absent. Head long and narrow. Arm suckers with crenelated distal margin,
the teeth separated by intervals as wide as the teeth. Large tentacle suckers
with rakelike dentition on the distal margin.

Only 3 specimens of this species have been collected in Naples, the young
animal described by Jatta (l896, Plate 7, Figure 4), and 2 adults resembling
the specimen in Figure 186 (one specimen in the collection of the Zoological
371 Station, the other in my collection). B. r i i s e i is apparently more common
in Messina. I have also a few specimens from the Adriatic which will be
mentioned elsewhere.

348

b. LITERATURE

1882 Steenstrup, Tracheloteuthis riisei (p. 294).

1882 Steenstrup, Tracheloteuthis behnii (p. 294).

1884 Rochebrune, Entomopsis velaini (p. 21, Plate 2).

1884 Rochebrune, Entomopsis clouei (p. 21, Plate 2).

1884 Pfeffer, Verrilliola gracilis (p. 22, Fig. 28).

1884 Pfeffer, Verrilliola nympha (p. 23, Fig. 29).

1886 Hoyle, Tracheloteuthis r i i se i (p. 164, Plate 29).

1886 Hoyle, Tracheloteuthis spec. (p. 166, Plate 31).

1889 Weiss, Tracheloteuthis behnii (p. 85, Plate 10).

1896 Lbnnberg, Tracheloteuthis riisei (p. 603).

1896 Jatta, Entomopsis velaini (p. 113, Plate 7,14).

1897 Fowler, Tracheloteuthis riisei (p. 525).

1898 Steenstrup, Tracheloteuthis riisei (p. Ill, Plate).

1898 Steenstrup, Tracheloteuthis behnii (p. Ill, Plate).

1900 Joubin, Entomopsis velaini (p. 91, Plate 11).

1900 Joubin, Entomopsis alicei (p. 92, Plate 11).

1900 Joubin, ? Tracheloteuthis s p e c. (p. 46, Plate 12).

1900 Pfeffer, Tracheloteuthis riisei (p. 175).

1904 Jatta, Entomopsis velaini (p. 201).

1905 Hoyle, Entomopsis velaini (p. 93, Plate 14).

1908 Pfeffer, Entomopsis velaini' (p. 79, Figs. 93— 95).

1908 Chun, Brachioteuthis riisei (p. 8, Fig. 1, p. 207, 214, 215).

1912 Pfeffer, Brachioteuthis (Tracheloteuthis) riisei (p. 355, Plate 26, Figs. 1-20, Plate 27,
Figs. 1-9).

1916 Naef (Syst.), Brachioteuthis riisei (p. 15).

1921 Naef (Syst.), Brachioteuthis riisei (p. 538).

1921 Grimpe (North Sea), Brachioteuthis riisei (p. 298).

c. STRUCTURE OF THE ADULT ANIMAL

I have no mature specimens, but they probably resemble the young
specimen in Figure 186. Markedly larger specimens have not been described.
At any rate, this species is a dwarf among the Oegopsida. My material
from Messina and the Adriatic is of similar size. The following description
is based on a well preserved specimen from Messina. This animal is
immature to judge from the small development of the ovaries and nidamental
glands. It would perhaps have grown to the size in the figure. The body
is membranous -muscular, not gelatinous; it is also very slender and
resembles the young forms of the related (l) Chiroteuthidae (Figure 190) but
is rarely found in other adult Oegopsida. Otherwise, the general habitus is
normal.

The gladius (Figure 106d) is very slender and still superficial, i. e. not
surrounded by the muscular mantle. The long, free, rhachis is therefore
visible dorsally as a dark line which widens gradually toward the anterior
margin of the mantle except for a short distance before the fins (Figure 186b).
The rhachis is visible again near the fins, widening suddenly posteriorly into
a lanceola between the separated fins, and then tapering toward the posterior
end of the body. The fin bases cover most of the cone flag dorsally. This
is visible again ventral of the base of the fins and curves visibly toward the
3 72 conical cone, which is stituated at the end; the projecting fleshy apex is
completely rudimentary.

349

FIGURE 186. Brachioteuthis riisei, female from Messina. 2x. Note the characteristic
proportions, the narrow head and mantle, the form of the cone which is visible through the skin,
the long funnel adductors, the typical length of the arms, and the arrangement of the suckers
on the tentacles, c) earliest larval stage, mentioned on p. 375.

The fins are of nearly typical form and both together are rhomboidal-
heart-shaped. They are connected only at the end and attached in their
greater part on the cone flag of the gladius, with which they are movably
articulated. Mantle and fins bear sparse chromatophores, the arrangement
of which is otherwise characteristic for juvenile forms.

The mantle margin shows the 3 normal corners and a narrow funnel
indentation. The funnel, though weakly developed, is quite typical but its
adductors are unusually long and distinct.

The long neck area of the younger stages is only indicated here, but
slightly smaller specimens still have a distinctly lengthened neck. On the
other hand, the head itself is very long. The eyes are of moderate size. As
in the older larvae, they are situated markedly anteriorly and only slightly
lateral, at any rate, they do not occupy the whole sides of the head. The

350

chromatophores of the head are very sparse and specifically arranged,

3 73 stressing the larval character. The eyelids are usually wide open and have
pointed sinus at the anterior margin. There is a luminous organ on the
ventral side of the eyeball, perhaps rudimentary, forming a raised stripe.
(Cf. Enoploteuthidae, Benthoteuthidae, Onychoteuthidae, Octopodoteuthidae,
Chiroteuthidae, Cranchiidae. ) Neck folds are not yet developed. This
depends not only on the condition of the specimen but also on the preser-
vation, since the first developed folds are only temporary and caused by the
contraction of certain skin muscles. According to Pfeffer, there is an
anterior transverse fold and a distinct second and third longitudinal fold;
the fourth fold is a tubercle and the third fold bears posteriorly the low,
rounded-triangular, lobe of the olfactory tubercle. Only the olfactory
tubercle is present in my specimens; it is situated far anteriorly on the
posterior part of the eyeball.

FIGURE 187. Situs of mantle cavity of Brachioteuthis riisei
(young female, 2 X). The funnel apparatus has the typical form. The
funnel retractors (2) are very large, bandlike. The larval visceral
complex, i.e. intestine, ink sac (12), liver (11), vena cava (3,13),
venous branches (4, 5), branchial hearts (9), posterior aorta (7),
posterior pallial veins (8), "anlagen" of the nidamental glands (6),
the long, narrow gills.

1 — funnel adductors; 10 — atrium of heart; 14 — olfactory tubercle;
15 — cone.

The arms show distinct differences in length. The two lateral pairs are
strong, of about equal length, although the third pair is shorter in younger
specimens. The first pair is very delicate and only about half as long. The
fourth pair is slightly longer and stronger, but also distinctly shorter than
the lateral arms. The 4 longest arms have blunt ends; they have swimming
margins their whole length which widen and become more distinct only
distally. A similar margin is also present on the distal part of the dorsal
arms. The outer margins of the ventral arms are weakly developed, usually
in the form of edges, which pass to the LV arms only at the base. Protective
margins are present everywhere, but only the ventral margins of the 3 dorsal
pairs are widened and have distinct supports. Elsewhere the supports are
only muscular thickenings and elevations of the edgelike margin. All arms
bear 2 rows of suckers with crenelated teeth on the distal margin of the
horny rings; the intervals between the teeth are about as wide as the teeth.

351

3 74 The tentacles are relatively strong, but their relative length and thickness
depend on the contraction. The stalk occupies less than half of the tentacle
and does not bear suckers. The first suckers, which are very small and
arranged in a loose zigzag row, are situated between the two edgelike
protective margins of the stalk part of the club. Further distally, the
suckers form 4, 8 and more rows where the inner surface of the club
widens into a hand part. This part bears about 16 rows of suckers, but the
pattern is not strictly regular or simple enough for a short formula. The
suckers become rapidly sparser further on; those of the marginal rows be-
come enlarged first, then those of the adjacent rows and the normal quadri-
serial condition appears toward the distal part of the club. As the marginal
rows of the distal part are apparently a continuation of the rows of the ab-
normal hand part, the numerous small suckers of the hand part give the
impression that they are pushed into an originally quadriserial pattern, but
this is obviously contrary to the development (cf. Pfeffer, 1912, pp. 359— 360,
Plate 27, Figures 1,2). In fact, all 4 primary longitudinal rows of the hand
part are divided temporarily into 2 and 4 secondary rows, and these are
collected again distally (p. 117).

The suckers of the hand part and especially those on the ventral margin of
the distal part are displaced markedly onto the wide, large protective
margins, the stalks of the suckers strengthening the margin. The distal
part bears also a typical swimming margin, which tapers rapidly into a
thin edge on the hand part. The suckers are bowl- shaped, usually with a
rakelike arrangement of teeth at the distal margin.

The buccal funnel consists of 7 parts. The mantle cavity (Figure 187)
shows a number of special characters connected with the form of the body.
The region behind the funnel is represented by the very large, bandlike
funnel retractors which seem to force the viscera into the posterior part of
the mantle sac. The posterior part has a normal, larval arrangement
(cf. Plates III — V; Figures 149, 157, etc.). The liver, however, is apparently
displaced posteriorly and down into the mantle cavity. The small ink sac
is typically embedded in the liver. Also typical in position are the "anlagen"
of the nidamental glands, the condition of which proves that the animal is
still very young.

The gills are very narrow and long, but otherwise typical.

An external sexual dimorphism is not evident in these immature speci-
ment; hectocotylization perhaps takes place later.

d. POSTEMBRYONIC DEVELOPMENT

Pfeffer (1912, p. 363, Plate 26, Figures 16, 17) described a form as the
youngest larva of this species, stressing its resemblance to Cranchia.
In fact, the specimen described is not a Brachioteuthis but a species
of Cranchiidae or a related form. My larva, which is slightly smaller
375 (Figure 186c), agrees better with the development of the species. It was
collected by the German Deep-Sea Expedition, and was labeled by Chun as
"undetermined larva"; I made a brief examination and drawing of it in
Leipzig. (The drawing is not suitable for publication and I have no other

352

early young stages of Brachioteuthis.) This is a rather typical early
larva of Oegopsida. Its LV and V arms are still stumplike. The tentacles
are the longest and bear suckers in their greater part. The two upper
arm pairs are still shorter than the width of the eye. The head is typical for
young larvae (slightly oblong). The long neck region is typical for the young
stages of Brachioteuthidae and the related Chiroteuthidae.

Further young stages were described by Chun (l910, Plate 29, Figures 9— 10;
Plate 30, Figure 2; Plate 31, Figure 6) and Pfeffer (1912, Plate 26,

Figures 12—15). Jatta (Plate 7, Figure 4; Plate 14, Figures 10 — 5).
illustrated "E n t o m o p s i s v e 1 a i n i i" which also belongs here. These
stages resemble the youngest stage in habitus; they have small, separated,
terminal fins, a short, cylindrical, sac -shaped mantle with only a few chroma -
tophores, a very long neck region and a typically larval head.

The club already bears more than 4 rows of suckers at a mantle length
of less than 1 cm; however, the multiserial part is at first terminal, because
the quadriserial end part develops later (cf. also Pfeffer, 1912, pp. 263—3 64).

353

376 Chapter 26

FAMILY CHIROTEUTHIDAE

Gray, 1849

a. DIAGNOSIS

Pelagic Oegopsida. Body more or less gelatinous. Funnel bonds small,
oval to ear-shaped, often complicated by tubercles and exceptionally fused
with the mantle bonds. Anterior part of gladius (before the fins) consisting
almost entirely of the rhachis; cone flag folded into a conical, very slender,
often tubular cone. Apex of cone projecting beyond the large fins into a tail-
like process of the mantle sac which may disappear during postembryonic
development. Third longitudinal neck fold reduced to the slender stalk of
the olfactory tubercle; other neck folds absent. Ventral buccal pillars with
median bonds. i

We place also Grimalditeuthis Joubin in this family, in contrast to
Pfeffer (1912, p. 540). The genera which agree with the above diagnosis
certainly form a natural group, as Chun (l910) proved. A close relative of
this group is the peculiar form "C h i r o t e u t h i s" portieri Joubin, but
its characters are so markedly different that I made it the representative
of a separate family (p. 237). Also after the exclusion of this type, with its
extraordinarily long arms and 6 rows of suckers on the arms (cf. Naef,

1922, Foss. Tintenf., p. 299), the family shows an unusual plasticity of
structure and development which has to be considered in view of its position
between the Brachioteuthidae and Cranchiidae.

b. TYPICAL STRUCTURE OF THE ADULT ANIMAL

The typical appearance of a representative of this family (without
tentacles) is shown in the drawing of "Loligopsis," i. e. Grimaldi-
euthis b o n p 1 a n d i, in Verany (1839) (cf. Try on, 1879, Plate 70,

Figure 272). The habitus of the family can also be derived from Figure 186,
if the posterior end and cone are thought to be lengthened and pointed and
the head widened by the enlargement of the eyes; the mantle sac is thought
to be widened and calyxlike anteriorly, and the tentacles lengthened like a
whip. Juvenile characters expressed in the scarcity of chromatophores
377 and the swollen body, the tendency to an increase of sucker rows on the

tentacle clubs and the elongation of the neck region, resemble the Brachio-
teuthidae. These characters form a type which does not differ greatly from

5110/2

354

the Cranchiidae and may be considered as a preceding stage of the Cran-
chiidae. However, the usually thick head does not fit the pattern of Cran-
chiidae, which probably had a longer funnel bond.

Already young Chiroteuthidae can be recognized by their short, rounded
funnel bonds, the primary form of which is still recognizable also after
subsequent fusion with the mantle (Grimalditeuthis). The cone is very
long, as in the Cranchiidae, and contains distinct remnants of the phragmo-
cone (p. 243). As in G o n a t u s, there are irregularly distributed watch-
glass-shaped septa, without a trace of a siphuncle (cf. Chun, 1910, Plate 41,
Figure 13). The elongated cone projects behind the fins into a tail-like
appendage,* as in the typical Cranchiidae. Also similar are the anterior
part of the gladius, which consists of a long free rhachis without remnants
of a flag (cf. Figure 106d), and the olfactory organ with a long stalk, the
formation of which is connected with the conditions in the Onychoteuthidae
(p. 302). The gelatinous, swollen skin and musculature may be considered
as distinctive characters of the family. The closer relationships are dis-
cussed in the following chapter and on p. 236. The more distant relation-
ships are shown by the structure of the buccal funnel: the ventral pillars
are attached to the median side of the ventral arms, as in the Onychoteuthidae,
Octopodoteuthidae, Brachioteuthidae and Cranchiidae (the same probably
applies also to the Joubiniteuthidae).

JUVENILE FORMS

The postembryonic stages of the Chiroteuthidae undergo early a very
peculiar development beginning with typical conditions, which leads to the
formation of the tail appendage, that contains the cone, partly widened fin
margins, and the large, oval form of the fins; there is also a marked
lengthening of the mantle, neck, head and tentacles ( Figure 191, 193). These
"D oratopsis " - like larvae resemble at first a larva of Brachioteu-
this (p. 372) and then a larva of Cranchiidae, but later differ from both,
assuming a very characteristic form (cf. Chun, 1910, Plate 46; Pfeffer,

1912, Plate 4 6). These developments indicate a longer planktonic life of
the larva, the typical characters of which are connected with those of the
later stages.

3 78 c. VARIATION OF THE TYPE OF CHIROTEUTHIDAE

The Chiroteuthidae are divided into 3 subfamilies (Chun, 1910, p. 219) with
the following characteristics:

* An analogous structure is found in the Pyroteuthinae (p. 266). This formation differs from the fleshy apex
of typical Oegopsida, which is suppressed here by the cone. It should also not be confused with an
appendage like that in Alloteuthis (p. 222), in which the rudimentary cone is restricted to the extreme
apex (p. 209). This is a distinctive formation which begins in the Brachioteuthidae (p. 272) and has
developed further in die Cranchiidae (Figure 193).

355

a) Funnel bonds well developed also in the adult. Ventral arms parti-
cularly strongly developed, longer than the others.

O') Fin circular to leaf-shaped, barely longer than the part of the mantle
sac anterior to it. A long, tail-like process which may have auxiliary
fins projecting beyond the fins, at least in the young stages. Luminous
organs: 1. on ventral side of eyeball; 2. embedded on each side of ink
sac; 3. on lateral margins of ventral arms; 4. on outer side of
tentacles, and on apex of club in the adult. Tentacle club with 4 rows
of suckers, those of the distal part with 7 longer teeth (median tooth the

longest) Subfamily 1. Chiroteuthinae

(3) Fin circular, leaf-shaped, elliptical to rhomboidal, usually longer than
part of mantle anterior to it. Tail-like posterior process projecting
beyond fins only in the unknown larvae, if at all. Luminous organs in
the above-mentioned positions absent. Tentacle club with more than

4 longitudinal rows of very small suckers

Subfamily 2. Mastigoteuthinae

b) Funnel bonds absent in the adult, because of fusion of the funnel with the
mantle in their position. Ventral arms delicate, shorter than the other
arms. Tentacles absent in the adult . . Subfamily 3. Grimalditeuthinae

d. SUBFAMILY CHIROTEUTHINAE Chun, 1910
DIAGNOSIS

Funnel cartilage well developed, oval to ear- shaped. Ventral arms
distinctly longer and stronger than the others. Fins forming a circular
to heart-shaped figure, barely longer than the part of the mantle anterior to
them. Body, at least in the young stages, with a tail-like appendage which
projects beyond the fins and often bears auxiliary fins. Luminous organs:

1. on ventral side of eyeball; 2. on ink sac; 3. on inner side of lateral
margins of ventral arms; 4. on outer side of tentacles, also on apex of club
in the adult. Tentacle club (but see p. 389.') with 4 rows of suckers; distal
suckers with 7 long teeth, median tooth the longest. Tentacle club with
knoblike terminal (luminous) organ after the normal distal part of the young
stages has been lost.

This subfamily contains the genera Chiroteuthis d'Orb. and Chiro-
thauma Chun. The other described forms, C h i r i d i o t e u th i s Pfeffer,
Planctoteuthis Pfeffer, D o r a t o p s i s Rochebrune, L e p t o t e u t h i s
Verrill are typical juvenile forms of the above or of other related but
unknown genera (cf. also C h i r o t e u t h o i d e s Berry, 1920, i. e. Figure 193f).

3 79 The typical appearance of Chiroteuthinae is shown more or less exactly
in Figure 190c, but the posterior part of the fins has to be thought slightly
leaflike and pointed. These are plankto-nektonic, bathypelagic, very peculiar
animals with gelatinous tissue. The typical tail-like process of the Chiro-
teuthidae projects beyond the apex of the fins (p. 377). The fin is wide,
heart-shaped to circular. The head is columnlike, thickened by the pro-
jecting eyes.

356

Neck folds are absent; the olfactory tubercle has a long stalk. The funnel
bond is oval to ear-shaped. Two tubercles project into the funnel pit, one
posteriorly, the other from the inside. The mantle bond bears a corres-
ponding noselike tubercle which fits into the pit of the funnel cartilage. Lu-
minous organs are situated on the ventral side of the eyeball. The eyelid
has only an indistinct sinus.

The ventral arms are extremely large. They are the longest and
thickest and have wide lateral margins with luminous tubercles along the
outer row of suckers, alternating with them. Formula; 4; 3, 2, 1.

The tentacle stalks are extremely long and bear a longitudinal row of
"glandular knobs" on the outer side which are certainly luminous organs.

The club bears 4 rows of suckers (but cf. p. 3891), the stalk a simple row at
the base, then a zigzag row and finally 2 rows. A carpal part is not
differentiated.

The situs of the mantle shows weak, bandlike funnel retractors and a wide
ink sac with a lens -shaped luminous organ on each side. The renal papillae
are situated very close to the base of the gills (Plate IV, Figure 3).

The typical development of the subfamily was described mainly by Ficalbi
(Figure 1 90 ), who proved that "Do rat op sis v e r m i c u 1 a r i s" is the
juvenile form of Chiroteuthis veranyi. This statement can be
generalized in the sense that the adult Chiroteuthinae have to pass through
a typical Doratopsis stage (Figures 191 and 192 show the Mediterranean
form). The Doratopsis stage has the following characteristics. 1. A
very long, thin, tail-like posterior end which develops at an early stage,
usually with lateral fin margins, and projects far beyond the fins; the margins
of the tail are often widened into auxiliary fins which may resemble the main
fins. 2. There is a long, thick neck region consisting of metameric
chambers filled with fluid between head and funnel apparatus.'’' 3. The arms
are short, delicate, based on a long pillar; the ventral arms become longer
than the others early on.

These characters, together with the extremely delicate, membranous -
gelatinous structure of the fantastically elongated body and the complete
transparency (only a few organs, liver, ink sac and eyes are opaque in life),
create a very peculiar planktonic larva. Later, at a length of about 20 cm,

380 begins a kind of metamorphosis in which the tail is partly or completely
lost (Figure 190), the neck region becomes shorter and the arm pillar dis-
appears as an external character, while the head and especially the eyes
grow enormously, the arms become longer and thicker, and the end part of
the tentacle club is apparently lost (p. 390).

The mantle cavity also does not show at first the more or less normal
conditions which develop later (Plate 4, Figure 3). My material did not
permit a detailed examination of these parts. The typical abdominal
complex of the young Doratopsis is apparently concentrated in the
anterior quarter of the mantle sac; the other part is occupied mainly by a
vesicular tissue which is later reduced. The vessels of mantle and fins
pass superficially over this modified posterior part of the body.

* A medioventral and a mediodorsal cord pass in this region beneath the skin, without penetrating into the
vesicular interior. The medioventral cord certainly contains the cephalic veins, the mediodorsal, the
cephalic aorta and esophagus.

357

Of particular interest is the youngest larva of Chiroteuthidae illustrated
by Chun, 1910, Plate 46, Figures 8 — 10 (mantle length 4 mm), mainly because
its entire structure differs less than any other known formi. from the typical
condition of the youngest Oegopsida. The tail, projecting beyond the fins, is
still very small, and the fins are therefore almost terminal. They are also
much wider than long, which is typical for the larvae (Figure 88). Mantle and
head have the typical early larval form, and the third pair of arms are still
stumps. Nevertheless, the character of the subfamily is distinct in the
following characters. 1. The tail is already present and even bears indi-
cations of auxiliary fins. 2. The neck region is lengthened, but it is not
as thick and long as in the older Doratopsis stages but thin and short,
as in the youngest Brachioteuthis (Figure 186c).* The arm pillar is
already rather long and slenderer than in the older larvae. 4. The ventral
arms are already fully developed and almost as long as the dorsal pairs,
not shorter as in the other early larvae of Oegopsida.

Another very juvenile larva (l. c. Plate 46, Figures 6 and 7) has a tail
which is still slightly lengthened, with a small auxiliary fin; the fins have
grown anteriorly, but each fin is about as wide as it is long. The neck
region resembles that shown in Figure 191, while the eyes protrude slightly
more laterally. This larva may be considered as a younger stage of
"Doratopsis vermicularis" or a related form. Similar or related
stages were recently illustrated by Issel (l920. Figures 9—12) which
certainly belong to the species described below.

There is a striking external resemblance between this form and the youngest stages of Brachioteuthidae
(cf. also Chun, 1910, Plate 30, Figure 1). This and other common characters in a similar direction suggest
a closer relationship (pp. 236,376,377).

358

381 Chapter 27

GENUS CHIROTEUTHIS
d'Orbigny, 1839

DIAGNOSIS

Tail of young forms long, rodlike, thin, often with leaf- shaped auxiliary
fins, which are later lost, so that the main fins are terminal. Luminous
organs on ventral side of eyeball not forming 3 longitudinal rows of distinct,
lens -shaped structures but partly fused into diffuse longitudinal stripes.
Suckers of tentacle club with thin stalks on pillarlike, terminally thickened
and fluted supports. Transverse supports of protective margins divided into
2—5 parallel branches in proximal part of club, simple, contiguous at the
base, tapering and diverging toward the margin in the distal part.

In addition to Chiroteuthis veranyi, this genus contains C. 1 a c e r -
tosa Verrill and probably other, closely related species with various
Doratopsi s-like larvae (p. 379) (cf. Pfeffer, 1912, pp. 541—608).

CHIROTEUTHIS VERANYI (Ferussac, 1835 )
d'Orb., 1839

DIAGNOSIS

Proximal part of tentacle club with dense "subparallel" supports of
protective margins separated at most by thin membranes at the free margin.
Suckers of arms toothed at the distal margin, smooth at the proximal
margin. Club of adults with about 100 suckers.

LITERATURE
a.. Juvenile form

1845 Riippell, Loligopsis ve rm ic ul a ri s.

1849 Gray, Loligopsis vermicularis (p. 40).

1851 Verany, Loligopsis vermicularis (p. 123, Plate 40).
1884 Rochebrune, Doratopsis vermicularis (p. 18).

1884 Rochebrune, Doratopsis riippellii (p. 19).

1884 Pferrer, Hyaloteuthis vermicularis (p. 28, Fig. 30).
1886 Hoyle, Doratopsis vermicularis (p. 43).

359

1889 Carus, Doratopsis vermicularis (p. 451).

1889 Weiss, Doratopsis vermicularis (p. 80, Plate 9).

1896 Jatta, Doratopsis vermicularis (p. 108, Plates 7, 14, Fig. on p. 22).

1899 Ficalbi, Doratopsis ve r m i c ul a r is (p. 83). (Recognizes that it belongs to Chi roteu this
veranyi I).

1899 Ficalbi, Doratopsis vermicularis (pp. 93“118, Plate).

1899 Joubin, Doratopsis vermicularis. [Page number garbled in German original.]

3 8 2 1900 Joubin, Doratopsis vermicularis (p. 95).

1900 Pfeffer, Doratopsis vermicularis (p. 186).

1902 Ficalbi, Doratopsis vermicularis (p. 37).

1904 Jatta, Doratopsis vermicularis (pp. 193, 203).

1906 Hoyle, Doratopsis vermicularis (p. 161, Fig. 3).

1909 Massy, Doratopsis vermicularis (p. 33).

1910 Chun, Doratopsis vermicularis (pp. 285, 288, 293, Plate 47).

1912 Pfeffer, Doratopsis vermicularis (pp. 555-569, Plate 46).

1916 Naef, Chiroteuthis veranyi (p. 18).

1921 Issel, Chiroteuthis veranyi (Figs. 9—12).

b. Adult form

1835 Ferussac, Loligopsis veranyi (Mag. Zool. Plate 65).

1835 Ferussac, Loligopsis veranyi (Reg ne animal, Plate 6).

1839 D’Orbigny, Chiroteuthis veranyi (p. 325, Loligopsis, Plate 2).

1849 Gray, Chiroteuthis veranyi (p. 43).

1851 Verany, Loligopsis veranyi (p. 120, Plates 38,39).

1869 Targioni-Tozzetti, Chiroteuthis veranyi (p. 49).

1885 Hoyle, Chiroteuthis veranyi (p. 330).

1889 Carus, Chiroteuthis veranyi (p. 450).

1889 Weiss, Chiroteuthis veranyi (p. 77).

1893 Joubin, Chiroteuthis veranyi (pp.l— 13, Figs. 1—12).

1894 Joubin, Chiroteuthis veranyi (p. 63).

1899 Joubin, Chiroteuthis veranyi (p. 72).

1899 Ficalbi, Chiroteuthis veranyi (p. 93, Plate).

1900 Pfeffer, Chiroteuthis veranyi (p. 185).

1902 Ficalbi, Chiroteuthis veranyi (p. 37).

1903 Chun, Chiroteuthis veranyi (p. 67).

1904 Jatta, Chiroteuthis veranyi (p. 193).

1910 Chun, Chiroteuthis veranyi (pp. 240-281, Plates 40, 42, 44).

1912 Pfeffer, Chiroteuthis veranyi (pp. 594— 606, Plate 44,45).

1916 Naef (Syst.), Chiroteuthis veranyi (p. 16).

1921 Naef (Syst.), Chiroteuthis veranyi (p. 538).

STRUCTURE OF THE ADULT ANIMAL

This species resembles the Histioteuthidae (cf. Figure 180) in the
enormously developed arms and head, and in the relatively small, calyx-
shaped posterior part of the body with the round fins projecting beyond the
mantle. This similarity is probably connected with a common mode of life
but does not prove any closer morphological relationship.

The fins are apparently very variable, if all the forms considered as this
species belong to it. It is often more or less circular (Plate IV, Figure 3);
Pfeffer (1912, pp. 594—595) considers the relatively large width of the fins
as a species character. In the large specimen from Naples, however, the
fin is distinctly longer than wide, markedly pointed posteriorly and its

360

(383)

361

i

FIGURE 188. Chiroteuthis veranyi, collected in Naples. Young female, half natural size. Drawn by Merculiano after a freshly preserved animal
(September 1899?). (Some parts of this drawing are shown in Figure 189.) Note the form of mantle and fins, the enormous head and ventral arms, the
wide outer margins of the ventral arms, and the arrangement of glandular knobs on the tentacle stalks (the clubs are absent).

whole form leaf- to heart-shaped (Figures 188, 189). The differences
between Ch. veranyi and Ch. lacertosa Verrill thus become even
more doubtful. The base of the fin always forms small but distinct "ear-
lobes" anteriorly. As in the typical Cranchiidae, and therefore also in the
adult Chiroteuthidae, the process of the mantle sac supported by the cone
occupies more than four -fifths of the fins. Less than the anterior fifth of
the fins is actually attached to the mantle sac, i. e. before the opening of
the cone. This is another resemblance to the Histioteuthidae, although the
topographically corresponding "apex" between the fins has a different mor-
phological value: it is situated from the beginning on the cone and is not
filled by it.

The mantle sac is calyx -shaped and widens uniformly anteriorly; its
anterior part narrows slightly to a cylindrical to bell- shaped form. The
funnel indentation and the ventral corners are indistinct but the dorsomedian
corner is distinct.

(384)

FIGURE 189. Parts of C h i ro t e u t h i s veranyi shown in Figure 188, drawn to scale
after old alcohol preparations (slightly diagrammatic):

a — ventral view of right third arm; b — lateral view of tentacle stalk with glandular
knobs; c — part of middle of ventral arm, viewed from above (from the inside),
showing suckers and luminous organs; d — posterior end of mantle and fins, ventral;
e — left funnel bond.

383 Only the olfactory tubercle is left of the neck folds. It is situated on a
long stalk ventrally on each side between eye and funnel. The olfactory
organ is itself probably the small, flask-shaped swelling at the end of the
process (Plate IV, Figure 3). There is no distinct funnel pit. The funnel is
strongly and almost regularly recurved (always?). The funnel is connected
with the head on each side by two adductors which are inserted together on
the head. The funnel bond is small, ear-shaped. A large tubercle ("tragus")
always projects into its pit from the median side, a smaller tubercle
("antitragus") (Plate IV, Figure 3; Figure 189) sometimes from the
posterior side. The neck is oblong- rectangular, with rounded corners.

The eyes are very large, with a rounded lid margin and a weakly
indicated sinus. They are larger in volume, than the whole visceral complex.
The ventral surface of the eyeball has luminous organs in diffuse distri-
bution. On each eye there are 2 brownish yellow, slightly raised, vaguely
delimited longitudinal stripes (corresponding to the outer rows of isolated

384 organs in C h i r o t h a u m a), and a rounded, isolated organ before and behind
the stripe. (My material was not suitable for a more detailed examination.)
The lens is correspondingly large and projects markedly from the head in
well preserved specimens (Plate IV).

362

The arm apparatus exceeds body and head together in mass. The arms
have the typical proportions of length of the subfamily (formula: 4; 3, 2, l).
All arms are connected at the base by small outer membranes which form
pockets (cf. p. 345; Histiotheuthidae). According to Chun, a swimming margin
is indicated on the 4 dorsal arms. At any rate, such a margin is present along
the entire third arm, it widens in the proximal half and ends proximally on
the ventral outer edge of the arm, continuing into the membrane of the
fourth arm. The fourth arms have large lateral margins which extend to the
apical part, so that the arm appears even more massive. These margins
bear round, flat tubercles with a strongly pigmented marginal zone of brown-
violet chromatophores, apparently luminous organs, situated along and alter-
nating with the outer row of suckers. Protective margins are distinct
everywhere, but rather delicate. Their supports are almost completely
fused with the conical pillars of the suckers so that they form thin mem-
branes between the pillars (Figure 189a). The suckers are usually biserial;
on the 6 upper arms they form oblique pairs and are loose proximally but
rather dense in the distal part. The suckers of the ventral arms are
slightly smaller and form a loose zigzag row which becomes distally a more
or less single row in the proximal half (according to Pfeffer, 1912, the
ventral arms bear 42 suckers and about 180 on the other arms). The
suckers show the form typical for the Oegopsida (further details in Pfeffer,
1912, pp. 598—599'). The extended tentacle stalks are about 12 times as long
as the mantle sac; they are thin, almost stringlike, and slightly flattened
dorsoventrally. The stalk is occupied by "glandular knobs" its whole length
on the outer side (Figures 188 and 189). These do not form a single row
as they do at first according to Ficalbi (Figure 190), but a more or less
385 regular zigzag pattern, at least in the proximal part. The knobs slightly
resemble suckers in form, but have nothing in common with the stalked
suckers of the Doratopsis stage, which are situated on the inner side
and have been lost. The knobs are rounded, flattened bodies situated on
slight swellings and have a slightly concave outer surface. Their margin
is darkly pigmented, the concavity light. The knobs are probably luminous
organs which emit light from the concave surface.

The clubs are missing in my specimen. According to data and illustra-
tions (especially by Pfeffer, 1900), the club is slender and lanceolate
(Figure 190ei) so that the distal part of the tentacle is widened mainly by
the protective margins. The club is more than half as long as the mantle
sac. The greater part of the inside of the club is occupied by 4 rows of
suckers which decrease slightly in size proximally and slightly more in
the distal part but are otherwise rather uniform. The suckers become
biserial at the base. The base of the club is without suckers. There is a
"terminal knob" at the apex (cf. p. 390) probably a modified "glandular knob"
(Chun, 1910, p. 248) with a short apex (Figure 190ei). Like the other
glandular knobs, it opens toward the outer side. A number of rudimentary
glandular knobs are situated on the club in the outer median line
(Figure 190di).

The protective margins of the proximal part of the club differ markedly
from those of the distal part (Figure 190ei), and both parts are separated
by a slight constriction. The number of transverse supports in the distal
part is equal to that of the marginal suckers. These supports are so wide

363

at the base that they are in contact with each other, and they taper gradually
toward the margin. The supports of the proximal part are markedly more
dense; they are wide to their end and leave little or nothing of the margin
itself. The supports are not simple but are parallel branches of supports
which are fused proximally. The protective margins reach the terminal
knob in the distal part of the club (Figure 190ei). On the absence of a
swimming margin and of the whole distal part of the club see p. 390.

The buccal funnel consists of 7 parts. Its attachments end separately
from the proximal ends of the protective margins. There are 6 buccal
pockets.

The mantle situs shows a number of special characters (Plate IV, Figure 3)
caused by the form of the mantle and the general proportions of the body.

The funnel retractors are very weakly developed, bandlike; the ink sac is
very wide and forms lateral lobes which bear lens -shaped luminous organs
homologous to those of Onychoteuthis and Octopodoteuthis
(pp. 320 and 341). The gills are short, compact; the renal papillae are
closely situated to the base of the gills. The nidamental glands are situated
in the typical position. Corresponding to the funnel bonds is the mantle
bond, a noselike process which fits into the pit of the funnel cartilage and is
surrounded by a shallow, triangular depression which reaches close to the
margin of the mantle.

386 Chiroteuthis has thus 4 types of luminous organs: 1. on the ink sac;

2. the "glandular knobs" on the tentacles; 3. lens -shaped organs on the
ventral arms; 4. organs on the eyeball.

Figure 7 of Plate XVII shows the jaws; the radula is shown in Figure 4
on Plate XV.

Sexual dimorphism has not been observed, except for the genital organs
and the nidamental glands.

On the coloration of the fresh animal see Verany, 1851, p. 122. The
whitish, translucent body is densely covered with chromatophores which
probably produce a dark brown-violet tone in life (cf. also Pfeffer, 1912,
p. 604). Blue, orange and carmine tones are also present, with a strong
metallic sheen.

POSTEMBRYONIC DEVELOPMENT

The development was described by Ficalbi (1899) (Figure 190), who thus
contributed considerably to the morphological knowledge of the whole family.
Pfeffer's reservations (1912, p. 551), on the other hand, are not correct (see
below). Ficalbi found that the so-called "Doratopsis v e r m i c u la r i s"
(Riippell) is the juvenile form of Chiroteuthis verany i. The transi-
tional stage described by Ficalbi (Figure 190c) makes this very likely. How-
ever, the description and drawing of the interesting material of Ficalbi are
not exact enough to be accepted as proof, and Pfeffer (l900) and Hoyle (1909)
therefore rejected it. We shall show on the basis of my observations that
these objections are not justified.

364

(387)

FIGURE 190. Development of Chiroteuthis veranyi according to Ficalbi (1899): his figures are
reproduced here half their size. Figures e^ and ez diagrammatic, after Pfeffer (1912, Plates 44 and 45).

a, b, c,d - 4 stages of Chiroteuthis veranyi from Messina; b^, c^, dj - left tentacle clubs of spe-
cimens b,c,d “ dorsal; b2,C2,d2 ~ tentacle knobs of these specimens with their stalks; e^ — tentacle
club of an adult, diagrammatic, after Pfeffer (Plate 45); 62 — terminal part of club of an adult, after
Plate 44 . Magnification: a-d - 0.5X; bj — 2.75x; cj - 1.5X; dj - 1.5X; e^ - Ix; eg - 2x.

Very young Doratopsis larvae of the Mediterranean species have been
recently described by Issel (cf. p. 388). The smallest of these forms is a
third smaller than the specimen in Figure 191, and the fins, tail and arms
are also smaller. The fins are still wider than long and have a narrow base,
but the habitus of this and related stages closely resembles that in Figure 191.
This specimen was collected in the deep-sea plankton near Capri. It was
glassy, transparent in life, with only a few chromatophores which have now
faded. It shows well the typical relationship between gladius and muscular

365

mantle. The rhachis of the gladius is visible as a dark line along the whole
dorsal midline of the body; it widens to form the conical posterior part
(cone flag) before the fins. The cone forms posteriorly a very thin tube
which passes through the fin region and tail as a supporting rod and reaches
the end of the tail. The margins of the tail form 2 small, asymmetrical

FIGURE 191. Young stage of Chiroteuthis
veranyi ("Doratopsis vermicula-
r is "), caught on 30 June 1910 near Capri.

5 X; slightly reconstructed and extended to
correct the consequences of insufficient pre-
servation. Note the characteristic habitus,
tail end, form of fins and mantle, trans-
lucent gladius and cone, wormlike neck region,
the typical larval head with the long "head
column" or "arm pillar," and the form of the
arms and tentacles.

widenings or auxiliary fins. They pass
anteriorly into the membranous part of the
fins (indicated by a dotted line), so that
these are slightly pointed, but otherwise
form a transverse oval. The mantle
sac is cylindrical but with a pointed
posterior part; its slightly bell-shaped
anterior margin has 3 typical but weakly
developed corners.

The funnel apparatus is typical for
the Teuthoidea, except that the funnel and
neck bonds already show the characters
of Chiroteuthis. The adductors are
distinct. The neck region is very long,
separated from the head by a slight con-
striction. The head is not wider than
the neck; if the neck is extended, the head
appears even narrower. The head has
the typical early larval form with a
relatively narrow eye region and wide
cheeks. The olfactory tubercles are
situated on the ventral side of the cheeks,
slightly behind the eyes (Figure 186), in
form of 2 projecting warts. The stato-
cysts project behind them into the last
chamber of the neck.

The "arm pillar," i. e. the muscular
base of the whole arm apparatus is pro-
jecting and only its wide, gelatinous skin
makes it less distinct than in some other
larvae of Chiroteuthidae and Cranchiidae.
The arms are still very short, with a few
biserial suckers. The third pair is the
shortest, stumplike; the ventral arms are
the longest (formula: 4, 2, 1, 3). The
tentacles, however, are already strong
but still undifferentiated, without protec-
tive or swimming margins. The suckers
form a loose row on their proximal half
beginning at the base (at about one ninth)
with a very large sucker. In about the
middle of the tentacle the suckers become
biserial and then rapidly quadriserial.
Each of these 4 rows consists of about
30 suckers, which are so small at the end
that they cannot be seen under the mag-
nifying lens.

366

388

FIGURE 192. Young form of C h i ro t e u th i s veranyi
from Naples ("Doratopsis vermicular! s") 2x.
Slightly reconstructed, after the specimen in the collection
of the Zoological Station in Naples. The dotted posterior
end is missing; the fins are torn from the tail; the mantle
margin is folded back and curved dorsally; the end of the
left club is missing; the mantle sac is probably shown
shorter than in the live animal because of contraction.

Note the changed form of the fins, the enlargement of the
eyes, the growth of the arms, which show already the typical
proportions of the adult, especially the enormous length of
the ventral arms and tentacles. The smaller figures show:

b - lateral view of head; c - deformed dorsal margin of
mantle; d - anterior part of base of fins, magnified;
e — posterior part of base of fins with damaged area.

367

The specimens illustrated by Pfeffer (Plate 4 6, Figures 3—5) and Chun
(1910, Plate 47) more or less resemble this stage, but it is not certain that
they all belong to the same species. At any rate, tail and auxiliary fins
have to be assumed to show a marked variation. The specimens of Issel
(1920, Figures 11 — 12) certainly belong to this species.

Marked progress is evident in a specimen from Naples (Figure 192).

The more developed arm apparatus, the larger eyes and the form of
the fins resemble those of the adult. The fins are wide, leaf-shaped,

389 slightly pointed posteriorly, with rounded, only indicated lateral corners
and typical "earlobes" which are partly attached to the body by a trans-
parent skin fold. The fins have grown further along the mantle, while
previously (Figure 191 ) they were attached only to the tail. The tail pro-
jects as a rod beyond the fins, and is broken posteriorly; its continuation
and the auxiliary fins are reconstructed. The neck region is shrunken,
with crushed and torn chambers; The olfactory tubercles are situated
further posteriorly than previously. The cheeks are receding, so that the
eyes are more prominent. The arms have grown; the third pair is now
longer than the second and the first pair is distinctly shorter. The fourth
pair is more than twice as long as the third and has distinct lateral margins,
so that the arms appear much wider.

The tentacles are also longer and already whiplike, as in the adult. The
hand part is only slightly widened; differentiated swimming and protective
margins are still absent. The arrangement of suckers resembles in
principle that of the previous stage: a larger sucker near the base,
followed by isolated small suckers and then an increasingly dense zigzag
row. The biserial pattern becomes quadriserial at the base of the hand part
and this extends over the entire wide part and end part of the club. How-
ever, the longitudinal rows on the hand part become zigzag rows because the
oblique transverse rows are slightly displaced and alternating. This is
thus not a simple quadriserial arrangement but a transition between a
quadriserial and an octoserial arrangement, which apparently later reverts
to the basic quadriserial type (?). I consider this as a developmental
tendency as in the Mastigoteuthinae and the related Brachioteuthidae, i. e.
a reversion to more primitive conditions or an echo of them (p. 378). A
tubercle on the outer side of the base of the tentacle is the "anlage" of the
"glandular knob" (Figure 192b). The oldest Doratopsis stages follow
this stage (cf., for example, Pfeffer, Plate 46, Figures 1 and 2; also Ficalbi,
i. e. Figure 190c). A similar, finger-long specimen with the tail already
lost so that there is only a pointed apex, is in the collection of the Vienna
Museum and should be more thoroughly studied. The transition between
"D o r a t o p s i s" and Chiroteuthis is well shown in the drawing of
Verrill of " L e p t o t e u t h i s d i a ph a n a" (l884, Trans. Connecticut,

Plate 32, Figure l). This is certainly the juvenile stage of a closely related
form, C.lacertosa Verrill, which has slightly narrower fins (cf . p. 382 ).

The inner side of the ventral arms of these transitional stages bears
luminous organs. On the outer side of the tentacle stalk are the "glandular
knobs" which I also consider as luminous organs. The club has protective
margins and a swimming margin in the distal part. The distal part of the
club bears also larger suckers, especially in the ventral median roV and is
therefore apparently of particular functional importance (Figure 186). The

368

luminous organs on the ventral side of the eyeball begin to show the form
typical for Chiroteuthis veranyi. The stalks of the suckers of the

390 tentacle already have the characteristic structure of this species (Figure 190),

but only in the proximal part. Pfeffer (1912, p. 552) stresses that the club of
Doratopsis vermicularis differs sharply from that of Chiroteu-
this veranyi; 1. Doratopsis bears more than 60 groups of 4 suckers,
Chiroteuthis only 2 6— 27. 2. The suckers in the proximal part of the

club of Doratopsis differ sharply in size from those in the distal region,
whereas those in Chiroteuthis are uniform and decrease only slightly

in size toward base and apex. 3. In contrast to D o r a t o p s i s, there is no
enlargement of the suckers of the ventral median row in Chiroteuthis.

4. The stalks of the distal suckers in Doratopsis are threadlike and do
not have a pillarlike base, but all suckers of Chiroteuthis are thus
attached. Furthermore, the club of Chiroteuthis has no swimming
margin. These differences, which Pfeffer considers decisive, can be reduced
to a single distinction, i. e. that a differentiated distal part of the club is
present in Doratopsis and not in Chiroteuthis. There are no further
morphological differences which could not occur in a direct transition.

Pfeffer realizes that the mantle of these delicate forms is too much changed
by contraction and fixation to be taken as a measure of size, age and stage
of development, especially in the gelatinous types which shrink considerably.

It is therefore not important that the stage of Ficalbi (Figure 190c) has a
slightly longer mantle than the youngest stage of Chiroteuthis (d) and
this may also be due to individual variation. This is even more important
because this stage would have to be considered as related to Doratopsis
since it has an identical club with a distal part with a swimming margin
and enlarged suckers, according to Ficalbi's drawings and description
(Figure 190c). Since Pfeffer (pp. 605—606) placed this specimen in Chiro-
teuthis, the gap is bridged and further discussion is unnecessary. In my
opinion, the matter can be explained as follows. We must assume that such
a "L e p t o t e u t h i s" stage precedes the developed Chiroteuthis. This
condition is caused by the loss of the end part of the tentacle club. At least,
it is not clear that the loss of the distal part of the tentacle club eliminates
the contrast between D. V e r m i c u 1 a r i s and C h i r o t e u t h i s veranyi;
the difference disappears altogether if it is assumed that the stalked suckers
of Doratopsis (which escaped the notice of Pfeffer) later disappear and
the characteristic terminal knob of the tentacle club develops instead of
the distal part. To check this view, let us examine Figure^ 190c. Although
not very good, this drawing clearly shows an intermediate form between the
typical adjacent figures of Doratopsis and Chiroteuthis. The tail
is apparently lost, but the specimen differs from Doratopsis also in the
wider mantle sac and head, the shorter neck, the larger eyes and the pro-
gressive development of the arms. But there is no marked difference
between the two stages, and there is no reason to doubt that the second

391 developed from the first. The tentacle club bears the typical distal part

of Doratopsis (Figure 190b), but at the same time, there are unmistakable
characters of C h i r o t e u t h i s, though only roughly indicated: there are
"glandular knobs" along the whole outer side of the tentacle, and protective
margins which are wider in the proximal part and become narrower distally.
The artist must have been impressed at this point by the larger transverse

369

supports, which are indicated in Figure 190c. The part of the club with
these structures is situated at the end in Chir ot euthi s, but it is here
followed by a distal part, which appears as an appendage in the drawing and
has a typical swimming margin and apparently enlarged suckers but no
glandular knobs. To explain how this club can be transformed into the
typical club of Chiroteuthis of the next stage (Figure 190d), in which a
swimming margin and a true distal part are already absent, there are two
possibilities. Either the whole distal part has been detached or perhaps
lost, like the suckers of Onychoteuthis, the tentacles of Octopodo-
teuthis or the dorsal arms of Tremoctopus; in this process, the last
glandular knob would become terminal and could develop into a terminal
knob. Or, the small terminal swelling of the preceding stage has developed
further, the substance of the whole distal part has been resorbed and the
suckers of this part have fallen off so that the typical terminal knob of the
club of Chiroteuthis was formed. New material or a re-examination
of the material of Ficalbi would be necessary to settle this point. The
available evidence is in favor of the first possibility, however.

The young C.veranyi shown in Figure 190d is closely related to the
above stage and shows the typical external characters of the species. The
mantle is widened further, the neck shortened, the eyes markedly enlarged,
and the arm apparatus stronger. The changes described above are already
complete on the tentacle club. The half-grown animal is shown in Plate IV,
Figure 3. The specimen in Figure 188 is still markedly immature but shows
the outline of the adult Chiroteuthis.

370

392 Chapter 28

FAMILY CRANCHIIDAE

Gray, 1849*

a. DIAGNOSIS

Funnel bonds and neck bonds fused with the mantle. Arms short, the
third or fourth pair the longest, the first pair remaining the shortest. Outer
funnel adductors or valves absent. Hand part of club with 4 rows of "an-
lagen" of suckers, of which the two median rows develop into hooks during
postembryonic development. Gladius consisting mainly of a long free
rhachis without widened lateral plates, and a cone flag with a tubular,
conical or spoon-shaped cone.

I had no material of Cranchiidae in Naples and the following description
was added later and is based mainly on examination of museum material.

This family could not be omitted from a synoptic treatment, because it
is necessary for the understanding of the relationships between the families
described above. Furthermore, 3 species of Cranchiidae probably occur in
the Mediterranean, one of which embodies all the typical characters of the
family.

b. TYPICAL STRUCTURE OF THE ADULT ANIMAL

The Cranchiidae show a large, almost fantastic variety of forms as well
as a number of peculiar and distinct characters. Chun (l910) therefore
opposed this group as "Oegopsida consuta" to all other Oegopsida ("Oegop-
sida libera"). Pfeffer (1912, p. 636) accepted this with some reservations
and did not include it in his classification. At any rate, Chun's concept is
not valid morphologically. The Cranchiidae are, on the contrary, a member
of a larger group of Oegopsida which also includes the Brachioteuthidae,

393 Joubiniteuthidae and Chiroteuthidae, as we have stated repeatedly (pp.236, 377).
This appears clearly if we attempt to determine the typical characters and to
unite them in a general picture. The sharp differences from the general
type of Oegopsida are obviously important, but even in the most marked
differences the resemblance to the above-mentioned families is evident.

The natural relationships would have to be disregarded if Chun's view were
accepted.

« The family was recognized and named in its typical character simultaneously by Gray and Prosch. Gray
(1847) writes; "Cranchiae" Prosch (1847): "Cranchina"; both list the Cranchidae as a family in 1849.

371

FIGURE 193a, b. Mantle sac of G a 1 i t e u t h i s armata from Messina, with translucent gladius. V2 x,
slightly diagrammatic. This form is typical for the family and shows its relationship with Chiroteuthis
(Figure 191) and Brachioteuthis (p. 372). The cone (Co) is very long, as in Chiroteuthis and
related forms. Note also the attachment of the fins, which are only partly fused on the cone flag (Fa).

This is visible dorsally (b) and forms a typical "lanceola" together with the adjacent part of the wall of the
cone, which is visible through the skin between the fins.

The other figures explain the typical relationships:

c — typical posterior end of a Metateuthoid, ventral; d — mantle sac (as far as not covered by c), dorsal
(cf. p. 153); e — young stage of Enoptroteuthis spinicauda after Berry (1920, Plate 16, Figure 6, 2 x).
This animal is a peculiar combination of the characteristics of the Cranchiidae and those of the Chiroteu-
thidae; I therefore placed it in the Grimalditeuthinae (1921, p. 535). The eye stalks, arm pillar and proportions
of the arms are of the type of Cranchiidae, the funnel bond is stated to resemble that of the Chiroteuthidae
(West Atlantic); f — also a planktonic young stage, "Chiroteuthoid es hastula" (West Atlantic) (Berry,
Figure 3); this certainly belongs to the Chiroteuthidae (ventral arms, fins), but does not have a long neck or a
tail (Mastigoteuthinae? ) (p. 378); g — a stage of Pyrgopsis (from the same area) (P. lemur Berry,

Figure 5) of Lea chi a (cf. p. 406); h - a typical early stage of Cranchiidae (cf. p. 396) with retracted head;
i - typical posterior end of a later stage of Chanchiidae, still with the general outline of the young Meta-
teuthoidea from which develop the conditions shown in Figure c; there is also an extreme lengthening of the
cone in the Chiroteuthidae and Cranchiidae (cf. Figure 187 on p. 373 and Figure 199 on p. 404).

372

394 The typical general appearance of Cranchiidae is shown by Galiteu-

this, which may be considered in many respects as the ancestral form of the
family (Figure 196). It is characterized by its general outline and by the
membranous gelatinous consistency of the body. The musculature is less
developed than in the Onychoteuthidae, Gonatidae and Ommatostrephidae.

The skin is swollen. All forms are small and delicate. The small number
of chromatophores makes the body more or less transparent, especially in
young specimens.

FIGURE 194. Diagram of the relationships between funnel and mantle in the Cranchiidae (b) for
comparison with typical Oegopsida (a). (Compare also Figure 32 on p. 100 for an understanding
of the typical relationships.) The reduction of the shell does not change these relationships in
principle. As in most adult Oegopsida, however, the funnel retractor is inserted on the mantle,
since the narrowed flag of the gladius has become retracted from the primary point of insertion.

The characteristic conditions in the Cranchiidae are caused by the formation of 4 sutures (nj— 04)
between funnel and mantle. Sutures n3 and n4 correspond to the boundaries of the lost funnel
cartilage, n^ marks the position of the nape cartilage; ng is a continuation of n4, i.e. of the con-
nection between mantle and funnel septum (and therefore of the origin of the funnel retractor).

This connection accentuates the division of the mantle cavity into a laterodorsal and a ventral
part, a condition already indicated in the typical forms. The thick line "y" is the margin of the
inner septum, i.e. the edge of the bandlike funnel retractor. It marks the opening between the
two parts of the mantle cavity, in which the gills are situated, thoroughly washed by the flow of
water. The locomotory mechanism is simplified because a retraction of the funnel in the mantle
sac (p. 93) is obviously no longer possible.

The mantle sac is very narrow, spindle-shaped (Figure 193) to calyx-
shaped. It has a long, thin apex, which is occupied by the very long cone;
the fleshy apex of typical Oegopsida is absent (cf. p.377). The gladius
forms a leaf-shaped cone flag, but otherwise consists almost entirely of a
narrow, long rhachis which is visible in the dorsal midline. The fins are
separated or only partly fused; together they form a pointed, leaflike figure
into which the lanceola is wedged anteriorly. It extends from about the
widest part of the cone flag to the end of the cone, but it is represented only
by margins near the needle -thin apex. The anterior margin of the mantle
forms the 3 indistinct corners. Characteristic cartilaginous ridges extend
from these corners posteriorly on the mantle sac; there is a single dorsal
ridge and two ventral ridges, which are connected anteriorly, diverge and
bear several tubercles posteriorly (Figure 202).

373

The head consists mainly of the large, protruding eyes. Neck folds are
absent, and the olfactory tubercle is a thin, stalklike process behind the
eye. The funnel pit and the outer funnel adductors are apparently absent,
and there is no sinus on the lid margin. The ventral side of the eyeball
bears numerous lens -shaped luminous organs, surrounded by a shining
golden margin.

The arm apparatus is weakly developed so that there is a marked
constriction before the eyes. The arms are short and delicate, although
they are the most muscular part of the body (formula: 4, 3, 2, 1 or 3, 4, 2, 1).
The swimming margins are rudimentary, but the protective margins are
well developed, with distinct transverse supports which correspond in
number and arrangement to those of the respective rows of suckers. The
outer edges of the adjacent arms are united at the base (p. 384) into small
membranes, except between the V arms. All arms bear 2 rows of suckers.

The tentacles are relatively long. The stalk part bears 2 rows of small
suckers which form zigzag rows toward the hand part; and there are several
knobs between them in the distal part. The suckers become larger and
quadriserial on the carpal part and the knobs become more distinct. The
hand part of the young animal bears 4 rows of enlarged suckers; the two
median rows finally develop into hooks. The distal part is short and also
bears 4 rows. The protective and swimming margins are very delicate and
indistinct. The buccal funnel consists of 7 parts; the ventral attachments
are median.

395 The funnel is rather strong and characteristically modified. 1. The
neck bond is replaced by a wide, solid fusion which does not form a neck
ligament as in the Sepiolinae (Chapter 42); the neck cartilage has disappeared
2. The adhesion surface between the funnel bond and the mantle bond is
also replaced by a slender triangular concrescence with the pointed angle

in the mantle corner. The funnel pocket is thus dorsally and ventrally
attached to the mantle, as in the Octopoda, by a longitudinal suture on each
side of the median line. The ventral wall of the funnel is now attached to
the mantle on each side, instead on the funnel cartilage. Because of this
connection between funnel and mantle, the funnel retractor originates on the
mantle, at least in its lateral part, which normally originates on the funnel
cartilage by its connection with the mantle (Figure 47, on p. 124): The
funnel retractor now connects mainly 2 parts of the mantle, forming on each
side a septum which divides the mantle cavity incompletely into a paired
laterodorsal and a single ventral part (Figure 194). These relationships
are not as peculiar as Chun thinks; the main importance of the functional
change is the narrowing of the slit between the laterodorsal part and the
ventral part of the mantle cavity, caused by the formation of the suture n2,
which can extend still further posteriorly and reduces the slit to a so-called
spiraculum. There is no change in principle in the flow of water through
the mantle cavity because of these changes. The funnel gland consists

396 typically of 3 parts. The bilobed median part has 3 tongue-shaped processes
one median and two on the lateral lobes. They are very characteristic for
the family, and so is the absence of a funnel valve.

The mantle cavity shows juvenile characters.: the gills are short, small
and little segmented; the hind intestine is also short; the anus is therefore
situated far posteriorly; the ink sac is small and median in position, and one
of the ventral arms of the male is more or less modified.

374

c. POSTEMBRYONIC DEVELOPMENT OF THE
TYPICAL CRANCHIIDAE

The earliest young stages differ little from those of other Oegopsida.'i' The
characteristic relationships between funnel and mantle cannot be assumed
to exist at the time of hatching. Further advanced larvae have stalked
eyes and an arm pillar which projects between the eye stalks. The
tentacles are relatively long and strong, the other arms are stumplike,
almost rudimentary. Suckers are present on the whole length of the
tentacles; the terminal suckers are at first arranged in 2 rows, as usual , but
later become quadriserial. The animal shows otherwise the typical
characters of the Oegopsida: the posterior end is at first blunt, with a
spoon-shaped cone which bears small, rounded fins, which are more or less
widely separated. The posterior end becomes later more pointed, the fins
become contiguous in the middle and gradually assume their typical form.

The tentacles are usually lost, so that there are always only stumps in some
species.

FIGURE 195. Typical young
stages of Cranchiidae. a)
with a long arm pillar;
b) with a short arm pillar.
Note the stumplike arms,
the long tentacles, the con-
crescence of the funnel, and
the larval fins, which are
separated at first. After
Chun, 1910, Plate LXI.

a — 5X; b — 3X.

397 d. VARIATION OF THE TYPE

Chun (1910) divided the family into the following subfamilies:

A. Species with cartilaginous ridges originating on the concrescences of
the funnel on the dorsal or ventral side of the mantle or on both,
each ridge with a row of cartilaginous tubercles. Ventral side of eye-
ball with at least one longitudinal row of 4 or more luminous organs
1 . Subfamily Cranchiinae Pfeffer

* Cf. Figure 472; but the arms are particularly retarded.

375

Genera: Leachia, Pyrgopsis, Liocranchia, Cranchia, Li-
gu r i e 11 a.

B. Species in which the ventral side of the eyeball bears only one luminous
organ or two crescent-shaped organs, the larger surrounding the
smaller. Cartilaginous ridges with tubercles either absent or re-
presented by a minute rudiment (Toxeuma) or a number of branched
tubercles (C r i s t a 1 1 o t e u t h i s) .... 2. Subfamily Taoniinae Pfeffer

Genera: Taonius, Galiteuthis, Desmoteuthis, Megalocran-
chia, Phasmatopsis, Phasmatoteuthion, Toxeuma, Cristallo-
teuthis, Corynomma, Taonidium, Teuthowenia, Bathothauma.

Some of these genera are juvenile forms (Pyrgopsis, Liguriella,
Corynomma, Taonidium, Teuthowenia).

376

398 Chapter 29

GENUS GALITEUTHIS

Joubin, 1898

GALITEUTHIS ARMATA Joubin, 1898
a. DIAGNOSIS

Posterior end of mantle sac produced into a long thin apex which con-
tains the cone and sometimes projects beyond the fins. Fins forming a
narrow, leaf-shaped figure on the tapering part of the mantle sac. Eyes
large, spherical, still markedly protruding but not stalked in the adult.
Suckers of arms with smooth chitinous rings (in contrast to Taonius).
Median rows of tentacle club developing into hooks during postembryonic
development. Chromatophores brownish, less numerous than in Taonius,
especially on the mantle, so that the animals are pale.

b. LITERATURE

Joubin's original specimen is from Nice, but this species is apparently
cosmopolitan. No significant differences were found between the following
species.

1898 Joubin, Galiteuthis armata (pp. 279,292, Figs. 1-9).

1910 Chun, Galiteuthis (Taonidium) suhmii (p. 382, Plate 59, Figs. 1-11).

1911 Berry, Galiteuthis phyllura (p. 592).

1912 Berry, Galiteuthis phyllura (pp. 315-317, Plate 46, Figs. 1-3, Plate 54, Figs. 5-6, Plate 56).

1912 Pfeffer, Galiteuthis armata (pp. 731-736).

1916 Naef, Galiteuthis armata (p. 15).

1921 Naef, Galiteuthis armata (p. 538).

1920 Issel, Galiteuthis armata (pp. 12-17, Plate, Figs. 13-21).

Figures 193 and 196—198 show further specimens from Messina. I saw
a large specimen from the Atlantic in Pfeffer 's collection in Hamburg.

No specimen of Galiteuthis or other Cranchiidae, young or adult, have
ever been caught in Naples, as far as I know.

377

399 c. STRUCTURE OF THE ADULT ANIMAL

Galiteuthis has a pale coloration, because of the very scarce chroma-
tophores on fins, mantle sac and arms and a gelatinous -membranous con-
sistency of the body, caused by the delicate musculature. The general
appearance is typical for the subfamily, and resembles that of Taonius
pa VO (Les.). The mantle sac is very long and the cone forms a tail-like
appendage ("tail filament") which projects beyond the fins, at least in the
smaller specimens. The fins form a narrow, leaf-shaped figure with
pointed posterior end. The fins of Berry's large specimen are about as
long as the mantle area anterior to them.

FIGURE 196. Galiteuthis phyllura*
(= G . a r m a t a Joubin?) , after Berry
(1912, Plate 56). Half natural size. Note
the outline of mantle sac, fins, head and
arms. The protective margins are wide,
and there is a stalked olfactory tubercle
behind the eye. This is the largest de-
scribed specimen of the genus.

* It is not certain that the Californian and Mediterranean form belong to the same species. Berry’s

specimen has more delicate tentacles than Joubin's original specimen and fewer suckers than the specimen
described by Chun. The "tail filament" is apparently absent. On the other hand, these characters may be
due in part to age differences, or perhaps to damage. At any rate, this is a closely related species which
represents the general appearance of a typical adult Galiteuthis.

378

The eyes are very prominent, spherical, but no longer stalked. The
larval arm pillar is reduced to a constriction in front of the eyes. The
ventral side of the eyeball bears a large, lens-shaped luminous organ

(Figure 198). A knoblike, stalked olfactory tubercle
is situated behind the eye.

The arms are short but quite strong (formula:

4^ 3, 2, 1). The third pair of arms has a swimming
margin, the other arms apparently not. All arms
have wide protective margins with strong transverse
supports. An inner membrane is probably present
between the 6 dorsal arms, as in Cranchia (after
Chun, 1910, Plate 1, Figure l). All suckers have
smooth rings.

The tentacles are rather long and strong and pro-
vide the most important characters. We know little
on the tentacles of other adult Cranchiidae, as
practically only juvenile specimens are known and
adult Leachia and Taonius have always lost
their tentacles. The stalk part is flattened on the
inner side, which has a thin longitudinal groove
between the loosely arranged pairs of small suckers
which alternate regularly with pairs of adhesive
knobs. This pattern passes into a denser zigzag row
which is divided further on the carpal part into the
originally quadriserial arrangement on the hand part.
The carpal part consists of a group of about 8 slightly
larger suckers with an equal number of knobs
between them. Only the median rows are preserved
on the club, with about 12 hooks; the suckers of the
marginal rows have been lost. The distal part bears
about 16 suckers in 4 rows. Protective margins are
present on the club, but a swimming margin is absent.

d. POSTEMBRYONIC DEVELOPMENT

The typical early stage shown in Figure 199c
probably belongs to Galiteuthis. This identification is the most probable
because of the relative frequency of this species in Messina. Issel (l920)
has recently published descriptions and drawings of similar stages which
definitely belong to this genus. The smallest of them, with the habitus
shown in Figure 199c, has a dorsal mantle length of only 6.5 mm. The
following stages rapidly approximate the condition shown in Figure 199d.

At a mantle length of only 8 mm (issel. Figure 15), the fins are already
'401 longer and have an almost transverse posterior margin, resembling the fins
in Figure 199c, d. They are about 0.7mm long. The mantle sac (cone) has
already become longer and more pointed, and the eye stalks are very slender
and later become gradually shorter. The oldest stage described by Issel
(Figure 19) is about five-sixth the size of the specimen shown in Figure 199d.
Chun described similar stages from the Guinea Current. The specimen

FIGURE 197. Juvenile
form of Galiteuthis,
after Chun (1910, Plate 59,
Figure 3), natural size.

The fins are still short.
Note the short "tail fila-
ment," the stalked eyes,
and the arm pillar with
the rudimentary arms.-
" Galiteuthis (Taoni-
dium) suhmii."

379

FIGURE 198. Young G a liteu this, natural
size, after Chun (1910, Plate 59, Figure 1,2)
"Galiteuthis (Taonidium) suhmii."
Note the further development of the tail fila-
ment, fins, eyes, arms and tentacles.

380

shown in Figure 197 has rounded fins, beyond which projects the "tail" fila-
ment or perhaps the cone which has pierced it. The eyes are stalked and
directed anteriorly. The arms are very short and delicate and are situated
on a long pillar. The tentacles are disproportionately long, as in all larvae
of Cranchiidae, and already show the beginning of the characteristic trans-
formation of the club: the hand part still bears 4 rows of adhesive organs
(most of them still true suckers); 8 organs in the median rows already
bear developing hooks but they still resemble suckers (Chun, Plate 59,
Figure 6.7). The carpal part bears a number of knobs but these are still
absent on the stalk.

The older specimen shown in Figure 198 distinctly resembles the
preceding stage, except that the fins and tail filament are larger and the
eyes are no longer stalked, although still markedly prominent and situated
as in the younger animal. The arm pillar is shorter, but the arms are
longer. The tentacles are relatively shorter, and their transformation is
apparently complete: the marginal rows of the hand part have been lost,
and the median rows consist of 2X6 large hooks. A large, rounded
luminous organ is present on the ventral side of each eye in both specimens.

381

402 Chapter 30

GENUS LIOCRANCHIA

Pfeffer, 1884

DIAGNOSIS

Mantle sac pointed posteriorly but not long, calyx-shaped; fleshy apex of
body reduced to a frenulum between the fins. Fins rounded, fused in the
middle and projecting distinctly beyond the short cone. Mantle sac with a
few regularly arranged chromatophores and 2 ventral cartilaginous ridges
on each side which are contiguous at the corners of the margin and bear a
row of tubercles but no scattered cartilaginous tubercles. Ventral side of
eyeball with at least one row of 4 similar, rounded luminous organs.

Suckers of arms with smooth rings; suckers of median rows of tentacle club
with several peglike teeth at the distal margin.

TYPICAL STRUCTURE OF THE ADULT ANIMAL

The typical general appearance of the animal is shown in Figure 199a,b.
The body is membranous, transparent and delicate, only slightly larger than
the drawing (cf. Chun, 1910, Plate 48). The sparse chromatophores create
a slight brownish tone on the mantle; the chromatophores of arms and head
are slightly more densely arranged.

The gladius shows a typical, thin, delicate rhachis in its anterior half,
while the posterior part consists of a slightly widened flag with a short,
conical cone. As in the Histioteuthidae, the cone continues posteriorly in
a short, reduced, thin fleshy apex which is attached as a frenulum medio -
ventrally to the suture of fusion between the almost circular fins.

The mantle sac is narrow, calyx- shaped, without scattered cartilaginous
tubercles. It bears a number of cartilaginous ridges, one dorsomedian on
the gladius, and 2 ventral ridges on each side w^hich originate at the position
of the funnel corner and diverge posteriorly; each ridge bears an alternating
row of larger and smaller tubercles. The ventral ridges occupy slightly
more than the anterior quarter of the mantle; the dorsal ridge may extend
posteriorly to the last quarter during postembryonic development.

403 The eyes are of medium size, directed slightly anteriorly, and project only
slightly. The ventral side of the eyeball bears rounded luminous tubercles,
at least 4 of which are situated at the insertion of the eye and form a regular
longitudinal row (Figure 199a). The arms are short and delicate, with
12—30 suckers , and are connected by a delicate membrane, except on the
ventral arms (formula: 3, 2, 4, l).

382

The tentacles are strong. The clubs are short, slightly widened, with
4 rows of suckers. The stalk part bears a zigzag row of very small
suckers, of which the proximal suckers are gradually lost. The suckers are
larger on the carpal part, of the adhesive type, and there are a few indistinct
knobs between them (Chun, 1910, Plate 51, Figure 12). Protective margins
are present everywhere as narrow edges, but a swimming margin is only
present at the apex of the club and on the third pair of arms.

Hectocotylization was observed by Lonnberg (1896) and Chun (l910). It
consists of a modification of the suckers and their stalks on the left ventral
arm (Pfeffer, 1912, p. 67l).

POSTEMBRYONIC DEVELOPMENT

The general appearance of the genus is already distinctly recognizable
in young forms only 9 mm long (Chun, 1910, Plate 51, p. 345). The eyes are
already lateral, situated inside the orbit, only slightly prominent and without
stalks. An arm pillar is not visible, and the arms are relatively strong.

The ventral cartilaginous ridges on the mantle are already distinct, but still
without tubercles. Liocranchia differs from corresponding stages of
Cranchia in the absence of stellate tubercles on the mantle. Both genera
apparently show the characters of young Cranchiidae (arm pillar, eye stalks)
only temporarily and little marked.

LIOCRANCHIA REINHARDTI (Steenstrup, 1856)

Pfeffer, 1884

DIAGNOSIS

Dorsal cartilaginous ridge extending over the greater part of the mantle
length and with numerous tubercles which are not quite uniformly developed
and distributed. Both ventral cartilaginous ridges of one side end in a double
tubercle with 2 apices, one behind the other, on the margin of the mantle.

LITERATURE

1856 Steenstrup, Leachia reinhardti (p. 200).

1861 Steenstrup, Cranchia reinhardti (p. 76).

1879 Tryon, Loligopsis reinhardti (p. 165).

1832 Brock, Cranchia reinhardti (p. 605; Plate 37, Fig. 4).

1884 Rochebrune, Perothis reinhardti (p. 25).

1884 Pfeffer, Liocranchia reinhardti (p. 29, Fig. 35).

1884 Pfeffer, Liocranchia brockii (p. 25, Fig. 33).

1884 Hoyle, Cranchia (Liocranchia) reinhardti (p. 184, Plate 31, Figs. 11-14, Plate 32, Figs. 1-4).
404 1892 Girard, Cranchia reinhardti (p. 217).

1896 Lonnberg, Cranchia reinhardti (p. 607, Figs. 1-4).

1900 Pfeffer, Liocranchia reinhardti (p. 194).

1906 Chun, Liocranchia reinhardti (p. 84).

383

1908 Issel, Liocranchia reinhardti (p. 218, Plate 9, Figs. 24— 26, Plate 10, Fig. 27).
1908 Issel, Liocranchia elongata (p. 220, Plate 10, Figs. 18— 32).

1910 Chun, Liocranchia reinhardti (p. 336, Plate 51, Figs. 5-7).

1912 Pfeffer, Liocranchia reinhardti (p. 676, Plate 48, Figs. 19-21).

1921 Naef, Liocranchia reinhardti (p. 538).

FIGURE 199.

a, b — Lio c r a nc h i a reinhardti (from Messina?). Well preserved (half grown) specimen from the
Leipzig Collection, collected by Carus. Reconstructed by straightening of the curved gladius and therefore
longer than in the preparation. Note the form and attachment of gladius and fins, the cartilaginous
ridges with tubercles on the mantle sac, and the luminous organs on the compact head. The larval
characters (arm pillar, eye stalk) have disappeared.

c — Typical juvenile stage of Cranchiidae (Galiteuthis?) from Messina (Leipzig Collection). 4x.

The absence of specific characters prevents more exact determination. The young forms of Leachia
and Galiteuthis (Figure 195 on p. 396), are probably similar, especially in the typical form of the
posterior end (a character of the Teuthoidea), the stalked eyes and the presence of an arm pillar with
short, delicate arms (characters of the Cranchiidae), and the simple tentacles, with suckers almost their
whole length, as is characteristic for young Oegopsida. The optic nerve and ganglion are visible in the
eye stalks. 2x.

CiiC2,C3 - ventral view of posterior ends of young Cranchiidae, the first younger and more highly
magnified, the third older and less magnified than Cj. Cone and fins begin to grow (Figure 193 on p. 393).
d - young Galiteuthis armata from Messina, natural size, from the Zoological Collection in Leipzig.
This older stage differs distinctly from Leachia in the pointed posterior end and the single large luminous
organ on the ventral side of the eyeball. The sparse chromatophores also suggest Galiteuthis.

384

THE ADULT ANIMAL

This species has also not been collected in Naples and it is very rare
in Messina. However, I made a detailed examination and drawings in
405 Leipzig of a well preserved specimen from the Cams collection, stated to
be from Messina'’' (Figure 199).

The habitus of the animal is typical for the genus (Figure 199). Gladius
and fins are as described on p.402. In large specimens, the fins reach one-
seventh of the mantle length according to Lonnberg (1896). The cartilaginous
tubercles of the ventral side are pointed, slightly compressed laterally and
twice as high as wide at the base; the smaller tubercles form more or less
irregular rows of 14—19, which grow longer during development. The two
rows end on the mantle margin in a double tubercle with 2 points, one behind
the other. The anterior end of the dorsal cartilaginous ridge is similar; it
extends posteriorly to about half of the length of the lanceola and bears
40—50 tubercles. The head has the typical form of young Teuthoidea. The
eyes are of medium size, slightly prominent and directed slightly anteriorly.
The olfactory organ forms a tubercle in the posterior part of the eye area.
The ventral side of the eyeball bears about 9 translucent luminous organs
of which 5 larger organs form a longitudinal row between the base and
equator of the eyeball; the other organs are less regularly arranged near
the pupil. The third arm is the longest, the first the shortest, the fourth and
second pairs of about equal length. The longest arms bear about 30 suckers,
the other arms fewer. The suckers of the ventral arms are very small but
more numerous. The suckers on the hand part of the club are rather uni-
form in size and bear several peglike teeth on the distal margin.

The largest specimen measures 46 mm from the anterior margin of the
mantle to the posterior margin of the fins. Because of the strong contraction
which always takes place in these Cranchiidae, it is only about 1.5 times as
large as the specimen in Figure 199; at any rate, it is smaller than the
drawing. This is a male, with distinct hectocotylization of the left ventral
arm: the suckers (24—26 pairs) rapidly decrease in size toward the apex
and appear reduced; the apex bears only about 16 transverse papillae
(basal pads). Pfeffer (1912, p. 671) attributes this condition to accidental
loss of the suckers and thinks that there is no difference in the hectocoty-
lization of L. reinhardti and L. valdiviae (Chun, 1910, Plate 51,

Figures 8,9). The hectocotylus of L. valdiviae is only slightly modified,
shorter, and its distal suckers are larger and uniserial.

I have not seen younger stages, but they probably show the characters of
the genus described above.

* This origin is certain according to Dr. G. Grimpe, Leipzig, who placed the specimen at my disposal.

I could not confirm it because Dr. Grimpe had already replaced the original label. There is, however,
some doubt. Also if this is an alien form, its description may show' new aspects of the typical relation-
ships in this family which are our main object. This is particularly necessary because most authors stress
more the specific characters and the characters of the family than the relationship to a more general
type. Cf. p. 292.

385

406 Chapter 31

GENUS LEACHIA

Lesueur, 1821

a. DIAGNOSIS

Body gelatinous. Ventral surface of mantle with only one (median?)
cartilaginous ridge with stellate tubercles on each side. Cone very slender,
pointed, reaching to the end of the body. Apex of cone at level of posterior
margin of fins or projecting slightly beyond. Eyes small, projecting, with
long stalks in the young stages. Arm pillar distinct, slender, lengthened in
youth.

The description of the genus is included here because Pfeffer (1912,
p. 656 ff. ) was of the opinion that "Loligopsis z y g a e n a" Verany
(cf. Figure 202) belongs to "Pyrgopsis." Although this cannot be accepted
or rejected at present, there is no doubt that the P y r g o p s i s-like young
forms develop into Leachia-like adults, and are their predecessors, in the
same way as the Doratopsis larvae develop into Chiroteuthis (p.387).
The only differences between them are either juvenile characters (arm
pillar, eye stalks) which do not justify the establishment of a genus, or
characteristics of habitus (form of the fins) which are at most species
characters. The eye stalk and arm pillar are later reduced, as in the
Cranchiidae (p. 39 6). Pyrgopsis are therefore only young Leachia, and
it is possible that the larval characters persist longer in some species
than in others or never disappear completely (cf. Figure 201 on p.408). This
does not affect the generic position. We therefore give a brief description
of L e a c h i a, based mainly on Pfeffer (1912, p. 656). Older, determined
stages are not known from the Mediterranean.

b. TYPICAL STRUCTURE OF THE ADULT ANIMAL

Adult Leachia has a slender, delicate, gelatinous body with numerous
dark but easily bleached chromatophores which, together with the iridocytes,
produce a violet tone.

The mantle sac is spindle- or calyx-shaped. It tapers posteriorly to a
narrow apex in which the cone is situated. The gladius is typical. The fins
are short, wide, separated in the middle and form together a transversely
407 oval or heart-shaped figure with a small posterior apex. The mantle bears
ventrally 2 cartilaginous ridges with smaller or larger, stellate, cartilaginous

5110/2

386

tubercles. The larger tubercles have the form of a transverse rhombus
with a tubercle in the corners and middle which may be simple or with
2 or 3 apices (Pfeffer, 1912, p. 647). There are intermediate forms between
the simple and divided tubercles. The head is small, with moderately large,
markedly protruding eyes with a longitudinal row of rounded luminous
tubercles directed ventrally and toward the middle. There may be other
luminous organs near the margin of the pupil (they form a second longi-
tudinal row in L. cyclura Les.). There is a knoblike, stalked olfactory
organ behind the eye.

FIGURE 200. Leachia cyclura (a,b)and Pyrgopsis schneehageni (c, d), after Pfeffer
1912, Plate 47); natural size. The juvenile form belongs to a closely related species, if not to
L. cyclura. Note the absence of tentacles in the adult; the projecting eyes; the cartilaginous
ridges on the mantle; the short arms on the slender pillar; calyx- shaped mantle; narrow gladius
(cf. Figure 106b); rounded fins (cf. Figure 393).

387

The arms are gelatinous, rather short, with very weak swimming margins
only in the distal part, and with weak but distinct protective margins. There
are 20—30 pairs of suckers which form 2 rows. As in many other Oegopsida
(pp. 323, 339), the tentacles are probably later lost, leaving only stumps.

408 The buccal funnel consists normally of 7 parts. The funnel is very large and
relatively strong; these are apparently good swimmers of the deep sea.

Two species are well known, L. cyclura is of greater interest since
the the following juvenile form probably belongs to it (Figure 202 on p. 410).

c. YOUNG FORMS

I consider the animals described by Pfeffer (1912, p. 656) as Pyrgopsis
Rochebrune to be young forms of Leachia. Synonyms: Zygaenopsis
Rochebrune, 1884; Pfeffer, 1900; Issel, 1908; Zygocranchia Hoyle,

1909; Euzygaena Chun, 1910.

FIGURE 201. Leachia pacifica (Issel, 1908), juvenile specimen, natural size,
after Chun, 1910, Plate 52, Figure 1. This stage is approaching sexual maturity,
since the left ventral arm is already hectocotylized (Chun, Plate 52, Figure 3);
the animal resembles adult Leachia in the following characters: the eye stalks
and arm pillar are thicker and shorter, the arms longer and the fins rounded. It
undoubtedly belongs to a species closely related to Leachia cyclura
(cf. Figure 200). If the larval habitus of the head were considered as a permanent
generic character, the name given by Chun Euzygaena pacifica would have
to be maintained and Euzygaena would have to be made a subgenus of
Leachia.

This is a small animal, with a gelatinous body which is certainly
completely transparent in life (Figure 200, c,d). The mantle sac is like
that of the adult; the cartilaginous ridges bear simple and stellate tubercles.
The fins are terminal and form together a transverse oval or trapezoidal,
slightly pointed figure with lateral, posteriorly displaced corners. The
eyes have stalks of different length which are thin at the base. "Anlagen" of
luminous organs are present on the ventral side of the eyeball. The arm
pillar is very long and thin, the arms are very short and the tentacles are

388

moderately long and delicate. The hand part of the club is widened; the
median rows consist of distinctly enlarged suckers with long teeth. Typical
swimming and protective margins are present. In large males, (Figure 201),
one of the ventral arms begins to become hectocotylized (Chun, 1911;

Pfeffer, 1912, p. 663).

409 LEACHIA CYCLURA Lesueur, 1821
DL\GNOSIS

Ventral side of eyeball with a longitudinal row of 4 (or 5) rounded
luminous organs of medium size; there is also another small organ situated
on the line connecting the last organ of the row with the pupil (Pfeffer,

1912, Plate 47). Cartilaginous ridges on ventral side of mantle less than
one third of mantle length.

The well preserved specimen of Pfeffer (1912, p. 653) has regularly
distributed dark spots on a whitish violet background.

We place the following juvenile stage in this species with some reservation
and use the original name as it cannot be identified with certainty.

LITERATURE

1821 Lesueur, Leachia cyclura (p. 90, Plate 6).

1833 4athke, Perothis dubia (p. 170? ).

1835 Grant, Loligopsis guttata (p. 24, Plate 2).

1839 D'Orbigny, Loligopsis guttata (Plates 3, 4, Figs. 9-16).

1839 D’Orbigny, Loligopsis cyclura (p. 322).

1845 Adams and Reeve, Loligopsis ellipsoptera (p. 2, Mollusca, Plate 1, Fig. 1).

1849 Gray, Loligopsis ellipsoptera (p. 40).

1849 Gray, Loligopsis cyclura (p. 41).

1861 Steenstrup, Leachia cyclura (p. 82).

1861 Steenstrup, Leachia ellipsoptera (p. 80).

1879 Tryon, Loligopsis ellipsoptera (p. 163, Plate 68, Fig. 254).

1879 Tryon, Loligopsis cyclura (p. 163, Plate 69, Fig. 255).

1879 Tryon, Loligopsis guttata (p. 164, Plate 70, Figs. 259-264).

1884 Rochebrune, Loligopsis cyclura (p. 12).

1884 Rochebrune, Dictydiopsis ellipsoptera (p. 16).

1905 Joubin, Leachia cyclura (Figs. 1-2).

1912 Pfeffer, Leachia cyclura (Plate 47, Figs. 2-10).

PYRGOPSIS ZYGAENA (Krohn, fide Verany, 1851) Pfeffer, 1912

LITERATURE

1851 Verany, Loligopsis zygaena (p. 125, Plate 40, Fig. C).

1879 Tryon, Loligopsis zygaena (p. 164, Plate 69, Fig. 257).

1884 Rochebrune, Zygaenopsis zygaena (p. 20).

1900 Pfeffer, Zygaenopsis zygaena (p. 193, part.).

1909 Hoyle, Zygocranchia zygaena (p. 276).

1912 Pfeffer, Pyrgopsis zygaena (p, 660).

1916 Naef, Pyrgopsis zygaena (p. 15).

1921 Naef, Leachia cyclura (p. 538).

389

A diagnosis cannot be given since there are no characters except transient
characters of development. The single known specimen was caught by
Krohn near Messina and described by Verany (1851 ).

The animal (Figure 202) shows the typical habitus of young Cranchiidae
but no generic characters, except the form of the fins.* The widening of the
fin toward the transverse, posterior margin with 3 corners resembles that
410 of the younger Pyrgopsis (Figure 200c, d) and confirms the determination
by Pfeffer. On the other hand, also in the youngest Galiteuthis (the only
genus of Cranchiidae which undoubtedly occurs in the Mediterranean) the fins
are not as pointed posteriorly as in the older forms (Figure 199c). One
could assume, therefore, that the fins of specimens of Galiteuthis only
slightly smaller than Pyrgopsis zygaena would have the same form.

In this case, Pfeffer 's argument would become very weak. The larva other-
wise resembles a young Galiteuthis: the arms are of almost equal
length (Verany states that the first pair is slightly shorter than the others),
while in Leachia (Figure 200) the LV arms soon become longer than the
others; the tentacles are too strong for Leachia but resemble those of
young Galiteuthis (Figure 197). The head pillar is also relatively short,
as in Galiteuthis, but these can be considered as juvenile characters, and
are not significant systematically. The cartilaginous ridges on the mantle
may have been overlooked by Verany. However, they are certainly missing
in the youngest stages of Leachia, and their absence would not prove that
this is not a Leachia. A re-examination may make a definite identification
possible and more intensive collection of plankton in Messina could provide
further material on the development of the Cranchiidae. Further studies on
the development of Leachia could also give more information.

FIGURE 202. "Pyrgopsis zygaena," after Verany (1851,
Plate 40, Figure 2), natural size. This is a typical stage of
Cranchiidae, except perhaps for the form of the fins, especially
their transverse posterior margin, a specific character of Pyr-
gopsis (or Leachia). Otherwise, the animal could well
belong to Galiteuthis (cf. Figures 197 and 199). The carti-
laginous ridges on the mantle (Figure 201) may have been over-
looked by Verany; they could also still be absent in such a
young stage.

Pyrgopsis r hy n c h o p h o r u s Rochebrune (1884, p. 23, Plate 2,
Figures 2—6) probably belongs to the same species (cf. Pfeffer, p. 660).

The animal is delicate, gelatinous and transparent. Its mantle is slender,
spindle-shaped, 22 mm long. The fins are slightly triangular, with a straight
anterior margin from the blunt lateral corner. The eyes are small, with
long stalks. The arm pillar is stalklike. The first pair of arms is shorter
than the others. The tentacles are as long as the mantle and show the typical
arrangement of suckers; they are smallest and most loosely distributed in
the middle of the stalk. The funnel is very large and projects anteriorly
beyond the eye stalks.

* This is the only objection against Issel's view (1920, p. 15) that the larva belongs to Galiteuthis,
which certainly occurs in Messina. The fins of a stage of Galiteuthis of similar size already show
the typical form shown in Figure 197, as Issel himself has shown (Figures 17,19) (cf. Figure 199d).

390

411 Chapter 32

FAMILY OMMATOSTREPHIDAE
Gill, 1871

a. DIAGNOSIS

Funnel bond broadly triangular, with a longitudinal groove which becomes
deeper posteriorly and bears strong tubercles, and a shallower transverse
groove along the posterior margin, which fit exactly into the corresponding
formations of the mantle bond so that they are firmly connected. Funnel
pit with a sharp edge which may cover the strong anterior funnel adductors
more or less completely; they are otherwise exposed and widely separated.
Gladius consisting mainly of a rhachis with 3 longitudinal ridges and with
rudimentary lateral plates; posterior part of gladius consisting of a small,
lanceolate cone flag with a conical cone. Fins firmly fused in the middle
and forming together a transverse rhombus. Neck with well developed,
longitudinal and transverse folds. Lid margin with pointed anterior sinus.
Arms and tentacles without hooks. Branchial pockets perforated posteriorly
during postembryonal development and then remaining wide open. Tentacle
stalks fusing during embryonic development into a snoutlike formation, but
later dividing again, and forming the "anlagen" of the clubs. Hectocotyii-
zation consisting of an enlargement of the arm suckers and a characteristic
modification of one or both ventral arms.

Together with the Thysanoteuthidae and other groups, the Ommato-
strephidae are a branch of the Oegopsida (p. 236) and form a well defined
group which resembles the type of Oegopsida in habitus and many other
characters, but differ from it in a number of marked specializations (funnel
cartilage, pockets of gill base, gladius, funnel pit) and in the reduction of
some structures (hooks). Particularly characteristic for the early stages
is the "snout," which results from the fusion of the tentacle stalks
(R h y n c h o t e u t h i s" stages).

The Ommatostrephidae are the only Oegopsida which occur in large num-
bers in the Mediterranean during a large part of the year (but only in some
age groups). They are also common in other seas and predominant in

412 littoral and surface waters. Sthenoteuthis bartrami is widely distri-
buted in warmer seas. Sthenoteuthis is the most powerful and active
type of the family and among the Oegopsida in general (cf. pp. 164, 225).

The muscular mantle is extremely strong and solid, a valuable commercial
property. On the other hand, the shell is reduced to a delicate, pliable
backbone.

391

b. TYPICAL STRUCTURE OF THE ADULT
OMMATOSTREPHIDAE

Typical Ommatostrephidae have the typical habitus of Oegopsida
(Figures 114,228), but the gladius is markedly atypical (Figure 231 ). It is
completely surrounded by the muscular mantle, except at the posterior end
("cone flag"). As in the Loliginidae, the gladius is found on the inner side
of the mantle sac during dissection. The posterior part of the gladius is not
situated superficially under the skin, but is covered by the muscular fins,
which are firmly connected with the muscular mantle. The rhachis occupies
the greater part of the gladius; the rudimentary lateral plates (flag) cannot
be delimited with certainty (cf. Pfeffer, 1912, pp. 376— 377). The lanceolate
"cone flag" is attached laterally and passes to the posterior part of the
rhachis posteriorly into the always distinct cone. The conical cone shows a
fine longitudinal striation toward the apex and is surrounded by layers which
correspond morphologically to a sheath (pp. 166, 22 6) and continue into a
small, blunt rostrum. The rhachis is stalklike on the cone flag, about as
thick as wide. It widens anteriorly, rapidly at first, then more gradually,
forming a thin plate which ends anteriorly in a pointed angle and is
supported by a median and 2 lateral ridges which originate on the stalklike
posterior part. These ridges become suddenly narrower near the anterior
end and disappear without reaching the margin.

The fins are very strong. They are fused firmly in the middle, forming
a very efficient structure. This fusion reaches to near the anterior
attachment. This plate has the typical "rhomboidal-heart-shaped" form.

The anterior margin forms a simple curve from the "earlobe" to the blunt
lateral corners (Figure 216 on p. 431). The posterior margin forms a
slightly more complicated curve: the line connecting the lateral corner
with the apex is more sinuate laterally and less so near the apex; the
margin in between is concave, forming a distinct apex.

The mantle sac is cylindrical in the anterior half, spindle-shaped in the
posterior half and forms an apex which ends with the fins and is firmly
connected with them. This terminal region requires a closer analysis. The
presence of a fleshy apex situated behind the cone which does not belong to
413 the muscular mantle is characteristic for the Oegopsida (p. 22 6). This
structure is present here also; however, it is not attached on the outside
of the mantle sac but forms its posterior part. It also connects the fins,
which are not contiguous in the posterior part of the body but are attached
laterally to this apex. These relationships are illustrated in Figure 203.

This median section shows that the fleshy apex is a separate structure
which is situated on and behind the cone and is apparently a continuation
of the mantle sac. The mantle sac ends primarily (p. 153) in the cone, and the
muscular mantle originates at its margin. As the muscular mantle is
attached to the cone flag further on but does not surround it, the posterior
end of the gladius would be dorsally exposed if the fins were not fused
above it. This took place directly in the anterior part of the section, and
further posteriorly by the fin cartilages, which are fused into a single
structure and reach the fleshy apex posteriorly. The fins are attached
laterally to the fleshy apex in the posterior part of the body, so that a
structure is formed which is very complex internally, but compact on the

392

outside, attached by strong skin musculature and a tough skin. The pocket
of the fin base, i. e. the articulation between fins, gladius and mantle, is
reduced, as it would not fulfill the requirements of a strong, swimmer. The
mantle margin shows 3 small but distinct corners and a shallow funnel
incision between the two ventral corners; it is surrounded by a narrow,
membranous margin (Figure 243), as in other muscular Oegopsida.

FIGURE 203. Sagittal section through the posterior end of the body oflllex coindeti
(enlarged, diagrammatic). The dark line is the rhachis (Ra) with the conical terminal
part (Co); the muscular mantle (Mm) is attached ventrally to the margin of the cone
(at Ma). Behind the cone (Co) is the posterior apex (Sp), anteriorly, the fin cartilage (Kn)
and the fin (FI), which here reaches the middle.

Co - coelom; Mh - mantle cavity; Mu - subcutaneous musculature; Ss - shell sac;

Ep — epidermis; Fu and Fo — lower and upper muscular layer of the fin.

The neck folds are typical (cf. Figure 230 with Figure 53) and reach the
margin of the funnel pit with a sharp process. The posterior end of the
funnel margin forms a sharp corner. The anterior transverse neck fold
originates on it, passes around the head, forms a blunt posterior corner in
the middle or the dorsal side and then extends posteriorly. The posterior
transverse fold occupies only the lateral zones; at any rate, its lateral parts
are not united dorsally (Figure 205). The longitudinal folds are situated
414 between the two transverse folds, one dorsal, one lateral and two ventral
on each side. The third longitudinal fold bears a flat, oval, indistinct ol-
factory papilla (p. 13 6) in the typical position, posteriorly at the base of the
fold. The fourth longitudinal fold separates the neck region from the funnel
pit but it is therefore more transverse than longitudinal. The funnel pit is
very deep (Figure 204) and the funnel can be more or less completely
situated inside it. The funnel is attached by the always covered inner
adductors and the strong outer adductors. These are very characteristic
for the family and are always visible if the funnel pit is not very deeply or
strongly contracted (Figure 20 622). These muscles form 2 thick cords
which connect the lateral parts of the funnel with the funnel pit. This
attachment of the funnel and its deep location must be connected with the
motoric power of the animal. They concentrate the expelled water in the
axis of movement of the animal. On the other hand, the structure of these
organs and the powerful muscular mantle permit strong thrusts (p.412).

The neck folds, particularly the large anterior transverse fold, and the
margin of the funnel pit make the head an almost cylindrical continuation
of the mantle tube which is adapted to the movement. The lid opening has

393

a more or less straight, transverse posterior margin with a deep, narrow,
anterior sinus. The posterior margin usually covers half of the markedly
convex lens in preserved specimens. Windows are present on the ventral
and dorsal side of the head (Figure 204) but their significance is not clear
since the translucent eyeball bears no luminous organs (p. 159). The
lateral pairs of arms are of about equal length and much longer than the
median arms, which are also of similar length. Large, muscular swimming
margins are present along the whole lateral arms. On the second pair the
margins occupy the ventral outer edge of the arm, they are widest near
the middle, become gradually narrower toward the proximal and distal end,
and pass into the membrane between the second and third pair of arms.

The swimming margins of the third arms are situated in the middle of the
rounded outer side, they are widest behind the middle where they form a
corner and end free at the base between the outer membranes between the
arms of the second and fourth pair. Such connecting membranes are also
indicated between the other arms. The dorsal arms have no swimming
margins but have prominent outer edges. The ventral arms have typical
outer margins.

The inner side of the arms bears 2 rows of toothed suckers and typical
protective margins the supports of which correspond in number and position
to the suckers of the adjacent row. The protective margins of the fourth
arms are least developed; those of the first, second and third pairs are
successively larger. On the lateral pairs the ventral margin is usually
much wider than the dorsal, to the extreme condition shown in Figure 249.
The protective margins reach the buccal funnel, at least as ridges (Plate VI,
Figure 3). The suckers on the lateral arms are larger than on the dorsal
arms; the suckers are smallest on the ventral arms and the proximal

415 suckers are always different from the distal suckers (p. 434). The dentition
becomes indistinct toward the base of the arm.

The tentacles are moderately long, with a rather short stalk of about
half of their total length. The cross section of the tentacle is about oval
at the base. The basal ligament disappears rapidly, but a typical stalk
margin is present on the outer side; the inner side is slightly flattened, with
two edges, of which the ventral is more marked. These edges pass into
the protective margins further distally. The suckers which are still present
in the adult begin farther along the margins (Figure 242). It must be
assumed therefore that the proximal suckers of the stalk part were lost.

The distal part always bears 2 rows of small suckers which rapidly pass
into a quadriserial arrangement in which the suckers of the median rows
are enlarged and pass into the hand part without forming a distinct
boundary to the carpal part. A distinct carpal part is apparently absent and
the hand part begins where the biserial pattern ends. The hand part bears
2 median rows of larger suckers and 2 marginal rows of smaller suckers.
The suckers of the dorsal rows are always slightly larger than the corres-
ponding suckers of the ventral rows. The largest dorsal suckers always
have a characteristic dentition; the others resemble those of the arms,
according to their proximal or distal position. The differences in form and

416 dentition are constant and very characteristic for the particular area and
species; the differentiation may be very marked; cf. p. 431.

394

(415)

FIGURE 204. Anterior part of body of O m m a t o s t r e p h e s sagittatus. Note mantle margin,
neck folds, funnel pit (which is very deep), window (Fe),lid margin, iris, lenses of eyes (Li); the
membranes on the arms, swimming margins (Ss), protective margins (Sh) and suckers. On the ten-
tacles: stalk margin (Sm), swimming margin (Sw), protective margin (Sc).

St - support of protective margin; Fo - foveola of the funnel pit; x - marginal corner of funnel pit;
Ta — funnel adductor; L1.3 — longitudinal neck folds; Qy.Qh — anterior and posterior transverse
folds of neck; Md,Me — dorsal and ventral corners of mantle margin.

Between the proximal dorsal marginal suckers of the hand part are
situated 2—5 alternating knobs which correspond to the suckers on the
opposite club, and form the only developed part of the adhesive apparatus
(Figure 242). The distal part differs from the hand part in that its suckers
become suddenly smaller, especially on the right club. The ventral of the
4 rows of suckers which become gradually smaller, contains the largest
suckers, but the difference is not very marked.

The protective margins have as many supports as suckers, i. e. each
marginal sucker corresponds to 2 supports in the quadriserial hand part.
They are not cirruslike, but wide, shovel-like and form rounded projections
on the margin, with indentations between them (Figure 242). This is not
recognizable on the distal part, which bears a typical swimming margin
along the dorsal edge and passes proximally to the outer side of the hand
part. The proximal end of this margin marks the beginning of the stalk
margin; this is not a direct continuation of the swimming margin, but the
stalk margin is displaced slightly ventrally.

395

There are beginning connections between the protective margins of the
arms and the buccal funnel; the protective margins end proximally in simple
edges or ridges and the 8 attachments of the buccal funnel are connected with
them, at least in part (Figure 113). The 6 buccal pockets are arranged
between these attachments and show a marked variation in the family. The
pockets are large and* are separated only by thin membranes deep below
the attachments (Figure 113b). Even the ventral pockets are contiguous
deep inside, although their openings are widely separated, so that not only
the superficial part of the buccal cone but also its inner mass is washed
by water, and the inner mass is separated from the arms. The buccal
supports and corners are not very strong and without suckers.

The funnel is not very large, but strong and firmly anchored. This is
evident not only from the formation of the outer adductors and the deep
funnel pit, but also from the neck bond and especially the funnel bond.

This excludes a rhythmic retraction of the anterior part of the body into
the mantle opening, which has to be assumed for most other Oegopsida
during swimming (p. 395). This increases the stability of the form of the
body and the control of movement. The improved efficiency of mantle and
funnel apparatus more than compensates for the lost range for contraction
and dilatation of the mantle cavity.

FIGURE 205. Neck region of a large, almost mature female of
Ommatostrephes sagittatus. 0.5 x. The body has been
removed from the mantle and shell in the M- shaped posterior zone
which contains the stellate ganglia (St) and the nerves of the fins
extending from them. A narrow slit corresponding to the dorsal mantle
cavity separates this posterior zone from the neck bond (Nk). This
projects beyond the posterior margin of the funnel pockets (Tt) which
are laterally attached to it. The funnel bond (Tk) projects beyond the
posterior (and lateral) margin of the funnel pockets, and the funnel
retractors (Tr) are inserted on it. Anterior to the funnel pockets is a
necklike constriction, and then follows the zone of neck folds with the
posterior transverse fold (Hq),the anterior transverse fold (Vq),the
dorsal longitudinal fold (Dl) and the lateral longitudinal fold (LI).

The figure shows the typical structure of this region in the Teuthoidea
in general (cf. Figure 85).

According to Pfeffer (p. 370), the funnel bond resembles that of the
Onychoteuthidae, i. e. it is of the typical form of the Oegopsida. It has to
be assumed therefore, that the funnel bond has become wider and triangular
posteriorly (Figure on p. 419), while thepockethas become deeper and
constricted by two projecting tubercles in the posterior part; the wide part
of the pocket remains as a shallow transverse groove behind the tubercles.
The mantle bond widens correspondingly posteriorly to a large process
which then narrows suddenly between deep pockets corresponding to the
tubercles, and is connected with a transverse, edgelike posteriorly sloping
elevation. The firmness of this connection is still recognizable in pre-
served specimens. Considerable force has to be used to separate the parts
in large, fresh specimens, and it is doubtful whether the animal is able to do
it. On the other hand, one may ask why the articulation between funnel and

396

mantle persists if it does not permit movement? This can only be understood
historically: the firm connection between funnel and mantle developed from
a typical preceding apparatus of closure which had the character of a gliding
sledge. A similar condition exists in the Chiroteuthidae (p. 377) and it is
no accident that Grimalditeuthis has replaced the cartilage with direct
fusion. Such a concrescence of funnel and mantle cartilages is present
also in the Ommatostrephidae (S y m p 1 e c t o t e u t h i s), without obliterating
their complex formation. At any rate, these phenomena prove that the
possibility to loosen the connection between funnel and mantle is not
important any more.

(418)

FIGURE 206. Young O m m a t os t r e p h e s sagittatus,

3 X, with opened mantle. The animal shows some characters
of the genus, (position of the appendage of the branchial
heart (2), the crescent- shaped fold of the funnel pit (21),
the long tentacle club). Most of the characters, however,
are those of the family. Note form and size of fins;
posterior end of body, where the apex of the cone is
visible at 43; the strong mantle; mantle bonds (35,36) and
funnel bonds (34); inner margin of funnel; outer funnel
adductors (22); funnel opening with valve (20); neck folds
and funnel pit (30); form of the head, which is more rounded
than in the adult; eyelid (24), lens of eye (25) and iris (29);
arm apparatus with indistinct protective margins and the
incompletely differentiated clubs. Note particularly the
organs of the mantle cavity; the gill ligaments (44) which
have grown toward the mantle margin, the lamellae are more
numerous, and the base formed as described on p. 426.

Also shown are entrance and exit of the main vessels, and
the posterior opening of the pocket of the gill base. The
visceral complex is longer than shown in Figure 215, but the
narrow form of the posterior part is caused by the fact that
stomach, caecum and coelom are empty and this is not
constant (cf. Figure 243). Note the renal openings (11),
hind intestine (12), anal papilla, ink sac (46), vena cava (17),
the characteristic pockets outside the renal pores, inside
of the funnel retractors (14) ("lateral pockets"). The
following structures are visible: venous branches (7),
branchial hearts (3) with appendages (2), lateral pallial
veins (48), posterior pallial veins and arteries (41) (fin
vessels). The fin nerves (42) are visible near the narrow
posterior part of the body. Neck folds: anterior transverse
fold (23), fourth longitudinal fold (31), third longitudinal
fold (32), second longitudinal fold (33).

26 - membrane between the 4th and 3rd arms: 27 - outer
median edge of the fourth arm; 28 — sinus of eyelid;

19 - longitudinal groove; 18 - transverse groove; 34 -
median tubercle of funnel bond; 35 - longitudinal ridge;

36 — transverse ridge of mantle bond; 45 — branchial gland;
15 - membranous connection of oblique posterior part of
funnel retractor with body; 10, 9 - venous appendages
before branching of vena cava (cf. Figure 109a); 5 - venous
appendage at base of posterior pallial evine; 6 — heart;

1 - branchial artery; 8 - lateral pallial vien (continuation
at 4); 37 — posterior aorta; 38 — median pallial artery;

39 - median pallial septum; 40 — posterior pallial artery
(stem).

397

The neck cartilage is also not adapted for a gliding movement. Such a
movement is definitely excluded in the closely related Thysanoteuthidae
(Jatta, 1896, Plate 9, Figure 5) which show a clearly more advanced stage of
the same basic form.

(419)

FIGURE 207. Organs of mantle cavity of Ommatostrephes sagittatus. 0.5 x. a) juvenile, b) almost
mature female, a) Note the typical characters of Ommatostrephidae: posterior opening of the pockets of the
gill base, lateral pockets (x),wide renal pores (Np), funnel bonds and mantle bonds. The complex of organs
behind the gills is here filled (in contrast to Figure 206) but only attached in a narrow stripe along the gladius.
The median pallial septum is preserved on the left (median pallial artery visible on its free margin, and
2 accessory pallial arteries join it further on). The other parts are as in Figure 206. Note also the "anlagen"
of the narrow nidamental glands (Nd); their anterior end is attached anteriorly by a suspensory ligament (Bd).
The renal pores are contracted to papillae in a). - b) Note the typical form of the half- mature nidamental
glands with their anteriorly wide open slit which opens upward and appears darker. Also visible are the sus-
pensory ligament of the nidamental glands and the wide, round renal openings. Near the nidamental glands
are the oviduct glands (Ei),the opening of which projects into the open pocket of the gill base. The posterior
aorta is situated between the nidamental glands; the posterior pallial veins are visible farther posteriorly.

The musculus rectus abdominis (dotted) is situated between the anterior part of the nidamental glands. For
explanations compare also p. 183.

The typical structure of the mantle cavity is shown in Figure 206 and
243 (cf. also Plate V, Figure 1 ).

The structure of funnel and mantle bonds is shown in Figure 210; see
also p.42 6 on the posterior opening of the pockets of the gill base. Note

398

418 particularly the following: 1. the renal openings, usually in the form of wide
round pores connecting the renal sacs with the mantle cavity, but which can
contract to low papillae by the action of their circular muscles. 2. the
"lateral pockets" which are characteristic depressions of the skin near the
body and inside of the funnel retractors, near the posterior part of the gills
(Figure 207) and which extend dorsally toward the mantle. The significance
of these pockets can perhaps be understood by the conditions in Thysano-
teuthis (p.470).* The other characters are typical for the Teuthoidea or
Oegopsida. On the funnel retractors, see p. 426.

The jaws differ distinctly from those of other large Oegopsida (Plate XVIl),
but resemble those of the Loliginidae. The biting process of the upper jaw
is longer and more pointed than in the Loliginidae, and behind it is a distinct
"lateral tooth," a strengthened part of the biting edge formed at the junction

419 of the inner and outer plates. This tooth is always dark (Plate XVII,

Figure 8: a. Figure 10: c) and is separated by a light stripe from the still
darker biting process (Plate XVII, Figure 10: b; cf. Figures 8— ll). This
formation is especially large inOmmatostrephes; in the Loliginidae
(plate XVII, Figure 3 (a)) and some other Oegopsida (Figure 6), it is present
in a less developed form, but it is indicated always in almost all the jaws

of the Dibranchiata. It is always distinct in the Ommatostrephidae and
also in the Thysanoteuthidae (Plate XVII, Figure 12). The lower jaw also
shows a similar structure in these two families (Plate XVII, Figure 10: d);
and also an important negative character; the inner plate lacks the
strengthening ridge observed in other Oegopsida (Plate XVII, Figure 6: e).

420 The radula of the Ommatostrephidae is better known than in other
families. Targioni-Tozzetti (1863, Plate 8) and Girard (1890, Figure 3;
cf. Plate XIV) have given good illustrations. However, it shows few
specific characters, except perhaps the presence of large lateral teeth in
the 3 median rows.

Sexual dimorphism is present in the mantle cavity: the nidamental
glands of the female develop in their typical position (Figure 207), while
the oviduct glands are displaced ventrally and posteriorly, so that they are
always present in the mantle situs in the later stages (b) (Ei). Their
opening often shows their old position near the pocket of the gill base,
into which the glands grow from behind (p,42 6; but see also Figure 221).

The male genital process and appendix (cf. Chun, 1910, p. 35, Figure 12)
develop similarly on the left side and the opening of the genital pocket is
situated so far posteriorly that it is visible in the opening of the gill base;
the development of the male genital process takes place at a position
corresponding to that of the female organ (cf. Figure 155). Sexual
dimorphism is expressed externally in the following characters, Male:
l) enlargement of the suckers of the lateral arms; 2) general strengthening
of arm apparatus and head, except for the tentacles; 3) hectocotylization of
one or both ventral arms by modifications of the protective margins,
suckers and their supports, the general type of which has not been deter-
mined. Female: l) extension and widening of the body by the enlargement
of the genital organs; 2) particularly strong development of the tentacles
(p. 126, Figures 216, 218 on pp. 431, 433 ).

These pockets have perhaps a more general occurrence. They are also present in the Onychoteuthidae
(Figure 1573),and in at least similar form even in the Octopoda (Figures 427—429).

399

c. TYPICAL POSTEMBRYONIC DEVELOPMENT

The Ommatostrephidae have very peculiar young stages, for which Chun
(1903) proposed the nanae R h y n c h ot e u t h i s. This term is preoccupied
for jaws of fossil Tetrabranchiata; d'Orbigny, 1845, cf. 1855), and Pfeffer
(1908) replaced it with Rhynchoteuthion. Both are fictitious generic
names for uncertainly determined objects. We speak only of "Rhyncho-
teuthis" stages of Ommatostrephidae. The older of these stages can be
determined more or less definitely. Such "larvae" apparently occur in all
genera; their embryogenesis can be described in detail (Volume II,

Plates IX— XII). Some freshy hatched specimens are shown in color on
Plate XIX (Figures 1 and 2); another is shown in slightly reconstructed
form in Figure 208, which also illustrates some typical states of contraction.

FIGURE 208. Freshly hatched larva of Ommatostrephidae
20 X.

c - contracted, head and funnel deeply retracted into the
mantle sac and visible through it; snout markedly shortened;
b - anterior part of head with extended "snout." a — normal
swimming position; snout of medium length, mantle maxi-
mally extended during respiration. Note the form of fins,
mantle sac, head and funnel, the attachment of the funnel
retractors. The only arms present, i.e. the dorsal two pairs,
have only one sucker; the fused stalks of the tentacles
("snout") have clubs with 4 suckers (3 suckers visible).

FIGURE 209. Youngest stage of
Rhynchoteuthis* from the deep
plankton of Naples (13 April 1916).

15 X. Dorsal and ventral aspects re-
constructed after drawings of the live
animal; the lateral aspect shows the
deformation caused by preservation.
Gills, funnel pockets, funnel and funnel
retractors are exposed by contraction of
the mantle. The ventral view shows the
translucent inner margin of the funnel,
the anal papilla, hind intestine, ink sac,
and gills in their natural position.

The animal was examined while still alive. It had a glasslike transparency, except in a few parts. The
chromatophores appeared black in transmitted light; in incident light, chromatophores and eyeballs were
brownish red to scarlet, partly orange, the ink sac black and the liver light pink. The argentea was thus
still rudimentary. There were 2 light yellowish brown chromatophores on the posterior part of the head.
Fat droplets were found in the stomach. The heart, was situated transversely and of typical juvenile form
(Naef, 1910,p. 325); it pulsated excitedly and irregularly usually alternating with the branchial hearts.
The gills had 2 lamellae on each side. The clubs of the tentacles were extended as in Figures 208b.

400

The habitus of these larvae is that of very young Oegopsida, except for
the "snout" (cf. p. 233). The mantle sac is short and blunt posteriorly,
with a small, spoon-shaped cone which continues dorsally in a short lanceola
but ends anteriorly in a narrow rhachis. The muscular mantle is at first
attached also here to the margin of the gladius, but it later rapidly and
completely surrounds the anterior part of the gladius (p. 412). The mantle
and funnel apparatus of these animals are very movable (Figure 209).

421 The fins have the form of narrow wings, markedly projecting laterally
and with a rounded, widened end. They beat rapidly. The fins are always
much shorter in preserved specimens (measured laterally) than in the
live animal. They are attached posteriorly to the gladius, but are still
narrowly separated in the middle. The mantle margin is still simple,
without corners.

The head is typically larval in form, without special characters. The eye
region is very narrow, the cheeks are rather wide. The lid margin is
rounded, without a sinus. The arm apparatus is typical; only the 2 dorsal
pairs are developed, they are short, whiplike in the live animal, still shorter
in preserved specimens, each with a single sucker in about the middle.

There is still no trace of the LV arms; the fourth pair is represented by
small, papilliform "anlagen."

The tentacles form a "snout." Their stalks are fused along the entire
inner surface, but the rudimentary clubs diverge, exposing a small area
with suckers. This area is very short, slightly spoonlike (this is indistinct

422 in preparations ~ Figure 209), and bears 4 suckers which are arranged not
quite symmetrically, but so that the two clubs appear congruent; later
development shows that the 4 suckers form the beginning of a dorsal and a
ventral row. There is no trace of the buccal funnel (Figure 88). The
funnel apparatus still has no funnel bonds, and the gills are quite unde-
veloped, in the form of papillae with indications of a lamella (cf. Figures 209

. and 88).

The youngest planktonic larvae show little progress. These forms are
among the more frequent pelagic stages of Oegopsida, but they are not
common. My large collection has been built up during several decades.

The larva shown in Figure 209 does not differ much from the previous
stage, but differs so markedly in its degree of development, that they
cannot be considered as belonging to the same species.

The eyes are larger, and the head is distinctly wider in the eye region
than in the cheek region. This is normal, but some older larvae have
relatively less prominent eyes (Figure 211). The fins are not as narrow.
The "anlagen" of the gills are displaced markedly anteriorly and have at
least one lamella on each side; there is another "anlage" of a lamella
distally. The funnel bond is still absent. The arrangement of the chroma -
tophores on the head is typical; their arrangement on the mantle apparently
varies widely, even in the same batch, so that no rule can be established.
Hind intestine and anus are situated typically median.

Another larva, resembling the preceding but larger and further developed,
is shown in Figure 210.

This stage shows marked progress compared with the preceding stage,
l) there are distinct funnel bonds, and their rounded triangular form begins
to resemble the definitive form; 2) the third pair of arms, represented by

401

small tubercles in the preceding stage, is now distinctly longer than the
fourth pair and bears a larger sucker; 3) a number of smaller suckers have
been added to the two already present on the two other pairs of arms, and
further "anlagen” of suckers are recognizable distally; 4) the mantle organs
423 have developed further, particularly the gills, which now have several
lamellae on each side.

(422)

FIGURE 210. Rhynchoteuthis
stage from the deep plankton of
Naples; caught on 12 February 1911,
10 X, slightly deformed, but it could
be easily reconstructed according
to Figure 208. Note the develop-
ment of the suckers on the arms, of
the third pair of arms, of the funnel
bonds, and of the olfactory tubercle.
Other external characters as in the
earlier stages.

FIGURE 211. Two stages of Rhyn-
choteuthis. 4.8 X. The upper
specimen was caught on 23 August
1905 at a depth of 100 m from the
deep plankton of Naples, the lower
specimen is from the Messina plank-
ton. Both show characteristic de-
formations. The lower specimen
is strongly contracted: the right
eye of the upper specimen pro-
jects from the orbit and the eyelid
is contracted behind it. Note the
development of fins and funnel
(adductors).

FIGURE 212. Rhyncho-
teuthis stage from the
Naples plankton, 29 May
1913. 5x. Note the parti-
cular the funnel- mantle and
neck-mantle bonds, and the
distribution of the chroma-
tophores.

The larva illustrated in Figure 211 resembles the preceding but is
further developed. The third pair of arms is almost as long as the second
arms and already bears numerous suckers in 2 rows, like the dorsal pairs
of arms; a larger sucker is now present on the fourth pair. The markedly
different appearance of these larvae is due almost entirely to contraction.
The fins of the upper larva are slightly more advanced than those of the
lower specimen: they are relatively longer and have begun to grow along
the mantle. On the other hand, the fin bases are contiguous posteriorly and
cover the fleshy apex, which is already present as an "anlage." This is

402

more distinct in the lower larva. The fins diverge from the median line
anteriorly. The mantle margin shows the 3 weak typical corners. The
inner adductors of the funnel are bandlike and situated nearer to the median
plane, the wider, less distinct outer adductors are
posterior and external to the inner. (Note the 4 inter-
ruptions of the boundary of the funnel in the upper
right figure.) The head, olfactory tubercle and eyes
are typically larval. Figure 212 shows a still older
stage with about the same morphology. Only the
outer adductors of the funnel are visible, and more
distinct. The ventral arms bear a larger number of
suckers. The folded-over mantle margin shows the
cartilages of mantle and neck. The distribution of
chromatophores on the head resembles that of the
previous stages, as far as they are visible in pre-
served specimens. Already in this stage, as in
Figure 213, the chromatophores extend to the inner
side of the mantle margin, where they form a single
row.

Figure 213 shows a larva of about the same age but
in the typical protective contraction, as often happens
during preservation. The arm apparatus is as
described above, the olfactory tubercles are already
displaced slightly posteriorly and outward. The funnel
bond has now reached the typical form of Ommato-
strephidae (p.417). The mantle cavity shows the
normal relationships of older larvae: the gills,
folded over by contraction, have a greater number of
lamellae, the general condition resembles that shown
in Figure 215, and the filled stomach occupies the
posterior part of the visceral complex.

The following stage closely resembles the above
specimen but shows some details better. Distinct
swimming margins are now present on the second and
especially the third pair of arms. The formula is
still: 2, 3, 1, 4 (the view of the mouth area shows the
second pair too shortened, compared with the third pair). The tentacles are
still fused their whole length, but their thickened bases is now connected
only by an extended membrane which tears in the next stage (Figure 215).

The "anlage” of the buccal funnel is now recognizable between arms and
mouth cone. It consists of the 5 upper buccal points (or "buccal arms")
connected by narrow ridges. The buccal points are papillae at this stage,
shrunken by preservation, but probably movable in the live animal. True
attachments are still not developed, but an attachment is apparently
beginning to develop at an abnormal point, between the second and third
arms. (This is perhaps the rudiment of another buccal point?) The ventral
points are still absent, but the base of the snout would cover them in any case
in this view, since they develop ventrally of the base of the tentacles and the
connection with the other parts of the buccal funnel can only take place after
the bases of the tentacles have become separated (Figures 235 and 245;

FIGURE 213. Rhyncho-
teuthis stage from the
surface plankton near
Capri, caught on 15 June
1914. 8x. The ventral
half of the mantle is shown
transparent to show the
form of head, funnel and
mantle complex. Note
particularly the anal pa-
pilla, gills, and branchial
hearts and between them
the venous appendages and
stomach. Also shown are
funnel cartilage, olfactory
tubercle, eyes and arm
apparatus. Typical pro-
tective contraction.

403

these relationships are shown better and more exactly on Plate VI, Figure l).
The mouth cone is typical. There is an outer and an inner lip; the apex
of the lower jaw is visible inside the mouth. Protective membranes are
indicated between the bases of the upper 3 pairs of arms; also the membrane
425 between the third and fourth arms has the form typical for young larvae of
Oegopsida. A ridge extends in a wide outward curve from the outer edge of
the ventral arm, but does not reach the third arm at this stage. The dis-
placement of the lid opening to the dorsal side of the head is interesting.

This is not constant but the result of abnormal contraction. (The already
markedly enlarged eyes are situated normally, the lenses directed laterally.)

(426)

FIGURE 214. Older stage of Rhynchoteuthis from the plankton in Naples. (July 1914), 5 X;
below: mouth region, slightly diagrammatic, 25 x. The upper drawing shows that the lid opening
is displaced to a completely dorsal position; the third pair of arms now have true swimming
margins, the eyes are larger, and the outer funnel adductors are distinct. The mouth region shows
the development of the arms (I— IV), of the buccal funnel, and the "snout, " i.e. the tentacles.

1-4 - buccal points; Uk - lower jaw; Ht - membrane between base of tentacles (Ba); Nt -
suture between the stalks of tentacles; Sh - membranous connection from the 4th to the 3rd arm;
II — inner lip; A1 — outer lip.

404

Figure 215 shows a fully developed Rhynchoteuthis, shortly before the
transition to the state of typical Ommatostrephidae. The ventral part of the
mantle is removed to the margin to show the mantle situs. Fins and apex of
body are still larval, also the form and margin of the mantle. The head is
further developed. The eyes are much enlarged; there are distinct outer
(and inner, covered) funnel adductors, and the "anlage" of the third longi-
tudinal neck fold is already situated near the olfactory tubercle. The lid
margin shows the indication of a sinus. The arms are further developed but
their proportions are the same (2, 3, 1, 4) and the fourth pair is still very
small. The protective membrane between the third and fourth arms is now
present but still very low, and the tentacles begin to assume their normal
position by becoming separated at the base and forming a kind of arch,
below which the already distinct "anlagen" of the ventral buccal corners
become connected with the dorsal part of the buccal funnel (Figures 235
and 245). The greater part of the tentacle stalks is still fused but the suture
is recognizable, as usual in the younger stages. The end of the tentacles
corresponding to the club still bears the 4 larval suckes, the distal sucker
is enlarged.

FIGURE 215. Stage of Rhynchoteuthis at the peak of its
development from the plankton of Naples, date unknown. 29/4 x.
Note the mantle situs and the inner part of the funnel with the
bonds, hind intestine, anal papilla, ink sac, vena cava, venous
branches, branchial hearts (translucent), posterior aorta, posterior
pallial veins, median pallial septum, gills, branchial gland, gill
ligaments. Behind the gills, the passage of the lateral pallial veins
to the mantle in a wide arch which is connected with the gill base.
The funnel retractors have a thick, muscular anterior part and a
thin, tendonlike posterior part. Note the following external structures:
fins, apex of body, funnel adductors, olfactory tubercle and adjacent
neck fold, form of head with very large eyes, the arms, and the
beginning separation of the tentacles. The smaller figure shows a
lamella of the gill (2) placed vertically. The lamella is plicated
(cf. Figure 88; Plate IV, Figure 2; Plate V, Figure 1).

1 — Effferent branchial vessel; 3 — branchial gland.

Of particular interest are the mantle cavity and the inner part of the
funnel. The latter resembles that of the adult in the form of the inner
margin and in the structure of the funnel bonds, to which the mantle bonds
correspond. The organs of the mantle cavity are of the type of the Ommato-
strephidae, especially the gill base, which is connected with the lateral
42 6 pallial vein, bridges the lateral part of the mantle cavity and is attached

laterally on the mantle, forming a ligament. This condition is made distinct

405

by the widening of the mantle (cf. Figure 207), and is obviously not primary,
but the attachment of the gill base in the younger stages passes continuously
from the body to the mantle, is then gradually detached from them by the
formation of a thin connecting fold which finally tears as it has no function,
(cf. Volume II, Plate XVIl). Another character of Ommatostrephidae at this
stage are the funnel adductors, which differ markedly in the anterior and
posterior part: the posterior part is strong and muscular, the anterior thin
and tendonlike. The thin part of the adductors becomes detached from the
body, forming a thin connecting membrane (Figures 233 and 243). The other
characters are typical for the Teuthoidea, both in the structure of the super-
ficial parts and in the topography of the translucent parts. Especially the
course and branching of the vena cava and posterior aorta is typical for the
Oegopsida. The smaller figure shows that the branchial lamellae are
undulate, preparatory to the later featherlike structure (Plate IV, Figure 1;
Figure 88).

The animals become later more slender and the fins grow anteriorly along
the m'antle so that the juvenile form shown in Figure 222 develops. The fins
now form a transverse oval because of the growth of the posterior apex, and
427 this oval slowly becomes rhomboidal-heart-shaped, as characteristic for the
older stages (Plate V) and develops lateral corners, a terminal point and
"earlobes." The three corners of the mantle margin are distinct but not
very marked. Further neck folds develop (Figure 206), reaching their full
height already when the animal is less than half-grown; the funnel pit and its
margin also develop. The olfactory papilla remains in the position shown in
Figure 215, i. e. at the posterior end of the third longitudinal neck fold, on the
upper side of the fold near its attachment on the neck. The olfactory papilla
is an oval wart which no longer projects since it is partly embedded in the
swollen skin. The eyes are now lateral, and the eyelid has the typical
outline; its posterior margin is later transverse, and the posterior margin
forms a narrow sinus. The arms develop swimming margins, especially on
the second and third pairs, and protective margins develop on their inner
sides. The tentacles become separated from behind (Figure 222). Their
anterior end grows further and develops new suckers in addition to the
4 existing suckers, at first in 2, then in 4 rows (Figure 245). Finally,
swimming margins develop on the distal part of the clubs and a thin stalk
margin proximally. The "anlagen" of the genitalia appear and develop.

The ancestral form probably had luminous organs, at least on the ventral
side of the mantle, as in the recent Hyaloteuthis (Pfeffer, Plate 42,
Figure 5) and other forms, perhaps also on the ventral side of the eyeball.

The male shows changes of the ventral arms and other phenomena of
hectocotylization already when half-grown.

428 d. VARIATION OF THE TYPE OF OMMATOSTREPHIDAE

The Ommatostrephidae are divided into 2 subfamilies according to the
following characters.

A. Funnel pit without crescent-shaped or longitudinal folds. Tentacle
stalk as long as or longer than club (Figure 223). Buccal pockets
shallow, separated 1. Subfamily Illicinae Posselt, 1890

406

Genera: 111 ex and Todaropsis (see below).

B. Funnel pit with crescent- shaped and longitudinal folds at the anterior
margin (Figure 204). Tentacle stalk very short (tentacle with suckers
or with a protective margin near the base). Buccal pockets very deep,
connected with each other below the bonds, forming a peribuccal space

around the mouth cone

2. Subfamily Ommatostrephinae Posselt, 1890

Pfeffer separated another subfamily, the Sthenoteuthinae, from the Omma-
tostrephinae but does not justify this sufficiently. It only confuses the close
relationship between several genera and the sharp contrast between the
2 subfamilies of Posselt. The Illicinae are apparently the stem group of the
Ommatostrephinae.

Genera: O m m a t o s t r e p h e s D'Orb., N o t o d a r u s Pfeff., Hy a 1 o t e u -
this Gray, E u c 1 e o t e u t h i s Berry, S y m p 1 e c t o t e u t hi s Pfeff., Do si -
dicus Steenstr. and S t h e n o t e u t h i s Verr.

407

429 Chapter 33

GENUS ILLEX

Steenstrup, 1880

a. DIAGNOSIS

Distal part of tentacle club with 8 rows of very small suckers. Largest
suckers of arms with characteristically rounded teeth (Figure 219), some-
times with one large pointed tooth in the middle of the distal margin. Right
ventral arm hectocotylized as often as left ventral arm.

In addition to Illex coindeti, the genus contains the American species
Illex illecebrosus Lesueur, 1821, in which the fins are longer, about two-
fifths of the mantle length. Largest suckers on the third arm of female
about as large as those on tentacle club, or smaller. Sexual dimorphism
less marked than in I. coindeti. (Compare Verrill's drawing of a mature
male with Figure 218.) Head of mature male much narrower, arms much
more delicate. I consider I. coindeti a valid species. Otherwise, it
would have to be regarded as a variety of I. illecebrosus. Except for
the above characters, I. i 1 1 e c e b r o s u s closely resembles the European
species. The description of the genus will therefore be restricted to a
description of I. coindeti.

b. ILLEX COINDETI (Verany, 1837) Steenstrup, 1880
DIAGNOSIS

Fins slightly longer than a third of the mantle length (36—37%). Largest
suckers of arms distinctly larger than largest suckers of tentacles also in
the female. Width of head before eyes barely one -fifth of dorsal mantle
length in mature females, more than one quarter in mature males.

The main reasons for distinguishing this species from the previously
described Illex illecebrosus are the different proportions of mature
males (width of head, size of arms), the size of the suckers, and other
characters. Specimens twice as large as Figure 218 differ markedly from
the mature male of 1. illecebrosus of Verrill (l881 ).

408

430 c. LITERATURE

1837 Verany, Loligo coindetii (p. 94, Plate 4).

1839 Ferussac and D'Orbigny, Om m a t o s tr e ph es sagittatus (p. 345, Lo 1 ig o. Plates 4, 5, 7, O m m a t o -
strephes, Plate 1).

1851 Verany, Loligo sagittata female (p. 10 6, Plate 32).

1851 Verany, Loligo coindetii (p. 110, Plate 36).

1851 Verany, Loligo pillae (p. 112, Plate 36).

1853 Forbes and Stanley, Omma tostrephes sagittatus (Vol. 4, p. 231, Plate RRR).

1862 Feffreys, Om matostrephes sagittatus (Vol. 5, p.l29).

1867 Fischer, Ommatostrephes sagittatus (p. 14).

1869 Targioni-Tozzetti, Ommatostrephes sagittatus (p. 52).

1880 Stossich, Ommatostrephes sagittatus (p. 159).

1880 Steenstrup, Illex illecebrosus (p. 82).

1880 Steenstrup, Illex coinde'ti (ibid.).

1881 Verrill, Ommatostrephes illecebrosa (p. 268, Plates 23, 29, 37, 39, p. 293, Plates 18-20, p. 412).
1885 Ninni, Ommatostrephes sagittatus (p. 160).

1890 Girard, Ommatostrephes coindetii (p. 260, Fig. 3^).

1890 Girard, Ommatostrephes illecebrosa (p. 261, Fig. 3^).

1890 Posselt, Ommatostrephes coindetii (pp. 346, 348, Plate 8).

1890 Norman, Ommatostrephes illex coindetii (p. 476).

1890 Cams, Ommatostrephes illex coindetii (p. 447).

1891 Lonnberg, Ommatostrephes illex coindetii (p. 34).

1892 Girard, Ommastrephes coindetii (p. 38).

1896 Jatta, Illex coindetii (p. 71, Plates 2, 11, 12).

1896 Jatta, Todaropsis veranyi (Plate 2, Figure 6). (male!)

1900 Pfeffer, Illex illecebrosus (p. 179).

1902 Hoyle, Illex illecebrosus (p. 199).

1908 Pfeffer, Illex illecebrosus (p. 89, Figs. 96-99).

1912 Pfeffer, Illex illecebrosus (pp. 390-405, Plates 28, 29).

. 1916 Naef (Syst.), Illex coindeti (p. 15).

1921 Naef (Syst.), Illex coindeti (p. 538).

1921 Grimpe (North Sea), Illex coindeti (p. 299).

d. STRUCTURE OF THE ADULT ANIMAL

The half-grown Illex resembles the type of Ommatostrephidae in
general appearance and in the structure of most parts.

The fin is transversely rhomboidal, its length is 36—3 7% of the mantle
length but never 42% as in the American species.

Mantle margin, funnel pit and neck folds are typical. As typical for the
Illicinae (Figures 223—225), the funnel pit is rather widely open and not
completely occupied by the funnel, so that the strong outer funnel adductors
are distinctly visible. The lateral and median pairs of arms differ
markedly in length, the second pair is only slightly longer than the third
and the fourth is slightly longer than the first (2, 3; 4, l). The suckers are
much denser on the median than on the lateral arms, but those of the median
arms (especially on the fourth pair) are distinctly smaller than those on
the second and third pair. The protective margins are solid but not wide;
they form angular projections at the points of the massive supports in the
area with large suckers.

The tentacles are well developed, the clubs forming about the distal half.
The only special character of stalk and hand part is the variation of the

409

(431)

FIGURE 216. Half-grown female II lex c oind e ti.natural size. Note the outline of mantle and fins,
mantle margin, neck folds, funnel pit, funnel adductors, eyelid, length of arms, swimming and protective
margins, the tentacles with stalk and swimming margins, and the suckers. The largest mature female is
exactly twice as large, only its posterior body is 2-3 mm longer.

410

adhesion apparatus. This is often nearly not recognizable; in other cases,
there are 2, 1, or (in a large male) as many as 5 knobs. On the distal part
are 5—8 suckers in oblique transverse rows, while the greater part is

occupied by 8 regular longitudinal rows, each
with about 50 very small suckers. Of particular
interest is the polymorphism of the suckers of
which there are at least 9 categories, which could
be further subdivided as follows: .

1. Arm base. The first formed suckers
have a horny ring with smooth proximal margin
and several blunt, broadly rounded teeth at the
distal margin. Adhesive ring indistinct
(Figure 219i9).

2. Middle of arm. The suckers are large
(Plate XII, Figure 4), with low, blunt, broadly
rounded teeth all around, including a large,
pointed, mediodistal main tooth. Adhesive ring
narrow and indistinct, or absent.

3. Distal part of arm (Plate XII, Figure 7).

The suckers are claw- shaped, caused by the
markedly eccentric position of the stalk and
particularly the dentition of the horny ring,
which bears flat, rudimentary teeth and a large
"buttress" at the proximal margin; the distal
margin has usually 7 (fewer in the smallest
specimens) large, pointed teeth, the middle
tooth especially long and strong. Adhesive ring
well developed, with papillalike elevations.

4. Apex of arm. The small suckers in this
region have a small opening as usual and a
relatively wide adhesive ring with papillae.

The horny rings have teeth all around which
are larger at the distal margin, the middle tooth
particularly long and strong. These suckers
are closely related to the ontogenetic pre-
ceding stage of all categories.

5. Suckers at base of club, and on proximal part of dorsal marginal row
(near the adhesion apparatus). The suckers have a smooth proximal
margin and 5—7 long, blunt, crenelated teeth at the distal margin. Adhesive
ring with papillae.

6. The large suckers of the median rows of the tentacle have flat, broad,
low teeth, sometimes only at the distal margin, but they may be indistinct.

The stalk is almost central, the adhesive ring very weakly developed.

These suckers are cuplike, i. e. deep, with very large suction pads.

7. The accompanying suckers of the marginal rows have teeth all around
which are long and pointed, especially at the distal margin; the proximal
margin has a weak edge. These suckers are flat, bowl-shaped to
hemispherical; pads smaller and adhesive rings distinct.

8. The suckers situated distal to the large suckers of the hand part, i. e.
at the base of the distal part. They resemble those of category 7 but have

a well-developed edge and rudimentary dentition, also on the proximal margin.

FIGURE 217. Small sucker of apex
ofarmoflllex coindeti,
transparent, highly magnified,
drawn at rest from the live animal.
Compare with Figure 46 on p.l21,
and note the eccentric attachment
of the stalk (St) into the outer pit
(Gr) of the suction pad (Sk).

Hr — "horny ring" with main tooth
(Hz) and ventral tooth (Vz); Hf -
adhesive ring, contracted; Ar —
marginal ring, ready for adhesion;
Bp — basal pad.

411

9. The smaller suckers on the distal part of the club are like those on the
apex of the arms (category 4).

FIGURE 218. Mature male of II lex coindeti. 0.5x, Compare with Figure 431, and note
especially the marked sexual dimorphism of the proportions of head, arms, tentacles, suckers,
mantle sac and fins (p. 434).

The buccal funnel (p. 240, Figure 113), the funnel apparatus and the organs
of the mantle cavity are typical. The jaws do not differ from those of

412

Todaropsis, which have a similar range of variation. However, they differ
distinctly from those of the Ommatostrephinae (see Plate XVII: compare
Figures 8 and 9 with Figures 10 and ll). The radula also differs little from
that of Todaropsis, while the differences from Ommatostrephes and
especially from S t h e n o t e ut h i s are distinct (Plate XIV: compare
Figures 3—5 with 6—7). More important is the sexual dimorphism which
is distinct already in the general proportions of the adult animal. The
mantle sac of the female (Figure 216) is relatively much longer and nearly
cylindrical, that of the male (Figure 218) very short, resembling a pointed
cone. The head of the female appears as a continuation of the mantle, i. e.
it is about as wide as the moderately contracted mantle. The head of the
male forms a continuation of the conical mantle; it is much thicker, mainly
434 because of the strongly developed arm apparatus. The sexual dimorphism
is also evident in the proportions of the fins, which have the same proportion
to the mantle length in both sexes. (The width of the head before the eyes
is a fifth of the dorsal mantle length in the mature female, barely a quarter
in the male; the difference is smaller in specimens which are not full
grown.) There is a marked difference in the arm apparatus: the adult
female resembles the juvenile female in this respect,* while all the arms of
the male are much larger, especially the lateral arms, and the suckers are
markedly enlarged. On the other hand, the tentacles of the male are rela-
tively more delicate (cf. the figures). This modification is apparently only
partly associated with copulation (see Sepiolidae, Chapter 42). One ventral
arm, which is modified into a hectocotylus, transfers the spermatophores.

FIGURE 219. Hectocotylus (right ventral arm) of II 1 e X coindeti,
natural size. Left, sucker from the proximal (9) and distal (10) part,
highly magnified (especially the upper sucker) to show the different
dentition. Note the 3 regions of the arm, of which only the basal is
more or less normal, the middle region bears smaller suckers and modi-
fied protective margins, while the suckers of the distal region are absent
and their basal pad is transformed into characteristic papillae.

1 — lateral protective margin; 2 — normal support of protective margin,
i. e. reduced in the manner characteristic for II lex; 3 - modified
supports; 4 — bases of supports; 5 - modified supports of distal part;

6 - same of median row; 7 - outer margin; 8 - median swelling; 9 -
tooth of proximal sucker; 10 - adhesive ring of distal sucker; 11 -
buttress of distal sucker.

Figure 216 is drawn twice natural size from a half-grown specimen. The picture resembles the largest
female in my collection so closely (Figure 221) that only exact measurements show the difference.

413

(435)

FIGURE 220. Mantle cavity of a half-grown female of 111 ex
coindeti. 0.5x. The typical characters of Ommatostrephidae
are evident in the eye region, neck folds, funnel pit, funnel appa-
ratus, mantle bond, hind intestine, vena cava, funnel retractors,
gills, posterior opening of the pocket of the gill base, posterior
pallial veins and artery, median pallial artery, mantle, trans-
lucent cone and fins. Genitalia: the nidamental glands (Nd)
are connected in the middle and form a long, free, anterior
point; the oviduct glands (Dr) project laterally and also form a
point, but their opening is directed ventrally. The undulate
oviduct (Od) is visible behind them to the inner opening.

Ov - ovary; Am - median pallial artery.

This hectocotylization (in the strict sense) affects equally frequently the
left or the right arm of the 4th pair; 1 found it on the left side in 24 and on
the right side in 2 6 specimens. The base of the hectocotylus is normal;
the first 8 suckers resemble those of the opposite arm, but the following
suckers become suddenly smaller. The 11th to 13th sucker are already
distinctly different; the 14th or 15th is the smallest. Further on, the
suckers become slightly larger, slightly more in the inner than in the outer
row, until the 19th or 20th sucker; then follow uniform suckers to the apex.
The proximal two-thirds of the arm bear about 3 6—40 suckers; the strongly
modified apical zone bears no suckers but their bases form a zigzag row
of papillae (cf. p. 184) which are connected by a longitudinal comb. On the
median side of the arm the papillae have the form of pointed tubercles,
while all except the first papilla of the lateral side are rounded, transversely
flattened lamellae. There are about 40—45 such papillae in each row; some
435 of the last papillae may still bear small suckers; the proximal papillae form
a rudimentary support of the protective margin in the form of a small point
directed outward. The protective margins are narrow but distinct on the
part bearing suckers; they are normal in the proximal part. The supports
in the middle of the arm are large, flattened processes with rounded outline
and serrated margin. The first processes bear several small warts

414

on the side directed obliquely to the arm; the following are smooth and
longitudinally placed. All these details are strictly constant for the species.
The surface of the tentacle is often constricted in the area of the small
proximal suckers but the suckers and their bases diverge like the edges of a
spoon further distally. This part apparently holds the spermatophore mass
during the transfer. A comblike elevation of irregular form (8) is situated
between the suckers in the constricted area. The dentition of the horny rings
corresponds almost exactly to that of the other ventral arm: the proximal
suckers bear blunt teeth (9) at the distal margin, while the small distal
suckers are clawlike (Plate XII, Figure 7) and have sharp teeth. These
characters are also constant for the species and show an advanced adap-
tation of the sexes. The spermatophores are attached to the inner side of
the mantle near the gill base. They form a cluster resembling a dense,
rounded,grassy pad (Figure 221, Sp). Several copulations are apparently
performed successively, as there may be as many as 5 such pads in the
mantle cavity, sometimes in a condition which indicates prolonged invervals
between the acts.

FIGURE 221. Opened mantle cavity of a very large, mature female oflllex coindeti (half natural size).
Note the genitalia: the ovary in the upper part of the mantle sac, and free eggs situated in the coelom; further
anteriorly the filled undulate oviducts (Od), which lead to the projecting oviduct glands (Dr) and reach the ex-
terior through them; (Sp — a cluster of spermatophores near the right gland); the large nidamental glands, which
are contiguous in the middle; the median pallial artery passes between them (Am). The crossed lines are the
arteries of the glands; the veins are visible laterally. The small figure (No) shows the freely projecting part of
the right nidamental gland and its dorsally directed opening.

Ms — mantle septum ; Vp — posterior pallial vein; Np — position of renal opening , covered by the nidamental gland .

The male genital organs in the mantle cavity show in general the
structure typical for the Oegopsida. The wide opening of the genital pocket
is situated in the posterior opening of the pocket of the gill base (p. 420).

The structures projecting from it, the genital process ("penis") and appendix,
are normally visible above the gill base if they grow into the open pocket of
the gill base, but they often extend anteriorly and ventrally from the gill
base and project free in the opened mantle cavity from the point where they
project from the genital pocket. The female genital processes of 111 ex
(Figures 220 and 221 ), i. e. the projecting ends of the oviduct glands,
always occupy such a position (cf. Figure 207), so that the typical relation-
ships between the coelom exists and the pockets of the gill base are changed
(Figure 35 on p. 105). The projecting structures with their long point
(Figure 22 1) are typical, except for their position. However, their slitlike
opening is directed ventrally because of the displacement. The nidamental
glands resemble them closely; they also have a slender point and their
opening is also similar but directed dorsally. Otherwise, they have the
normal form and are contiguous in the middle in their greater part, as in
Loligo (p. 185). The mature female also shows an enormous enlargement
of these organs and of the ovary, which explains the changed form of the body
and particularly the lengthening of the mantle (p. 434).

437 The coloration of freshly killed specimens is shown in Jatta (1896,

Plate 2, Figures 1 and 6). The live animal is colored almost exactly like
Todaropsis and has a bright metallic, iridescent sheen (Plate XIX,

Figure 5).

e. POSTEMBRYONIC DEVELOPMENT

Figure 222 shows the youngest specimen which could be determined with
certainty.

FIGURE 222. Young stage oflllex coindeti from the plankton of the Bay
of Naples. x. The following characters are specific: arrangement of
chromatophores on the mantle and the structure of the tentacle club and stalk.
Otherwise, the specimen shows the typical characters of the family in such
stages, i.e. form of fins, apex of body, mantle margin, outer funnel adductors,
longitudinal and transverse neck folds, eye region, arm apparatus, swimming
margins, etc.

416

However, this form has only been determined by elimination of other
species (cf. Todaropsis): the crescent-shaped fold is absent, in contrast
to other species of Ommatostrephidae in Naples, in which this structure is
already recognizable (Figure 24 6). Similar stages of Ommatostrephes
and Sthenoteuthi s are well known (q. v. ) and differ also in the distri-
bution of the chromatophores (Sthenoteuthis) or in their general pro-
portions (Ommatostrephes, Plate V, Figure 2 ), but mainly in the structure
of the tentacles.

Of particular interest is the development of the neck folds. The anterior
transverse fold is just indicated, and the only longiutidnal folds present are
the third, with the olfactory tubercle, and the second (lateral) fold. The
eyelid already has a sinus. The ventral arms are still very delicate, the
dorsal arms relatively well developed; swimming margins are present on
the 3rd pair. Only the distal half of the tentacle bears suckers which
rapidly decrease in size until their "anlagen" are no longer recognizable
on the terminal part. The suckers or their "anlagen" are arranged proxi-
mally at first in 2 and then in 4 rows. The tentacles are very delicate and
have apparently just become separated; they are probably not functional at
this stage. The development of the distal part of the club begins in slightly
older stages, in which 8 rows of "anlagen" of suckers appear. These animals
resemble the adult 1 11 ex also in habitus, and their determination is self
evident (see also Pfeffer, Plate 29, Figures 3 and 4).

The animals already closely resemble the adult female when the mantle
is only about 6 cm long (Figure 216).

417

438 Chapter 34

GENUS TODAROPSIS

Girard, 1890

TODAROPSIS EBLANAE (Ball, 1841 ) Hoyle, 1892
a. DIAGNOSIS

Distal part of tentacle club with 4 rows of suckers. Large suckers of
lateral arms with pointed, triangular teeth. Both ventral arms hecto-
cotylized. Fins nearly half (over 42%) of dorsal mantle length.

b. LITERATURE

1841 Ball, Loligo e b 1 a n a e (p.363, Figs.l— 7).

1849 Gray, Loligo eblanae(p. 65)

1851 Verany, Loligo sagittata (male.') (Plate 31).

1853 Forbes and Stanley, Om mastrephes ebla na e (Plate SSS).

1856 Thompson, Loligo e b 1 a n a e (Vol. 4, p.270) (N. Hist. Ireland).
1869 Jeffreys, Ommastrephes eblanae (Vol. 5, p.l30).

1880 Steenstrup, Ommatostrephes eblanae (p.97).

1890 Girard, Todaropsis veranyi (p.204).

1890 Girard, Todaropsis v er a n y i (Vol.l, p.261, Vol.2, p.43 Photogr.).
1890 Norman, Ommastrephes eblanae (p.476).

1890 Posselt, Todaropsis veranyi (p. 357).

1891 Girard, Todaropsis v er a n y i (p.43).

1892 Hoyle, 1 11 e X eb la na e (p.l89).

1892 Girard, Todaropsis veranyi (p.lO).

1893 Posselt, Todaropsis eblanae (p.l).

1896 Jatta, Todaropsis v e ra ny i (p.76, Plates 2, 12).

1900 Nichols, Todaropsis ebla na e (p. 495).

1900 Pfeffer, Todaropsis eblanae (p.l79).

1903 Hoyle, Todaropsis eblanae (p.l, Plate).

1908 Pfeffer, Todaropsis ebla na e (p. 92, Figs. 100— 103).

1912 Pfeffer, Todaropsis eblanae (pp.423— 433, Plate 30).

1916 Naef, Todaropsis eblanae (p.l5).

1921 Naef, Todaropsis eblanae (p.538).

1921 Grimpe (North Sea), Todaropsis ebla na e (p. 299).

5110/2

418

c. STRUCTURE OF THE ADULT ANIMAL

I have not been able to examine mature, well preserved specimens and
mature specimens have never been observed off Naples. They probably
inhabit deeper waters. I obtained remains of a male with a filled sperma-
tophore pocket from the stomach of a shark; the macerated arms of this
specimen are shown in Figure 22 6. The arms show that the animal is
about as^large as an adult 111 ex (Figure 218), or at least more than
3 times as large as the specimen shown in Figures 223 and 224. In habitus
439 the adult probably resembles the forms in the above figures, but its body

was probably still more compact and its head thicker (p. 434). The following
description is based largely on the well preserved specimen in Figures 223
and 224, a young male which shows distinct evidence of hectocotylization but
otherwise differs little from females of similar size (Figure 225).

FIGURE 223, Young male of T o d a ro p s is eblanae. Half natural size. The animal was preserved
with the ventral arms folded back, to show the inner side of the arms and the mouth region. Note the
proportions of fins and mantle sac; the neck folds; and mainly the arm apparatus (tentacle stalk and
club, adhesive knobs, swimming and protective margins on all arms); points, attachments and supports
of buccal funnel; buccal cone with outer and inner lip; apex of lower jaw.

The habitus of Todaropsis closely resembles that of a mature male of
Illex coindeti (Figure 218) in the thick head, strongly developed arms,
and the relatively short body. Verany (1851 ) therefore regarded this species

419

441

as a male of "Loligo s a g i tt a t a, " i. e. Illex coindeti. Verany
probably knew the male of the much more common Illex, but accidentally
took a Todaropsis for his description with which he confused it. Jatta

(1896, Plate 2, Figure 6) made the opposite error
in depicting a typical male of Illex coindeti
as a Todaropsis. The coloration of the live
animal is shown in Plate XIX, Figure 5.

The important character for the distinction of
these species is the fin/ mantle ratio. In well
preserved specimens of medium size (Figure 224)
the part of the dorsal mantle before the fins is
only one -sixth to one -quarter greater than the
length of the fins; the corresponding ratio in
Illex is always more than a third. The fin of
Todaropsis is of typical form.

The mantle sac is wide, cylindrical anteriorly,
pointed and spindle-shaped in the posterior part;
its anterior margin is usually slightly bell-shaped.
The neck folds are typical, but the anterior trans-
verse fold is often more or less indistinct and
depends on the state of contraction.

The powerful arm apparatus is even more
developed than in mature males of Illex and
represents an extreme condition in the family.

In well preserved, not markedly contracted spe-
cimens the longest arm (2nd) is almost as long
as the mantle (still longer in the live animal, see
Plate XIX, Figure 5). The relative lengths of the
arms are typical (formula: 2, 3, 4, 1 ), except for
the hectocotylization of the ventral arms; also
typical is the structure of the swimming margins,
but the corner of the margin of the 3rd arm is
situated slightly more distal than in Illex, i. e.
just before the middle. The protective margins
are also typically developed, weakly developed
on the 4th arm and large elsewhere, especially
on the ventral edges of the lateral arms.

The tentacles are long, the stalk longer than
the club. The club usually bears 7 pairs of large
median suckers and 2 adhesive knobs, the distal
part 4 regular rows. The large suckers have
sharp teeth all around, separated by intervals wider
than the teeth. The largest suckers of the arms
are situated slightly distal to the middle; the
suckers of the DL and VL arms are larger than
those of the D arms; they are smallest on the
V arms. There is no marked difference between
the suckers of the proximal and distal parts as
in Illex. The zone of the large suckers has apparently extended distally,
losing some of its character. The distal margin of the large suckers bears
pointed, triangular, separated teeth, which become more crenelated laterally.

FIGURE 224. Todaropsis
eblanae, young male, half
natural size. The animal in
Figure 223 is shown here in
the natural swimming position
but with slightly opened arms.
Note the general proportions
of fins, mantle sac and arm
apparatus, also details of funnel
pit, lid margin, windows, and
swimming and protective
margins.

420

Smaller, secondary teeth are situated between the large teeth. The adhesive
ring is rudimentary, but becomes increasingly distinct in the distal suckers.
The buccal funnel (Figure 223) is typical, as in 111 ex.

The mantle cavity shows the conditions typical for the family, but the
length/width ratios are slightly different because of the compact body
(cf. Figure 225 with Figure 220).

FIGURE 225. Half-grown female of Todaropsis
eblanae with opened mantle, natural size. The
funnel is also opened by removal of its lower wall
to show the funnel valve (k) and the dorsal part of
the "funnel gland" (d). The half-developed nida-
mental glands are widely separated in the middle
in the posterior part of the body. Note also the
branched posterior aorta, the branchial hearts, the
lateral pallial vein, posterior pallial vein, gills,
venous appendages, vena cava, renal papillae (n),
hind intestine, funnel retractors, lateral sinuses,
funnel and mantle cartilages, neck folds, funnel pit,
funnel adductors, eyelids and window (f);

1 and li — ligaments of the nidamental glands.

FIGURE 226. Ventral arms of a large male
of Todaropsis eblanae from the
stomach of a Heptanchus, natural size.
The animal was partly digested and had a
filled spermatophore pocket. On the distal
part of the arm (Di), suckers and skin are
reduced to shreds, but the muscular axis is
well preserved and presents a sharp contrast
to the proximal part (Ba), which bears on
each side 8 "scales," the last of them only
just indicated. Note the margin of the larger
outer scales (Sai_4) and the relative length
of the arms.

Sii_4 — median scales; Te — tentacle stalk;
Bu — ventral attachment of buccal funnel.

421

The jaws are as in 111 ex. I could not find any distinct differences
(p. 433) but the lateral teeth of upper and lower jaws are reduced and
rounded, those of the lower jaw often absent (Plate XVII, Figure 9). The
radula shows the characteristics of the family (Plate XIV, Figure 5).

The sexual dimorphism is incompletely known, because of the lack of
well preserved mature animals (p.438).

Mature females have apparently never been studied and they could not be
obtained at Naples. They probably have a dorsal mantle length of 15— 20 cm.
My data are therefore also based on half-grown animals and on a partly
digested, mature male from the stomach of a Heptanchus; the ventral
arms of this male are shown in Figure 22 6.

The nidamental glands are typically situated; in contrast to 111 ex, they
are widely separated in the middle. Their anterior part projects free
anteriorly and outward and becomes fastened at an early stage (see also

442 Figure 243) to the body by an anterior and a lateral attachment. The
anterior attachment probably develops into a true suspensory ligament
(as in Figure 207). In contrast to 111 ex, the genital process has a typical
position in both sexes, i. e. it extends into the pocket of the gill base and
projects from it anteriorly. The wide, slitlike opening of the genital pocket
is also visible (p. 420).

Of particular interest is the hectocotylization. This causes a slight
enlargement of the suckers of the lateral arms but affects mainly the ventral
arms. A distinction has to be made between the changes in the proximal and
distal parts of the arm which become increasingly distinct during growth:
the changes of the proximal part of the arms develop symmetrically on both
arms but the distal part is modified only on the right arm; as far as we know.
It begins from the base of the arms, which are normal in the youngest spe-
cimens, and the basal pads of the suckers become modified, mainly in the
outer rows and the suckers themselves are finally lost. This modification
creates hard, muscular scales, directed anteriorly and with a pointed pro-
jecting corner at the inner side of the margin. The scales of the outer row
have characteristic indentations. The part of the arm which bears these
structures becomes very thick and muscular; it probably performs some
forceful action during copulation (widening of the mantle cavity, etc.).

Figure 22 6 shows these conditions at the peak of their development, but partly
damaged.

This modification apparently affects 4 or 5 bases of the suckers of the
outer row and 3 or 4 of the inner row. Modification of the distal part of the
arm is restricted mainly or entirely to the outer row of the right arm.

443 This cannot be considered as a continuation of the scale formation but as
a change corresponding to that on the same part in the male of Illex
(Figure 219), because traces of scale formation are also evident in the
proximal part of the arm of Illex. The suckers disappear early in this
species, before formation of scales is complete (Figure 223), and the basal
pads develop into papillae with a serrated, projecting lateral margin.

Similar but more weakly developed papillae are also formed in the inner
row, but the suckers persist or are lost only much later (cf. Hoyle, 1902).

The distal part of the left arm resembles that of the female; the ventral arms
are always of different length: the left arm is longer in the well preserved
specimen in Figure 223, the right arm in the macerated, mature specimen in
Figure 226. The completely hectocotylized arm is perhaps always the
shorter, and this is a variation similar to the one in Illex (p. 434).

422

d. POSTEMBRYONIC DEVELOPMENT

Figure 227 shows the youngest stage that can be identified with certainty
as this species. Much smaller specimens are difficult to distinguish from
Ommatostrephes of the same size (at first I placed here the specimen
shown in Figure 23 6, since intermediate stages were not available). Like
the other, slightly larger specimens, this animal shows a very characteristic
habitus and proportions, but it differs from the half -grown Todaropsis
in a number of characters: the plump, compact body, the short apex of the
mantle, the short, wide fins. The head is still rounded as in young speci-
mens and the arms and tentacles are still relatively short. The fins are
444 of typically juvenile form as in the stages following that shown in Figure 222
and later in Figure 206 (Plate V). Funnel and funnel pit show the typical
characters of the Illicinae. The same is the case with the eye region
(bulbus, lid margin, window), although the sinus is still less deeply incised.
The protective and swimming margins of the arms are still less developed
than in the adult; those of the 3rd pair of arms lack the sharp lateral
corner, and those of the 2nd pair the widening in the middle of the arm
(Figure 223a). At this stage the tentacle club does not yet have adhesive
knobs.

(443)

FIGURE 227. Young Todaropsis e b la na e, natural size. Note the plump body
(of. Figure 206), form of fins and mantle, translucent subterminal cone, funnel
adductors, funnel pit, neck folds, eye region, arm apparatus (especially the swimming
and protective margins, tentacle stalk and club) and the distribution of chromatophores.

The following stages lead rapidly to the conditions described above.
Both these and the younger forms differ from the slenderer stages of
1 11 ex in the quadriserial pattern on the apex of the club (p.430).

423

445 Chapter 35

GENUS OMMATOSTREPHES

d'Orbigny, 1835

DIAGNOSIS

The suckers occupy more than half the length of the tentacles. Accessory
folds ("lateral folds") absent lateral to crescent-shaped fold of the funnel
pit. Lateral ridges of gladius ending anteriorly without dividing (cf. Sthe -
noteuthis. Chapter 36).

O. sagittatus may be considered as typical for the genus.

OMMATOSTREPHES SAGITTATUS (Lam., 1798)
d'Orbigny, 1835

a. DIAGNOSIS

The suckers occupy at least three-quarters of the length of the tentacles.
Stalk part and hand part of club together with 70—80 suckers.

b. LITERATURE

1799 Lamarck, Loligo sagittatus (Vol. 7, p. 663).

1814 Rafinesque, Loligo toda rus (p.29).

1824 Carus, Loligo sagittata (p.318, Plate 30).

1834 Delle Chiaje, Loligo to d a r us (Mem. 4, p. 59, Plate 60),

1839 Ferussac and D'Orbigny, Ommastrephes todarus (p.349, Loligo Plate 1,0 m ma strephes
Plate 2).

1849 Gray, Ommastrephes todarus (p.60).

1851 V€rany, Loligo todarus (p. 101, Plate 33).

1851 Verany, Loligo aequipoda (p.l05, Plate 35, Fig.a and b juv.).

1853 Forbes and Stanley, Ommastrephes sagittatus (Vol. 4, p.233, Plate RRR).

1862 Jeffreys, Ommastrephes todarus (Vol.5, p.l28),

1869 Targioni-Tozzetti, Ommastrephes todarus (p.54,Fig.).

1878 Sars, Ommastrephes todarus (p.334, Plate 30).

1880 Steenstrup, Todarodes sagittatus (p. 73, Fig.4).

1884 Pfeffer, Todarodes sagittatus (p.28),

1885 Herzens tein, O m m a tost r eph es todarus (p.713),

1885 Steenstrup, Toda rod es sagittatus (p.l26),

1886 Hoyle, Todarodes sagittatus (pp, 34, 163).

424

1889 Posselt, Todarodes sagittatus( p.l44).

1889 Girard, Ommatostrephes sagittatus (p.264, Fig.3).

1890 Girard, Ommatostrephes sagittatus (p.214).

1890 Posselt, Todarodes sagittatus (p. 301, Plate 8).

1890 Norman, T od a rod es sagittatus (p,477).

1890 Carus, Todarodes sagittatus (p.447).

1890 Lonnberg, Todarodes sagittatus (p.33).

1892 Girard, Ommatostrephes sagittatus (pp.214, 220).

1894 Joubin, Todarodes sagittatus (p.214).

1896 Jatta, Todarodes sa gitta tus (p.81, Plates 1, 10, 11).

1896 Grieg, Todarodes sagittatus (p. 23).

1900 Nichols, Todarodes s a g i t ta gus (p.495).

1900 Pfeffer, Ommatostrephes sagittatus (p.l79),

1902 Hoyle, Ommatostrephes sagittatus (p,197).

1903 Lo Bianco, Todarodes sagittatus (p.l71).

1904Jatta, Todarodes sagittatus (p.l98).

1908 Pfeffer, Ommatostrephes sagittatus (p. 94, Figs. 104—1 08),
1908 Massy, Ommatostrephes sagittatus (p.32),

1912 Pfeffer, Ommatostrephes sagittatus (pp.439-“451, Plate).
1916 Naef (Syst.), Ommatostrephes sagittatus (p. 16). (32, 33),
1921 Naef (Syst.), Ommatostrephes sagittatus (p.538).

1921 Grimpe (Nords.), Ommatostrephes sagittatus (pp. 299, 302).

446 c. STRUCTURE OF THE ADULT ANIMAL

The general outline is typical for the family but the fins taper distinctly
into a long point, and the mantle sac therefore has a narrow tapering end,
which is slightly laterally compressed, but less than in the Ommatostrephidae
described above (about the gladius see p. 432). Mantle margin, neck folds
and funnel pit are typical; the funnel pit is deep, as is characteristic for the
Ommatostrephinae, so that the funnel is situated almost completely inside it
and the lateral margin of the pit projects above the funnel at least in the
posterior part. The strongly developed lateral funnel adductors are there™
fore not visible. In relaxed specimens, extension of the pit produces a
picture similar to that in Sthenoteuthi s (Figure 240). The anterior
part shows the crescent-shaped fold (Figure 240) around the 9 — 13 irregularly
parallel longitudinal ridges or folds, which are separated by deep incisions.

FIGURE 228. Lateral view of a young O m m a t o s t re ph es sagittatus swimming position. Note the
base of the fins, posterior end of body, form of mantle, neck folds, funnel pit, funnel; eye region with lid
margin, sinus, iris fold (black), lens and window; the bases of the arms with the connecting membranes,
swimming and outer margins; tentacles with stalk margin, swimming margin and protective margin. This
figure shows the appearance of the live animal when it is about to catch the prey, swimming forward with
the arms slightly opened (I have observed this not in this species but in II lex and Loligo).

Funnel apparatus, head and eyes are typical (p. 415). The protective
margins of the arms are typical for the family, i. e. they do not show such

425

(447)

FIGURE 229. Ventral view of O m m a -
strephes sagitta tu s.half natural size,
after a young specimen from Capri. Note
the pointed fins and posterior end. Other-
wise the proportions are typical; the deep
funnel pit with crescent-shaped and longi-
tudinal folds; the margin of the pit which
touches the anterior transverse neck fold
at a right angle and covers the outer funnel
adductor. The eye region is typical, and
also the arms with protective and swimming
margins. The tentacles have stalk margins,
swimming margins and protective margins.
Note the differentiation of the suckers.

FIGURE 230. Dorsal
view of the same
O. sagittatus. Note
the form of the fins,
"earlobes," mantle
and head. The arms
are typical, especially
the swimming margins
and the long tentacle
club.

FIGURE 231. Gladius
of a half-grown O m -
matostrephes sa-
gittatus, natural
size ( cf. Figure 238),
This is a typical shell
of Ommatostrephidae
(p.413) with a weakly
developed cone. The
lateral ridges are un-
divided anteriorly.

426

an extreme degree of development as the following genus; the suckers bear
a distinct adhesive ring with regular radial striation which is particularly
well marked if the chitinous parts are stained reddish brown from preser-
vation in for mol -alcohol. The chitinous coat of this zone has apparently
been dissolved into narrow stripes. The adhesive ring has become detached
from the toothed ring and has become narrow and slightly interrupted so that
the stripes are not radial but oblique to each other (Plate XII, Figure 5). The
large suckers of the arms bear 9 (7—9) pointed, isolated teeth at the distal
margin; the middle tooth is the largest, and the adjacent teeth are smaller
than the lateral teeth.

The tentacles have an enormous club which occupies more than three-
quarters of the contracted tentacle (Figure 232). All parts of the club are
448 lengthened, especially the stalk part and hand part. The boundary between
these two areas is not distinct (p.415). The biserial part is considered as
the stalk part, and the quadriserial part with larger suckers as the hand
part. However, there is a transitional zone, the indistinct carpal part, in
which the pairs of suckers are alternately displaced. The boundary of
this part varies according to the state of contraction; the hand part begins
at the first enlarged sucker of a median row and ends at the last large
sucker of this row. Each longitudinal row of the hand part then contains
13 — 17 suckers.

FIGURE 232. Right tentacle of Ommatostrephes s a gi 1 1 a t us, half natural size.

Note the length of the club; the gradual transition from the stalk part to the hand part,
which is characterized by the larger suckers in the median rows; the relatively short
distal part. A small part of the stalk margin and swimming margin; the protective
margins and their supports. This drawing could be inserted in Figure 229 as it is drawn
from the same specimen. The differentiation of the tentacles is reduced and the tentacles
resemble ordinary arms with 4 rows of suckers.

The suckers of the stalk part, the carpal part, the proximal zone of the
hand part and the adjacent suckers of the marginal rows are like those on
the arms and bear 5—7 teeth at the distal margin. The median suckers of
the hand part are larger, cuplike and toothed all around. The teeth (15— 20)
are widely separated and are pointed conical; those at the proximal
margin are smaller than those at the distal margin, which usually include
one particularly strong tooth (Plate XII, Figure 2). Lower, rounded,
secondary teeth may be present between the larger teeth. The marginal
suckers on the larger distal part of the club are similar, but the secondary
teeth are more distinct, and pointed at the aboral margin.

The suckers of the tentacle show the same characteristic differentiation
of the adhesive ring as on the arms. This differentiation is, however.

427

modified in the large median suckers (Plate XII, Figure 2). The adhesive
ring avoids the teeth by fitting into gaps.

The mantle cavity shows the typical conditions of the family (p. 418,
Figures 233 and 207).

(449)

FIGURE 233, O m m a t o s t r e ph es sagittatus with
opened mantle. Young female, half natural size.

FI — fin; Ap — posterior pallial artery; Tc — truncus
communis; Aa — accessory pallial artery; Vp — posterior
pallial vein; Am— median pallial vein; Nd — nidamental
gland; Kh — branchial heart; Kt — posterior opening of
pocket of gill base; Vb — branchial vein; Br — bridge
over the perforated pocket of the gill base; n — renal
pore; t — ink sac; d — hind intestine; s — "lateral
pocket"; v — vena cava; r — funnel retractor; Mm — cut
surface of muscular mantle; Kb — gill ligament; Mh —
mantle bond; a — anus; Th — funnel bond; Ta — outer
funnel adductor; x — corner of funnel pit; Qv and Qh —
anterior and posterior transverse neck folds; L2,L3 — second
and third longitudinal neck fold; Tr — funnel; Tt — funnel
pocket; Tg — margin of funnel pit; Fo — foveola with
longitudinal and crescent-shaped folds; Fe — window;

Li — lens; PI — primary lid; Ir — iris margin; Si — eye
sinus; Ht — membrane between the 3rd and 4th arm;

Ls — lateral margin of the 4th arm; Mk - median outer
edge of 4th arm; Sw — swimming margin; Ax — axis of
arm; Sh — protective margin; St — support of this;

St — tentacle stalk; Sm — stalk margin; K1 — club;

Ts — protective margin; Swj — swimming margin of
tentacle.

428

The jaws closely resemble those of Sthenoteuthis; the differences
shown in Figures 10 and 11 and Plate XVII do not exceed the range of

449 variation and age differences. For example, the inner plate of the upper
jaw of Ommatostrephes (Figure 10a) bears a dark strengthening stripe
which increases with age. This stripe is less developed in young animals
and is only indicated in the smaller Sthenoteuthis in Figure 11. The
lateral tooth is supported in both species by a posteriorly widened dark
stripe, which is distinct in Figure 10b, c, d, especially on the upper jaw.

450 The radula (Plate XIV, Figure 6) is difficult to distinguish from that of
Todaropsis and II lex but differs distinctly from that of Stheno-
teuthis: the median teeth are longer than the lateral teeth.

There is no description of hectocotylization in mature, well preserved
specimens (cf. Pfeffer, p. 444) and I have been unable to obtain such a
specimen. Hectocotylization affects particularly the end of one ventral
arm, but the proximal suckers of the same arm are already slightly smaller
than those on the other arm. The end of the arm lacks suckers; its basal
pads and protective margins are modified (cf. also Ishikawa’s description
of the hectocotylized arm of the Pacific O. sloanei in Zool* Anz., 1913;
also Sasaki, 1915, Plate 4, Figure 4: C. v o 1 a t i 1 i s). Especially O. vola-
tilis resembles 1 11 ex (Figure 434) in the reduced median suckers and
the formation of terminal papillae. The female genitalia are of typical
form (p. 419).

Merculiano (in Jatta, 1896, Plate 1, Figure l) shows the coloration of a
fresh specimen, without, however, its sheen and iridescence; the live animal
has still more intensive colors, mainly a luminous brownish red.

d. POSTEMBRYONIC DEVELOPMENT

O. sagittatus is not common near Naples; 111 ex and Sthenoteuthis
are much more frequently found on the market, but most of the young stages
obtained belong to O. sagittatus. Figure 234 shows the youngest stage
which can be determined with more or less certainty. It does not show the
important characters of the genus or species; its position is determined by
its resemblance to the following stages and exclusion of the 3 other species
which occur in the bay: Similar stages of the closely related Sthenoteu-
this bartrami still have tentacles like the Rhynchoteuthis larvae
451 (Figure 244); the chromatophores are much more densely arranged than

in the young Sthenoteuthis (cf. Figures 237, 244 and 246). The chroma-
tophores of the youngest, only slightly larger stages of 111 ex are still
sparser (Figure 222) and their habitus is markedly different. Young
Illex (with a mantle length of less than 80mm) are very rare, while the
similar stages of Ommatostrephes are found frequently at the surface.
The stage shown in Figure 234 was caught together with several slightly
older specimens which could be determined without any doubt. This speci-
men shows the typical transition from Rhynchoteuthis to a young spe-
cimen of Ommatostrephidae.

429

(450)

FIGURE 234. Juvenile stage of O m m a t o s t r e ph es sagitta tus showing the
transition from a typical Rh y n ch o t e u t h i s to a young specimen of Ommato-
strephidae (caught at the "Amontatura” on 25 April 1913 at 250m depth). 4.8x.

The tentacle stalks are almost completely separated, and the development of clubs
is advanced (Figure 235). Note the development of the arms; presence of swimming
margins; eyelid with sinus; olfactory tubercle with adjacent neck fold; funnel
adductors; form of mantle; posterior end of body; distribution of chromatophores.

FIGURE 235. Mouth area of the young Ommatostrephes
sagitta tus in Figure 234. 20 X. Note the bases of the
arms (I-IV); the edges on ihe inner surfaces; the just indi-
cated membranes, especially between the 3rd and 4th arms
(,v), which delimit the shallow tentacle pockets; the circular
buccal cone, surrounded by the buccal funnel with 7 distinct
points (1—4), of which the 5 upper points already have attach-
ments (h2,h3); in the middle, the outer (a) and inner (i) lips,
with the end of the jaw inside. The development of the
ventral arms and ventral buccal corners has caused the
tentacle stalks now to be separated except for a short part
(n) before the club. Note the growth of the club, with the
formation of new suckers (cf. Figure 245).

ti-3 — buccal pockets; f — membrane connecting the ten-
tacles at the point of separation.

The almost complete separation of the tentacle stalks deserve attention.
The changes in the mouth region connected with it are shown better in
Figure 235. The fins are in a stage of transition; their base has become
longer. The formation of the fleshy apex has also begun, but the indentation
on the posterior margin of the fins is still present so that the fins form at
first a transverse oval and later a rhombus, which is typical for the juvenile
form (Plate V). The mantle sac is particularly wide. Both pairs of funnel
adductors are developed; the inner adductors are covered in the figure.

430

whereas the outer are visible but still quite delicate. The flat olfactory
tubercle is situated next to the 3rd longitudinal neck fold. The lid margin
has a pointed sinus. The lateral arms bear swimming margins, but that of
the 3rd pair is still without a lateral corner. The
protective margins are only indicated. The tentacle
stalks are united in only a short distal part. Their base
is situated in shallow pockets distant from the middle,
and the bases have apparently been pushed apart by the
secondary development of the ventral arms. The club
shows new parts in development which were absent in
the early larvae (Figure 215 on p.427). The 8 larval
suckers are still the largest but numerous new suckers
develop on the growing end, the first still in 2, the
following already in 4 rows; the youngest "anlagen" are
again arranged in 2 rows and finally in a single row
(cf. Plates V and Vl) and indicate the typical develop-
ment (p. 127).

The buccal funnel of this stage consists of 7 parts;
the 2 ventral points occupy the space between the bases
of the tentacles (cf. Plate VI, Figure 1, also Figure 214).
The development of the 2 ventral points and the ventral
arms facilitates the separation of the tentacles (cf.

Figure 245). The attachment of the connecting buccal
membrane is already displaced to the buccal cone, and
the attachments of the 5 dorsal points are displaced out-
ward, toward the bases of the arms. These conditions
resemble the definitive conditions in the Ommatostre-
phidae (Figure 223), in which the "anlage" of the 6 typical
buccal pockets forms a slitlike invagination around th^
buccal cone. The complicated buccal funnel thus develops
from a circle of papillae (rudimentary arms) which
surrounds the buccal cone.

Figure 236 shows a similar stage. The habitus is different, but this is
due to contraction and there is only a slight further development of the
tentacles and fins. Older stages can be easily determined as Ommato-
strephinae: they have a crescent-shaped fold even before the funnel pit is
sharply delimited. In contrast to the wine red to reddish brown coloration
of Sthenoteuthis, these forms are mainly yellowish brown and the
chromatophores are loosely arranged. Figure 237 shows such a specimen
from Messina (cf. Figure 246).

A similar specimen illustrated in Plate V, Figures 2—3 shows the typical
characters of young Ommatostrephidae. The fins are slightly longer,
transversely oval, with a slightly indicated posterior indentation. The
453 ventral (4th) and lateral (2nd) longitudinal neck folds are already present,
and also the indistinct anterior transverse fold. The funnel pit begins to be
delimited, and a crescent- shaped fold develops on the still indistinct anterior
margin. The arm apparatus already resembles the definitive state, except
that the ventral arms and tentacles are still retarded. Protective margins are
indicated on all arms, at least in the form of narrow ridges with small
evaginations at the point of each support. The lateral arms have swimming

(452)

452

FIGURE 236. Young
O m ma tostr ephes
sagittatus. The
tentacles are com-
pletely separated and
the fins are slightly
longer. This specimen
closely resembles that
in Figure 234, but it is
strongly contracted.
4.8X.

431

454

margins with an indicated lateral corner on the 3rd pair. A drawing of the
arms of a specimen of similar size (Figure 3 of Plate VI, highly magnified)
shows the above characters and also the arrangement of the suckers on the

arms, the membrane between the 2nd
and 3rd arms (a characteristic of the
Ommatostrephidae) and especially the
differentiation of the mouth area, which
is intermediate between the conditions
in Figures 235 and 223. The following
are distinct: the 7 points and bonds of
the buccal funnel, the developing con-
nection between the bonds and the
protective margins, the 6 buccal
pockets around the buccal cone, including
the ventral pockets between the buccal
funnel and the adductors of the 3rd arms,
and finally, the thin outer lip, the thick
inner lip and the apex of the lower jaw.

The tentacles (Figure 237) are much
larger and the "anlagen" of most
suckers are present but neither suckers
nor club are completely developed yet.
The protective margins are only edges,
and the whole tentacle still resembles
an arm. There is still no trace of a
swimming margin and the tentacles are
still shorter and thinner than the 3rd
pair of arms.

A more advanced stage is shown in
Plate V, Figure 1 (cf. Figure 206).

The mantle cavity of this specimen is
opened, but the habitus of the intact
animal can be visualized: the mantle sac
is slender and the posterior apex is
longer, but mainly, the fins have become
transformed from a transverse oval
to a rhombus, with distinct lateral and
terminal corners and weakly marked "earlobes." The neck folds are
almost complete, and also the funnel pit with the crescent-shaped fold. The
eyes are now lateral and the head is still slightly more rounded than later.
The arms have typical swimming and protective margins; the latter are
still little developed but are already present on the tentacles as well. The
swimming margins of the tentacles are still absent or only slightly indi-
cated at the end. The club begins to become differentiated and the median
suckers have become larger. The tentacles are now longer and thicker than
the 3rd arms, while the ventral arms are still slightly shorter than the
dorsal arms.

The older stages now rapidly begin to resemble the adult stage. They can
always be distinguished readily from Sthenoteuthis, even if the chroma-
tophores have become bleached, by the structure of the tentacles, which have
a very short stalk and bear suckers on their greater part.

FIGURE 237. Young O m m a t o s tr eph es sa-
gittatus from Messina. 4.5x. Note form of
fins: apex of body, mantle margin, funnel adduc-
tors, longitudinal and transverse neck folds, funnel
pit, tentacles, arms, swimming margins and es-
pecially the distribution of the chromatophores
and the presence of darker and ligher chromato-
phores.

432

I have not examined a mature specimen; the largest animal, caught with
a "latero, " has a dorsal mantle length of about 33 cm (cf. Figure 207 b).

On the other hand, half-grown and younger stages are not rare on the fish
market, where they are sold together with 1 11 ex as "totero verace." No
distinction is made from 111 ex, but they are distinguished from Sthe no-
te ut his which has tougher flesh. O. sagittatus is frequently caught
together with Sthenoteuthis near Capri in September and October. The
description on p. 44 6 applies to a typical specimen with a mantle length of
18 cm.

433

455 Chapter 36

GENUS STHENOTEUTHIS

Verrill, 1880

DIAGNOSIS

Suckers present only in the distal half of the tentacle. Accessory folds
("lateral folds") present lateral to crescent-shaped fold of the funnel pit.
Lateral ridges of gladius ending anteriorly in 2 separate ridges. Large
suckers of tentacle with 4 teeth forming a cross. Ventral protective margin
of 3rd arms several times broader than the arms themselves. End of arms
not whiplike.

The genus contains only the cosmopolitan species S.bartrami.

STHENOTEUTHIS B A R T R A M I (Lesueur, 182 1 )

Verrill, 1910

a. DIAGNOSIS

Ventral protective margins of 3rd arms at most 4 times as broad as the
arms. Tentacle clubs with 3—4 rows of adhesive suckers and with the same
number of knobs; 4—7 suckers situated proximally before the first knob.

b. LIT ERATUR.E

1821 Lesueur, Loligo bartrami (p. 90, Plate 2),

1835 D'Orbigny, Ommastrephes bartrami (p.55).

1839 Ferussac and D’Orbigny, Ommastrephes bartrami (p.347, Lo li go Plates 2, 21,0 mmastrephes
Plate 2).

1849 Gray, Ommastrephes bartrami (p.62).

1852 Gould, O ny choteuthis brevimanus (p.483, Plate 50).

1853 D’Orbigny, Ommastrephes bartrami (p.59).

1880 Verrill, Stenoteuthis p t e ro p u s (p.228, Plates 27, 36).

1880 Verrill, Stenoteuthis bartrami (p.322),

1880 Verrill, Stenoteuthis pteropus (p. 3 17, Plates 7, 17).

1880 Verrill, Stenoteuthis bartrami (p.322).

1880 Steenstrup, Ommatostrephes b a r t ra m i (pp.73, 79, 81, Figs.2, 3).

1884 Verrill, Stenoteuthis bartrami (p.l06).

1886 Hoyle, Ommatostrephes bartrami (p.32).

1889 Carus, Ommatostrephes bartrami (p.446).

434

1890 Girards O m m a t o St r e ph es bartrami (p.265s Fig.3).

1890 Posselt, Om matostrephes bartrami (p.301).

1896 Jatta, Ommatostrephes bartrami (p=64, Plate 10, Testfigs.8, 12, 19).

1897 Lonnberg, Ommatostrephes bartrami (var, sinuosa) (p.702),

1900 Pfeffer, Stenoteuthis bartrami (p.l80).

1906 Nichols, Stenoteuthis pteropus (p.54).

1907 McIntosh, Stenoteuthis bartra mi (p.l72,Plate 7).

1908 Hoyle, Stenoteuthis b a r t ra m i (p-132).

1908 Pfeffer, Stenoteuthis bartrami (p.97, Figs.l09--114),

1909 Massy, Stenoteuthis pteropus (p.32).

1915 Pfeffer, Stenoteuthis bartrami (pp.462— 482, Plates 35, 36, 37, 39).

1916 Naef, Stenoteuthis bartrami (p.l6).

1921 Naef, Stenoteuthis ba rtra mi (p.538).

1921 Grimpe, Stenoteuthis ba r t r a m i (p.299).

456 c. STRUCTURE OF THE ADULT ANIMAL

Sthenoteuthis has a slender body with very powerfully developed
muscles of mantle and fins, making it the strongest swimmer of all known
Cephalopoda (p. 412). The mantle is unusually thick, tough and hard, so that
it is very unpalatable.

The fins are very large; their relative length in growing specimens with
a dorsal mantle length of over 6 cm is one-third to one-half this length and
this ratio increases slowly but constantly. The anterior margin of the fins

457 has a thin, almost membranous stripe separated from the muscular part,
which is more distinct than in other Ommatostrephidae. This stripe is
usually folded ventrally during preservation (in Figure 238 marked with a
dotted line). The membranous stripe widens laterally, but its boundary
becomes less distinct. The anterior margin of both fins forms an almost
regular curve, the center of which is situated slightly before the posterior
end of the body. The outline of the fin is otherwise typical, not pointed as
in Ommatostrephes (p.447).

The gladius differs from that of Ommatostrephes sagittatus in
2 characters: l) the ventral length of the cone, which is about one-fifth of
the cone flag and 2) the division of the lateral ridges at the anterior margin
of the rhachis; the outer branch (Ar in Figure 239) is situated closer to the
margin than the inner (jr).

Neck folds and funnel adductors are typical. The funnel pit encloses
the funnel particularly closely when the animal is not completely relaxed
(Figure 240a). The crescent^shaped fold encloses 6—9 variable longitudinal
ridges. There are 3 — 6 oblique, irregularly arranged transverse folds lateral
to it ("lateral folds") which form shallow pockets directed anteriorly.

The arms are rather short because of contraction during preservation in
formol (cf. Figure 228). The outer edges of the dorsal arms are so sharp
that they could be considered as swimming margins, especially the lateral
edges. The general structure is typical, but the 3rd arms are longer than
the 2nd (formula: 3, 2, 4, 1; in the largest specimens: 4, 3, 2, l).

The protective margins are characteristic, especially the ventral margins
of the lateral arms. They are extraordinarily wide and have an almost

435

straight margin (Figure 241 ), and the supports project into slight corners
only at the base. The supports are relatively delicate and often do not
project distinctly from the connecting membrane. The ventral margin of
458 the 3rd arm is very wide but no exact measurements can be given, because
of the marked contractility. A width of the margin 4 times that of the arm
may be considered as maximal. The other protective margins are normal
but very strong, those of the dorsal arms stronger than on the ventral arms.

(456)

FIGURE 238. Young Sthenoteuthis bartrami from Capri, half natural size. Note the slender,
compact body: the size and form of the fins; the short arms; the deep funnel pit. Note particularly
the anterior margin of the fins, the firmly surrounded funnel, the strong ventral protective margins of
the lateral arms, and the characteristic form of the dorsal longitudinal folds of the neck.

The tentacle club, i. e. the part of the tentacle which bears suckers
(Figure 241) occupies only about half of the tentacle. There is a rudi-
mentary stalk part with a zigzag pattern of small suckers which pass into

436

the indistinct carpal part (pp.415, 558). The adhesion apparatus consists
of 3 or 4 suckers and knobs. Where the proximal element is a sucker
(Figure 242), there are 7 suckers before the following knob. This is the
usual condition. The adhesion apparatus marks the transition from the
carpal to the hand part, which bears about 12 pairs of distinctly enlarged
suckers. The distal part bears 25 or 26 oblique rows of 4 suckers in which
the ventral sucker is always larger, except in the 2 proximal rows of four.
The protective, swimming and stalk margins are typical, but the protective
margins extend proximally markedly beyond the first suckers. The ventral
protective margin is especially wide, with large transverse supports. This
margin extends to the base of the tentacle, while the dorsal margin soon
tapers to a thin edge or a narrow fold and shows denser and more delicate
transverse supports only further distally. This part of the tentacle,
considered as belonging to the stalk (Pfeffer, 1912) because of its form and
the absence of suckers, belongs morphologically to the stalk part of the club
459 and has lost its suckers during the postembryonic development. This ex-
plains the presence of true protective margins with transverse supports.

It makes the close relationship with Ommatostrephes more probable,
because the whole club and its stalk part are very long in this genus. I
cannot prove this ontogenetically, as I have not made systematic obser-
vations, but I have transitional forms in which the lost suckers are still
partly present.

(456)

FIGURE 239. Gladius of a young Sthenoteuthis b a r t ra m i, natural size.
Note the long, striated and laterally compressed cone. The lateral ridges (Sr)
are divided anteriorly (see Figure 231 on p.447).

Vr - anterior margin of rhachis; Jr, Ar - branches of lateral ridges (Sr); Mr -
median ridge; PI — plate of rhachis; St ~ stalklike, narrow part of rhachis;

Fa — cone flag, b) lateral view of posterior end of gladius.

437

(457)

(458)

FIGURE 240. Funnel and head of a large Sthe-
noteuthis bartrami, Half natural size.
Drawing b shows the anterior end of the funnel
pit (Tg) of another specimen of about the same
size, with the crescent-shaped fold (Hb),the longi-
tudinal ridges (Lf) and grooves enclosed by the
fold, and the lateral folds (St). Figure a shows
also neck folds (L2 — L4) funnel adductors (Ae),
funnel opening with valve, funnel bonds and fun-
nel pockets (Tt).

Si — sinus of eyelid; Li — lens; Ls — longitudinal
ridge; Ti — inner tubercle; Te — outer tubercle;

Ri — longitudinal groove of funnel bond.

FIGURE 241. Right arm of 3rd pair in Stheno-
teuthis bartrami. Half natural size. From
a very large specimen about 80 cm long without
tentacles. Collection of the Zoological Station.
Note the large swimming margin; arrangement
of suckers at the base; the attachment (Bh) of the
removed buccal funnel. The dorsal protective
margin (Ds) is normal, the ventral (Vs) markedly
widened. The supports (St) are shown too distinctly
on the ventral margin; they may actually not be
.visible in some places.

If — inner surface.

The suckers resemble in general those of Ommatostrephes. Their
adhesion ring is also distinct, as in that genus but with a weaker radial
striation, at least on the large suckers (Plate XII, Figure 6). The relative
size varies with age. In young specimens the suckers of the ventral arms
are the smallest, and those of the two lateral pairs the largest. The suckers
of the DL arms are the largest in the male (or in both sexes?), and those of
the V arms correspond to the growth of the arm and are the second largest.

The largest suckers of the arms (Plate XII, Figure 8) have a large,
median tooth at the distal margin and 2—3 smaller but also pointed triangular
lateral teeth; then follow smaller teeth, at first sloping, then blunt toward
the proximal margin. There are lower, blunter, secondary teeth irregularly
interspersed between the large teeth. The smaller distal suckers of the
arms have relatively longer and more pointed teeth at the distal margin,
while such teeth are absent on the proximal margin. The proximal suckers

438

of the club resemble those of the arms; they bear rounded teeth further
distally. The suckers of the adhesion apparatus arenas usual, without teeth.
The dentition of the large suckers of the hand part is very characteristic:
the 4 main teeth form a cross and the distal tooth is the largest; there are
about 5 smaller teeth in the intervals and secondary teeth, arranged more
or less regularly, in the intervals between the smaller teeth. This arrange-
ment is not marked on the first suckers of the hand zone, but is transitional
to that of the stalk zone in the form and dentition of the suckers. The
marginal suckers of the hand part resemble those of the distal part: they
are of the type of the median suckers of the arms and resemble those of
the proximal part of the club.

(458)

FIGURE 242. Right tentacle of a half-grown S th e n o t eti th i s bartrami.
Half natural size. Note the arrangement and size of the suckers; the "cross-
teeth" of the largest suckers; the adhesive knobs (Kn); the swimming margin.
The protective margins particularly merit attention. They are typical on the
widened hand part, but they contain only as many supports as marginal suckers
near the adhesion apparatus. Note particularly that the protective margins
extend proximally to the stalk (Sm) where they have distinct supports (St2)
although suckers are absent.

Sn — suckers of adhesion apparatus (indicated by arrows); Stj — widened sup-
ports of hand part of dorsal protective margin.

The funnel is typical, but funnel and neck bonds are more sharply marked
than in Ommatostrephes. The anterior end of the longitudinal groove
of the funnel bond is particularly deep and is surrounded by larger ridges
("infrabasal cartilages" of Pfeffer) (Figure 240, Ls). In addition to the
formations present in Ommatostrephes (Figure 204), the neck bond
may bear 2 lateral ridges and a thin ridge in the middle of the groove.

The mantle cavity shows the typical characters of the family, but also
interesting special characters. The branches of the dorsal part of the
funnel gland extend down to the funnel retractors so that they are visible
also if the inner opening of the funnel is not damaged ( Figure 243). The
funnel retractors are divided distinctly into a thin, tendonlike posterior

439

part and a thicker, muscular anterior part. The renal opening is very
interesting: a tapering tube projects from the opening and is fused firmly
with the outer margin of the pore (see Naef, 1912, p. 332; 1913, p. 442).

460 This is an abnormally long "pericardial funnel" (renal injector, reno-peri-
cardial duct). Instead of opening in the kidney, near the renal aperture,
this duct has grown out of the kidney, so that there is no longer an open
communication between coelom ("pericardium") and renal sac. The
"anlagen" of the female organs are typical (Figure 243) but I could not
obtain mature females. In the young male, the spermatophore apparatus
is situated in the position of the duct apparatus of the female (Ov).

FIGURE 243. Young female S th e no t eu th i s bartrami with opened mantle cavity.
Half normal size. Note particularly the posteriorly projecting parts of the funnel gland;
the funnel retractors; the tubular pericardial funnel (Pt) which projects from the renal
opening; the posterior perforation of the gill pocket of the base which is surrounded by a
fold and from which the proximal apex of the juvenile oviduct gland (Ov) projects. Nearer
the middle are the typical "anlagen" of the nidamental glands and posteriorly the undulate
oviduct (Od). The following details are also typical: mantle, fins, septum of mantle cavity,
mantle and funnel cartilages, gills, lateral pockets, vena cava, hind intestine, lateral pallial
vein (Vp), posterior pallial vein, median pallial artery, posterior pallial artery.

440

The jaws resemble those of Ommatostrephes (Plate XVII, Figure 1 1 ;
p. 448). The radula is typical for the Ommatostrephidae, but the median
row contains shorter teeth than in the adjacent rows (Plate XIV, Figure 7).

Hectocotylization in the available immature specimens affects one of
the ventral arms (always the right?) (cf. Pfeffer, pp. 471—472).

The coloration resembles that of Ommatostrephes (Jatta, 1896,

Plate 1; see also d'Orbigny, 1839), but it is usually much darker, with deep
black-blue- to brown- violet tones, especially in the middle of the dorsal
side; it is bluish, if the chromatophores are contracted. There is a bright,
usually silvery metallic sheen on the mantle and head.

461 d. POSTEMBRYONIC DEVELOPMENT

Several juvenile stages are illustrated by Pfeffer (1912, Plate 37;

Figures 2 — 7 and 14 — 15 belong almost certainly to this species). Figure 244

shows a definitely determined, very young
stage. The tentacles of this specimen are
still of the Rhynchoteuthis type as their
stalks are fused in their greater part.
Otherwise it is more advanced than the
oldest Rhynchoteuthis stage of other
Ommatostrephidae, particularly Omma-
tostrephes sagittatus (Figure 234),
but smaller than this. The fins are
relatively long, while the form of the mantle
resembles that of the later stages. The
outer funnel adductors and the latero-
ventral neck fold are situated very close
together, as in the later stages of Stheno-
teuthis (Figure 246, 237). The dorsal
arms are almost as long as the lateral
arms, but the ventral arms are still very
short and stumplike. Especially charac-
teristic is the distribution of chromato-
phores. Comparison with slightly older
juvenile forms (Figures 246, 237) shows
that the chromatophores of Sthenoteu-
this are much more densely arranged and
more differentiated than those of Omma-
tostrephes, and this is distinct already
here. Figure 245 is a drawing of a slightly
older specimen with deformed (badly pre-
served) outline.

The arm apparatus and mouth region (Figure 245) have developed further;
the tentacle stalks are further separated, and the clubs and ends of the arms
have grown further (cf; the similar relationships in Figure 235). A further
young stage (Figure 246) shows the differences from an Ommatostrephes
of the same size: the fins are longer (over one-fifth of the mantle length.

FIGURE 244, Rh yn ch o t e u th i s stage
of Sthenoteuthis bartrami,
4.8x(from the deep-sea plankton of
Naples; old material). Compare with
Figure 234. The tentacles are relatively
retarded; the chromatophores are more
densely arranged despite the smaller size
of the specimen, and this distinguishes
the larvae from Ommatostrephes.
The slender form of the body is probably
due to the strong contraction of the
mantle.

441

(4 62)

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(462)

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442

the arms are shorter, and the tentacles much less developed, with a short,
pointed club. The funnel pit has a distinct margin and the crespent-shaped
fold is present; the margin of the funnel pit is connected with the anterior
transverse neck fold at an acute angle. The anterior transverse neck fold
forms further on a characteristic angle, and the longitudinal neck fold with
the olfactory tubercle is situated close to the funnel adductor so that the
median longitudinal fold between them is no longer visible. The young
Sthenoteuthis can then be readily distinguished from Ommato-
strephes by its form. The distinction is even easier if the chromato-
phores are preserved in color. Not only are they more densely arranged,
they also show characters which are absent in Ommatostrephes. All
chromatophores of Ommatostrephes are rather uniformly yellowish
4 62 brown, (except the light yellow and a few dark brown chromatophores on
the head). The chromatophores of Sthenoteuthis form 3 distinct
categories (if extended), as in Loligo: light brown, dark brown and wine
red. Especially the latter two groups give the animal a striking reddish
coloration at an early stage (dark reddish brown in the dorsal midline)
which gradually changes later into that of the adult animal.

As in Ommatostrephes, this species rapidly assumes the appearance
of the adult animal already in juvenile stages and a detailed description and
illustration of the following stages can therefore be omitted.

443

63 Chapter 37

FAMILY THYSANOTEUTHIDAE
Keferstein, 1866

GENUS THYS ANOTEUTHIS Troschel, 1857
DIAGNOSIS

Funnel bond broadly triangular, with a longitudinal groove along the outer
margin from which a wide transverse groove extends toward the median side
of the bond behind a large tubercle. Fins separated and extending almost to
the anterior margin of the mantle during postembryonic development.

Funnel pit and neck folds resembling those of the Ommatostrephidae,
except the 1st (dorsal) longitudinal fold. Lateral funnel adductors less
developed than in the Ommatostrephidae and displaced laterally. Distal
part of tentacle club with a double row of alternating suckers and knobs.
Gladius without cone but bearing a wide flag with arrowlike wings directed
anteriorly and projecting into the mantle sac (enclosing the liver). Three
dorsal pairs of arms with very wide ventral protective margins with dense
transverse supports.

Pfeffer (1912, p. 537) distinguishes two species of Thysanoteuthis:

T. rhombus and the new species T. nuchalis. I am not sure that the
two are different. 1) According to Pfeffer, the fins ofTh. nuchalis do
not reach to the posterior end, in contrast to those ofTh. rhombus; yet
this species has similar fins. 2) The posterior transverse neck fold bears
a crescent- shaped widening between the 2nd and 3rd longitudinal fold, a
character without sufficient significance. 3) The folds and swellings of
the "afterneck" are stated to be different from those ofTh. rhombus.
However, they differ even from one side to the other and the distinctive
characters are hardly constant. 4) The sculpture on the dorsal side of the
head, as described by Pfeffer for T h . rhombus, differs from that of
Th. nuchalis, but this sculpture is not constant inTh. rhombus but
depends on the contraction of the head musculature. 5) There is an inner
attachment between the dorsal arms of Th . nuchalis, i. e. a fusion of
464 the protective margins, but this is present also elsewhere as a minor

abnormality caused by contraction of certain cutaneous muscles connected
with the dorsal attachment of the buccal funnel. 6) All attachments of the
buccal funnel of Th . nuchalis are perforated, so that a circular
peribuccal cavity is formed which is bridged by these attachments and is
formed in our opinion by a fusion of the 6 normal buccal pockets. Th.
rhombus shows exactly the same conditions (see p. 468). The other
differences are still more doubtful.

444

I therefore assume that Th. nuchalis and Th. rhombus belong
to the same species, and the largest and only known mature male is then that
ofTh. rhombus. Th. nuchalis is probably a very closely related
form, perhaps a variety (2nd character). The genus Thysanoteuthis
and the family Thysanoteuthidae are characterized below on the basis of
Th. rh o m bu s , the only valid species.

THYSANOTEUTHIS RHOMBUS Troschel, 1857
a. DIAGNOSIS

Posterior transverse fold of neck not markedly projecting as a crescent-
shaped lobe between the 2nd and 3rd longitudinal folds. Fins reaching
posterior end but only as narrow ridges (cf. Loligo).

b. LITERATURE

1857 Troschel (p. 70, Plate 4.5), Thysanoteuthis rhombus.

1857 Troschel (p. 74, Plate 4), Thysanoteuthis elegans,

1881 Vigelius (pp. 150— 161), Thysanoteuthis rhombus.

1889 Weiss (p. 91), Thysanoteuthis rhombus.

1889 Carus (p. 445), Thysanoteuthis rhombus.

1889 Carus (p. 445), Thysanoteuthis elegans.

1896 Jatta (p. 56, Plate 9, Fig., p. 3), Thysanoteuthis rhombus.

1900 Pfeffer (p. 182), Thysanoteuthis rhombus.

1912 Pfeffer (pp. 523— 531, Plate 27), Thysanoteuthis rhombus.

1912 Pfeffer (pp. 531 — 534), Thysanoteuthis nuchalis (probably identical.*).
1916 Naef (p. 16), Thysanoteuthis rhombus.

1921 Naef (p. 538), Thysanoteuthis rhombus.

c. STRUCTURE OF THE ADULT ANIMAL

The general habitus of Thysanoteuthis resembles that of the
Loliginidae in the form and length of the fins (see Sepioteuthis) and in
the proportions of mantle sac, head and arms, but there is no special
relationship. Thysanoteuthis is a true oegopsid, closely related to the
Ommatostrephidae (p. 236; cf. Figures 238 and 249, Pfeffer 1912, Plate 27).

To understand this relationship, it must be realized that the
Ommatostrephidae and Thysanoteuthidae are highly specialized in many
characters and that a common type would resemble the general type of
Oegopsida more closely than the above families. The recently described
genus P s y c h r o t e u t hi s (p. 237) is intermediate in many characters,

465 and its gladius still has a typical flag (Thiele, 1921, Plate 53, Figure 9).
However, the shells shown in Figures 230 and 247 have diverged from
this condition.

The gladius of Thysanoteuthis (Figure 247) is of a very characteris-
tic form and gives the morphologist much material for thought. How is such
a structure at all possible in the Teuthoidea? The strongly modified anterior

445

parts have to be traced back (also ontogenetic ally) to normal preceding
conditions (Figure 63 on p. 150 and Figure 65 on p. 153) and have to be
incorporated in the framework of the soft body. Their secondary
modification can only be caused by a marked change of the correlations.

As in the Ommatostrephidae, the gladius is situated on the inner side of the
mantle sac and is surrounded by the muscular mantle, but the surrounding
is still incomplete anteriorly and the rhachis is visible (Figure 249b). In
contrast to the Ommatostrephidae, however, the cone is absent, at least in
the adult, although it certainly has to be assumed to be present in early
youth. The gladius of Thysanoteuthis has instead a wide, leaf- shaped
flag of abnormal form, the wing- shaped anterior processes of which project
beyond its connection with the rhachis. The spread flag is as shown in
Figure 247a. In the animal it has the form shown in Figure 247b, with the

466 wings curved upward, and detached from the muscular mantle (see p. 470),
not as in Figure 247a or as illustrated in Jatta (1896, Plate 9, Figure 10).
This whole abnormal part of the flag is a new development which separates
Thysanoteuthis distinctly from all other Oegopsida. Otherwise, the
shell resembles that of Loligo, although rostrum and cone are absent

(p. 150). The rhachis is a simple, narrow, uniformly thickened median ridge,
with an angular anterior end as in the Ommatostrephidae. Marginal ridges
strengthen the posterior part of the flag. The fins extend along the entire
side of the mantle except for a small anterior and posterior part. They
form together an oblong rhombus the margins of which are curved as in the
Ommatostrephidae. The fins are widely separated and situated almost on the
sides of the mantle sac (Figure 249b). The apex of the fins is particularly
characteristic. This part, however, is damaged in most of the specimens.
After years of storage in alcohol in an upright position, the posterior end
has been crushed, but the fins apparently do not reach the posterior end
but are subterminal (Figures 249, 251). However, the groove between fin
and mantle continues ventrally to the apex of the body, and a ridge forms
an incomplete continuation of the fin to the posterior end or to near the end

467 (cf. Loligo, p. 200). The anterior end of the fin forms "earlobes." The
mantle sac is cylindrical to spindle-shaped; it tapers rapidly posteriorly
to a blunt apex which becomes more slender in larger animals. The

3 typical corners of the anterior margin are weakly developed.

Funnel pit and neck folds resemble those in the Ommatostrephidae, but
the 1st (dorsal) longitudinal fold is absent and the 2nd (lateral) fold projects
slightly beyond the transverse fold. There is a deep pocket in the funnel
pit on each side near the funnel at the level of the neck folds (Figures 248
and 249). The funnel adductors are visible in the pit near the anterior
attachment of the funnel if they are folded back; the inner adductors are
normal, the outer adductors are displaced laterally, but much weaker
than in the Ommatostrephidae. The head becomes increasingly cylindrical
with age. The lid margin, eyes and window also resemble those of the
Ommatostrephidae. The arms, on the other hand, are different (Figure 249).

The 3rd pair of arms is by far the longest; then follows the 2nd pair and
then the 1st and 4th pair, the latter two being of almost equal length
(formula: 3, 2, 1, 4). The dorsal arms bear swimming margins which
widen rapidly from the base, forming an angular projection, and then taper
gradually. This is very characteristic for the family. The swimming

446

margins of the 2nd and 3rd pair are as in the Ommatostrephidae: those
of the 2nd arms correspond to the widened ventral outer edge and taper
slowly toward the distal end, and the margin of the 3rd arm is situated
on the outer side and forms a sharply projecting lateral corner near the
base.

(465)

(465)

FIGURE 247. Gladius of Thysanoteuthis
rhombus. 0.5X. a) flattened; b) curved
together as in the animal. The anterior
wings are then detached from the muscular
mantle.

FIGURE 248. Neck and head of Thysanoteuthis
rhombus, natural size, after a large specimen from the
Collection of Naples University. Mantle length 26 cm.
Note; mantle margin; recurved funnel and valve; funnel
pit with lateral pockets; funnel adductors; 3 longitudinal
folds and anterior transverse fold of neck; typical form of
eyes.

The inner surface of the arms bears 2 rows of small, densely arranged
suckers, which are toothed all around. The teeth on the distal margin are
longer than on the proximal margin; the teeth resemble a rake in form and
arrangement. The protective margins are well developed everywhere,
especially the ventral margins of the 3 dorsal pairs of arms; they are
extremely wide, exceeding the width of an arm on the 1st pair, twice as
wide as an arm on the 2nd pair, and 3 times as wide on the 3rd pair. They
are strengthened by very dense transverse supports which project like
ridges on the upper side. All protective margins taper rapidly toward the
base of the arm, as in the Ommatostrephidae (Figure 2 23), and reach the
buccal funnel as simple edges.

447

FIGURE 249. Y oung Thysanoteuthis rhombus from the collection of the Zoological Station in
Naples. Reconstructed after the specimen illustrated by Jatta (1896, p. 61), natural size. Note the
form of mantle and fins; the rhachis, which is still visible anteriorly; neck folds and funnel pit; the
rounded head; relative length of the arms; development of swimming and protective margins on the
different arms; the delicate tentacles and the structure of the clubs. The two small crosses near the
club indicate the location of the two larger distal adhesive knobs. These are still absent on the stalk
part, but the small tubercle close to and in front of each sucker apparently belongs to the basal pad
and perhaps replaces a knob (?).

The tentacles (Figure 250) are very delicate, with a narrow stalk margin
on the outer side. A zigzag row of small suckers occupies about the distal
half of the "stalk" so that this zone forms morphologically the "stalk part"
of the club. The base of the widened part of the club bears 4 slightly larger,
still biserial suckers. This is the carpal part. Then follows the uniformly
quadriserial hand part, on which the suckers are larger. The suckers of
the median rows of this part are slightly but distinctly larger than those of
the marginal rows. The suckers of the distal part become abruptly smaller,
especially in the dorsal marginal row; the change is least marked in the
ventral rows. An adhesion apparatus is present only on the stalk part and
hand part. In older specimens, the suckers of each longitudinal row of the
stalk part alternate with low adhesive knobs surrounded by a deepened ring.
Such knobs are (permanently?) absent on the carpal part. The first two

448

468 particularly small suckers of the dorsal marginal row of the hand part
alternate with 2 larger adhesive knobs as a continuation of the adhesion
apparatus.

FIGURE 250. Right tentacle of Th y s a n ot e u th i s rhombus.
0.5 X. Drawn after a large specimen of the University Museum in
Naples. Dorsal mantle length 26 cm (cf. Figure 248). Note the
stalk without suckers; the zigzag pattern on the stalk part of the
club; the 4 carpal suckers; the enlarged suckers of the hand part
and the distal part with the small, quadriserial suckers; swimming
margin; protective margin, 'adhesion apparatus of stalk part and
hand part (the intercalated adhesive knobs in the dorsal row of the
proximal part of the hand part).

The largest suckers of the hand part are uniformly toothed all around
and are always smaller than those of the 3rd arms. Protective margins
are present already on the distal part of the stalk part. On the hand part,
the margin forms rounded projections caused by the widening of the
supports. Each support corresponds to a marginal sucker. The larger
suckers bear outward -directed papillae or fringes on the free, soft marginal
ring, at least in large specimens (as in L o li go and Sthenoteuthis).

The swimming margin ends proximally near the end of the stalk margin,
slightly dorsal to it.

The buccal funnel is as in the Ommatostrephidae in all important
characters. At least the 4 dorsal buccal pockets communicate directly
below the attachments, but this could not be determined with certainty for
the ventral pockets (p. 464).

The inner margin of the funnel and the funnel and mantle bonds differ
distinctly from those in the Ommatostrephidae (Figure 251). The projecting
part of the funnel is more conical, narrow and delicate than in the
Ommatostrephidae. The inner margin of the funnel projects anteriorly in
an angle (Figures 251 and 243). The funnel bond is situated close to the
posterior end of each arm of the angle, forming a characteristic oblong
triangle with the longest side laterally, the shortest side posteriorly-
inward and the third side anteriorly -inward. Along the outer side passes
the main groove, from the middle of which begins a wide transverse
groove which runs toward the middle. Before this transverse groove
projects a large inner tubercle which constricts the main groove;

449

* behind the tubercle is a lesser elevation. A similar pattern is present on
469 the mantle bond: there is a longitudinal edge and a wide transverse edge
extending from it; a deep depression is situated before it and a shallower
depression behind it. This closure apparatus can be compared with that of
the Ommatostrephidae if it is assumed that the outer tubercle of the funnel
bond is lost and the transverse groove has become more complex. This
apparatus is not as firm as in the Ommatostrephidae, but has the same
function in principle (p. 417). Thysanoteuthis is also a good swimmer.
This is proved by the extremely well developed muscular mantle
(Figure 251) and by the neck bond (Jatta, 1896, Plate 9, Figure 5) which is
comparable to that of the Ommatostrephidae (Figure 205) if it is assumed
that the posterior part has become narrower, while the ridges along the
median longitudinal groove have become contracted and the marginal
grooves have reached the posterior end. On the other hand, the anterior
part is still wider, and the two small elevations lateral to the median
ridges in the Ommatostrephidae have developed into large tubercles which
reach to the median ridges. This is probably the most solid neck bond in
the Decapoda, except for the concrescences (p. 417).

FIGURE 251. Situs of mantle cavity of Thysanoteuthis
rhombus. Collection of the Zoological Station in
Naples. 0.5 x. The mantle is opened in the middle; note
its unusual thickness. The figure shows the posterior part
of the head with funnel pit, neck folds, funnel, and funnel
and mantle bonds. Note particularly the organs of the
mantle cavity: funnel retractors (m), funnel organ, and
papilla; posteriorly; hind intestine and terminal part of
ink sac, on the right: vena cava; between body and funnel
retractors "lateral pockets" (y), renal pores (n), gill base
with the posteriorly open pockets, narrow gills with the
ligament; further posteriorly; veins of the fins (v) with
the cutaneous bridges which support them, median pallial
artery (a), median pallial septum;

z — marginal pan of the gladius, translucent; g — genital
process; d — funnel gland; x — point of anterior part of
lateral margin of gladius; t — larger tubercle of funnel bond;
tj — smaller tubercle of same; q — transverse ridge; 1 —
longitudinal ridge of mantle bond.

The jaws are very characteristic (Plate XVII, Figure 12). In contrast to
the Ommatostrephidae, the biting processes are short and thick. The
process of the lower jaw is also curved in a very unusual manner and is
apparently a cracking device, because of the large, pointed lateral tooth.

5110/2

450

The radula is of the type of the Ommatostrephidae (Plate XIV), but the teeth
are very short and strong. I have not seen fresh specimens. The coloration
of live animals probably resembles that of Ommatostrephes or
Sthenoteuthis.

470 Many details of the mantle cavity resemble those of the Ommato-
strephidae, and others are specific for the genus (Figure 251). Except for
the funnel and mantle bonds, the following parts resemble those of
Ommatostrephidae: 1) the form of the funnel retractors; 2) the posteriorly
open pockets of the gill base; 3) (?) the formation of "lateral pockets"

(p. 418). All these three characters are explained by the structure of the
gladius and the characteristic relationships between gladius and soft body.

As in Loligo, the wide flag of the shell (Figure 251, z) is only covered
by the thin primary mantle and reaches the mantle cavity, projecting
laterally beyond the soft body; the gill base and the adjacent lateral
pallial vein have to cross this part of the gladius to reach the muscular
mantle (cf. Figures 206 and 95). The posterior vessels of the mantle follow
the same pattern, but they are markedly displaced anteriorly. The pocket
of the gill base breaks through posteriorly along the gladius, mechanically
stretched by its form. The abnormal lateral margin of the gladius is
detached from the mantle, beginning from the gill base, projecting towards
the funnel retractor and then following its posterior continuation. The
funnel retractor can therefore not be strongly contracted because it is
firmly connected to the margin of the gladius, and this explains the
characteristic form and position of the shell and the reduction of the
posterior part of the funnel retractors. It may be assumed on systematic -
morphological grounds that a similar condition probably existed also in the
Ommatostrephidae, and resulted in a similar formation. Not only the
reduction of the funnel retractors, but even more the perforation of the
pockets of the gill base and the (?) formation of the lateral pockets in both
families must be considered on the assumption that the shell was primarily
widened (cf. Figures 251 and 243).

The wings of the gladius extend far anteriorly and could be recognized
by touch, for example, at x in Figure 251. However, they are apparently no
longer firmly connected with the funnel retractors so that they have the
small room for action they need. The cephalopodium (p. 417) is no longer
rhythmically retracted and the activity of movement is concentrated in the
muscular mantle. The otherwise probably (p. 185) thick muscular cords of
the funnel retractors are here only moderately movable supports of the
funnel. The perforation of the pockets of the gill base, the reduction of the
funnel retractors and the specific form of the gladius of Thysanoteuthis
are thus apparently correlated to a modified mechanism of movement
comparable to that in the Ommatostrephidae (p. 417) and possibly explain
its development.

The fin vessels (posterior pallial veins and arteries) must be considered
in the same context. They must cross the margins of the flag (cf. Loligo,
Figure 82), forming cutaneous bridges but no perforations. Because of

* Cf. Sepiolidae (Chapter 42). As in the Sepiolidae, the consequences of the ancient form of the shell are
apparently preserved here, after the reduction of the shell. Ontogenetic studies may give information on
the subject. The topographical relationships suggest a secondary change; the entire posterior region of
the gills (Figure 215 pn p. 427) of young Ommatostrephidae appears to be torn apart by a wide gladius
which no longer exists, and also the lateral pockets could have developed by such a lateral stretching.

451

471 the growth of the fins anteriorly, these bridges are displaced anteriorly,
as in the Sepiolidae (Figure 339), and form pockets which open anteriorly;
the common trunk of the fin arteries, a typical character of the Oegopsida,
probably disappeared for the same reason (cf. Figure 243 on p. 460).

The gills are typical but very long and narrow. The renal pores
resemble those of the Ommatostrephidae (Figure 207; the animal in
Figure 251 has a wide round opening on the right side* and a low papilla
on the left formed by the contraction of the circular musculature). The
median pallial septum extends far anteriorly. Hind intestine and ink sac
are situated as in the Ommatostrephidae: the hind intestine is displaced
to the right, the ink sac to the left, so that they are situated next to each
other. The branches of the dorsal part of the funnel gland are visible inside
the funnel opening and extend posteriorly to the funnel retractors, as in
S t h e n o t e u t h i s (Figure 243) and Loligo (Figure 95).

Luminous organs have not been found. The female genitalia could not be
examined. As in the Loliginidae (Figures 80, 95), the male genital process
is fused with the body for some distance and becomes free only some
distance from the gill base. This process is present only on the left side.
Pfeffer (1912, p. 533) described the hectocotylization of "Th. nuchalis."
If this form is a valid species (p. 463), Th. rhombus would show a very
similar condition, i. e., a slightly modified distal half of the left ventral arm.

d. POSTEMBRYONIC DEVELOPMENT

I was unable to obtain very young stages; they probably resemble
Figure 209 but certainly have normal tentacles as in Figure 118. I could
not acquire new material in Naples, but the embryo illustrated on Plate VIII
in Vol. II and the freshly hatched animal in Figure 67 probably belong here,
but this was not mentioned in the legends. Later stages are illustrated by
Pfeffer (1912, Plate 27, Figures 26 — 33); Issel (1920, Plates 1—7) recently
illustrated similar forms. These publications give a clear picture of the
postembryonic development. Small specimens with a mantle length of
about 4 cm have very short, typically larval, transversely produced fins of
about V4 of the mantle length. The protective margins of the arms are still
very narrow, the tentacles armlike, uniformly covered with suckers
almost their whole length. Head and mantle are unusually wide, almost as
wide as the mantle is long (3.3 and 3.8 mm). These young stages are very
plump (Pfeffer, Figures 26 — 27; Issel, Figures 1—4).

At a mantle length of 7 mm, the semicircular fins occupy already half
of the sides of the mantle; the ventral protective margins are markedly
widened, especially on the well -developed LV arms. The V arms resemble
the dorsal arms in length and form and have wide outer margins.

472 Formula: 3, 2, 1, 4. The tentacle club is beginning to be formed and
probably differs little from its later structure (Figure 250; see also
Pfeffer, loc. cit.. Figures 28 — 31; Issel, Figure 5).

* Because of an unsolicited "improvement," this detail is missing in Figure 251; the original shows an
ink spot symmetrical to n.

452

At a mantle length of 14 — 16 mm the animal already shows the general
habitus of the adult (Figure 249). The fins occupy about Va of fho sides of
the mantle and have distinct but still very blunt lateral corners. The
anterior protective margins of the dorsal arms are very wide, the
tentacles almost completely developed (Pfeffer, Figures 32— 33; Issel,
Figure 7).

The above description is based largely on a young specimen
from the collection of the Zoological Station in Naples (Figure 249),
with a dorsal mantle length of 59 mm. It shows distinctly definitive
characters, except for the sexual differentiation and the later lengthening
of the body. Observations on larger specimens (Figures 248 and 251)
were given for comparison.

453

473 Chapter 38

SUBORDER SEPIOIDEA
Naef, 1916

(==Myopsida s e p i a e f o r m e s Naef, 1913)

Contents: a. Diagnosis. — b. Establishment of the suborder. — c. Typical structure of the adult animal
(p. 47). — d. Individual development of typical Sepioidea (p. 490).— e. Variation of the type of Sepioidea
(p. 492).

a. DIAGNOSIS

Phragmocone (if not absent by reduction) curved ventrally at the
posterior end. Free lower margin of shell situated in the body; muscular
mantle attached to outer side of sheath, on swollen ridges, edges or flat
elevations (Figures 255, 257, 268). Fins not united in the middle at the
posterior end (Figures 275, 303). Axis of gills not perforated, i. e.
without longitudinal canal between the afferent and efferent vessels
(Figure 17b). Teeth of radula always with a single, blunt cusp. All forms,
except Spirula, live near the bottom and burrow into the sand or mud.

The characterization by an ancestral form is particularly necessary
because only this can establish the relationships between the families. The
reader should study Figures 255 and 260, which show the typical relation-
ships of the shell and their correlations with the soft body.

b. ESTABLISHMENT OF THE SUBORDER

I discussed the general reasons for my classification of the
Dibranchiata, published in 1916, in 1921 (pp. 528 — 534), especially for
the suborders of the Decapoda.^'' As this is decisive for the general review
and morphological interpretation of the most important and best known
types and also discusses the nature and position of many fossil forms, it
would be useful to describe how I obtained results which apparently
474 escaped many specialists. The progress was based on two factors:

1) fresh material was examined; 2) the methodical study of the typical
relationships has resulted in a logical synthesis of the existing variety
of forms.

I had false assumptions when I began work in 1910. One of the most
important was the apparently natural classification of the recent

* See also my monograph on fossil Cephalopoda, Jena, G. Fischer, 1922, p. 38 ff.

454

Decapoda into Myopsida and Oegopsida (this was rejected on pp. 149 and 155,
and the unification of the Loliginidae with the Oegopsida proposed).

However, I still confused the Loliginidae with the other ’’Myopsida" and
placed these two groups in opposition to the Oegopsida. Against the
prevalent view (cf. Naef, 1911), I had come to the conclusion that the
Oegopsida are derived forms, while the Loliginidae are the primitive form
of Decapoda. As Was shown above (p. 155), this conclusion was not quite
wrong and could be supported by anatomical evidence. However, it did not
recognize the true nature of the systematic relationships as the problem
was incorrectly posed. It was necessary to discuss the phylogenetic
(systematic) relationships of the recent Decapoda in general. The
morphologist, however, cannot easily rid himself of deeply rooted concepts
because the infinite variety of phenomena becomes confusing. The structure
of the Myopsida had to be traced back to a fossil type, particularly the
Belemnoidea, mainly based on the shell. This group, however, proved very
difficult: the "Myopsida" (Sepiidae, Sepiolidae, Loliginidae, Idiosepiidae)
are so different that a common origin (type of shell) seemed highly unlikely.
The Idiosepiidae have no shell; the Sepiolidae and Loliginidae have a not
calcified "gladius," and the Sepiidae have a calcified shell. The shell of
the Sepiidae seemed most promising as a source of morphologic al-
phylogenetic information, especially during ontogenetic development. The
chambered, calcified shell could be related morphologically to the
Belemnoidea.

The development of the shell of the Sepiidae did not show closer relation-
ships to the Belemnoidea, but to the recent Spirula, which had been
incorrectly classified as belonging to the Belemnoidea. In addition to typical
characters (chambering, parts of siphuncle, etc.), the embryonic shells of
Sepia (Figure 301) also show regular eccentric growth and an at first
marked curvature which gradually decreases during development
(Figure 314). I realized that this shell had to be compared with the
"flattened" shells of snails (Naef, 1911, pp. 91, 98), in which the reduced
curvature no longer permits a typical spiral coiling. The relationship
between Sepia and Spirula corresponds to that between Hali ot i s
and Polytremaria, and the contrast between the two shells was
apparently more external and secondary, even though it was so striking.

The comparison of shells of young and embryonic Sepia was facilitated
475 by the description (Joubin, 1910) of a young Spirula with 5 air chambers,
a homologous stage which finally proved that the shell of the very young
Spirula is completely internal, like that of other Dibranchiata. This had
been suspected previously, but the speculations of Pelseneer (1895) made it
doubtful (see Lang (Hescheler), 1900, p. 72). The two shells of "Myopsida"
were similar not only in their general form but also in details of their finer
structure (cf. Appellof, 1894) and in that the air chambers were displaced
into the dorsal part of the animal in both cases. The proostracum was
absent already in the early stages with a few chambers. Chun (1910) showed
that Spirula is "a true myopsid." Particularly striking was Chun's
drawing of the situs of the mantle cavity, which is reproduced here slightly
diagrammatic ally in Figure 280. A number of characters strikingly
resemble those of Sepia: particularly the structure of the gills and the
position of the renal papillae with their specific structure. I concluded that
Spirula is more closely related to Sepia than to any other "Myopsida".
At any rate, the two genera had to be considered as closely related and it

455

could be assumed that their shells could be traced back to a common
ancestral form despite their apparent differences. This could only be done
by intermediate forms among the fossil Spirulirostridae and Belosepiidae,
as I suggested in 1911 (p. 91). This was later confirmed.

FIGURE 252. Diagrammatic median section through a young Spirula, 14 x, after a series of sections of
C. Chun. Leipzig (see Naef, 1913, pp. 454—461; Chun, Plate 73, Figure 1). Note the general form of the
shell and its position in the body, especially the position of the opening of the shell between the viscera.
The atypical origin of the muscular mantle on the outer side of the air chambers, ventrally, behind the
curved initial chamber. The absence of the proostracum is striking; it permitted an atypical displacement
of the dorsal margin of the shell into the inside of the body. The proostracum is replaced by the muscular
mantle, which is otherwise absent in the dorsal midline (see Figure 23 on p. 91 and Figure 33 on p. 110).

1 — 6 — air chambers; Ps — prosiphuncle; Sa — first caecum of siphuncle; Sd — septal neck; Sw — shell
septum; Sc — siphonal coelom; Si — fleshy siphuncle; Fg — interstitial tissue; Co^- C63 — parts of coelom;
Go — gonad; X — ventral margin of the shell; Ma — stomach; Bl — caecum; Hz — heart; Gg — gastric
ganglion; Ni — kidneys; Mm — muscular mantle; Ed — hind intestine; Tb — ink sac; Af — anus; Mv —
mantle cavity; Vc — vena cava; Lb — liver; Ao — aorta; Md — dorsal mantle cavity; Oe — esophagus;

Gd — poison gland; St — statocyst; Vg — visceral ganglion; Nk — neck bond; Nh — collar bond; Cg —
cerebral ganglion; Pg — pedal ganglion; Ob — dorsal buccal ganglion; Ub — ventral buccal ganglion;

Dr — funnel gland; Tk — funnel valve; Tr — funnel; A1 —outer lip; Sr — subradular organ; J1 — inner lip;
Uk — lower jaw; Ob — upper jaw; Rd — pocket of radula.

476 By the courtesy of Chun, I was able to examine younger and older stages
of Spirula in Leipzig in 1912 and 1913; Figures 252, 253 and 281 are
based on these specimens. The information obtained permitted a detailed
comparison of Spirula and Sepia and explained the nature of a type of
Decapoda which was unknown until then in its specific character. This type
(Figure 252) requires closer attention because of its fundamental
importance: it shows a number of more or less general characters which
are not present in recent Dibranchiata. 1. The general proportions of the
chambering which are typical for all Cephalopoda: the inflated initial
chamber; the blind, caplike initial part of the siphuncle, which is not
calcified, like the prosiphuncle, so that a direct connection would be
formed between the initial chamber and the siphuncle during fossilization;
the structure of the siphuncle with the central coelomic canal; the general
arrangement of the viscera, funnel and organs of the head. 2. Characters
of Dibranchiata are the envelopment of the shell by the shell fold and the
development of a special muscular mantle. 3. The presence of the neck

456

bond, funnel valve, coelom and a chambered shell proves that Spirula
belongs to the Decapoda. 4. The type of the Sepioidea is shown in a) the
ventral curvature of the phragmocone; b) the displacement of the origin of
the muscular mantle to the outer side of the shell, far from the free margin
of the shell; c) the penetration of the free margin of the shell into the soft
body. The penetration of the shell caused marked changes in the soft body;
in particular, the part of the coelom situated in the shell opening (living
chamber) becomes more or less separated (Naef, 1913, p. 459). This
separation created a narrow medium pore in this preparation, although Chun
(1915, p. 474) doubts that the pore is open (it is partly clogged in the
preparation''^). The genital ligament (p. 58, Figure 9), which originates in
the dorsal part of coelom, is also cut off so that the gonad remains
attached to the stomach only on one side. (This takes place not only in
Spirula but also in the Idiosepiidae and Sepiolidae, its closest recent
relatives, although these have no shell, at least in the adult.) 5. A special
character of this shell is the absence of a proostracum: the two halves
of the muscular mantle are apparently connected in the dorsal midline and
have thrust off or squeezed out the dorsal part of the shell (Figure 253a).

The muscular mantle is therefore inserted on the dorsal margin of the
phragmocone, but is apparently subsequently displaced to the outer side
of the shell also in this case. Thus, even the dorsal mantle cavity could
extend posteriorly beyond the margin of the shell, as the figure shows
(see Chapter 39). The resulting conditions strikingly resemble those in the

477 Octopoda in that the muscular mantle forms a continuous wall like a barrel
which also dorsally encloses a large part of the mantle cavity and permits
extensive movements.

The distinct characteristics and correlative importance of the above type
of shell became clear already in 1913, when I established the group of
Sepiaeformes as opposed to the Loliginiformes within the "Myopsida."
Although the gladius of Loligo could not be easily related to the shell
of Spirula, I did not at the time attempt to abolish this artificial
connection, mainly because of the similarity of the embryonic forms of
the "Myopsida" (see p. 151, also Vol. II, Plates II, XV, XXIII). However,
sharp contrasts between the Sepioidea and Teuthoidea have since been found
(Vol. II) also in the embryological development.

Later (1914) I studied shells of Belosepiidae and Spirulirostridae in
the State Paleontological Collection in Munich and obtained a clear concept
of the relationships between the young forms (Figure 253b, c). Both fossil
forms are clearly intermediate between Spirula and Sepia in the sense
that Spirulirostra provided the key to the understanding of the other
forms so that their more or less direct derivation could be considered.
Belosepia, wrongly considered as a transition to the Belemnoidea
(Lang, 1900, p. 99, Figure 107), has a true shell of Sepiidae but resembles
Spirulirostra in the distinct ventral curvature of the initial part, the

478 markedly convex first chamber and the typical formation of the first
septal neck.

This is not essential in this case: the homology of the parts of the coelom which Chun did not recognize
is evident from the topography of the whole complex of organs (Figures 252 and 23), and not from this
connection, which probably later becomes loosened (?), although this would deprive the siphonal coelom of
an outlet.

457

(477)

FIGURE 253. Morphological relationships between S pi r u 1 a and Sepia. Juvenile stages of S p ir u 1 a (a),
S piru lir ostr a (b), Be lose pi a (c) and Sepia (d), diagrammatic, a — a young planktonic animal;
d — an embryo (Vol. II, Plate XIX); b and c are reconstructions after nuclei of shells (juvenile shells) of
fossil shells. The morphological comparison should begin from b, only in which a typical proostracum is
preserved.

The muscular mantle and cephalopodium are dotted in the figure, while the shell fold is shown transparent,
as it is in young S piru la and embryos of Sepia, so that the shell and its chambering are clearly visible
through the thin skin. (See also figure in "Fossile Tintenfische, " p. 39.)

These relationships became of even greater interest because the inner
organs are markedly affected by the form of the Spiral a -like shell and
its atypical position in the soft body, as the examination of Spirula had
shown (Figure 252). This is naturally also true for Spirulirostra and
the forms derived directly or indirectly from it. This particular form of
the shell prevented not only the primary attachment of the muscular mantle
at the free margin of the shell, and caused its penetration into the soft body,
but it also led to other important anatomical changes (see p. 486, also
Naef, 1913, p. 456). This resulted in important changes of the primary type
of Decapoda, so that these forms were placed in a separate position. I also
found a number of related fossil types the shells of which showed the
characters of Spirula described on p. 476: 1) strong ventral curvature,
especially in the initial part of the phragmocone; 2) penetration of the
phragmocone into the mantle sac, recognized by the origins of the muscular
mantle and the impressions of vessels on the outer side of the shell. This
left no doubt that the Sepioidea are a systematic unit within the fossil and
recent Decapoda and can be derived from a common ancestral form (p. 480).

It became clear now that the Loliginidae could no longer be placed together
with these forms and a problem became urgent of which I had already
become aware during the early anatomical study of the recent Cephalopoda
(p. 149), i. e. whether the resemblance between the Oegopsida and the
Loliginidae does not show a closer relationship between these groups. This
was decided by re-examination of the material, especially on the basis of
paleontological data. The forms grouped in 1916 as "Sepioidea" were related
to certain fossil types of Cephalopoda, and this was also the case with the
"Teuthoidea, " consisting of the Loliginidae and Oegopsida. The Teuthoidea
479 could be traced back through the series: Teuthopsis, Beloteuthis,

( Tr achyteuthi s ) , Geoteuthis, P a r a pi e s i o t e u t h i s to forms

458

closely resembling the Belemnoidea, although in a different manner in
principle than the Sepioidea, so that a mixing of the two series was
excluded. Already the earliest Teuthoidea had no phragmocone, while it
has been preserved in the Sepioidea (Spirula, Sepia), although in a
modified form. Such typical Teuthoidea occur at least from the Lower
Jurassic (Lias 7); on the other hand, Sepioidea are absent in the Jurassic
and Cretaceous, and appear only in the Paleocene.'!'

FIGURE 254. Characterization of the Sepioidea and their systematic-phylogenetic predecessors in
diagrammatic median sections of hypothetical young stages with 2 air chambers, a) Type of Cephalopoda
(Orthoceras); b) type of Dibranchiata and Decapoda (Belemnoidea); c) type of Sepioidea. Figure b
shows a higher stage, because the shell is already surrounded and partly reduced, and the muscular mantle
is developing; c shows a further advanced stage, caused by displacement of the typical parts in b. The
curvature, which extends to the proostracum, is slightly exaggerated (cf. Fossile Tintenfische, p. 42).

1, 2 — first septa; 3 — first caecum of siphuncle; 4 — 2nd septal neck; 5 — dorsal wall; 5a — proostracum;
6 — primary mantle (dorsal); 7 — anal papilla; 8 — shell wall (ventral); 9 — primary mantle (ventral);

10, 10a — free margin of shell (ventral); 11 — shell fold, point of closure; 12 — muscular mantle; 13 —
displaced insertion of the muscular mantle.

The old classification of Decapoda was thus proved invalid and the new
classification proposed was confirmed.

c. TYPICAL STRUCTURE OF THE ADULT ANIMAL

None of the known species could be considered as the ancestral form of
the Sepioidea, and therefore I established in 1915 an ideal type by methodical
comparison of all known recent and fossil forms (cf. "P rotosepioides,"
Figure 255). The parts not shown in this figure are assumed to resemble
those of Spirulirostra (Figure 260), as both forms are based on data
obtained from the recent Sepioidea. I have later seen a drawing of a fossil
species of Sepioidea (Figure 256) which fulfills all requirements as an
ancestral form of the group and resembles my construction in all important
characters. The ideal type will be described because of its essential
interest.

As the unchanged explanation of Figure 255 shows, I considered at first
the possibility of a blunt rostrum after I had found a new species which

It is interesting historically that Blainville (1827) placed Beloptera belemnitoidea demonstratively
next to Sepia (Plate 1), so that the general homologies became distinct. D'Orbigny (1826) also used the
name of 'Sepia" parisiensis for B. belemnitoidea. He (1842) created the family Spirulidae,
consisting of the fossil Belopteridae, Spirulirostridae and Spirula, and placed this family next to the
Sepiidae, which also included the Sepiolidae (1845). D'Orbigny thus understood the natural relationships
established here, although he did not formulate them clearly (cf. p. 149).

459

differed from my construction only in the blunt rostrum (Naef, 1921, p. 530;
also Figures 259, 263). Still more similar to my construction, however,
is the above-mentioned fossil, Belemnosella americana (Meyer
and Aldrich, 1886) Naef, 1922 (Foss. Tint., p. 49), placed at my disposal
by von Billow (1920). This proves that morphological constructions (p. 10)
can in fact prove the existence of real forms.

Like almost all forms derived from it, the extinct Belemnosella
(="P rotosepioide s") is only a few centimeters long (mantle sac about
6 cm long) and has also to be considered as a bottom form (exception;
Spirula). Belemnosella probably also burrowed in the sand or mud,
480 like the recent Sepiidae and Sepiolidae.'’' This fossil may also have been a
good swimmer, like the recent Sepia, because the modification of the
typical shell of Belemnoidea which is present in Belemnosella is a
marked improvement of the hydrostatic apparatus (see p. 483).

The general habitus probably resembled that of the recent Idiosepius
(Figure 275), because this shows the least atypical changes of the external
morphology (Sepiidae and Sepiolidae show secondary modifications of the
form and position of the fins and eyelid (cornea); their mantle becomes very
plump and the head thick. The marked curvature of the shell of Spirula
and the loss of the rostrum also cause secondary changes of the typical
form). The form of the shell causes a distinct lateral compression of the
pointed posterior end and forms a natural base for the small, probably
subterminal fins (Figure 260). In other aspects the form probably
resembled a small, slender Sepia.

FIGURE 255. Posterior part of body of Protosepioides:

a — lateral, with translucent profile of shell; b — median section. This drawing can be combined with
Figure 260 and the missing parts added. The proportions have to be slightly changed only in the posterior
part to make the reconstruction complete. The rostrum could also be considered to be blunt, about as in
Belopterina (Figure 264). However, this closely related genus probably already showed specializations
which lead to Beloptera, while the other forms derived from Protosepioides have a pointed rostrum.
Note particularly: 1) insertion of muscular mantle on the sheath near the phragmocone; 2) position of fins
on outer side of sheath; 3) uniform ventral curvature of phragmocone; 4) position of shell opening, deep
inside the visceral sac which is a basic contrast to the typical conditions in the Decapoda and Dibranchiata
in general (Figure 23 on p. 91 and Figure 37 on p. 110). This drawing has been completed and slightly
corrected in Figure 260 (cf. p. 479).

* They live almost exclusively on sand or mud bottoms. They burrow into the ground with the posterior end,
using the fins, until only the eyes remain exposed. They spend most of their life in this position, hiding
from enemies and waiting for prey, becoming more active only at night.

460

The shell shows the parts of the shell of Decapoda in general but it is not
completely known. 1. The delicate proostracum is broken off, as in all
fossil relatives, including most Belemnoidea; its existence can only be
concluded from the point of the break, as in the Spirulirostridae (Figure 269)
and Belopteridae (Figure 265), and from the presence of a proostracum in the
481 recent derived forms (cf. Sepiolidae). The form of the proostracum is not
exactly known, but it was probably also pliable, i. e., not completely
calcified, as in the Sepiolidae. 2) The phragmocone may be assumed to
resemble that of Spirula but much less curved and with more densely
arranged septa, and its diameter probably increased more rapidly. The
form of the phragmocone, the position of the septa and initial chamber are
proved by the corresponding cavity (alveole) of the fossil. 3. This consists
of rests of the conotheca and of a periostracum or sheath which forms a
thick layer on the initial part which becomes gradually thinner anteriorly.

The posterior end is pointed; a capitulum develops around the initial
cham'ber (Figure 260) the homologies of which will be discussed later.

The phragmocone bears large laterally swollen ridges or edges ("lateral
edges") which extend anteriorly and disappear rapidly toward the ventral
side. The latter, which is only partly preserved in the fossil, has a thin coat
of sheath substance which we name, together with the adhering conotheca,
the "ventral wall."

FIGURE 256. Belem nosella americana (Meyer and Aldrich, 1886) Naef,
1922. After the original drawing, completed (Plate II, Figures 26 and 26a);
added parts are dotted. Natural size. Compare with Figure 255, which resembles
this drawing, although its explanation was written long before this typical form
of Sepioidea became known. The smaller upper figures show Belemnosis
a noma la (de Sow.), natural size. (Fossile Tintenfische, p. 49.)

If we attempt to fit this shell into the typical mantle sac of a decapod, it
appears that the muscular mantle cannot be attached in a "belemnoid"
manner, i. e., on the free margin of the phragmocone; the shell would then
form a nonsensical process at the posterior end. The phragmocone must
be placed in a position as in the young Spirula (Figure 252 on p. 475),
and the attachment of the muscular mantle must be displaced from the typical
position in the anterior region, i. e., from the margin of the proostracum,
to the posteriorly adjacent lateral swollen ridges to the capitulum, where
both sides meet. This is obviously a markedly modified decapod shell.

The shell itself and also its relationships to the muscular mantle, fins and
viscera (p. 478) appear as the product of a metamorphosis which is difficult
to understand and has to be traced back to the typical conditions of Decapoda

461

482 and Dibranchiata illustrated in Figure 23 on p. 91. An interpretation of
these peculiar conditions is only possible if we consider the functional
relationships. We will first determine the special characteristics.

1. The muscular mantle of the typical Decapoda (Figure 37 on p. 110)
is attached on the primary margin of the shell, i. e., conotheca and
proostracum, whereas inProtosepioides it is displaced to the outside
of the phragmocone covered by a thick sheath. 2. This causes differentia-
tions for the insertion of the musculature on the corresponding parts of the
sheath, mainly on the "lateral edges" or "lateral swollen ridges" along
the phragmocone which become of great importance in the further
development of the shell of the Sepioidea (Figures 257, 268). 3. The opening
of the shell (i. e., the margin of the conotheca) is therefore necessarily
displaced from its typical position inside the muscular mantle to the
interior of the visceral complex, which creates marked changes in the
internal topography (Figure 252 on p. 475). 4. The phragmocone shows
distinctly typical characters (position of the siphuncle, structure of the
septal necks and prosiphuncle, etc.), but it shows a marked ventral curvature
which is atypical for the Decapoda but very characteristic for the Sepioidea.
5. The phragmocone is enveloped posteriorly by a massive rostrum, this
being apparently connected with the life of the animal on the bottom, which
makes an added weight of the posterior end and compensation for the
buoyancy necessary. 6. The fins, which are primarily situated near the
transition of the phragmocone into the proostracum, are displaced closer
to the posterior part of the shell, and become based on the outer side of
the thickened sheath (Figures 257 and 260).

FIGURE 257. Diagrammatic cross section of the fin region ofProtosepioides slightly obliquely from
a posterior -upper to an anterior-ventral point. Note the connection of the base of the fin with the sheath;
the structure of the lateral edges for the insertion of the muscular mantle on the sheath; the position of the
phragmocone in the soft body, the parts of which are situated ventral to the shell opening instead of being
surrounded by it (Figure 56 on p. 139);

Flj, Fh”" dorsal and ventral muscular layer of fin; Wt— articulation capsule of fin base; Co — conotheca;
Kr — fin cartilage; Si — siphuncle; Sk — lateral edge of sheath; Mm — muscular mantle; Ct — coelothelium;
Vk — ventral edge; Mh — mantle cavity.

462

These 6 characters form a very characteristic type of Decapoda and
suggest the possibility of further changes of the morphological equilibrium
to be discussed later. They give the clue for an understanding of the most
important recent Decapoda (Spi rula. Sepia and Sepiola).

483 One or the other of these phenomena can be considered as primary and
as the cause of the others. Thus, the ventral curvature may be the cause
of the displacement of the shell opening into the body. We do not accept this
but assume that the whole complex of changes developed together under
certain preconditions. They are mutually necessary, because the displace-
ment of the shell opening to the interior of the body (Figure 258c) is
impossible without a ventrally curved phragmocone, and such a curvature
cannot exist without a displacement of the attachment of the mantle. This
becomes most distinct in the young stages in which the sheath is still little
developed. The growth of the Spirulirostridae, Belopteridae and Sepiidae
shows that the sheath of all Sepioidea probably develops at a later stage.

FIGURE 258. Young stages of Decapoda, explaining the connection between the ventral curvature of the
phragmocone and the displacement of the attachment of the muscular mantle to the outer side of the
phragmocone with a displacement of the shell opening to the interior of the visceral complex:

a — typical condition (restoration of a young stage of Belemnoidea); b — hypothetical transitional stage;
c — Protosepioides; d — Belopterina.

The following considerations may provide an explanation, i. e., proof that
this change is effective. Forms like Or t ho c e r a s , Protodibranchus,
Protodecapus and the Teuthoidea (in contrast to the Sepioidea) actively
maintain their equilibrium in the water, by movements of the fins and
direction of the water jets from the funnel because the small overweight
caused by the ventrally situated siphuncle can hardly prevent rotation of the
axis. A structure of the shell in which the air chambers are situated more
dorsally, like the swim bladder of fish, is therefore a marked advance.

484 Such a displacement was already present in Protosepioides,

eliminating the need for active maintenance of the equilibrium. This also
applies to the most recent Sepioidea, the Sepiidae, which maintain their
normal swimming position even after death, in contrast to other
Dibranchiata.

463

The formation of a strong sheath with a massive rostrum in the
Sepioidea is probably mainly of ecological significance. Like the rostrum
of the Belemnitidae, this structure acts as a counterweight to the air-
containing phragmocone and thus permits the animal to stay horizontal and
to swim horizontally without muscular action. If, as we assume,
Belemnosella burrowed into sand or mud with the posterior end first
(p. 480), the rostrum was a valuable tool (cf. p. 110).

Figure 255 shows that the curvature of the phragmocone inhibits the
growth of the rostrum. Any point at the end of the phragmocone is moved
constantly downward during development, and if this point bears the
"anlage" of the rostrum, there must be a progressive change in the position
of the rostrum with respect to the longitudinal axis of the body. This must
somehow be controlled in the swimming animal and this takes place, in fact,
in the Sepioidea as shown by the growth lines in Figure 255b. The sheath
thus grows eccentrically, and the shell substance is deposited dorsally more
rapidly than ventrally, so that the mass of the rostrum is not only displaced
in progressive compensation to the dorsal side but the apex is also displaced
dorsally, if the growth of the phragmocone tends to direct it ventrally. The
axis of growth of the whole structure is thus not straight but curved in a
direction opposite to the curvature of the phragmocone in order to maintain
the motor axis and the direction of the apex. The earliest stages of
Sepioidea with few chambers still have no rostrum, and its "anlage" is
therefore no longer situated in the continuation of the morphological axis
of the phragmocone (as in the Belemnitidae, Figures 38 and 39) but is
displaced more or less dorsally. In the Sepioidea this displacement is
accentuated in the degree in which the primary curvature increases. If the
"anlage" of the rostrum is situated as shown in Figure 253c, it would be
situated between the 3rd and 4th chambers, not on the 1st chamber as in the
Belemnitidae (or even still in Belemnosella, respectively P r o t o -
sepioides, provided that the degree of curvature shown in Figure 255b
applies to this form). On the other hand, the dorsal displacement of the
rostrum by eccentric growth is restricted. If the rostrum were very
long, deposition would create a lumpy formation in profile^!' and therefore
an increasing divergence of the apex. A greater curvature of the phragmo-
cone would have similar consequences. These possibilities can be controlled
485 by various means: the growth of the rostrum could be delayed if the

curvature were greater (Spirulirostra, Figure 271); if the rostrum
were very long, this could be compensated by a decrease in the ventral
curvature to which the rostrum cannot adjust itself (Beloptera, Figure 265).
Figure 265). An excessive curvature is only possible when the rostrum
is completely reduced (Spirula, Figure 277).

The curvature of the phragmocone necessarily decreases gradually as the
animal grows. This has nothing to do with the so-called biogenetic law and
does not permit the conclusion that the curvature of the phragmocone was
more marked in the ancestors.

* A thick rostrum is a general character of the older (fossil) Sepioidea and is certainly connected with the
ventral curvature, as explained above. The predecessors of the Sepioidea among the Belemnoidea
probably also had a thick rostrum.

464

FIGURE 259. Diagrams illustrating the development of the shells of Teuthoidea and Sepioidea and
their morphological contrast, a) ancestral form of Decapoda; b) ancestral form of Teuthoidea;
c) ancestral form of Metateuthoidea; d) form of Belemnoidea with a massive rostrum, from which
the type of Sepioidea must be assumed to have evolved; e) transition; f) ancestral form of Sepioidea,
better shown in Figure 260, i. e. with a pointed rostrum. (A similar corrected illustration is given
also in "Fossile Tintenfische, " 1922, p. 31.)

The following fossil genera correspond to these^diagrams: a — Nannobelus Pawlow; b —Para-
ple si ot eu t hi s Naef; c — Pa 1 a e ol ol i g o Naef; d — Diploconus Zitt.; f— Belemnosis
Edwards. The shell is shown white, transparent, the muscular mantle transversely striated. Note
in the Teuthoidea (b, c); the reduction of the phragmocone; in the Sepioidea (e, 0; the displacement
of the attachment of the muscular mantle to the sheath if the curvature of the chambered shell increases.

1 — Median asymptotes; 2 — lateral asymptotes; 3 — lateral plate; 4 — ventral attachment of mantle;

5 — lateral attachment of mantle (lateral edge); 6 — free margin of shell; 7 — proostracum.

We have no information about the transition from the typical form of the
Belemnoidea to such a form in the Decapoda. The closest form could be a
Diploconus - like form with a short but thick sheath and a short
phragmocone; this best explains the displacement of the attachment of the
muscular mantle from the margin of the conotheca to the outer side of the
sheath {Figure 259d). As the systematic position and classification of the
Sepioidea is the most important problem, the history of the group will not
be discussed at this stage."

* It must be remembered that also the typical Belemnitidae showed a marked ventral curvature of the
phragmocone and that there are similar consequences for the growth of the rostrum (cf. p. 484), i.e. a
predominantly dorsal deposition of shell substance was necessary to keep the continuously diverted
rostrum in the body axis. The resulting eccentricity and curvature of the "axis thread" increases the
stronger the curvature of the phragmocone and is usually distinct in the long rostrum of the Belemnitidae.
(Cf. Naef, 1922, Fossile Tintenfische, especially Figure 71.) However, they never reach a degree which
may indicate a position of the shell in the body as in the Sepioidea (cf. Figures 38, 39 on p. 111).

The ventral curvature of the phragmocone is necessary because swimming Decapoda need a spindle-
shaped mantle, and this causes the curvature of the proostracum (Figure 37 on p. 110). During the
ontogeny, the proostracum gradually develops into a conotheca and this assumes the curvature of the
phragmocone if the dorsal plate is fixed by calcification. If the dorsal plate remains thin and pliable
for a long time it can be straightened when it is incorporated in the phragmocone.

465

486 The anatomical changes in the soft body caused by the penetration of the
shell opening cannot be discussed here in detail. However, one point drew
my attention in Spirula (Naef, 1913, p. 458), although its general
significance was not clear at the time. The penetration is facilitated by the
large coelom in the posterior part of the body which permits the displace-
ment of the various parts (Figures 37 and 260). This causes some
difficulty for the attachment of the gonad, which is attached by the "genital
ligament," a remnant of the dorsal mesenterium, which is attached antero-
ventrally to the stomach and dorsoposteriorly to the shell above the
entrance of the siphuncle. The posterior part of the genital ligament is
detached when the shell margin cuts into the body (Figure 260). This
explains that the gonad of the typical Sepioidea (Spirula, Sepiola,
Idiosepius) is attached only to the stomach and not to the floor of the
coelom in the adult animal (This is secondarily changed in Sepia because
of other factors; see Figure 285.) This attachment of the gonads caused by
the specific structure of the shell of Sepioidea can persist (Sepiolidae,
Idiosepius) even if the shell has become rudimentary and no longer
requires it. The previous attachment persists as a symptom of the
relationships before the shell became reduced. This condition is also
connected with other factors, mainly modified embryonic conditions.

Many of the forms derived from Protosepioides have small,
rounded subterminal fins, which are typical for young Dibranchiata in
general, and we must assume that similar organs were present also in
Protosepioides (Figure 260). The exact position of the fins can be
determined as follows. 1) It has to be considered that some descendants
(Idiosepius and S p i r u 1 a) still have fins of typical form. 2) The
assumed position of the fins must agree with the typical course of the
vessels (posterior pallial artery and vein). 3) Special indications (vessel
imprints) in related fossils have to be considered (Figure 264). All these
considerations result in the picture described on p. 487.

Because of the changed structure and the displacement of the shell, the
articulation capsule of the fin base is displaced far posteriorly on the outer
side of the sheath to its lateral edges (Figures 257, 268), i.e. to a position
which is very suitable for it in Beloptera and Spirulirostra. The

487 structure of the fin and its articulation are apparently not changed from the
typical conditions in the Decapoda (p. 114).

The mantle margin of Protosepioides is typical for the Decapoda,
with 3 distinct corners, which are extremely well developed in Spirula
and should therefore be more marked in Figure 260 (cf. Figure 276).

The neck folds of the Teuthoidea are absent and, in contrast to the
Teuthoidea, which are much stronger swimmers, the funnel pit is shallow,
as the direction of the water jet is less important (cf. p. 414).

The olfactory organ is a round papilla in the youngest animals (Plate XIX,
Figure 7); its margin is later raised, so that the olfactory epithelium is
situated in a shallow depression (Plate XIX, Figure 5). Its position is
typical (Figure 37).

The funnel shows no special characteristics, but the neck bond of the
Sepioidea has to be assumed to have been less narrow than in the recent
Teuthoidea (Metateuthoidea). If it were to become narrower within the

488 group, this would be connected with a narrowing of the proostracum
because of the progressive development of the muscular mantle at the
expense of the shell (cf. Figures 64, 291, 323).

466

(487)

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467

ganglion; Bo — dorsal buccal ganglion; Bu — ventral buccal ganglion; Mb — mouth cavity; Ok — upper jaw; Z — tongue; S — subradular organ;
Nh — neck bond; Pr — proostracum; II — inner lip; A1 — outer lip; Sh — membrane between dorsal arms; B — buccal pillar (oral arm); Sa —
swimming margin of dorsal arm; Sc — swimming margin of tentacle club; Tk — funnel valve.

The primary eyelid of Protosepioides is assumed to be wide open,
as in Spirula and the Oegopsida (Figure 277). My experience with
Spirula and young Oegopsida (Plate V, Figure 5) shows that strong
stimuli (preservation) cause a contraction of the lid to a narrow pore which
persists at the position of the typical sinus (Figures 118, 119, on p. 254).

This may have been the permanent condition in the adult Proto-
sepioides, because its burrowing or benthic habits could result in
contamination of the orbit with sand grains or other material. The recent
Spirula has abandoned the benthic mode of life and has again become a
permanent swimmer; this is possibly the reason for abandoning the
permanent closure of the orbit. On the other hand, observations on young
abnormal specimens of Sepia with artificially produced eyes of
Oegopsida without closure of the cornea (inhibited development) did not
show any disadvantages.

The arms of Protosepioides resemble those of Protodecapus
(p. 117, cf. Rossia, Sepia, Spirula), with the 4 rows of suckers
on the arms and 8 on the tentacle clubs, at least on their greater part. The
club shows the following characters of the Sepioidea: 1) the swimming
margin extends to the base of the club and is situated close to the dorsal
protective margin along its shole length; 2) the club is always sickle-shaped
at rest, open upward and markedly widened in the middle, mainly because of
the greater development of the ventral protective margin. The stalks of the
suckers are fused with the margin and 2 or more ventral marginal rows of
suckers are displaced to the margin. These rows are longer than the dorsal
rows on this type of club, so that a typical and very characteristic pattern
is formed (cf. Sepiidae and Sepiolidae): the ventral marginal rows of the
Sepioidea are displaced into each other and the 8 rows are apparently
reduced to 6 — 7. (The pattern of 16 or 32 rows is correspondingly
modified, and the number of suckers in the ventral marginal rows is
correspondingly increased.)

The pockets of the tentacles still show the typical condition preserved in
the Sepiolidae (Figure 322) and Spirula- : they are narrow passages and
follow the stalk to its base behind the base of the ventral arms so that the
strongly contracted stalk can be partly retracted into the pocket. The
ligament on the inner side of the base of the stalk is pulled more or less
into the pocket and is no longer (cf. Figure 76 on p. 17 7) visible from the
outside if the tentacle is contracted. The vein of the tentacle passes on its
489 margin and opens in the circumoral venous ring, but not directly as in the
Teuthoidea but passes first on the inner wall of the pocket.

The situs of the mantle cavity shows no special characters, but the renal
papillae are lengthened like a chimney and displaced closer to the anus
(cf. Sepia and Spirula, Figures 280 and 300). An important character
of all recent Sepioidea is the absence of a longitudinal canal in the axis of
the gills between the afferent and efferent vessels. The Sepioidea probably
inherited this condition from the Belemnoidea so that the Teuthoidea show a
new development. However, the presence of a gill canal in the Octopoda
(Figure 392) suggests that this part could have been present in Proto -
dibranchus, from which it passed to the Octopoda and Teuthoidea, while
its development was suppressed in the Sepioidea. The small size of all

* The secondary expansion observed in the Teuthoidea (p. 139) is therefore absent, although it recurs
in a different form in the Sepiidae.

468

primitive Sepioidea, which also show a general disposition to simplification,
is in favor of this view.

The nidamental and accessory glands were probably typical (p. 125).

The gonoducts must be assumed to have been bilaterally symmetrical,
at least in the female, as in Idiosepius. The male gonoduct, on the
490 other hand, was probably only developed on the left side. The right gonoduct
of the male is absent in nearly all Dibranchiata, and this suggests that the
common ancestral form had at least a tendency to such an asymmetry
(p. 105). There is no right gonoduct in the male of the Sepioidea.

FIGURE 261. Situs of mantle cavity of typical Sepioidea/ natural size. As in
Figure 280, the ventral part of the muscular mantle is cut off; the cut passes
posteriorly close to the attachment of the muscular mantle to the sheath.
Comparison with Figure 272 shows the insertion of the shell; the inner side (lateral
edge) of the shell is not completely covered (as in Loligo, Figure 95 on p. 205).
This figure is based on the conditions in the Idiosepiidae, Sepiolidae and Sepiidae
(see below) and also on Figure 280 after elimination of the characters caused by the
specific form of the shell of Spirula. The body is much more slender, the organs
stretched but not markedly different otherwise; the gonoducts are visible and sym-
metrical, as in Idiosepius. Note the typical renal papillae (2), nidamental
glands (4), accessory glands (3), genital processes (1), funnel bonds and funnel
retractors, musculus rectus abdominalis (5), and posterior pallial arteries (9) and
veins (10), which extend towards the fins, branchial vein (6), lateral pallial veins
(7), median pallial artery (8).

The hectocotylization of Protosepioides probably resembled that of
Spirula, Idiosepius, some Sepiolidae and the Sepiidae in that only the
left ventral arm, or both ventral arms, were affected. It cannot have been
very marked, because the different forms of hectocotylization in the recent
Sepioidea (p. 232) do not show a uniform basic type; the only common
character is the reduction of the suckers.

The jaws of the Sepioidea, and therefore also of Protosepioides,
show in general the typical form of the Decapoda and closely resemble those
of the typical Teuthoidea (Loliginidae, Enoploteuthidae); the resemblance is so

* The rostrum is shown much too large. The drawing was originally intended to show Spirulirostra
(Figure 272) as a type of the Metasepioidea (but see p. 147). The proostracum, however, would be too
short for Spirulirostra (Figure 269), and the body is not slender enough. At any rate, the figure may
illustrate these relationships and can be corrected after Figures 260 and 256.

469

marked that no distinction is possible (Plates XVII and XVIII). On the other
hand, the radula of all Sepioidea examined (that of Spirula was not
available) has a specific structure: the marginal platelets are completely
reduced and usually not recognizable, while in the other 7 rows, the marginal
teeth are the longest and the median teeth the shortest. All teeth are simple,
i.e., without lateral cusps, as in most Teuthoidea and Octopoda (Plate XV;
also Figures 31 and 14).

d. INDIVIDUAL DEVELOPMENT OF TYPICAL SEPIOIDEA

Our knowledge (Figure 253) of the ontogeny of the fossil Belemnosidae,
Belopteridae and Spirulirostridae is based only on the structure of the shell,,
and the embryonic stages of the closely related recent forms (Spirulidae,
Idiosepiidae) are still unknown. We have therefore to restrict ourselves
to information on the development of other recent forms, the later stages of
Spirula, and the fossil shells.

The eggs of Sepioidea (p. 126) are relatively large and rich in yolk, so that
a major part of the ontogeny takes place inside the egg. In contrast to the
Teuthoidea, the young animal therefore externally resembles the adult to
some extent. This can be attributed also to the ancestral form, although the
state of the freshly hatched animal cannot be exactly determined (but cf.
Spirula, Figure 279). The earliest stages of development of the shell
have to be assumed to resemble that of other Decapoda or Dibranchiata
(Figure 36 on p. 107). At any rate, it may be assumed that the cone of
the embryonic shell (i. e. the still open initial chamber) shows the typical
correlation to the soft body (Figure 262a, b). The muscular mantle is
primarily attached to the free shell margin also here and follows its
entire circumference. This is confirmed by the comparative material
available at present (Sepiola and Sepia; Figures 287, 321); future
491 studies may show the condition in corresponding stages of Spirula, or
in a form more closely related to the type.-'=

It may be assumed with certainty for Spirula and according to
observations of other recent forms, that the insertion of the muscular
mantle is later displaced from the free margin of the shell (or from the
margin of the shell sac) to the outer side of the phragmocone (Sepia,

Figure 287a), i. e. to partly differentiated parts of the sheath or the
secondary shell epithelium. This must apply also to the typical form of
Sepioidea (Figure 26 2d) and must take place earlier because of the
accentuated ventral curvature of the phragmocone, which is otherwise of
typical structure (Figure 253 on p. 477 and Figure 258 on p. 283).

The arms show also evidence of direct development in the typical
Sepioidea. The arms develop rapidly a large number of suckers, usually
in 4 rows already in young embryos. The strictly larval conditions
observed in the Oegopsida (p. 233) and to a lesser extent in the Loliginidae
(p. 188) do not occur here. The clubs of the tentacles are also in a
completely developed stage at hatching, with 8 — 16 rows.

* I found a larva in Chun's material the mantle sac of which closely resembles that in Figure 362a. Unlike
Chun, I considered this larva as a young Spirula. Later examination of my drawing, however, made me
sceptical; this is probably a very young larva of Oegopsida (cf. Figure 472; also Naef, 1913, p. 455).

470

FIGURE 262. Typical development of the Sepioidea. Older embryos and median sections. Comparison with
Figure 36 on p. 107 shows that the conditions shown are mainly those of the Dibranchiata or Decapoda but
the curvature of the proostracum is more marked (slightly exaggerated), and the initial chamber is therefore
directed downward. The ventral margin of the shell penetrates into the soft body, and the origin of the
muscular mantle is displaced slightly to the outer side. These tendencies become necessarily more marked
later in proportion to the ventral curvature of the phragmocone. To arrive at the later stages, we must
assume that there was a similar metamorphosis in ontogenesis as in the systematic (phylogenetic) development
(Figure 254 on p. 478):

1 — margin of proostracum; 2 — olfactory papilla; 3 — primary eyelid; 4 — attachment of mantle on outer
side of phragmocone; 5 — attachment of mantle on ventral margin of shell; 6 — ventral margin of shell,
penetrating the soft body; 7 — displaced attachment of mantle; 8 — anterior margin of muscular mantle.

The ventral curvature is accentuated, about as in Spirulirostra; this is exaggerated for the ancestral
form. The muscular mantle is shown contracted; the posterior part of the body has to be assumed to be
almost vesicular.

492 e. VARIATION OF THE TYPE OF THE SEPIOIDEA

Contents: 1. Family Belemnosidae (p. 493). — 2. Family Belopteridae (p. 493). — 3. Family Spirulirostridae
(p. 496). — 4. Family Spirulirostrinidae (p. 501). — 5. Family Idiosepiidae (p. 503).

Although this work deals mainly with the recent Cephalopoda of the Bay
of Naples, another discussion of related fossil forms is necessary for a
better understanding of the natural relationship between the forms. The
relationships between the recent Sepioidea are still not clear, although the
establishment of the above type (p. 487) provides a general formula for
their understanding. The fossil Sepioidea also fail to give a complete
picture of the history of the suborder, which is surprising in view of its
relatively young age (Paleocene). The fragmentary paleontological

493 evidence is due to the small size, delicate structure and littoral habits of
the animals. With the exception of the very atypical family Sepiidae, the
recent and fossil forms of the suborder (Belemnosella, Belemnosis,
Belopterina, Beloptera, Spirulirostrina, Idiosepius and
most Sepiolidae) are less than 10 cm long. This is certainly connected
with the structure and mode of life of the group and it determines many
specific characters of their partly juvenile constitution. The fossil

471

Sepioidea are divided into the Belemnosidae, Belopteridae, Belosepiellidae,
Spirulirostridae and Spirulirostrinidae (Naef, 1921, p. 536). Also included
in the suborder are the fossil and recent Spirulidae and Sepiidae, and the
strictly recent Idiosepiidae and Sepiolidae, the shells of which are reduced
or lost. Fossils of these groups can hardly be expected.

(492)

FIGURE 263. Belemnosis cossmani Naef, reconstructed after Cossman (1895, Plate 1, "Belem nosis
a noma la"). 2x. a) Lateral view. The ventral wall (6) of the phragmocone, the outline of the mantle
(11) and the parts of the shell situated above the line of fracture (x) are reconstructed; the position of the
phragmocone inside the alveole is shown by dotted lines, b) Ventral view, completed anteriorly; the
greater part of the ventral wall of the phragmocone is missing posteriorly; the septa are broken off, but their
lines of attachment (4) are visible, c) Ideal cross section near the shell opening. The attachment of the
muscular mantle (9) on the "lateral incrassations" (5) of the sheath is reconstructed after Spirula and other
fossil Sepioidea (Figures 265 and 266) to illustrate the insertion of the shell into the mantle sac:

1 — proostracum; 2 — conotheca; 3 — marginal thickening of proostracum as a continuation of 5; 4 — line of
attachment of a septum; 5 — "lateral incrassations"; 6 — "ventral wall"; 7 — capitulum; 8 — blunt rostrum;

9 — muscular mantle; 10 — siphuncle; 11 — outline of mantle; 12 — skin.

I have published elsewhere a monograph of the fossil Sepioidea (1922,
Fossile Tintenfische) and we will deal here only with species which shed
light on the morphology of the recent forms.

1. The Belemnosidae (Naef, 1921) are represented by shells from the
Eocene which resemble more or less closely the ancestral form of the
Sepioidea (p. 487) (cf. Fossile Tintenfische, pp. 48 — 53). The family includes
Belemnosella americana (p. 481 ) and B e 1 e m n o s i s cossmanni

472

(Figure 263), which are very instructive. Before my acquaintance with
B. americana, I considered B. cossmanni for some time as the
ancestral form of the whole suborder (Figure 255 on p. 480 and Figure 259
on p. 485). Before I knew B. americana, B. cossmanni could be
considered as the first member of a series which leads through
S p i r u 1 i r o s t r i d i u m obtusum (Figure 266) to S p i r u 1 i r o s t r a
and the Sepiidae.

FIGURE 264. Belopterina le v e squ e i (d'Orb.), reconstructed after
a specimen from the State Collection in Munich. 2X. a) Lateral
view of the shell fragment with completed outlines and soft parts.

The reconstruction was made in comparison with Bel opt era (Fig-
ure 265) and according to the "lateral edges," i.e. the attachment of
the muscular mantle. The position of the typical fins of Sepioidea was
determined by the course of the fin vessels of Beloptera tFigure265),
in which the impressions are more distinct than in the less differentiated
Belopterina. The anterior part of the body shows the typical
characters of all Sepioidea. b) Median section through posterior part of
the body. Note the curvature of the phragmocone; position of rostrum;
the convexity above the embryonic chamber, corresponding to the
"capitulum" of Belem nos is (p. 487). c) Cross section through the
middle of the body, showing the attachment of the muscular mantle.

Note the assumed passage of the lateral edges to the margin of the
proostracum in comparison with Figure 266.

Pr — proostracum; Sk — lateral edge; VI — ventral ridge; W1 — capitulum;
Ro — rostrum; Ek — embryonic chamber; Sw — shell septum; Si — siphuncle.
From Fossile Tintenfische, p. 54.

2. The Belopteridae on the other hand (s. restr. Naef, 1921) are
completely isolated but they show a number of interesting characters,
because the predecessors of the recent Sepiolidae perhaps belonged to this
family (for further details see Vols. II and III). Species of Belopteridae
(Fossile Tintenfische, pp. 53 — 60) are Belopterina levesquei (d'Orb.)

194 and Beloptera longa Naef (Figure 265). Figure 264 shows a well

preserved specimen ofB. levesquei from the collection in Munich, with
distinct differentiation of the surface. This species has a plump, flask-
shaped, posteriorly truncate rostrum the outline of which closely resembles
that of Beloptera (Figure 265). An indicated capitulum forms a slight
swelling at the apex of the phragmocone at the position of the initial
chamber (cf. Protosepioides, p. 487). As in the young Beloptera,
and in Spirulirostra and Spirula, the typical ventral ridge of the
sheath runs in the median line at the ventral wall (Figure 257 Vk,

Figure 264 VI).

The lateral edges are of greater importance; their role as places of
attachment for the muscular mantle is quite clear (Figure 264c) and explains
the incrassations in the Belemnosidae (Figures 256 and 261). These edges
are thin lines which extend to the anterior margin of the fragment and the
attempt to reconstruct them suggests that they continue into the lateral
margins of the destroyed proostracum. The lateral edges end posteriorly
near the capitulum, and behind it is an indentation (Beloptera, Figure 265b).

473

495 We assume with good reason that the fin vessels pass through the mantle
at this point (p. 496). The soft parts are shown only diagrammatic ally and
they probably resemble Figure 260 in details. Belopterina is thus still
closely related to our concept of the ancestral form of the Sepioidea (p. 487).
On the other hand, the form of the rostrum of Belopterina resembles
that ofBeloptera so closely (p. 494) that we cannot consider B e 1 o p t e r i n a
as a direct transitional stage toSpirulirostra, Spirula and the Sepiidae.

(494)

FIGURE 265. Beloptera longa Naef, reconstructed after a specimen from the Munich Collection,
natural size, a) Median section, b) Ventral view of fragment, completed by a dotted line and placed in
the soft parts (mantle situs), c) Cross section showing the attachment of the muscular mantle to show the
maximal width of the lateral wings, d) Cross section in about middle of phragmocone. e) Cross section of
anterior end of phragmocone (i.e., the available fragment). Note the connection between phragmocone and
rostrum; the formation of the sheath of the phragmocone; attachment of lateral wings on both; position of
shell opening in the soft body; curvature of phragmocone, which decreases anteriorly; special form of the
rostrum; position of hypothetical "terminal organ"; passage of fin vessels (Vp, Ap) through the mantle
(cf. Figure 264) and the assumed position of the stellate ganglion.

Bt — buccal funnel; A1 — outer lip; 11 — inner lip; Ok — upper jaw; Zg — tongue; Ob — dorsal buccal
ganglion; Ub — ventral buccal ganglion; Cg — cerebral ganglion; Nk — neck bond; Sd — poison ("salivary")
gland; Pr — proostracum (which could also be narrower); Ph — phragmocone; Or — terminal organ (hypo-
thetical); Mm — muscular mantle; Hz — heart; Ni — kidneys. From Fossile Tintenfische, p. 57.

A variation on this reconstruction has to be described. The relative length of the proostracum is obviously
uncertain, while it can be determined more or less exactly in the Belemnitidae. The proostracum may be
much larger and the animal correspondingly more slender. Evidence for this is present in Spirulirostra
(Figure 269), in which the phragmocone projects markedly beyond the widened lateral edges. If we assume
the same here, it follows on the basis of recent knowledge that the body must be more slender and the mantle
sac proportionally widened because of the corresponding widening of the shell opening. The expanded
mantle sac can then be inserted naturally on the heel of the boot, instead of on the margin of the sole, because
it is curved around it. The "terminal organ" is then included in the frame of the muscular mantle, which
here probably serves as a buffer (cf. p. 495), as this region contributes little to contraction.

474

FIGURE 266. Spirulirostridium obtusum Naef (from Fossile Tintenfische, 1922, p. 62). A fossil
form of Spirulirostridae from the Lower Oligocene in Haring near Kufstein (Bavarian State Collection in
Munich), a) Ventral view; the dotted parts of the outline and the anterior part of phragmocone (2) and
proostracum (1) are reconstructions (Figure 269). b) Reconstructed median section through the posterior
part of the body, c) Actual median section; growth axis of rostrum (y) indicated by a dotted line; x —
remaining stone (cement marl), d) Cross section at the anterior end of c. The dotted half has not been
exposed. Muscular mantle (8) and skin (13) are indicated, e) Lateral view, given as a mirror image for
comparison with c and completed anteriorly (dotted line). All drawings 2x.

1 — proostracum; 2 — ventral wall of phragmocone; 3 — lateral edge; 4 — capitulum; 5 — rostrum;

6 — ventral ridge; 7 — mantle septum; 8 — muscular mantle; 9 — attachment at the capitulum;

10 — prosiphuncle; 11 — siphuncle (initial caecum); 12 — 2nd chamber; 13 — skin; 14 — lamella of
sheath; 15 — eroded part of rostrum; 16 — point of 11th septum; 17 — not marked; 18 — fragments
of ventral wall; 19 — siphuncle; 20 — swollen beginning of lateral edge (cf. Figure 263).

Beloptera (Figure 265) shows the natural development of the lateral
edges of Protosepioides or Belopterina. The lateral edges are
widened into "lateral wings." Beloptera represents an interesting
parallel toSpirulirostra in this respect. The form and the assumed
growth of the rostrum are particularly instructive as a guide to understanding
the type of Sepioidea. The rostrum of the Belopterina described above
and other species of Belopterina forms a rounded, still undifferentiated
mass, while that of Beloptera is characteristically formed like a boot.

An attempt to attribute this particular form to a species of Belemnoidea
gives a highly improbable picture. However, if the rostrum is considered
as an endoskeleton which does not protrude from the body, its interpretation
is not difficult. The depression of the "sole of the boot" before the "heel"
gives the impression that it was occupied by an organ which could have
served as a shock absorber during an impact. We associate this structure
with the terminal organ of the related Spiral a (q.v.), the terminal
glandular sac of Sepiella (cf. Sepiidae), and the terminal organ of young
Sepiolidae (q.v.). This hypothetical structure will also be named terminal
organ (Figure 265, Or). The rostrum tapers characteristically anteriorly.

475

where the apex of the phragmocone is situated so that the shell has the
496 appearance of an hourglass. The primary curvature of the phragmocone
can obviously not regulate the growth axis of this highly specialized and
long rostrum for a long time (p. 484). This explains that a thorough early
secondary straightening of the chambered shell was necessitated. It is also
striking that the rostrum of Beloptera is strongly curved ventrally in
comparison with Belopterina and other related fossils (Figure 265a)
and that it increases in mass posteriorly as an attempt to continue the
regulation as long as possible. The ventral curvature of the rostrum
suggests an attempt to increase the curvature of the phragmocone. This
must result also in a dorsoventral widening of the mantle sac, to make
place for the developing gonad.

FIGURE 267. Posterior end of Spirulirostra bellardii d'Orb., reconstructed
after Fragment A of Figure 270 (cf. Figure 260). a) Lateral view. Note the
attachment of muscular mantle and fins; compare with Figures 268 — 271.
b) Median section. The growth axis (as in Figure 266c) is shown markedly
eccentric. The juvenile rostrum is situated on the outer side of the 4th chamber.
From Fossile Tintenfische, p. 63.

The "lateral wings" are situated on each side between rostrum and
sheath of the phragmocone. The lateral wings apparently do not reach the
proostracum anteriorly, but they extend far beyond the initial chamber
posteriorly. The lateral wings end with an indentation which is more distinct
in other species and in which the fin vessels were apparently situated as
in Belopterina (p. 495), because branched impressions of vessels
extend from here on the otherwise polished dorsal side. The fins were
probably situated here although they could have been correlated also with
the lateral wings. At any rate, the lateral wings suggest an anterior
displacement of the articulation of the fins, as in the Sepiolidae (Figure 274).

3. The Spirulirostridae (Naef, 1921) are of much greater importance
for the morphology of the recent types. They certainly fuKill all
requirements for an intermediate stage between Protosepioides or
Belemnosella and the Sepiidae. Because of the close anatomical and
embryological relations between the Sepiidae and the other recent families,

I considered Spirulirostra in the first draft of this work (1916) as a
representative of all recent Sepioidea, for which I established the
Metasepioidea (p. 49). More detailed examination of the fossil material
showed, however, that this group is artificial for the following reasons:

497 1) Some forms of Spirulirostridae (like in Figure 266) are closely related
toBelemnosis, so that there is apparently a double way from one

476

family to the other (cf. Fossile Tintenfische, p. 35). 2) True Sepiidae occur
together with their morphological predecessors (i. e., Belo sepia in the
Paleocene). 3) It cannot be proved that the Sepiolidae and Idiosepiidae are
more closely related to the Sepiidae and Spirulidae than a species of Sepioidea
in general, i. e., the resemblance between these groups goes back to their
common ancestral form (Figure 260). The Metasepioidea were therefore
again abolished (see p. 147). It is evident, however, that the Belemnosidae
are the first link in a morphological series which passes through the
Spirulirostridae, Spirulirostrinidae, Belosepia to the recent Sepiidae.
Spirula is apparently a side branch of Spirulirostra. An inter-
mediate between the Belemnosidae and Sepiidae can only be visualized like
Spirulirostra; if this were not actually known (although I had only
fragments at first), I would have constructed it like Protosepioides
(p. 480) and Spirulisepia (Figure 282).

FIGURE 268. Reconstructed cross section of Spirulirostra, to show the typical relationships between
shell, muscular mantle and fins. Compare with Figures 41 on p. 114, 56 on p. 139, 257 on p. 482 for an i
understanding of the displacement of the parts of the shell. The fin base becomes situated on the lateral
edges of the sheath as a preparatory step to the conditions observed in Sepia (Figure 290a):

Flj and FI2 ~ muscle layers of fin; Sf — lateral edges; Mm — muscular mantle; Ct — coelothelium; Ms —
mantle septum; Gh — cavity of gonads; Hd — testis; Vk — ventral ridge of sheath; Si — siphuncle; Ss — shell
sac; Wt — articulation capsule of fin base; Kn — fin cartilage, (cf. Fossile Tintenfische, p. 34.)

It should be stressed, however, that this is not a pedigree or phylogenetic
series (p. 31) of Sepia; it is not even a paleontological sequence.
Belosepia already existed in the Eocene, the Spirulirostridae are known
only from the Oligocene and Miocene, together with true Sepiidae. Spiruli-
rostridae have since been found in the Eocene of Australia (Figure 269), but
Spirulirostrinidae older than the Miocene (Figure 273), i. e., transitional
forms between Spirulirostra and Belosepia, are still unknown.
However, a negative conclusion cannot be drawn, because the probability of
fossilization of such small, delicate shells is obviously very slight (p. 493)

477

and the absence of Paleocene material does not exclude the existence of
Spirulirostrina at that time. We have only a few accidental finds
of fossils a few cm long (p. 493). The Belosepiidae of the Eocene were
much larger; some of their shells are more than 20 cm long (cf. p. 499).

498 A detailed description of the fossil Spirulirostridae is beyond the scope
of this work (cf. Fossile Tintenfische, pp. 60 —70), but we will describe a
number of morphologically important forms of shells.

Diagnosis. The Spirulirostridae are Sepioidea (p. 473) with the
following characters: posterior part of phragmocone markedly curved
ventrally. Capitulum large. Lateral edges projecting in form of plates
which extend laterally from the capitulum to the lateral margins of the
proostracum.

FIGURE 269. a — c — S p i ru 1 i r os t r a hoernesi v.
Koenen 1867, from the Miocene of Westphalia. Natural
size. Slightly reconstructed after the illustrations and
data of the author. Apex of body and proostracum have
been completed; the dotted lines indicate the lines of
breaking. The phragmocone in a is placed in its natural
position; those of b and c are placed accordingly. The
reconstruction is negligible and has a sound basis. Only
the form and length of the proostracum are doubtful.

The original or a closely similar specimen was examined
in Berlin.

1 - proostracum; 2 - free ventral wall of shell; 3 -
phragmocone (ventral wall); 4 - lateral wings; 5 -
capitulum; 6 — rostrum; 7 — last septum; 8 — free dorsal
margin of conotheca; a - lateral; b - dorsal; c - ventral,
d-f- Spirulirostra curta Tate; 1893, Plate 1,
Figure 1, natural size, Eocene of Australia. From
Fossile Tintenfische, p. 64.

Figure 266 shows a new species of special interest. This form still
closely resembles Belemnosis (Figure 263), especially in the blunt
rostrum and the swollen lateral edges at the posterior end. The other
Spirulirostridae have a pointed rostrum.

A detailed description of the shell of Spirulirostra is not
necessary. Figure 269 shows Sp. hoernesi based on direct observation
and with a few small corrections (after A. v. Koenen, 1867,=^' cf. Naef, 1922,
Fossile Tintenfische, p. 64). The structure of the rostrum is well known.

* Slightly corrected and completed, partly from the original specimens.

478

The rostrum resembles in general that of the Belemnitidae in its structure
with radial fibers and concentric layers. A distinct apical line indicates the
growth axis, which is markedly eccentric (ventral) and curved (Figure 271,
below). The sheath was only a thin coat of the phragmocone in early youth
(Figure 253); then appeared the rostrum, first as a blunt tubercle situated

499 on the 4th — 5th chamber. The proostracum must have been thin and pliable
in these stages to become constantly adapted to the curvature of the
phragmocone. A more solid proostracum would be an obstacle to the

: curvature of the phragmocone and it necessarily acted toward a secondary

straightening. This straightening became still more distinct by the
relatively late formation of the apex of the rostrum and the development
of widened lateral edges, which fixed the posterior end in the position
attained (Figure 267). A regulation like that described on p. 484 was thus
I only possible in a restricted degree, as in Beloptera (p. 496). In

' contrast to the marked curvature of the juvenile shell, there is here a

rectilinear lengthening of the phragmocone, which results in an ever more
slender form of the body. This lengthening provided space for the viscera,
especially the developing gonad (p. 496). However, the lengthening could
not be continued indefinitely because of the fixed form of the posterior end
and the size of this type of Sepioidea was therefore restricted. ’^ Only the
transformation into a shell of Sepia permitted a new and relatively
unrestricted enlargement."-'

The form of the soft body of Spirulirostra depends so largely on
that of the shell (Figures 266 — 272) that there can be no doubt about the
manner in which the shell is inserted. An attempt to place the shell of
another belemnoid into it (p. 127) would be impossible. The form of the
animal can be imagined by combining Figures 267 and 260. (The fossil
shells agree so well with the general picture that this is a strong argument
for the validity of the suborder.) The anterior part of the body may be
assumed to be as in the above figures, and there is thus no justification for
the establishment of a special type. The form of the mantle sac differs
in the various species. In Spirulirostra bellardii and
S. sepioidea (Figure 271, above), the mantle sac is strongly compressed
laterally; that of S. hoernesi (Figure 269) is probably more cylindrical
and has lateral edges which are often markedly widened posteriorly
(Fossile Tintenfische, p. 64, Figure 22c) (cf. Figure 306) but this probably
varied according to age and sex.

The mode of life is probably as was assumed for the ancestral form of
the Sepioidea (p. 480). This is suggested by the large rostrum which

500 increases -the weight of the posterior end. The large rostrum has to be

* This may be connected with the fact that the eggs of all Sepioidea are relatively large and not

numerous, in contrast to those of the Teuthoidea. The young Sepioidea hatch in a markedly developed
condition and with a relatively large shell. Some Sepiolidae grow only about 5—6 times of their length
at hatching (Oegopsida a 1,000 times or more). On the other hand, the embryonic chamber of this minute
form is 0.5 mm long, like that of large Belemnitidae.

This is a general problem. Hard parts which do not consist of live tissue (as in vertebrates) can only grow
by apposition. (Intussusception plays no role in the growth of the shell of molluscs.) The constant
enlargement is also connected with certain conditions of the general form, as in the case of a conical
or snail-like form of the shell. Differentiation, however, continuously creates structures which are not
restricted by these preconditions. Many forms overcome this difficulty by a secondary breakdown of
finished parts.

479

FIGURE 270. Two fragments of shells of
Spirulirostra, A and B, from the
Bavarian State Collection in Munich. 2x.
Ventral and lateral;

A — S pirulirostra be Hard i d'Orb.;

B — Sp. sepioidea Naef. Note the
attachment of the lateral wings (10) on the
capitulum (8) and their end at the proostra-
cum (1), which is situated between them.
Fragment B shows a remnant of the thin
sheath of the ventral wall with the ventral
ridge; B^ shows the ventral process (5) with
a transverse edge which is connected with
the lateral wings (10) by tubercles as a
preparatory step to the unification of the
origins of the muscular mantle on the
shell (Figure 282).

FIGURE 271. Reconstruction of
Spirulirostra. Above — anterior
view of fragment B. slightly dia-
grammatic. (The transverse break is
actually irregular but shown here as
smooth.) Note the swollen transition
(12) of the lateral edges (9) into the
capitulum (3); also the apparently
adjacent ventral process transversely
below it. 11 — septum; 10 —
siphuncle; 8 — ridge on ventral wall.
Below — median section through
fragment B (cf. Figure 267). Note
the course of the apical line (6),
which is drawn after fragment A and
is probably attached more ventrally
on the phragmocone and is also more
strongly curved (Figure 266). 5 —
growth lamellae; 4 — ventral pro-
cess with insertion of muscular
mantle; 3 — capitulum, and the
gonad above it.

Figures 270 and 271 from Fossile
Tintenfische, p. 66.

FIGURE 272. Position of the
shell of Spirulirostra in
the mantle sac. Reconstruc-
tion of the shell situs after
Figures 64, 73 and 291. 2x.
The upper drawing shows
the anterior end with pro-
ostracum (a), neck bond and
stellate ganglia, and the fin
nerves which run posteriorly.
The lower drawing shows the
posterior end, with the cut
insertion of the muscular
mantle at the ventral pro-
cess, the adjacent lateral
edges (e) and the entrance
of the fin vessels into the
mantle (at d). The removed
middle part has to be
assumed to be longer than
the gap (Figure 269).

considered as an indispensable burrowing tool, because large fins like those
of the Sepiidae and Sepiolidae'' probably were not yet present (cf. Spirula,
Figure 281, and I d i o s e p i u s , Figure 275). The mode of life of the young

* These forms first stir up the sand or mud by powerful strokes of the fins, then burrow in rapidly with the
posterior end and therefore do not (no longer) need a sharp tool for digging in.

480

forms and their structure probably resembled that of Spirula (Figure 253
on p. 477) and this provides a possible explanation for the derivation of this
type,* in which the differentiation of the sheath was delayed, and the
proostracum became reduced, etc.

501 The two fragments evidently belong to different species; a) is Sp.
bellardii d'Orbigny, b) the new species S p . sepioidea (Fossile
Tintenfische, p. 67), The difference consists in the proportions of all
parts, particularly the presence of a transverse edge in B which appears
in profile (Bj) as a large process (5, ventral process) on the ventral side of
the capitulum where the muscular mantle is inserted (Figure 267);
fragment A shows only small rugosities at this point. This process is
connected with the lateral edges on the capitulum by irregular tubercles
as a preparatory step to a uniform point of insertion (Figure 272e). If this
area formed a continuous wide ridge, it would already be a kind of dorsal
shield as in the Sepiidae (Figure 282) and its margins would contain the
following parts: 1) the newly formed ventral process; 2) the rugosities
leading to the lateral edges; 3) the lateral edges; 4) the margins of the
proostracum which is adjacent anteriorly.

FIGURE 273. Fossil shell of S p ir u 1 i r ost r i n a lovisatoi Canavari, natural size, from the "Argille
fanghiane di Fangano presso Cagliari" (Miocene). Original in the Museum of Stuttgart, a) Dorsal; preserved
and exposed parts indicated by solid lines, reconstructed parts dotted. The shell is distinctly Sepia -like,
b) Lateral view showing phragmocone and rostrum; outline of terminal part of mantle sac indicated by dotted
lines, c) Ideal median section, constructed after b. d) Ideal ventral view, based on characters of the
related Sepiidae and Spirulirostridae (cf. Figures 269 and 282);

Pr — proostracum; Ls — last septum; Si — last septal necks; Vw — ventral wall; Vr — free margin of ventral
wall; Lk ~ air chamber; Vk — air chamber on the forked part; Si — siphuncle; Sh — lateral edge; Cp —
capitulum with first chamber; Mm — muscular mantle; Vf — ventral process; Do — spine. (From Fossile
Tintenfische, p. 76.)

The recent Spirula (see Chapter 39) is related to Spirulirostra.
The resemblance between the two genera was already recognized by

* The young Sepiolidae are also better swimmers than the adults, which explains the development of the
type of Heteroteuthis.

481

d'Orbigny (1842); Spirula is today considered as a Spirulirostra with
a reduced sheath but no detailed comparison has been made.

The derivation of the Sepiidae has to begin with Spirulirostra
sepiodea and continues to Spirulirostrina.

4. The family Spirulirostrinidae (Naef, 1921) contains the interesting
Spirulirostrina lovisatoi Canavari, 1892. I did not have
illustrations or well preserved specimens of this species until recently;
the published data do not give a good picture, or else it would not have been
nec essary to construct an ideal transitional form from Spirulirostra
toBelosepia (Figure 282), as is the case of P r o t o s e p i o i d e s (p. 479).

Diagnosis. Shell slender, delicate, intermediate between Spiruliros-
tra and Sepia. Rostrum, ventral process and capitulum still completely
developed but much smaller than in Spirulirostra. Lateral plates
502 forming narrow stripes along the phragmocone, which is straight anteriorly
and compressed dorsoventrally. Septa drawn posteriorly and ventrally and
therefore convex and oblique. First chambers still resembling those of
Spirulirostra but slightly compressed dorsoventrally and obliquely
depressed. (Further data belong to the reconstruction, see Figure 273; also
Naef, 1922, Fossile Tintenfische, p. 75). Spirulirostrina lovisatoi
Canavari 1892 is the only species of the family. It has been mentioned in
various publications (Zittel, Grundziige, 1921, p. 593), but it has never been
correctly described'^' or illustrated; I described it in detail from a specimen
from the Natural History Museum in Stuttgart (loc. cit.). Its general habitus
resembles Sepia. This is shown in Figure 273, which is reminiscent of a slen-
der shell of Sepia. However, Sepia officinalis L. is quite unsuitable
for comparison (see Figure 315). On the other hand, Spirulirostrina
resembles Spirulirostra in the following characters: 1) outline of
rostrum, ventral process and capitulum; 2) distinct ventral curvature of
posterior part of phragmocone; 3) structure of first chambers; 4) secondary
straightening of chambered shell; 5) general outline of lateral edges. All
this is recognizable if Figures 273 and 269 are compared. However, other
characters show a distinct resemblance to the Sepiidae: 1) the initial part
of the phragmocone is markedly dorsoventrally compressed, the first
chamber is also not rounded and vesicular; 2) the following chambers
become more and more flattened, and the septa are not situated on the
conotheca at a right angle but distinctly obliquely; the suture is curved
ventrally and posteriorly also in lateral view (Figure 273b); 3) the lateral
edges run straight along the straight part of the phragmocone; 4) rostrum
and capitulum are relatively smaller, as in the Sepiidae; 5) ventral process
curved posteriorly. Compare the respective considerations about
Spirulisepia (Figure 282a). As in S p i r u 1 i s e p i a , we assume here
a marked widening of the siphuncle, at least in the anterior part of the
phragmocone which corresponds topographically to the "incrassation"
in the Sepiidae and also a reduction of the ventral wall, although this cannot
yet be reduced to a fork.''”’'

* Sacco (1904, p. 6) assumes, for examples, that this form is represented only by isolated phragmocones of
Spirulirostra. As in this genus, the marl acids often destroy the sheath, so that only the chambered shells
are preserved. Spirulirostra, however, is usually embedded in sand and the rostrum is therefore better
preserved.

These details could have been established by preparation of the shell embedded in marl, but I was not
permitted to do this. This could perhaps be done with new and more abundant material from Cagliari.

5110/2

482

FIGURE 274. Typical form of Sepiolidae (a) and Idiosepiidae (b) (diagrammatic;
arms shown slightly too short). Compare with Figure 275, and note position of fins,
shell rudiment (1) and eye pore (2):

a — about IX; b — about 3 X .

503 Nothing is known about the soft parts of Spirulirostrina and we
assume therefore that it is as shown in Figure 260. The characteristics
of the series of Sepiidae are apparently as follows: 1) the secondary lid
encloses the orbital pore (in contrast to the Sepiolidae); 2) the ink sac is
displaced asymmetrically to the right of the hind intestine during
development (to the left in Spirula). Further details may be found in
the anatomical part.

FIGURE 275. Idiosepius parad oxu s Ortmann, 1888, male from the Bavarian State Collection in
Munich (collected by Doflein, 1904/5, obtained by H. Lauter from Todohokke (Hokkaido), Japan),
a, b; 4X; d; 7 X; h — hectocotylus. Note form and position of fins, posterior end and mantle margin,
biserial tentacles. Dorsal side: (a) an oval posterior area with glandular (opaque) skin above the
presumed place of the widened part of the shell. Ventral arms (c) with only one basal sucker, which is
characteristic for many males of the family; otherwise, only supports are preserved as low papillae. Left
ventral arm with a subapical lateral lobe and a slight median terminal widening. Fold of cornea
(primary lid) contracted, with small radial folds (pore?) surrounded by a secondary lid fold, which partly
covers the pupil.

Idiosepiidae Appellof, 1898.

Beloptera and Spirulirostra are related to Spirula and
Spirulirostrina and the Sepiidae; they are also related to the
Idiosepiidae and Sepiolidae, which are themselves obviously closely related
and probably of common origin (Appellof, 1898). Methodical comparison
shows that the habitus of the Idiosepiidae (i. e., form of arms, head, mantle

483

sac and fins) resembles the general type more closely, while the Sepiolidae
have a more derived, specialized character. We shall therefore discuss
the group as represented by Idiosepius. This is mainly characterized
by the marked reduction of the shell. The adult Idiosepius has not even
a remnant of a shell (Figure 275). In the Sepiolidae, on the other hand, a
not calcified "gladius" is usually preserved, i. e., a delicate proostracum
in the anterior part of the dorsal side (Figure 321b). The embryo, however,
has a large shell sac of the typical form and topography of the Sepioidea.

A phragmocone with rostrum and lateral plates is not developed, but the
"anlage" can be concluded from the form of the shell sac (Vol. II). The
ontogeny of these structures in Idiosepius would probably show still
504 more distinctly the typical characters of the shell of Sepioidea; at any rate,
reminiscences of previous conditions could be expected to occur there
in less disturbed form. (cf. p. 486).

The Sepiolidae have a typical adductor pallii med. This muscle is
present but weaker in Idiosepius (Figure 326), but it is absent in the
Loliginidae, Oegopsida, and other Sepioidea (but compare the Octopoda, and
also Lepidoteuthis Joubin, p. 166). This and other similarities (p.503)
make it necessary to infer that the Sepiolidae developed from an
Idiosepius - like predecessor which is characterized by a reduction of
the shell to a rudimentary proostracum and parts of the lateral edges as a
"gladius flag" and the development of a median mantle adductor. A special
character of the Sepiolidae is the exclusion of the orbital pore from the
secondary lid fold.

Some of these characters are probably connected with its very small
size. Idiosepius is a dwarf among the Cephalopoda; adults of several
species are only 2 cm long. Like other small types, it shows a reversion
to simplification, e. g., not only the arms but also the tentacles have
only 2 rows of suckers, 'i' and the buccal funnel is only indicated.

* This circumstance, and also the whole habitus, suggests that also "Cranchia minima" d’Orb., 1839
(Cranchia, Plate 1, Figures 4-5) belongs to Idiosepius. especially because d’Orbigny (1845,
p. 241) placed it in the Sepiidae, asCr. maculata, probably not without reason. On the other hand,
this could also be a young form of Oegopsida (p, 236) in which the clubs are also biserial at first;
Oegopsida have also similar juvenile forms (cf. also Figure 88 on p. 188 and Figure 89 on p. 190).

484

505 Chapter 39

FAMILY SPIRULIDAE

(d'Orbigny, 1826)

Contents: a. Diagnosis. — b. Introductory remarks. — c. Genus Spirula. — d. Spirula spirula.

a. DIAGNOSIS

Phragmocone with round cross section, with transverse septa, spirally
coiled, with small remnants of the sheath in the form of a thin coat. Pro-
ostracum reduced at an early stage and replaced topographically by the
muscular mantle; dorsal margin of shell penetrating the mantle sac.

b. PRELIMINARY REMARK

The only genus of the family is Spirula Lam., for which d'Orbigny
(1826) created the family "Spirulees." Later (1842) he included the fossil
Beloptera and Spirulirostra as Spirulidae (after Owen, 1836).

This classification is no longer acceptable, but it expresses the natural
relationships between these forms. The establishment of the "Sepioidea"
as a suborder of the Decapoda has now given these relationships their
definitive classification.

c. GENUS SPIRULA Lam., 1881

Diagnosis. Shell forming an almost closed spiral, at least in youth, with
2 V2 turns, of which the last half turn becomes detached (which shows the inherited
tendency to straightening, p.499). Posterior end developing postembryonally a
pointed tubercle around which a fleshy circular swelling develops later, forming
a suckerlike structure (terminal organ). Fins small, round, S e p i o 1 a-like,
attached obliquely to the posterior end and directed anteriorly and ventrally.
Funnel bond simple, oblong, extending along the whole funnel pockets. The
primary eyelid can be widely opened and is contracted only temporarily to
a small pore to protect the eye. Length of arms slightly increasing
ventrally (formula: 4, 3, 2, 1 ); each arm with 4 rows of small suckers, with
506 smooth rings.. Tentacle clubs with 16 rows of similar suckers. Bases of
all arms connected by a distinct protective membrane.

485

I could not obtain fossil species of Spiral a, although these are stated
to be common in the Miocene of Italy.* Only recent species: Spirula
s p i r u 1 a.

Live Spirula have not been found in the Mediterranean and their
occurrence is at least doubtful. However, empty shells of Spirula have
been found on Mediterranean coasts (Tunisia, Spain, the Balearic Islands;
cf. Girard, 1892, p. ll), where they could have been brought by the current
through the Strait of Gibraltar. However, Spirula will be treated here
because of its extreme importance for the systematic morphology of the
Dibranchiata. Spirula is apparently distributed in all oceans (Atlantic,
Pacific and Indian); wherever specimens rise to the surface, they are
isolated and dying. The genus inhabits the cold waters of the deep sea,
where it is probably common, to judge from the numbers of the dead shells
on the shore.** Only a few well preserved specimens are in collections.
Figure 276 shows a slightly reconstructed specimen.

d. SPIRULA SPIRULA (L., 1767) Hoyle, 1909

Contents: a. Diagnosis. - 2. Literature. - 3. Structure of the adult animal (p. 508). — 4. Postembryonic
juvenile stages (p. 515). - 5. Derivation of the type of Spirula (p. 516).

1. DIAGNOSIS

Shell with about 2V2 turns, last half turn slightly more distant from the
others.

All specimens of Spirula and shells apparently belong to the species
which Linne (1767) named "Sepia" s p i ru 1 a, despite their wide distri-
bution. The differences mentioned by Lonnberg (l896) concern only the
degree of preservation; the few specimens with a soft body that are known
are all in bad condition. As Joubin (l910), Naef (1913) and Chun (1915)
have shown, the young stages show striking variations. The main locality
in which the species was found is the Canaries Current, where it apparently
occurs in large numbers in greater depths. The first specimen was found
adrift near the Moluccas (Peron, I8OO).

507 2. LITERATURE

1767 Linne, Sepia spirula.

1799 Lamarck, Spirula (genus).

1807 Pdron, Spirulea prototypes.

1816 Lamarck, Spirula australis (same specimen).

1822 Lamarck, Spirula peronii (Vol. 7,p. 601; same specimen).
1848 Owen, Spirula reticulata (Plate 4, Figures 3, 9, 10).

*■ Errors of determination are not excluded. Only the phragmocone is usually preserved in some sites,

because the acid of the medium (CO2?) apparently dissolves the rostrum and the sheath of the Sepioidea.
Spirulirostra-type chambered shells could therefore be considered as Spirul a. Compare also
Spirulirostrina,p.502; the sheath of the specimen illustrated is strongly macerated.

** The light shells of dead animals rise to the surface, and currents may sweep them around the globe
(Walther, 1894, p. 514).

486

1865 Angas, Ammonia laevis (p. 157).

1879 Agassiz, Spirula australis (p. 298).

1886 Hoyle, Spirula peronii (ChalL, p. 122, Catal., p.26).

1896 Lonnberg, Spirula p e r o n i i, a us tr a 1 is, r e t ic u 1 a t a and blakei distinguished.
1906 Hoyle, Spirula spirula.

1910 Chun, S pi ru 1 a australis.

1915 Chun, Spirula australis.

1921 Naef, (Syst.), Spirula spirula (p. 538).

1922 Naef (Foss. Tint.), Spirula spirula (p. 70).

(Compare also the next part.)

FIGURE 276. Spirula spirula (L.). Adult female, natural size,
from East Africa (Vienna Royal Museum), slightly diagrammatic.
Compare the attachment of the mantle (m) on the shell with that
of the young specimen in Figure 281. The insertion is not changed
in principle, but the mantle sac is drawn back on each side (x) and
situated between the two saclike projections of the terminal organ
(t). The fins are attached obliquely dorsoventrally on the terminal
organ and completely independent of the muscular mantle. The
skin is worn, but the "windows" of the shell were not completely
exposed; the apparently tough shell sac is partly preserved.

d — position of dorsal margin of shell; si — supporting ridge of
penultimate septum; 1 - eye lens, covered by the primary eyelid
in its greater part, leaving only a small opening. (From Fossile
Tintenfische, p.69.)

Chun (l910, 1915) markedly improved our knowledge of the adult
Spirula. Since the first drawing was published (Peron, 1807), the animal
which shows a peculiar combination of the characters of fossil and recent
forms, has been dealt with by many authors. Roissy (1805, cf. Montfort)
recognized the decapod character of Spirula; Lamarck (1822) gave the
first detailed description of Peron's specimen. Robert (l836, C. R.,
pp. 322, 362) reported on several damaged specimens from the Canaries
and described the terminal organ ("bouton terminal"). He is quoted by
Blainville (1837, 1839), who studied the chambered shell and its relationship
to the soft parts in comparison with the fossil Cephalopoda. Blainville
described the attachment of the small fins, the oval skin windows on the
dorsal and ventral side of the mantle sac which form only a thin layer above
the shell (shell fold!), the two gills, the attachment of the cephalopodial

487

508 retractor on the free margin of the shell and showed (after Roissy) that the
shell is completely internal, as in other Dibranchiata. Gray (1845)
illustrated a well preserved specimen from New Zealand, which Owen
(l848) described again together with another specimen from the East Indies,
and identified the funnel valve, funnel cartilage, salivary gland, and a liver
consisting of 2 parts. Angas (1865) obtained a complete specimen, washed
ashore in South Australia. Owen (l879) described the anatomy of the female
described by Gray and in 1880 he described the only male ever found.
Agassiz (1879) obtained a well-preserved female in the Caribbean, which he
illustrated in 1888 (Figure 280). Huxley and Pelseneer (l895) gave in the
Challenger Reports a detailed description of a specimen caught near Banda
in 1878 and of two others, mentioned already by Giard (l893). Together
with much correct information, Pelseneer made some erroneous obser-
vations on its systematic position (Oegopsidal) and the relationships between
body and shell. He described the shell as only partly internal, exposed
dorsally and ventrally at the oval "windows," and considered that the
mantle surrounds the shell only during embryonic development (see Land,
1900, pp. 72— 73; Prell, 1921). Lonnberg (1896) corrected these views after
a study of a well preserved specimen from Madeira, but his findings were
not accepted. He considered Spirula as belonging to the Myopsida,
related to Idiosepius, Sepia and S e p i o 1 a, and that it has a completely
internal shell. Chun (1910—1915) made a detailed examination of an adult
female dredged near Sumatra by the German Deep-sea Expedition (Chun,
1915, Plates 44—50) and also of a half-grown specimen (Figure 277 here)
and a larger damaged specimen. By the courtesy of Chun, I was able to
examine all this material and the younger postembryonic stages in 1913,

The following description is thus based on personal experience and supple-
ments Chun's description. A new interpretation of the systematic -morpho-
logical position of the type of Spirula is given here, which Chun did not
attempt.

3. STRUCTURE OF THE ADULT ANIMAL

The general appearance of the adult Spirula (Figure 276) closely
resembles that of the related Idiosepius, Sepiola and Sepia. The
Sepiolidae are apparently most closely related to Spirula, and there is
also a resemblance to embryos of S e p i a in some characters (Vol. II,

Plate XIX, Figure 2). The coloration of Spirula is caused by the dense
reddish brown to violet brown chromatophores at the posterior end, on head,
arms, and anterior margin of the mantle. The middle of the mantle sac
has almost no chromatophores but shows a silky to silvery sheen, as in
some bathypelagic Oegopsida (Neoteuthis); this coloration is caused by
the iridocytes and the undulate-fibrous subcutaneous tissue (Chun, 1915,
p. 425).

509 The shell of the adult Spirula is well known and represented in many
collections. The shell was well described by Appellof (1894). We can
confirm most of his data, but we differ in the morphological interpretation.

488

FIGURE 277. Half-grown S p ir ul a, 2 x (Chun, 1915, p. 426, Figure 36). Drawn after the original specimen;
completed after a comparison with the material available, especially Chun's specimens and drawings and
the earlier illustrations of Owen (1878), Huxley (1895) and Lonnberg (1896). Note the Sepia-like habitus,
a) ventral; b) dorsal; c) lateral. Figure c is shown transparent to show muscular mantle, ventral skin, anal
papilla and terminal organ in optical section; shell and other internal parts are visible laterally (Chun, 1910,
Plate 1, Figure 1, corrected in some details).

tg — base of tentacle inside a simple pocket; va - ventral attachment of muscular mantle on the shell, of
which an oval area is visible through the thin skin; the posterior part (x,y) is covered by the terminal organ
(to); ir - iris; li -- eye lens; Id - primary lid, slightly contracted (open in Figures a and b); md — dorsal
median section through muscular mantle; rc - retractor capitis; wk - living chamber; da - dorsal attach-
ment of muscular mantle on the shell; ht - skin; fl - fin; st - terminal spine, enclosed in the terminal organ;
va - ventral attachment of muscular mantle; ms — mantle septum; mv - ventral median section through
muscular mantle; mh — mantle cavity; an - anal papilla; ki - gill; rt - funnel retractor; th - funnel bond;
tt - funnel pocket; nh - neck bond; ro - olfactory organ. (From:Fossile Tintenfische, p.71.)

A proostracum is absent in the postembryonic stage, as in Sepia
(cf. p.476). All other typical elements of the shell of Decapoda,— ostra-
cum, hypostracum, periostracum, septa, siphimcle and prosiphuncle — are
present, but the form and correlation of these parts are characteristically
changed. However, the type of Sepioidea is distinctly recognizable and the
close relationship with Spirulirostra is also distinct.

The marked spiral coiling is particularly striking in an internal shell
because it cannot be explained by comparison with the derivation of the
Nautilidae from straight Tetrabranchiata. Spirula and Gyroceras
are certainly similar, but the early chambers of Gyroceras are curved
upward, those of Spirula downward. But mainly the coiling takes
place in the interior of the soft body, in which are situated not only the
shell opening as in all Sepioidea, but also the apical parts.

The specimens examined have 2V2 turns of which the first two are
almost contiguous, with an interval of 0.1— 0.5 mm. The last half turn
510 becomes detached and shows a tendency to become straight. This resembles
Spirulirostra (p.499) and probably provides space for the developing
gonad. The following characters change during development: 1. The first
chamber is vesicular, almost spherical, more or less as in the primitive
Ammonoidea (diameter 0.7 mm), not flattened as in the true Nautiloidea.

2. There are moniliform constrictions between them which are most distinct
between the first and second chamber and become later gradually less
distinct. 3. The living chamber is usually shorter than a single air
chamber.

489

FIGURE 278. a) Median section through the last chambers
of a shell of Spiral a, after Appelldf (1894, Plate 8, Figure 1),
12 X (cf. Figure 277). b) section through part of the external
wall of the shell, showing the 3 typical layers; hypostracum
(hy),ostracum (os) and periostracum (po) (the first two strata
are united in the main figure).

x,y — anterior boundary of suture; ws — supporting ridge;
spi - last septum; sp2 - penultimate septum; wk - living
chamber; Iki - last air chamber; lk2 - penultimate air
chamber (they form narrow slits ventral to the siphuncle
(si)); kd - calcified neck; ch - conchiolin neck; pf - pillar
substance; rd - free shell margin of left side. (From Fossile
Tintenfische, p.74.)

The inner structure of the shell is shown in the median section
(Figure 278). Of particular interest are the initial parts of the shell
(Figure 279). The wall consists of the 3 normal layers in the whole region
or at least in the anterior part (Figure 2 78b). The ostracum is thin and
strongly retractile, the hypostracum thicker, with typically stratified
nacreous substance which is without sheen and only slightly retractile.

The periostracum is a thin, tuberculate coat which thickens markedly in
the initial part of the shell on the ventral side near the siphuncle; it forms
here a "ventral ridge" (Figure 279,24), as in the fossil Sepioidea
(Figure 2 69). Secondary development of the ventral ridge brings the first
1V2 turns closer together and strengthens the fragile juvenile part of the
shell.

The septa consist of mother-of-pearl with its characteristic sheen.

They are concave and pass directly into the long calcified necks on the
ventral side. Each neck ends in a sharp margin, inserted into the funnel
formed by the preceding neck. It continues into a neck of conchiolin which
lines the interior of the preceding calcified neck to near the next septum + 1
and is connected with it. A layer of transverse calcified rods prevents the
necks from coming in contact with each other, so that the osmotic connection
between the air chambers and the siphonal epithelium is preserved (l).

The septum is attached at the outer wall by its curved margin to form a
511 narrow suture and is connected with the inner nacreous layer. On the out-
side, however, the suture appears as a wide stripe (as in fossil shells)
because of the presence of a broad supporting ridge which is situated in the
adjacent corner of the preceding chamber, between the wall of the shell and
the dishlike septum. This bar consists of a nacreous but more retractile
substance, the stratification of which follows the margin of the septum and
disappears toward the air chamber. The last layer apparently passes into
the posterior layer of the septum. This is very typical (also observed in
Nautilus) and explains how the septa are formed, but differently than
described by Appellof (1894, p. 64). The septa are deposited not at one time.

490

but gradually, beginning from the annulus. The posterior shell epithelium
at the outer wall forms at first the posterior part of the supporting ridge
and strengthens it until the septum can be attached to the surface of the
ridge, (its posterior lamellae are soft, delicate and not strongly calcified
in Nautilus. They are a direct continuation of the last layers of the
supporting ridge, which are strongly calcified. This develops later into an
easily detached calcified membrane which forms a coat on the posterior
side of the septum. I could not determine whether a similar process takes
place in Spirula.) The pillars of the siphonal part develop simultaneously
with the supporting ridge and the calcified membrane.

The periostracum is relatively thicker in the initial part of the shell
(Figure 279) because it can be thickened continuously, while the primary
layers of the shell do not grow further (p. 510). Hypostracum and ostracum
probably are present, but they cannot be easily distinguished. The initial
chamber is nearly spherical and so markedly constricted at the site of the
first septum that there remains almost no room for it. It is therefore
rudimentary and passes into the first calcified neck without distinct
boundary. The first calcified neck is naturally not supported by a preceding
neck, but its free margin bears a weakly calcified process, the "conchiolin
cap" of the "initial caecum." This continues directly into the prosiphuncle,
a slightly calcified lamella of the shell which rests sagittal on the ventral
and anterior part of the wall of the initial chamber. The prosiphuncle is
strengthened at the free margin by a transverse support and is morpholo-
gically comparable to the pillars of the other parts of the siphuncle. The
second septum and its neck are almost normal and all parts can be dis-
tinguished. I assume that the neck of conchiolin has a blind posterior end
and provides a continuous lining for the conchiolin cap of the first neck,
as in the homologous parts of Nautilus.

(512)

$ p » h n

FIGURE 279. a) Median section through the initial part of a shell of
Spirula. 32 x. Based on data and illustrations of Appellof (1894),
especially Plate 9, Figure 1. Primary wall of shell (ostracum and hypo-
stracum) (hatched) (6), periostracum (7) dotted, septa and calcareous
necks also dotted (18,15,13, 5,4). Epithelium of soft part of siphuncle
dotted.

I - first chamber; 2 - prosiphuncle (transverse support); 2a - its
sagittal lamella; 3 - conchiolin cap as initial part of siphuncle;

4 - calcified neck of first septum (rudimentary), adjacent to it;

5 — conchiolin neck of second septum; 8 — beginning of soft siphuncle;
9 - ventral part of first neck and septum; 10 - septal knob of siphon;

II — pillar substance; 12 — conchiolin neck (beginning); 13 — calcified
neck to second septum; 14 - second chamber; 15 - second septum;

16 - supporting ridge; 17 - pillar to third neck; 18 - third calcified
neck; 19 - epithelium of siphuncle; 20 - third air chamber; 23 -
ventral part of third air chamber; 24 - ventral ridge of periostracum;

25 - second septal knob of siphuncle; 26 - ventral part of second
air chamber.

b) Hypothetical median section of a freshly hatched stage, 4 x (cf.
Figures 252,254):

m - muscular mantle; p - rest of proostracum; s - shell sac (shell
epithelium); h - mantle cavity; n - neck bond; w - position of
liver; a — anal papilla. (From Fossile Tintenfische, p.73.)

491

The insertion of the shell in the mantle sac will be described at first
for a young animal, in which the relationships are simpler (Figure 281).

The skin (shell fold) is still so thin that the parts of the shell and their
connections are clearly visible from the outside. About % of ^ turn are
visible in the median plane from the dorsal to the ventral side, so that about

512 half of the cross section protrudes. The muscular mantle is attached on
each side of the turn, forming 2 lobes,* which extend posteriorly. The line
of attachment is displaced dorsally and ventrally in the middle so that the
entire cross section is surrounded and the shell becomes surrounded by the
mantle sac.

The fins are attached to the posterior part of the shell, obliquely in an
anterior -ventral direction (Figure 260 on p. 487). The base of the fins is
situated close to the lateral lobes of the muscular mantle which surround
the shell, so that it could be assumed that the fins developed from these
lobes (cf. Vol. II; also p. 95). The form of the fins is typical for the
Sepioidea; as in the Idiosepiidae and Sepiolidae, they are rounded lobes with
a slightly narrower base and a sharp indentation forming an "earlobe" at
the anterior margin (Chun's figures show usually the shrunken, preserved
fins, without reconstruction, i. e. spreading). The obliquely attached surface
may act as a depth- rudder which compensates the buoyancy of the posterior
end.

The "terminal organ" develops in the half-grown and adult animal and
conceals the relationships described. Its central part is a terminal spine
(p.235) which is apparently deposited as a conical elevation on the tissue
which covers the shell. According to Chun (l910, p. 185; 1915, Plate 70,
Figure 5), the solid tissue of the spine is a luminous organ, and he considers

513 its parts as reflector, luminous body and lens. As in other true or alleged
"luminous organs," these interpretations are very speculative and partly
erroneous (reflectors are very thin mirror surfaces, not massive layers of
tissue).

I doubt whether this is a luminous organ. I consider it a shock absorber
which protects the shell, like similar organs of young Sepiolidae and their
embryos (Figure 321, Sp). A fleshy ridge develops later in Spirula around
the terminal spine, probably as a protective structure; its narrow muscular
margin faces the spine (Figure 277); the tissue diverges dorsally and
ventrally and covers the posterior end of the shell more firmly than the shell
fold (Figure 255). This thin layer is divided into 2 oval areas ("windows,"
see p. 507) on the dorsal and ventral side of the mantle sac. The oval
areas are thus obviously parts of the primary mantle, not gaps in the
muscular mantle.

The mantle margin bears 3 large processes which end in pointed arches
and are divided by deep, rounded incisions for eyes and funnel (cf. funnel
indentation in Sepiola au r a nt i a c a. Figure 358).

The head is large, Sepio la-like because of the large, laterally pro-
jecting eyes. The primary eyelid resembles that of the Oegopsida (P. 227)
and also tends to become contracted to a narrow pore at the position of the
sinus. This is the condition in most specimens preserved while alive.

" The above criticized concept of a secondary surrounding of the shell (p. 508), which does not take place in
the Dibranchiata (p. 107) is based on a confusion of the muscular mantle with the primary or skin mantle
(p. 92).

492

without anesthetization, the contraction is due to the effect of formol. A
secondary lid is not even indicated. The lens projects markedly laterally,
as in the Oegopsida. The olfactory organs are typically situated, round,
simple warts (Figure 277).

The funnel is situated deep inside the mantle sac in preserved specimens,
but probably glides extensively back and forth in the swimming animal
(pp. 93, 83). The funnel bonds are oblong, with rounded ends (Figure 280),
as in the Sepiolidae (Figure 326); they occupy the entire length of the funnel
pocket. The neck cartilage is also normal and has about the form of the
sole of a shoe. The collar and mantle bonds are simple longitudinal ridges
which begin slightly behind the corners of the mantle and gradually dis-
appear posteriorly.

The arm apparatus reaches its definitive development at a relatively late
stage. It is less developed in half -grown animals (Figure 277) than would
be assumed from Chun's figure (l915, Plate 67; 1910, Plate l). This is due
to the fact that the bases of the arms occupy the whole width of the head in
profile, but only part of the width ventrally and dorsally.

FIGURE 280. Spirula a u st r al i s, female, 2 x, with opened
mantle cavity. This slightly diagrammatic drawing is a combi-
nation of Chun’s preparations and figures (1910,1915). Note the
structure of the arms; large eyes; primary lids, contracted to
narrow pores (Au) (cf. Figure 276 on p. 507); olfactory organs
(Ro); structure of funnel, especially of the funnel bonds (Th);
anus (Ap) situated near the renal papillae (Np); gills and female
genitalia. Genital process present only on the left, covered by
the large nidamental (Nd) and accessory (Ac) glands. The mantle
has been cut off posteriorly at the boundary of the mantle cavity
(Mm); the shell (Sch) projects as a longitudinal oval from the
remaining part covered only by skin; the muscular mantle is
attached on this oval. Fins small, rounded, terminal (FI).

Ss — swimming margins of tentacle clubs; Sh — membrane
between the 3rd and 4th arm (A4); Tg - base of tentacle (in the
pocket); Li — lens, covered: Tt - funnel pocket; Mm - muscular
mantle.

The arms are short (formula: 4, 3, 2, l) and connected at the base by a
well developed membrane, which is absent between the ventral arms. The
swimming margins are distinct on the dorsal arms, weakly developed only
on the distal part on the two following pairs. The arms of the living animal

514 bear the 4 usual rows of suckers. Preservation in formol causes con-
traction, which may displace the suckers into an irregular pattern of 8 rows
(Chun, 1915, p. 423). The adhesive ring of each sucker has a distinct external
zone of rods, followed by a tuberculate zone (cf. Plate XIII, Figure 5). The
horny ring is finely denticulate with a slightly enlarged median tooth. The
protective margins are well developed, with indistinctly differentiated
supports.

The (contracted) tentacles are strong. Their attached base is rounded
but its inside is flattened, and becomes gradually wider distally (Figure 45
on p. 119:Fb). The club tapers gradually apically. It bears typical swim-
ming and protective margins, of which the ventral is wider, so that the club
appears slightly wider if it is extended. The contracted club bears about
12 irregular longitudinal rows of suckers but their typical arrangement is
not certain (8—16 rows; see p. 488).

The buccal funnel consists of 7 parts, without suckers on the points. The
radula has not been described; it probably has only unicuspid teeth as in all
Sepioidea. The jaws (Chun, 1915, Plate 68, Figure 7) have a characteristic
form. The upper and lower beak bears lateral teeth which reach to near the

515 biting process. The biting process of the lower jaw forms a sharp angle
(less than 90°).

The mantle situs shows a number of special characters caused by the
shell, which cuts into the soft body also ventrally from behind (Figure 280).
The viscera are therefore situated laterally, right and left to the shell and
the nidamental and accessory glands are pressed together. The gills are
short and compact and the renal papillae have a chimneylike form and are
situated near the anus. The small ink sac (Figure on p.475) is also
displaced here,- but to the left, not to the right, as in the Sepiidae. The
gonoduct is present only on the left side in both sexes and opens on a low
genital process near the anus.

Bothe ventral arms of the male are hectocotylized, i. e. without suckers
and longer than the other arms (Owen, 1880). The right arm is curved
inward at the end and longer than the left. Owen stated that the gonoduct
is situated on the right and opens in a short genital process. Steenstrup
(1881 ) confirmed the data on the arms and stressed the close resemblance
to Idiosepius in many characters (Figure 275).

4. POSTEMBRYONIC JUVENILE STAGES

Juvenile specimens of Spirula are available in large numbers. The
first specimen was described by Joubin (l910), further specimens by
myself (1913) and Chun (1914, 1915). Very young postembryonic stages
are unknown (Figure 279b). As the fully developed eggs are only 1.7 mm
long (Chun, 1915), they are probably very small compared with those of
other Sepioidea (p. 409) - probably about 4.2 mm long, to judge from the

" Obviously a reminiscence of earlier states, from which it developed more strongly.

494

proportions between eggs and young of Sepia officinalis (freshly
hatched Sepia officinalis are 15 mm long with arms but without
tentacles, and the egg is 6 mm long). The specimen in Figure 252 on p. 475
would be about twice as long in life. A freshly hatched specimen has
probably only 2 or 3 air chambers, because there would be no room for
more. The youngest stages already show the character of the Sepioidea
despite the strong curvature of the initial part of the shell and all other
young stages are probably similar.

FIGURE 281. Posterior end of mantle sac of a young Spiru la, after a specimen from Chun's collection,
4x (cf. Chun, 1915, p. 425, Figure 35). a) From right side; b) oblique ventral view. The muscular mantle
surrounds the shell laterally to the line of inse:tioh of the obliquely placed, Sepiola- like fins, which are
always shown shrunken in Chun's drawings. Ten chambers are shown. The terminal spine at the posterior
end (3) resembles a small wart.

1 - Attachment of muscular mantle on the shell; 2 ~ fin; 3 -- terminal spine; 4 - suture, slightly im-
pressed; 5 -- same.

Only a remnant of the proostracum can still be present, but younger
embryos probably show the general morphology of Decapoda, particularly
a typical proostracum or the corresponding anterior part of the shell sac,
which corresponds to it. The shell opening must also occupy the position
typical for all Dibranchiata, i. e. it must surround the posterior end and
form the basis for the attachment of the muscular mantle (Figure 3 6 on
p. 107).* We must assume, however, that the muscular mantle extends to
516 the outer side of the cone already during embryonic development

(Figure 262) in the degree in which the cone is curved ventrally and that
the position of the shell in the body after the formation of 2 air chambers

* I found a specimen in the material of the German Deep-sea Ekpedition which in my opinion belonged
to this stage (Naef, 1913, p. 456), but I could not examine it again and my drawings were not suitable
for publication. The specimen is freshly hatched, still with an internal yolk sac, and showing the general
habitus of the youngest larvae of Oegopsida.a markedly inflated "cone" and a broad "lanceola" which
ends in a pointed proostracum. G. Pfeffer showed me drawings of a similar larva and expressed the same
view. Thiele (1912, Plate 55, Figures 11—13) illustrated a larger specimen which he named "Para-
teuthis." See also Figure 33 on p. 101, and Vol.II, Plate VIII, Figures 7-8; Plate X, Figures 1-3. All
these young forms show the insertion of the juvenile shell typical for Dibranchiata (with a scoop- shaped
cone) in the mantle sac, and they belong to the Oegopsida. The cone of the Loliginidae becomes rudi-
mentary earlier and more markedly (Figure 87 on p. 187); in the Sepioidea these stages end during
embryonic development and in the Octopoda the shell is completely reduced. Addendum in the legend
to Figure 472.

495

is as shown in Figure 279b, p. 512, and is comparable to Figure 252 on
p.475 and Figure 262 on p.491. The reduction (inhibition of growth) of
the proostracum during embryonic development may create a dorsal contact
between the two halves of the mantle, at least in the anterior part, so that
these parts of the mantle remain attached to the anterior margin of the
shell. This would permit the mantle cavity to reach the shell margin behind
the neck cartilage (Figure 279) and finally extend beyond it, following the
insertion of the muscular mantle ( Figure 252 on p.475).

5. DERIVATION OF THE TYPE OF SPIRULA

A comparative study of the shell and its position in the soft body shows
a relationship to Sp i r u 1 i r o s t r a, and Spirula can be easily derived
from this fossil type (pp. 500, 501 ), because Spirulirostra has been
interpreted and reconstructed according to the recent genus. However,

517 Spirulirostra provides sufficient morphological data for a recon-
struction of the posterior part of the body. Spirula is apparently a
Spirulirostra in which the rostrum and the side wings of the perio-
stracum are reduced, and this has become reduced to a crust on the
phragmocone. The curvature of the phragmocone has formed a spiral,
which may not be very surprising at first. However, there is a marked
difference between a mere curvature and a regular coil and the distance
from Spirulirostra to Spirula should therefore not be underestimated.
More important is the loss of the proostracum which takes place not later
than the stage with 3 chambers and is necessarily connected with the coiling
(see p. 486). It is clear that a straight proostracum which extends from
the anterior dorsal margin of the shell anteriorly must be curved repeatedly
to permit a spiral growth. This is a mechanical impossibility if the dorsal
plate is solid, and a plate which is not solid would have no importance. We
therefore assume that the proostracum became at first rudimentary and
soft, was retarded in growth and was finally resorbed during the increasing
curvature of the phragmocone (cf. also Naef, 1922, Fossile Tintenfische,
p. 72). The spiral coiling is also necessarily connected with the reduction
of the periostracum, because a continued curvature of the phragmocone would
be impossible if the periostracum were as in Spirulirostra or more
strongly differentiated and projecting posteriorly (see p. 485).

These relationships are distinct but negative. The question is, what are
the causes of this characteristic metamorphosis? It has to be assumed that
the parts of all stages are naturally correlated and adapted to the environ-
ment. It is thus surprising that one part (periostracum) should disappear
almost completely, while the other (phragmocone) not only persists but
even markedly develops. The proportions of the chambered shell in
Figure 252 deviate considerably from the typical. This is particularly
striking because (as Steenstrup recognized in 1881), the closely related
Idiosepius no longer has a shell, at least in the adult, while the phrag-
mocone has disappeared in the Sepiolidae (p. 503).

An explanation of the problematic nature of this type can only be
connected with the mode of life, but it is difficult to obtain information on

496

these deep-sea forms. It has to be considered, however, that the air-filled,
disproportionately large phragmocone is situated at the posterior end and
is not compensated by a rostrum (p. IIO). The animal can therefore not
swim horizontally but only with the posterior end obliquely upward. Such
a position is not unique, the nektonic Heteroteuthinae, all young Sepiolidae,
and the juvenile stages of the Teuthoidea and Octopoda almost always swim
with the posterior end pointing obliquely upward, but not as steeply as must
be assumed in this case. The rhythmic contractions of the mantle often
result in characteristic hopping movements, as in Daphnia, because the
animals are heavier than water and sink after each jump. Spirula
518 probably moves like this; the air-filled shell permits effortless permanent
swimming, as in the early stages of the Sepioidea described and in the
Belemnoidea. Similar movement has also been assumed for many Ortho-
ceratidae (cf. Figure 89). We consider Spirula as a neotenic type with a
marked morphological development on the basis of a typically juvenile
structure (p. 2 61) and a juvenile mode of life. It probably feeds on plankton.

Addendum

Schmidt (l922) described live Spirula. Their habitus is identical with
that of the forms illustrated here, but the arms are shown slightly too short
in Figure 277 on p. 509 because of preservation in formoL The following
points are of interest: The subcutaneous tissue has an asbestoslike sheen
(cf. p.48). The resting position is with the head hanging down and the apex
of the funnel directed upwards. Rapid movements are usually backward, but
turning of the funnel can propel the animal also in jerks and laterally and
forward. The fins make fluttering movements, directed upward; they
counteract the buoyancy, like the movements of the funnel. The arms hold
on to the bottom or to solid objects on it (cf. p. 566). It reacts to strong
disturbance by ejecting small clouds of grayish ink. Terminal organ
luminous, emitting yellowish green light continuously for hours. They have
been found alive between 10 and 35°N in the Atlantic and nearshore parts of
the Pacific, at a depth of 300—500 m, not above 200 m. Bathypelagic.

Mature females: length of mantle sac to 44 mm, 40 air chambers. Mature
males: 38 mm, 34 air chambers. Sexual maturity at a length of 30 mm.
Floating shells apparently belong to old animals with 30—40 chambers.

Only such shells have been found so far in the Mediterranean.

497

519 Chapter 40

FAMILY SEPIIDAE
s. restr. Keferstein, 1866

Contents: a. Diagnosis. - b. Derivation of the shell of Sepiidae. - c. Typical structure of the Sepiidae
(p. 523). - d. Typical development of Sepiidae (p. 525). - e. Variation of the type of Sepiidae (p. 525).-
f. Subfamily Sepiinae (p. 527). - g. Variation of the type of Sepiinae (p. 541).

a. DIAGNOSIS

Periostracum forming a longitudinal oval, oval- awl-shaped to longi-
tudinal rhomboidal ‘'dorsal shield." "Lateral edges" displaced anteriorly
(Figure 282), completely suppressing the proostracum and replacing it
topographically. Rostrum absent or reduced to a large spine on posterior
part of dorsal shield. Phragmocone divided into a swollen part which
consists of dense leaflike chambers extending to anterior end of the shield,
and into a forked part situated ventral to the siphuncle in the posterior part
of the body. All septa of phragmocone situated markedly oblique, almost
longitudinal, in an anterodorsal-posteroventral direction.

b. DERIVATION OF THE SHELL OF SEPIIDAE

The shell of molluscs is the part which determines the whole. This rule
is particularly valid in the Sepiidae. The shell of the common Sepia is
a time-honored object of study, but its systematic morphological and
ecological significance is still incompletely understood because there has
been no basis for its understanding (cf. Chapters 38 and 39). We believe
that we were able to provide an instructive interpretation of this charac-
teristic structure. The typical shell of Sepiidae can be derived directly
from that of Spirulirostrina (p. 50l), and some changes have to be made
in the following text, which was written before this fossil became known.
However, it seems useful to give the new observations together with the
earlier morphological construction, which was illustrated for years (1915:
Figure 282). To facilitate the comparison between the adult shells of the
Sepiidae and Spirulirostra I established the ideal (hypothetical) inter-
520 mediate stage of Spirulisepia (see Fossile Tintenfische, 1922, p. 77;
also Figure 282 (left) and Figure 285a below). This intermediate shell
differs in the following points from that of Spirulirostra: 1. Already
the beginning of the phragmocone is dorsoventrally flattened, and the later
formed parts are increasingly flattened, 2, The later septa are very
densely arranged and are progressively sloping in an anterodorsal-postero-
ventral direction. 3, The siphuncle widens rapidly to an open funnel.

4. The transverse expansion of the funnel divides the phragmocone into a

498

dorsal swollen part which extends far anteriorly and a forked ventral part
which is restricted to the posterior end and is adjacent to the reduced ventral
wall. 5. The lateral edges extend far anteriorly on the lateral margin of
the proostracum, and the main mass of the periostracum forms a longitu-
dinal oval dorsal shield in which the modified phragmocone (l— 4) is only
slightly embedded. 6. The rostrum is relatively strongly reduced in mass.
7. The ventral process is curved posteriorly.

(521)

FIGURE 282. Derivation of the shell of Sepia (right)
from the structure demonstrated for Spirulirostra
(p. 498) by the ideal transitional form "Spirulisepia"
(left). This hypothetical form resembles in principle the
later found Spirulirostrina (p.501), but it is more closely
related to Sepia in many characters and may be con-
sidered as a more advanced link of the series of Sepia
(p. 497) which has not yet been found. Further expla-
nations in the text.

Ro - rostrum: Vw - ventral wall, belonging to the fork
(Gb) in Sepia; Si - siphuncle, widened; Ls ~ last sep-
tum; Sh - lateral edges; Pr — proostracum.

FIGURE 283. Median section through shells of Sepiidae
(diagrammatic), a) Section through the posterior part of the
shell of Belosepia sp. (fossil), which still closely resembles
that assumed for Spirulisepia and also the conditions in
S pir ulirostr a (pp. 522 and 500). b) Sepia orbignya-
na (juvenile). The ventral curvature is still very marked.
Note the phragmocone embedded in the periostracum, the
arrangement of the supporting pillars and interseptal lamellae,
and the form of the spine; identify the parts homologous to
the septal necks, c) Sepia officinalis. Note the form
of the rostrum. The typical conditions are later covered
(Figure 309) by a secondary folding of the fork part over the
posterior part of the shield.

499

(521)

FIGURE 284. Cross section of the middle of a recent shell of Sepii, (diagrammatic):

Rp ~ dorsal shield; Sp — lateral edge; Sk — margin of lateral edge, always curved ventrally in reality
(Figure 290b); Co - thin conotheca around the phragmocone; Lk - oldest air chamber with "niches" between
pillars; Sw2 - second septum; Jn - intermediate septum; St — pillar; Ga - septa of fork, laterally adjacent to
the area of the siphuncle, which is not shown at its deepest part (Figure 283); Sw^ — last completely cut septum,
i.e. basis of cut part of siphuncle, which appears here raised instead of sunken; Si - septal necks of following septa.

(522)

FIGURE 285. Comparison of the shell of Sepiidae and its topographical relationships to the soft body with the
relationships known or assumed for Spirulirostra (cf. Figures 282 and 271).

a) Median section through the hypothetical Spirulisepia ;

b) median section through a typical Sepia.

a) The structure of the shell should be compared with that in Figure 273. The resemblance in principle is
evident. Consider the characters assumed for Spirulisepia (p. 520), especially points 1-4,6 and 7.
Identify particularly the ventral process (Vf) of the periostracum, which has become here part of the dorsal
shield; also the part of the shell (Ga) situated ventral to the siphuncle which forms the fork of the Sepiidae.
The modification of the shell opening prevents the detachment of the gonad from the dorsal wall of the
coelom (pp. 476,482) which is otherwise typical for other Sepioidea; the preservation of the connection
between gonad and shell (always a primary formation in embryogenesis - Figures 87, 321), which always
persists in typical Decapoda (Figure 53, on p. 136). The ink sac is displaced ventrally to the hind intestine
by turning around it to the right.

b) The changes are further developed in Sepia, as indicated on p. 521. Consider points 1-7, find the homo-
logies of ventral process, fork and siphuncle, and note the moderate displacement of the gonad to a part of the
shell which corresponds to an anterior (flat) part of the siphuncle. The ink sac (cf. Figure 289) is larger and
extends further posteriorly, causing a slightly asymmetrical displacement of the adjacent organs (posterior
aorta). The anterior septum extends beyond the neck cartilage, so that also this region is changed.

500

All these aspects essentially resemble the conditions observed in or
assumed for Spirulirostrina (p. 501 ) but Spirulisepia is evidently
more closely related to Sepia than to Spirulirostrina and represents
a more advanced link of the Sepia series.

This structure prepares the shell of Sepia by gradual accentuation of the
changes described here, exactly as the shell of Spirulisepia resembles
that of Sp i ru 1 i r o s t r a, without essential changes (p. 498). The shell of
Sepia has become different in the following characters: 1. The phragmo-
cone of recent Sepiidae (Figure 310), which advanced anteriorly, has
suppressed the proostracum completely or has left only traces (Figure 301).
2. The last septa are almost longitudinal and the ventral wall of the cono-
theca is markedly reduced ("fork”). 3. The necks of the septa are

521 markedly flattened; the later septa are no longer impressed but raised;

the siphuncle forms only a shallow groove which may exceed in area the
preserved part of the last septum (Figures 291, 306). 4. The rostrum is
reduced to a small spine and may disappear; at any rate, it no longer
encloses important parts of the shell. 5. The suppression of the proostra-
cum causes the dorsal shield to become a single, oval, moderately thick
plate which could be easily considered as the original conotheca,
especially when the part contained in the ventral wall of the phragmocone
disappears (Figure 309). 6. The widening of the siphuncle transforms the

ventral wall (Figure 282), i. e. the ventral part of the conotheca, together
with the adjacent remnants of the septa and septal necks into a forklike
structure, while the dorsal parts of the typical shell are converted into a
swollen incrassation. 7. The whole phragmocone is flat and only slightly

522 embedded in the periostracum (i. e. in the dorsal shield) from which it
can be easily detached by maceration in acid alcohol (which apparently took
place in cones of Belemnites in seawater). The incrassation remains
delimited by a special conotheca and separated from the dorsal shield.

This characteristic modification of the shell obviously affects its
relationship to the soft body, as shown in Figure 285.

523 c. TYPICAL STRUCTURE OF THE SEPIIDAE

The morphologically primary conditions in the Sepiidae can be deter-
mined from 3 different sources: 1. The development of the special
structure from other, more general preceding stages can be observed onto-
genetically. 2. Fossil Sepiidae, e. g. Belosepia (Eocene), have a shell
which still resembles the more general conditions in the suborder
(Figure 283). 3. Certain structural norms in many recent species have

a morphological primacy before others, because of the resemblance of their
ontogenetic or paleontological preceding conditions. The use of these three
sources of information (the form of the shell of Belosepia is known and
distinctly primary in its morphology) permits the reconstruction of a fossil
type, but its differentiation into separate species will not be discussed
here.* The establishment of an ideal basic form is therefore not necessary.

* Cf. Naef, 1922 , Fossile Tintenfische, p. 82.

501

FIGURE 286. Shell fragment of Belosepia (Munich Paleontological Museum), 0.5x. The dotted parts
are hypothetical reconstructions after Edwards and Wood (1849,1877).

a) Ventral. Note the ventral wall (Ga, ventral part of phragmocone); entrance to siphuncle; last septum;
posterior part of dorsal shield ("ventral process") with serrated margin and radial grooves, and the strong,
pointed, curved, laterally compressed spine.

b) Median section. The phragmocone is still as in Spirulirostra. Note the ventral curvature; the
marked convexity of the first chamber; the typical, but wide siphuncle (Si). The periostracum consists of
a large spine (Do), a tuberculate medio-dorsal crest (Cr) which gradually disappears anteriorly, and the
ventral process (Vf, Figure 271) in the posterior reconstruction of the shield.

Vw — homologue of capitulum.

The shell of Belosepia has been interpreted wrongly because of the
inadequate drawings of Munier-Chalmas (Zittel, Handbuch, 1885, p. 514, and
Fischer, Manuel, 1887, p. 557). Lang (1900, p. 99, Figure 107) placed this
genus incorrectly between the Belemnitidae and Sepia. Belosepia is in
fact a true species of Sepiidae which differs from the recent species of the
family in the direction toward Spirulirostra and the Belemnosidae and
not toward the Belemnitidae. This genus shows no special characters of
the Belemnitidae, as was shown by Edwards and Wood (1849, 1877) (see
Naef, 1922, Fossile Tintenfische, p. 83; also Chenu, 1859, p. 46). On the
other hand, the general resemblance to a shell of Sepia is distinct. The
524 shield is oblong-oval, more convex than in recent species of Sepia,

heavier and more solid and with a characteristic, jagged serration of the
posterior margin. I assume that there was also an adjacent, not calcified
marginal zone, as in the recent species of Sepia. The inner surface is
covered with radial grooves which correspond to the marginal teeth; such
grooves are also indicated in Sepia. The posterior margin of the shield
(i. e. the part corresponding to the ventral process) is curved posteriorly
along the spine, which is very large and curved dorsally as in Sepia
orbignyana (Figure 283 on p. 521). This shows traces of a displacement
of the longitudinal axis, caused by the strong ventral curvature during
development (pp. 484 and 499); a similar displacement takes place in Sepia,

502

but to a lesser degree (Figure 283 on p. 521 ). Anterior to the spine is a
large, tuberculate mediodorsal incrassation which gradually becomes
indistinct and resembles the weaker formation in Sepia elegans
(Figure 315a). All these parts closely resemble the structures in the
recent species of Sepia. On the other hand, the phragmocone still closely
resembles that of Spirulirostra, in the markedly convex initial
chambers (Figure 286 and 273), the still relatively narrow siphuncle, the
typical differentiation of the forked part (Ga), and the loosely arranged septa,
between which supporting lamellae are apparently only little developed
(Edwards and Wood). The forked part still contains air chambers so that
parts of septa and of septal necks, and the conotheca can be recognized
and probably also an outer coat belonging to the periostracum in well pre-
served specimens (Figure 287a).

FIGURE 287. Median sections through older embryos of Sepiidae.

b) Diagrammatic median section through an older embryo of Sepia of ficinalis.showing the embryonic
shell and its relationship to the soft body. The parts of the dorsal shield cannot be exactly distinguished from
those of the conotheca. There is a slightly raised indication of the fork part at the posterior end (Gt). Identify
the air chambers (1 --3), pillars (shown by simple lines) which perhaps replace the prosiphuncle (Ps); the depression
(Sa) in the first septum, which represents the initial caecum of the siphuncle; also the homologies of the
septal necks (Sd). Rostrum still absent.

a) Construction of a corresponding section through an embryo of Belosepia based on the initial parts of the
shell. Prosiphuncle, siphuncle and chambering are more normal and explain the atypical conditions in
Figure b. A distinct proostracum is assumed to be present. The yolk sac is shown very small so that it could
be included in the figure (cf. Figure 87 on p. 187).

503

525 Our knowledge of the soft body of Belosepia is limited to facts

obtained directly from its correlation with the shell, but the body certainly
closely resembled that of the recent species of Sepia. The convexity and
relative length of the shell suggest a slender animal (more or less as in
Figure 311) with a cylindrical or laterally compressed mantle sac, not
dorsoventrally flattened, which is more or less characteristic for the recent
species of Sepia. Belosepia was probably about 50cm long without
tentacles (the specimen in Figure 286 is about 45 cm long). The form of
this fossil type should be determined by adaptation of the form of the shell
to the general relationships shown in Figure 260 on p. 487. There is no
reason to transfer to it the special characters of the recent types (seep. 527).
The corresponding correlations of the recent Sepiidae should only be con-
sidered as a part of this adaptation to the specific shell of Belosepia.

The relationships between animal and shell then appear as a stage of the
metamorphosis illustrated in Figures 282, 283 and 285 and are shown in
general in Figure 287.

d. TYPICAL DEVELOPMENT OF THE SEPIIDAE
(BELOSEPIA)

The structure of the inner parts of the shell of Belosepia gives
valuable information on the ontogeny of the genus and proves its inter-
mediate position even more convincingly than the shell of the adult (p. 19).
Figure 287 is a comparison between a diagrammatic median section of an
embryo of Sepia officinalis and a corresponding section through an
ideal embryo of Belosepia; the relationships of the former were changed
as required by the typical form of the shell of Belosepia. The con-
struction of Belosepia shows that the young forms of the oldest Sepiidae
still closely resembled Spirulirostra (cf. p. 477) and even Spirula
(Figure 279b). On the other hand, the recent species of the family develop
much more directly, i. e. all parts of these forms reach their definitive
position and proportions much earlier, so that their ontogeny only little
resembles that of the typical Sepioidea. (On embryonic development, see
Vol. II; postembryonic development is described on p. 540.)

e. VARIATION OF THE TYPE OF SEPIIDAE

The recent Sepiidae are closely related to Belosepia but there are
some marked differences. This finds its systematic expression in the
establishment of two subfamilies:

1 . Shell with deep siphuncle which is narrow in its initial part; fork
raised, in form of a typical ventral margin in which air chambers, septa,
septal necks, conotheca and periostracum are still readily distinguishable.
Shell markedly convex in cross and longitudinal sections, suggesting a
526 laterally compressed mantle sac. Initial chambers particularly highly
convex, showing distinctly the form of a ventrally curved phragmocone.
Fossil Sepiidae (Paleocene and Eocene). Subfamily Belosepiinae Naef, 1921.

504

FIGURE 288. Shell of a Sepia
officinalis at the time of
hatching, ventral (left) and dorsal
(right). Compare with Figure 282.
The dorsal aspect shows distinctly
the eccentric growth. Only the
primary flat form prevents this
eccentric growth from creating a
ventral curvature of the initial
part. The small "anlage" of the
spine is situated posteriorly on the
initial chamber. Ventral view:
the narrow ventral process and the
widened lateral edges of the perio-
stracum can be distinguished from
the phragmocone. The narrow
fork of the phragmocone begins
to become separated from the
broad incrassation. Eight
flat, rapidly widened parts of
the siphuncle are recognizable.

FIGURE 289. Mantle situs of an
almost fully developed male
embryo of S e p i a officinalis,
5X:

1 - posterior pallial artery; 2 -
median pallial artery; 3 - mantle
septum; 4 — branchial heart,
translucent; 5 - ink sac; 6 - hind
intestine; 7 - renal papilla; 8 -
ligament of gill; 9 - branchial
gland; 10 — attachment of lamella
of gill; 11 “ funnel retractor;

12 “ stellate ganglion; 13 — funnel
bond; 14 — olfactory organ; 15 -
tentacle stalk in the pocket, trans-
lucent. Otherwise, the preparation
is opaque, so that the relief of the
mantle cavity is particularly
distinct.

2. Part of siphuncle flat, forming a depression and very wide already
at the beginning; part of fork low, its ventral wall visible only as a conti-
nuation of the septa of the ridge into the lamellae of the fork. Initial
chamber only shallowly convex, like a spoon, containing, like the later
chambers, a dense skeleton of pillars which support the septa. Form of
shell causing a more or less distinct dorsoventral flattening of the mantle
sac. Fossil (Miocene) and recent Sepiidae. Subfamily Sepiinae Naef, 1921.

527 f. SUBFAMILY SEPIINAE (Naef, 1921, System, p. 538)

1. DIAGNOSIS

Shell as in Sepia (p. 522). Already initial chamber very shallowly
convex. Septa supported by pillars on their whole circumference.

505

Siphuncle beginning as a flat, rapidly widening depression, the bottom of
which may show tubercles in the region of the incrassation. Fork part
more or less reduced, i. e. without distinct air chambers or septal and
siphonal parts. Body usually distinctly dor soventr ally flattened. Ventral
margin of mantle with rounded projections near the funnel indentation. Fins
occupying more than seven eighths of sides of mantle. Primary eyelid
contracted in the embryo to a narrow pore, forming a "cornea" surrounded
ventrally by a secondary lid fold which encloses the pore (cf. pp. 504 and
564). Arms with 2—4 and tentacle clubs with 4—8 rows of suckers. The
tentacles can be coiled and retracted completely into the pockets, which
occupy almost the whole ventral side of the head. Ventral arms with
muscular margins which extend on the ventrolateral side to the head (like
the swimming margins of the other arms). Ink sac very large, displaced
during ontogeny from its typical position above the hind intestine to the
right and growing toward the posterior end of the mantle sac (Figure 289).

2. TYPICAL STRUCTURE OF THE SEPIINAE

The typical habitus does not differ much from the form of Sepia offi-
cinalis, which may be a little plumper and is characterized by the
structure of the skin. The activity of the chromatophores creates a variety
of effects which are caused by the activity of the iridocytes and the
cutaneous muscles (cf. Octopodidae). The body is covered with typically
situated warts formed by circular and stellate cutaneous muscles,
especially on the sides of the mantle sac below the fins, and on the dorsal
surface (Figure 303b). The lateral lines are light stripes or ridges of
cutaneous muscles enclosing the ventral side of the mantle sac (Figures 307,
311, 317). If they are maximally raised and widened, they form the margins
of an incomplete sucker which permits the animal to adhere to the substrate
(sand, rock). This can be easily observed by keeping young Sepia in a
glass dish and creating a current with a pipette; the animals will not be
displaced unless they move voluntarily (cf. Bather, 1895).

528 The typical characters of the subfamily are most distinct in the species
related to S. aculeata. This applies particularly to the shell, which
still shows a raised fork, as in Belo sepia.* The Mediterranean species
S. orbignyana resembles the type most closely; S. officinalis is
more specialized in many parts.

The shell is oblong -oval. Its anterior margin is parabolic or slightly
produced in the middle, indicating a proostracum. (This is present in the
aculeata group, in the fossil species of the Miocene, and in embryonic
shells which are blunt in the later stages (see Figure 301 on p. 541). The
septa of the incrassation are moderately convex (Figure 309), very delicate,
much more densely arranged than in Belosepia; they leave anteriorly
only a very narrow free stripe on the dorsal shield. The depression of the

■> Cf. d'Orbigny, 1893, Sepia, Plate 5, bis; Plate 19, Figure 7; Plate 21, Figure 4; Plate 25, Figure 4; 1845,
Plate 13, Figures 11—12 (S. aculeata, rouxi, blainvillei); also Ortmann, 1890,S. microcoty -
ledon,S. framea; Hoyle, 1901, S. k6ttlitzi,etc.; Lang, 1900, p. 100, Figure 109.

506

siphuncle is longitudinally oval (Figure 309) and extends anteriorly to about
half the length of the shell. This depression is very shallow, especially
anteriorly, so that a secondary convexity may be formed (Figure 283). The
siphuncle becomes deep, like a funnel only posteriorly, dorsal to the fork
(Figure 285). The septal necks in the region of the incrassation can be
distinguished from the septa because they form narrow stripes which pass
from one septum to the preceding septum. The fork resembles that of the
aculeata group (Figure 282).

(529)

FIGURE 290. Diagrammatic cross sections through the posterior part of the body of Sepia officinalis,
illustrating the topographical relationships between shell (Sch), muscular mantle (Mm), fins (Fl),cephalo-
podial and funnel retractors (Mu), gills, etc. a) Posterior part; b) middle of shell. Compare with Figure 268
on p. 497. a) The muscular mantle is attached ventrally to the dorsal shield near the fork (Ga), which is
exposed toward the mantle sac; the fins are attached in their primary position to the outside of the shell
and are articulated with the shell (Figure 114). b) The muscular mantle is inserted at the inward curved
margin (Rd) of the shell, the fins are attached to the muscular mantle. The powerful retractors of the
cephalopodium (1) and funnel (2), are attached to the incrassation and to the inner side of the margin and the
liver is situated between them (Lb).

Kb -- ligament of gill; Ka - branchial artery; Kv - branchial vein; Km - branchial gland; B1 — branchial
lamellae; Bh -- their insertion; Tg — ink duct; Vc -- vena cava; Ed — hind intestine; Mh -- mantle cavity;
Fk -- fin cartilage; FI ~ capsule of articulation of fin; Ss - shell sac; Sp - shell septa.

The convexity of the dorsal shield is shallow anteriorly, but it increases
posteriorly, as in all Sepioidea (Figure 283b). We consider this as distinct
resemblance to older conditions. The posterior margin of the shield, i. e.

507

the homologue of the ventral process (Figure 285), is also sharply curved
posteriorly from the attachment of the fork. The derivation of this structure
is still recognizable in its development: its form and structure differ early
from those of the lateral edges (Figure 288; cf. Vol. II, Plate XX, Figure 4;
also Figures 279 and 301 ).

The rostrum is a long, pointed, laterally compressed spine. It is slightly
curved dorsally as in Be lose pi a (p.52l). The spine forms a terminal
point between the fins. It is covered by skin and mantle sac in the living
animal (contrary to Abel, 1916, p. 162) (cf. S.orbignyana, Figure 311).

In principle, the spine resembles that of Spirulirostra or of the
Belemnitidae as it consists of radial fibers and concentric layers.

These characters determine the position of the shell in the mantle sac.

The dorsal shield is, of course, decisive, because its margin must follow
the whole circumference of the muscular mantle (cf. p. 501 ). The complex
history of the dorsal shield is still expressed in the more detailed topo-
graphical relationships and even in the form of the margin itself. This is
solid and calcified only at the posterior end from the early stages.

(Figure 288); the lateral margins have a delicate, pliable zone which extends
to the anterior part. This "conchiolin margin" is always curved inward

529 ventrally, especially laterally. This curvature may be considered to be
caused by shrinking during drying. Cross sections (Figure 290) show,
however, that this is a natural structure and the muscular mantle is inserted
at the outer side of the inward curved lateral edges.

Figure 291 shows the topographical relationships between shell and soft
body. The different regions show a different formation and a simple formula
cannot be given. (The soft parts are not attached directly to the shell but to
the shell sac so that a strict coordination of the shell parts to certain
attachments of muscles is not necessary.) The anterior part of the shell
is almost directly adjacent to the dorsal part of the dorsal cavity; it is
covered only by the primary mantle, and adheres through it to the broad
neck cartilage. This applies especially to the zone that is hatched in
Figure 291, which is also slightly cartilaginous. The other part (cf.

Figure 312) is normally occupied by the soft body, which is attached in the
middle by strong muscles on a wide stripe on each side. These are the
retractors of the cephalopodium and funnel, which originate as a single
muscle from the shell; the cut surfaces are clearly recognizable in the
figure. (The dorsal and lateral cephalopodial retractors are divided by the
pallial nerve.) The muscular mantle is attached to the inward curved
margin of the dorsal shield in the entire anterior and middle zone, i. e.
to the dorsal side of the shell.

530 In the posterior part of the shell (the "wings," where these are
differentiated), the attachment of the muscular mantle is displaced to the
ventral side; the dorsal side would be covered only by the shell sac and the
skin if the fin muscles were not related to the shell. The base of the fins
is situated directly on the shell, without a capsule of articulation in the
posterior part, so that a concrescence with the shell sac can take place at
least at the spine and in its vicinity. The fork, depression of the siphuncle,
and a small corner (x) are without muscular attachments and are clearly
visible after the viscera are removed. Not only the free margin of the
dorsal shield but also a lateral stripe of the incrassation is occupied in the
middle by attachments of muscles.

508

FIGURE 291. Topographic relationships between shell and soft body in Sepia officinalis (male). The
viscera are removed; the cephalopodial retractors are cut, so that their attachments are distinct. The gill
ligaments (Kb) are cut near the mantle, also the posterior pallial veins (Vp), anterior pallial veins (VI) and
posterior pallial arteries (Ap). This figure is slightly corrected in "Fossile Tintenfische," 1922, p. 86. The
shell (shown separately in Figure 306a) is visible shining through at some points. The slightly curved line
(Gr) connects the stellate ganglia and shows the depth of the mantle cavity at this point. The area above this
line is only thinly covered by the shell sac and skin; it adheres to the neck cartilage in life. The hatched parts
of this area (Nk) are also cartilaginous. Median to the stellate ganglia, the thick muscles are cut which are
inserted on the free margin of the dorsal shield and the adjacent parts of the incrassation, or the corresponding
parts of the shell sac: retractors of head (1,2,3) and funnel (4). The free posterior part of the dorsal shield is
covered by the muscular mantle, which is attached here on the ventral side of the shield. The fork, the me-
dian parts of the siphonal depression, part of the last lamella of the incrassation and 2 small corners (x) of
the shield margin are free or covered only by the shell sac, without attachments of muscles. Note; 1) the
zones occupied by the large muscles; 2) the part occupied by the posterior part of the muscular mantle;

3) the anterior part of the shell, which is covered only by the primary mantle; the large nerves of the stellate
ganglion (cf. Figure 47 on p. 124); gill ligament; lateral pallial veins; posterior pallial vessels; translucent
spine; fins; mantle cartilage (cf. S pi ru la. Figure 277).

531 The different relationship of the muscular mantle to the posterior part
of the shell is perhaps connected with its derivation from the "ventral
proc ess " of the shell of Spirulirostra (Figure 2 7 1 ), in which the mantle
has to be attached in the area directed inward and anteriorly.

Near the shell, on the inner side of the muscular mantle, pass the
posterior pallial veins and arteries, which enter the muscular mantle;
further anteriorly are the lateral pallial veins and the gill ligaments, and
still further anteriorly are the stellate ganglia with their nerves.

The finer structure of the shell can be derived in part from its morpho-
logical derivation. Sections through the incrassation, fork and the dorsal
plate show the typical components, if they are not obliterated by differentia-
tion, but the general modification has caused secondary complications.

509

This is shown, for example, in the median sections in Figure 292. The
conotheca and periostracum would have to be distinguished first (after
532 Figure 260 on p. 487); this was not possible on the decalcified sections
through the embryonic shell, probably due to inadequate technique.*

Figure 292a shows the differentiation of the plate; however, the conotheca,
on which the pillars and septa are deposited (19, 18), is apparently very
delicate while the adjacent periostracum has time to develop. The further
differentiation of these layers becomes more distinct in the latter stages
(Figure 292c).

FIGURE 292. Median sections through the shell of
Sepia officinalis (interpreted and diagrammatic,
partly after Appellof, 1893).

a) Embryonic shell after formation of the first septum
with the siphonal depression (18). Already at this stage,
the primary shell (21,ostracum and hypostracum) and

the secondary shell (22, periostracum) can be distinguished,
which was impossible in preparations, or in decalcified
sections (see Appellof, Plate 6, Figure 2). The part of the
phragmocone, i.e. the flattened first chamber is already
completely embedded in the periostracum, which also
bears the recurved ventral process (20). 19 — pillar in
the area of the initial siphuncle, corresponding to the
prosiphuncle (Figure 279).

b) Polished section through the parts facing the siphonal
depression (cf. Appellof, Plate 2, Figure 1; Plate 5,

Figure 4). 1— septum; 2— septal neck of next septum;

3 — "filling substance of posterior corner of chamber";

4 - air chamber; 5 - pillar inside it; 6 - intermediate
septum.

c) Polished section through anterior end of shell (cf.
Appellof, Plate 2, Figure 2). 1 — outer layer of perio-
stracum ("dorsal plate of Appellof); 2 - inner layer of
periostracum ("middle plate"); 3 - boundary of calcifi-
cation; 4 - calcified part of middle plate; 5 - ostracum
("layer of pillars" of "inner plate"); 6 — hypostracum
("black stratified layer of inner plate"); 7 — growing
anterior margin, belonging to periostracum; 7-8 - growth
zone of inner layer of periostracum; 8—9 - growth zone
of ostracum; 9—10 — growth zone of hypostracum (a part
of the shell epithelium as matrix of development corres-
ponds to each of these zones). 11 (-16) - soft, gelatinous
stratified shell mass with embedded pillars (17), corres-
ponding to the following "cavity layer" which produces
cavities by the shrinking of the filling material, after
formation of the new septum; 12 - attachment of the
preceding septum (15) on the hypostracum; 13 - sup-
porting ridge ("filling substance of anterior corner of
chamber"); 14 - pillar; 15 (see 12); 16 (see 11); 17
(see 11); 17a - intermediate septum; k - last chamber.

• Riefstahl (1885) also did not find the "inner plate" below the spine.

510

A new development of the structure of the shell of Sepiidae is the inner
skeleton of the air chambers. The septa of the Sepiidae are very delicate
and vulnerable and are easily crushed. This is prevented by thin inter-
calated lamellae of calcified conchiolin, the interseptal and pillar lamellae.
The pillar lamellae are longitudinally arranged, perpendicular to the septa
and almost parallel to each other. Their attachment on the later formed
septum forms an almost regularly undulate line. In the lateral parts the
pillar lamellae are no longer sagittal but radiate from each other. The
lamellae show numerous structural variations which will not be described.

There are 1—4 interseptal lamellae in each air chamber. They are
more or less parallel to the septa and are only present in the later formed
air chambers. These lamellae strengthen the pillar lamellae and their
development is connected with that of the air chambers, which are formed
in stages, by secretion of aqueous, gelatinous layers of chitin which later
dry and are replaced with air (cf. Appellof, 1893). The interseptal lamellae
are the remnants of these layers of chitin. The interseptal lamellae do not
reach into the anterior part of the air chambers, in which the pillars are
stronger but less numerous (Lk in Figure 284). The pillars are similar in
structure to the prosiphuncle (see also p. 511) and also to the structures
between the septal necks of Spirula.

The form of the mantle sac is determined mainly by the shell in the
typical basic form and in the atypical variants. In addition to the dorso-
ventral compression (p. 525), the anterior margin of the dorsal shield causes
an anterior curvature of the dorsal mantle margin (Figure 316) because it
has to keep pace, so to say, with the growth of the air chambers (Figure 285).
This curvature ends laterally in a curve from which begins a blunt pro-
jection around the funnel indentation. This is shallow, with a transverse,
truncate posterior margin. There are no sharp corners.

(533)

FIGURE 293. Pupils of Sepiidae. a) Sepia officinalis; b) Sepia elegans;
c) Sepia orbignyana. Note the different form of the iris flap, which divides
the opening of the pupil and the field of vision into separate zones for objects
before, beside and behind the animal. This is most marked in c.

The fins occupy the greater part of the sides of the mantle, as in many
adult Teuthoidea (Ctenopteryx, Ancistrocheirus, Octopodo-
teuthis, Neoteuthis, Thysanoteuthis, S e pi o t eu th i s), but there
533 remains a small free part near the anterior margin, and also the middle of
the posterior end is not occupied by the fins (Figure 311). As in all
Sepioidea, the fins are separate or only apparently connected by cutaneous
muscles. They form wide margins which are widest near the posterior
end (Figures 302, 303). The fins end posteriorly in wide, rounded lobes
which project beyond the end of the body; they form typical "earlobes'*
anteriorly. There are no lateral corners at the posterior end. There is a
direct muscular connection between fins and shell, particularly with the
posterior spine (p. 530); otherwise, the base of the fin is articulated with

511

the fin cartilage and capsule its whole length. The articulation is situated
on the shell sac posteriorly, on the muscular mantle in the anterior part
(Figure 290).

The head is rounded. The olfactory organ (Figure 297) is a small, oval,
slightly raised area of epithelium, surrounded by a projecting margin which
is often contracted above it so that a typical olfactory pit is formed.

FIGURE 294. Part of the arm of S e p i a o f fi c i n a 1 i s, after removal of the
suckers. 2x. Note particularly the arrangement of the stalks of the suckers.

They are of simple form and normally arranged in the inner rows, longer and
united into a common stem with the supports of the protective margin in the
marginal rows, so that they are situated on the base of the margin. The short
accessory supports also originate on the common stem (see Figure 44 on p.ll8).

(535)

FIGURE 295. Tentacle club of an older embryo of Sepia officinalis
(cf. Vol.II; Plate XVIII, Figure 5). Note the still completely octoserial
arrangement and the beginning differentiation of a stalk part and a hand
part. The arrangement in the adult has to be interpreted on the basis of
this condition. Protective margins are still absent, and the swim-
ming margin (on the left) is restricted to the distal part. The suckers are
only "anlagen" in form of papillae, 40 x.

The eye is covered with a transparent "cornea," as in the Loliginidae
(p. 173). The eye chamber opens to the outside only by a small pore which
is often covered by the anterior margin of a secondary lid fold around the
cornea. This lid forms dorsally only a narrow ridge or is recognizable
only by the contrast between the delicate cornea and the tough normal skin
(Figure 303b, dotted). Anteriorly, posteriorly and especially ventrally, this
lid forms a free fold which can be drawn over the eye from below and usually
creates a kind of pocket. The cornea hardly deserves its name; it is very
soft and pliable and probably elastic in the live animal; it can be protruded
and extended if the eye chamber is completely filled with water (as in
L o 1 ig o).

The pupil is very characteristic. The widened upper margin of the iris
(the iris flap) is normally bilobed, the lower margin indented so that a
w -shaped opening is formed.

Arm formula: 4, 3, 2, 1. The dorsal arms are of almost equal length,
otherwise similar, the ventral arms are markedly longer and of special

512

character. Distinct external membranes connect the arms which are
absent only between the ventral arms, but they are connected by cutaneous

534 muscles the contraction of which can form a similar skin fold. The outer
side of the 3 dorsal pairs of arms is rounded; swimming margins are
present in the middle in form of thin longitudinal crests which may form
folds. The swimming margins continue proximally as thin "glandular lines"
(p. 562, Figure 317b) which extend on the head. The lateral margins of the
ventral arms have a special structure. They form wide muscular folds on
the outer side of the arm; they are finlike at the base (Figure 303) and
extend to the ventral side of the head to near the eyes where they suddenly
end. This is specific for the Sepiidae; they serve as a rudder, like the
"swimming margins" of the 3rd pair of arms in the Teuthoidea.

The inner surface of the arms is occupied by distinct, wide, uniform
protective margins and the outer rows of suckers are situated on the base of
these margins; their pillars (p. 118) are fused with the supports of the mar-
gin, forming a common basal part so that the free terminal part of the stalk
apparently originates on the support (Figure 294). On the same basal part
originate shorter accessory supports (p. 118), rudimentary supports
belonging to the inner rows of suckers, so that similar conditions are formed
as on the quadriserial tentacles and arms of the Teuthoidea (Figure 90 on
p. 195). The protective margins of both sides are united at the base of each
arm; this occurs only inthe Sepioidea, and also on the tentacle clubs in the
Sepiolidae (Figure 339).

The tentacles have very long stalks and short clubs, as in the Sepiolidae.
The stalk is not rounded but has 3 blunt longitudinal edges, one on the outer
side and two at the borders of the inner surface. These two edges continue
into the protective margins of the club or lead directly to them; the
efferent vein is often visible between them as a thin light line.

The clubs are formed in principle as in the Sepioidea (p.488) (Figure 296).
Swimming margins are present along the entire length of the club close to
the dorsal protective margins, to which they adhere. A deep groove divides
the two folds and extends to between the muscular bases of the marginal
suckers. The ventral protective margins are very wide and enlarge the
clubs so that they appear leaf-shaped. The ventral protective margins are
covered with suckers which extend to the marginal zone and their lengthened
stalks are fused with the margins. The supports of the two protective
margins are not very distinct and their arrangement and numbers corres-
pond to those of the marginal suckers.

535 The two protective margins converge markedly toward the base of the
club. However, they do not become united as in the Sepiolidae, but end toward
the inner margins of the stalk. The arrangement of the suckers can usually
be derived embryologically from a regular octoserial arrangement
(Figure 295). However, they become rapidly biserial at the base and quadri-
serial at the apex. The club is widest in the middle, and the suckers are
largest, but the longitudinal rows differ in this respect. The largest suckers
are situated in the third row from above, but they may be only slightly
larger than the others, and this is certainly typical (Figures 339, 295, p. 487).
In many Sepiinae, however (cf. Sepia), the largest suckers (i. e. a few in

the 3rd row) are still larger than shown in Figure 296. The marginal rows
of the ventral side also show a tendency to displacement and apparently to

513

reduction (p. 488), but the regular octoserial arrangement often persists,
either as in Figure 295, or still more regular, as shown in Plate XXI,

Figure 9 in Vol. II (see also Hoyle, 1886, Plate 22, Figure 1 , club of Sepiella
maindroni de Rochebr.). Figure 296 shows the more specialized club of
Sepia officinalis to characterize the direction of development.

(536)

FIGURE 296. Left tentacle club of S e p i a officinalis. 2x.
Diagrammatic. The numerals indicate the number of the longi-
tudinal row after an assumed primary pattern of 8 rows (Figure 295).
The suckers of some rows are enlarged. Note also the arrangement
in the basal and distal parts; the formation of protective margins
(Ds, Vs) and the swimming margin (Sm); the terminal swelling (Ek)
and the adjacent suckers (xx); cross section of stalk and efferent vein.
This figure is reproduced by mistake in this form (cf. Figure 305).

The numbering of the marginal rows on the right is wrong. The
oblique rows should bear the following numbers from below:
a) 1—6; b) 1, 2, 3, 4, 5, 7, 6, 8, etc.; 7 and 6 are always interchanged.
The outer connecting lines always enclose the last 3 suckers of an
8th row (7,6,8 should read: 6, 7, 8).

The very large pockets of the tentacles occupy nearly the whole ventral
side of the head (Figures 289,297). The tentacles are completely retracted
into the pocket at rest and only their relatively narrow entrance between
the buccal funnel, the 3rd and 4th arms and the interbrachial membranes
indicates their position (Plate VI, Figure 2). The retracted tentacle is
coiled into an S-curve (Figure 297); It is extended (Figure 297b) when the
animal intends to attack (Figure 307); a sudden lengthening of the stalk
throws the club forward (p. 119). The stalk is normally attached, but the
weak ligament is no longer superficial as in the type of Decapoda (Figure 45
on p. 119) but situated deep inside the pocket.

5110/2

514

536 The suckers of Sepiidae usually have an irregular^ crenelated dentition
so that the smooth margin of the projecting ring is apparently divided into
"teeth" by narrow, more or less densely arranged incisions (Figure 298).
These incisions, however, may also be incomplete or absent in some places
so that the teeth are replaced by a continuous sharp edge. The adhesive
ring is finely granulate or covered with small papillae which are strikingly
like teeth in the small distal suckers, at least near the distal margin
(Plate XIII, Figures 4, 6, 8, 9).

The suckers are usually cup-shaped and more spherical at the apex.

The marginal ring is smooth on the tentacle clubs, it is fluted or divided
into rounded elevations on the arms.

The suckers of the median rows of the arms often have narrowed
openings which permits them to adhere to narrow surfaces, like the hooks.
This is also a division of labor as described on p. 128, and it is especially
marked in the male (Figure 299, also p. 539).

537

FIGURE 297. Head and funnel of Sepia elegans. The funnel has been cut off and the
pocket of the tentacle has been opened after removal of the ventral arms (A4):

a) the stalks of the tentacles are coiled inside the pockets; the right club has been re-
moved, the left is drawn out. Note the relationship of the base of the tentacle (Tt) to that
of the axis of the ventral arms (A4); ligament (x) and club.

b) the tentacles are assumed to be extended forward, leaving the pockets empty. The
pockets have a narrow opening, bordered externally by the thick membrane between the
3rd and 4th arms. Also shown: funnel, its pockets and bonds; olfactory organ (Ro); buccal
funnel with its points, pillars and attachments; protective margin; suckers. Ai-A4 - arms;

B2-B3 — buccal pillars.

The buccal funnel is typical in general, i. e. with 7 pillars, points and
attachments. The second attachment ends opposite the dorsal protective
margin of the 2nd arm, the third, against the ventral protective margin of
the 3rd arm, the fourth, opposite the inner margin of the 4th arm. All
these attachments are unconnected with the respective margins. In Sepia
aculeata (p. 546), the points still bear a few small suckers at the apex.
The 6 buccal pockets are shallow (Plate VI, Figure 2); the mouth cone is
typical.

515

The funnel shows a number of characteristics (Figure 312): l) the
pockets are connected with the head by a projecting skin muscle which I
name "adductor infundibuli lateralis"; this muscle has been observed so
far only in the Sepiolidae and has been confused there with the adductor
infundibuli externus of the Ommatostrephidae (p. 414). 2) The funnel bond

is short, oval to ear -shaped and occupies only about the posterior half of the
funnel pockets; the mantle bond is correspondingly a short, rounded longi-
tudinal comb. 3) The neck bond corresponds to the opposite adhesion surface
of the mantle (Figure 291), it is very wide, shieldlike, of characteristic form,
with a narrow median longitudinal groove (Figure 322). The other conditions
are typical.

(538)

FIGURE 298. Large sucker of the arm of a mature female of Sepia officinalis,
highly magnified, a) Opening from above; b) median section. Teeth, suction pad,
adhesive and marginal ring (fluted) .horny ring, circular muscle, and pocket of stalk
are visible (see Figure 121).

FIGURE 299. Sucker of a median row of the 3rd arm of a mature male of Sepia
officinalis. 36 x. a) Opening: b) median section. Compare with Figures 298
and 325. Note the general form, the transverse contraction of the opening, the
strengthening of the circular muscle (Rm),the invagination (Sg) of the stalk (St) the
deep pocket (Gr) of the suction pad (Sk),the narrow opening, the fusion of the teeth
into a continuous marginal edge (Sc, Vr) which is widened in the distal part, and the
narrower, higher form of the suction chamber and of the supporting horny ring.

Tr - stalk of sucker, basal pad; Nv - nerve; Hr - adhesive ring; Ca - fluted part
of outer side (Au).

The mantle situs (Figure 312) shows some special characters: the form
of the broad funnel retractors is determined by the form of the shell, the
renal papillae are long and situated near the anal papilla, as in Spirula

516

(p.514, see also Sepiolidae, Figures 364,366). Another character present
also in the Sepiolidae and many Teuthoidea is the incomplete penetration
of the mantle cavity into the posterior part of the mantle sac. Anal papilla,
gills, and juvenile genitalia of the female (nidamental glands, accessory
glands) are typical. On the other hand, the ink sac is characteristically
turned around the hind intestine (Figure 289) and extends into the posterior
part of the mantle sac under the skin (VoL II, Plates XVIII and XIX),
displacing the posterior aorta from its original median position to the left,
so that it has to form a curve around the ink sac to reach the middle again
and to divide into 3 branches.

(539)

FIGURE 300. Development of the female genitalia of Sepia officinalis.

a) Female, a few weeks old, 10 x. The nidamental glands are small, short sacs and the glandular lamellae are
little developed. The accessory glands are still separated, rounded, with radial ridges. Cf. Figures 48,49

on p. 125.

b) Female, a few months old, natural size. The nidamental glands are still small but typically differentiated.
The accessory glands are much larger and in contact in the middle; a zone of radial ridges is still present in
the anterior and central parts (p.l25); the larger middle part is occupied by dense glandular pores, the openings
of short tubules. The radial ridges are still visible but the tubules have grown near the lateral and median
margins to form a compact glandular mass. The opening area begins to become concentrated.

c) The process has developed further. The area of the openings has become more contracted, while the tubules
under the skin form a pad. The radial ridges persist only between the two areas of opening and are recognizable
in the arrangement of the last formed tubules. (Half-grown specimen, 8/9 x.)

d) (see also Plate VII). The organs are mature (2/3 x). The accessory glands are united in the middle into

a single zone surrounded by the pits of opening of the nidamental glands which are separated in the middle and
concentrated near the openings of the nidamental glands. Note also: anal and renal papillae, ink sac, posterior
aorta, gills, and the protruding part of the oviduct (genital process).

517

538 The jaws are of the typical form of Decapoda and closely resemble those
of the Teuthoidea (e. g. Loliginidae) (cf. Plates XVII, XVIIl). However,
they are much stronger and almost uniformly brown at an early stage,
while only the biting processes are brown in the Loliginidae, the other parts
being yellowish brown. The radula is typical for the Sepioidea, without
marginal plates and with only unicuspid teeth (Figure 31 on p. lOO).

The skin is characteristic for the Sepiidae. The chromatophores can be
divided into 3 categories: more or less dark brown, yellowish brown and
orange to ochre yellow (see S. o f f i c i n a 1 i s, p. 549 ). Of particular interest
is the surface, where groups of cutaneous muscles form transient papillae
in a constant position, particularly the larger papillae. They are con-
centrated on the dorsal side of head and body and laterally on the ventral
surface of the mantle. Groups of chromatophores associated with the
papillae form recurring spots. Like the Octopoda and other benthic forms,
the Sepiidae produce a stereotype pattern of skin and coloration resembling
the ground as a reaction to certain stimuli, e. g. the "lateral lines, " narrow,
light stripes on the ventral side, lateral to a median zone in which the chroma-
tophores are sparser (p. 527). These lines should not be confused with the
2 more lateral rows of light spots or papillae (Figure 307).

The sexes differ also in coloration and pattern. The mature male is
slightly more brightly colored than the female (see S. officinalis). Of
special importance is the hectocotylization of the arms in the maturing
male. This is not restricted to the left ventral arm but also affects the

539 suckers in the median rows of the lateral arms (Figure 299, cf. Figure 298).
The suckers become more spherical, as in the Sepiolidae (Figure 325),
rather than cup-shaped; the circular muscle becomes very strong, while
the stalk is inserted in a deep invagination and the pocket in the suction pad
becomes deeper. The horny ring of the sucker is very narrow, its opening
is narrowed and the teeth are replaced by a continuous, sharp, flat edge.

These changes apparently enable the male to hold the female during copu-

540 lation without injury. The spermatophores are attached in the medio -
ventral part of the buccal funnel, between the two ventral supports of the
funnel and the folded skin of the inner side. The left ventral arm is the
hectocotylus and some of its suckers become reduced.

The female genitalia correspond to the width of the mantle sac and
resemble those of the Sepiolidae (Figure 340 ). The nidamental glands are
broadly oval (Plate VII, Figure 2; Figure 300) in contrast to those of
Teuthoidea (Figure 84 on p. 185). The large accessory glands are situated
before their opening; they are broadly fused in the middle. The middle of
the fused glands projects and seems to be a separate formation. The area
of opening of each gland forms an impressed stripe lateral to this apparently
single structure. The two zones of opening are nearly contiguous anteriorly.
The accessory glands are visible during dissection also in the mature
animal (Figure 318), because the nidamental glands do not extend as far
anteriorly as in the Teuthoidea (Figure 84 on p. 185). The protruding end of
the gonoduct ("genital process") is typically situated in both sexes (Plate VIl),
projecting from the branchial pocket and fused with the body for some
distance in the middle. The female process is of characteristic form,
with a flasklike basal thickening caused by the development of the oviduct
gland. It is narrower distally and opens in a lateral slit.

518

The shell (cT Figure 306 and p. 201), differs markedly in both sexes,
at least in the adult (cf. Figure 306), but only in its proportions. The
posterior marginal zone of the dorsal shield near the fork is broader in
the female. This is due to the enlargement of the ovary and to the plumper
form of the whole posterior part of the body.

3. TYPICAL DEVELOPMENT OF THE SEPIINAE

The Sepiidae, like the Sepiolidae and other Sepioidea, reach a high degree
of differentiation inside the egg, because of the large amount of yolk. The
newly hatched animal already closely resembles the adult, and the post-
embryonic development shows no special characters. From the egg of
Sepia officinalis hatches a typical Sepia which can be identified
to the subgenus. However, it shows some juvenile characters: the body
is relatively shorter and more compact; the arms are relatively shorter;
the suckers are little differentiated, especially on the tentacle club and the
541 fins are still short and mainly developed at the posterior end, and "ear-
lobes" are still absent (Figure 307).

FIGURE 301. Embryonic shell of S e p i a officinalis L.

a) Median section; 1 - ventral process; 2 - initial part of siphuncle; 3 -- chambers with pillars;

I— IV - 4 septa at the transition to the siphonal part; 4 - septal neck part of 3rd septum; 5 ■“

siphonal depression of 4th septum; 6 - pillar for 5th septum; 7 ~ conotheca in part of proostracum;

8 “ periostracum (dorsal shield).

b) Lateral view; 9 — border of calcified parts; la-IVa - sutures.

c) Dorsal view; 10 - comer between lateral edge and "ventral process."

d) Ventral view; I—IV siphonal lines of the 4 septa; x — indication of proostracum. 10 x.

The juvenile shell is wider, the septa less numerous. The shield has a
rudimentary posterior part (Figure 301 ). The distinction between fork
and marginal incrassation is less marked, and the rostrum is only a small
"anlage" which develops relatively late. These juvenile characters are
gradually replaced by definitive characters, but traces of them persist for
a long time.

The club of the freshly hatched animal (Figure 295; VoL II, Plate XVIII,
Figure 6) resembles that of the adult, but its stalk part is characteristic.

519

This stage would resemble the definitive form if a suppression of the
small suckers of the stalk were assumed. However, the process is quite
different: the small suckers of the stalk part grow gradually, the zone on
which they are situated becomes gradually wider proximally, they are in-
cluded in the hand part and become rapidly the main suckers of the club,
while the suckers which appear at first as such form the distal part. New
suckers do not develop after hatching.

4. VARIATION OF THE SEPIINAE

The variety of forms in the Sepiinae, as in the Loliginidae (p. 191), is not
completely known. The typical species were placed in the genus Sepia,

542 and the more different forms were considered as subgenera or genera.

A general revision* is urgently necessary, but this cannot be made here.

The principles on which the characteristics of the different species should
be based have been discussed above. We describe here only the major
variations of the general type. Further study of these variations and their
correlations is necessary before a natural classification of the Sepiidae can
be made. Such a study has to be based on the shell, which shows distinct
characters:

1. The dorsal shield may be short oval, oblong oval (ancestral form),
oblong rhomboidal, short rhomboidal, straight or forming a sharp angle
posteriorly, more or less ventrally curved, winged (Figures 310, 315) or
without wings. 2. The spine may be long and strong, short or absent,
sinuate or straight (Figure 283; see also S. el eg a ns). 3. The ventral
process may be curved posteriorly, reduced or directed downward; it may
surround the posterior end of the mantle sac like a scoop or the cone of
the Teuthoidea. 4. The fork may be low and like a ridge posteriorly or ver-
tical as in B e 1 o s e p i a, or it may adhere to the marginal incrassation like
a pocket. It may also be folded over posteriorly (Figures 308 and 309).

5. The siphonal depression may be restricted to the posterior half or dis-
appear anteriorly (in old animals) on the marginal incrassations. 6. The
posterior boundary of the lamellae of the marginal incrassation may form

a free curve or an almost transverse line in young forms (Figures 308, 314);
or it may be changed atypically by the formation of lateral incisions
(Figure 314). 7. The lamellae of the marginal incrassations may form

shallow convexities or very steep angles, and form a thick median comb
which may be swollen or keeled (S.lefebrei d’Orb. ).

Other important characters are found in the club. The 8 rows of suckers
can increase to 16—32 rows with small, uniform suckers or remain in a
pattern of 8 regular rows of small suckers. The suckers may also form
8—6 rows (Figure 296) in which the 3rd row from above includes a number
of more or less enlarged suckers (the usual condition**).

The two lobes of the iris flap may be simple, rounded and little marked
as in Sepia officinalis; they are produced into more or less long and
narrow corners in other forms (S. e 1 e g a n s, S. o r b i g ny a n a). If the flap

*■ Including the fossil shells (see Fossile Tintenfische, pp. 82 and 92; also p. 546 below).

** All these variations must be taken into account in descriptions, which should give precise data. This could

make a natural classification possible. Shells should be stored in alcohol.

520

is lowered, it does not form a narrow, uo -shaped slit as in S. officinalis
but an opening divided into 3 parts, i. e. a division of the field of vision into
3 zones situated before, beside and behind the animal (Figure 293). The
543 following key is based on some less general characters suitable for the
definition of smaller groups.

1. Posterior end with a glandular pore into large pocket above the

posterior end of the shell. Shell narrowly oval, with posterior wings;
spine absent. Tentacle club with 8 regular rows of small suckers . . .
Genus 1 . Sepiella Gray

2. A row of glandular pores on each side of ventral surface of mantle.

Shell very delicate, broadly oval with oblique, conical marginal
incrassations which leave large part of the dorsal plate exposed

I ....... Genus 2. Hemisepius Steenstr.

3. Glandular pores on mantle sac absent

Genus 3. Sepia L. s. restr. Keferstein, 1866

Rochebrune (1884) attempted a further division of the genus Sepia
without the necessary knowledge and he could therefore not establish a
natural classification.

521

544 Chapter 41

GENUS SEPIA

L., 1758, s. restr. Keferstein, 1866

Contents: a. Diagnosis. — b. Typical structure and development of S e pi a c. Variation of type of
Sepia. — d. Sepia (Eusepia) officinalis (p. 547). — e. Sepia (Parasepia) orbignyana
(p. 554). — f. Sepia (Parasepia) e le ga ns (p. 560).

a. DIAGNOSIS

Mantle sac without glandular pores (p. 543). Shell without marked
deformation of marginal incrassations. Shell wide to narrowly oval or
slender, rounded rhomboidal to awl-shaped, with or without distinct wing-
like widenings at the posterior end. Spine sometimes absent.

b. TYPICAL STRUCTURE AND DEVELOPMENT OF SEPIA

I The characters of the subfamily (pp. 527 — 541) apply to the genus
Sepia, because the different forms (p. 542) have been excluded. This
applies to the adult and the juvenile stages.

c. VARIATION OF THE TYPE OF SEPIA

Many species are not well known and a comprehensive classification can
therefore not be given. The principles on which such a classification should
be based are given on p. 542. This would result in a number of subgenera.
We shall describe only those species which explain the relationships of the
Mediterranean species. These species form two related groups which can
be characterized as follows: Shell oval, not tapering markedly posteriorly.
Apex of arms normal, not whiplike. Hectocotylization affecting the base to
the middle of the left ventral arm, which bears at least one pair of normal
suckers at the base and has a normal apex; the middle is modified, the
545 suckers are smaller and broad stripes of wrinkled skin pass between the
rows. The third row from above of the tentacle club bears markedly
enlarged suckers (Figure 296), and the adjacent suckers are less enlarged.

The two groups are distinguished as follows,

a) Body broad, markedly flattened. Shell broadly oval, indistinctly
winged (Figure 309a). Siphonal stripes near lateral margin markedly
curved from the beginning, without pointed angles projecting sharply

522

posteriorly. Hectocotylus normal at least in distal half. Subgenus 1.
Eusepia n, subgen. Type species: S. officinalis L.

b) Body more cylindrical. Shell of adult forming a slender oval, with
distinct posterior wings (Figure 310). Siphonal stripes with pointed angles
projecting posteriorly near lateral margin; forming very shallow curves
in the juvenile part. Hectocotylus normal only at the apex. Subgenus 2.
Parasepia n. subgen. Type species: S. orbignyana Fer.

DIAGNOSIS OF SUBGENUS EUSEPIA

Mantle sac markedly flattened. Shell broadly oval. Dorsal shield curved
posteriorly, with indistinct wings; fork gradually disappearing on dorsal
shield. Siphonal stripes laterally without sharp, posteriorly projecting
angles which would form a regular longitudinal edge. Tentacle club: third
row from above with a few markedly enlarged suckers; swimming margin
always shorter than zone with suckers. Hectocotylus resembling a normal
arm at least in distal half. Iris flap (Figure 293) divided into 2 small rounded
lobes at the margin. Coloration usually gray to dark brown.

This group contains several species related to S. officinalis, which is
the type species. Some of its characteristics mentioned in the diagnosis
may have to be omitted in a definition of the group.

DIAGNOSIS OF SUBGENUS PARASEPIA

Mantle sac only slightly flattened, almost cylindrical. Shell with distinct
posterior wings; posterior boundaries of lamellae of marginal incrassations
proj.ecting posteriorly into a regular row of lateral corners (Figure 310).
Part of hectocotylus resembling a normal arm not extending beyond distal
third. Iris flap (Figure 293) produced into 2 narrow points at the margin.
Swimming margin of tentacle club longer than zone with suckers . Coloration
usually yellowish brown to reddish brown.

The Mediterranean species S. orbignyana Fer. and S. elegans
d’Orb. belong to this subgenus. My observations, and the incomplete
published descriptions show that these two species resemble a number of
546 species from other regions and form a natural group. Some of the above
characters may have to be omitted in a definition of this group, but they
have to be considered.

The following characters are typical: 1. Presence of distinct "wings”
on the dorsal shield. 2. Posterior boundary of the lamellae of the marginal
incrassations. 3. Structure of tentacle clubs. Not only the marked
widening of the swimming margins is characteristic but also the form of the
proximal part of the protective margin, especially of the ventral margin,
which (Figures 311, 317) curves around the part with suckers and delimits
it sharply proximally; the ventral margin passes close to the dorsal margin,
and then turns suddenly toward the stalk and disappears. The club of
S. orbignyana and S . elegans thus differs distinctly from that of
S. officinalis.

523

Exotic subgenera:

c) Body and siphonal stripes as in Eusepia. Rostrum very large,
forming a terminal process between the fins. Dorsal shield narrowly oval;
ventral process weakly developed, projecting downward, not curved
posteriorly along the rostrum. Fork large, projecting. Club without
markedly enlarged suckers as in Figure 296 but with very small, almost
uniform suckers in more than 8 longitudinal rows. Hectocotylus unknown.
Type species: S. aculeata Hasselt (pp. 528, 537), East Asia.

Subgenus 3; Ac antho sepia de Rochebrune, 1885 (nom, emend.).

d) Body (and siphonal stripes?) as in Parasepia but still more
slender, with a narrow awl-shaped shell the lateral edges of which converge
posteriorly at an acute angle to a ventrally curved end with small,
projecting wings; posterior margin projecting and with a large spine which
protrudes externally as a terminal tubercle or process. Arms more or less
whiplike at the end; ends of arms markedly lengthened on one or both dorsal
pairs in some cases. Hectocotylization restricted to apex of arm.

Subgenus 4. Doratosepia de Rochebrune, 1885 (nom. emend) (cf.

Wiilker, 1910, p. 17 ff.). Type species: S. and re ana Steenstr.

e) Body and siphonal stripes as in Eusepia. Shell broadly oval.
Rostrum large, forming a more or less distinct tubercle at the posterior end
(as in c). Ventral process very blunt, closely adjacent to rostrum (as in c).
Fork low, partly folded over. Club with ’’anlage” of 16 — 8 rows, with or
without markedly enlarged suckers (transition from Acanthosepia to
Eusepia, etc.). Subgenus 5. Platysepia nov. Type species:

S. esculenta Hoyle (1886, Plate 17).

f) Shell without spine; marginal incrassations raised into a large median
comb as in S . lefebrei d'Orb. Subgenus 6. Lophosepia de
Rochebrune, 1885 (nom. emend.).

g) Shell without spine, markedly rhomboidal. Subgenus 7. Metasepia
Hoyle, 1884. Type species: M. pfefferi Hoyle, 1886 (Plate XXI).

547 d. SEPIA OFFICINALIS L., 1758
1. DIAGNOSIS

Arms constantly with 4 longitudinal rows of suckers. Tentacle clubs
with 110 — 130''”!' suckers, including 5 — 6 markedly enlarged suckers in the
third row. Shell with a large sharp spine and a posteriorly curved ventral
process of the dorsal shield; fork flat, gradually disappearing. Hectocotylus
with 4 normal rows of suckers in the larger distal half.

2. LITERATURE

1758 Linne, S. officinalis.

1554 Rondelet (Lib. XVII, pp. 497 — 506, Figs.), Sepia.

* A general revision of the recent and fossil Sepia will be given elsewhere.
Confirmed only for the form in Naples. Varieties are possible.

524

1718 Ruysch (Lib. IV. p. 9. Plate I). Sepia.

1738 Swammerdam (p. 876), Sepia.

1758 Seba (p, 6, Plate 3), Sepia.

1758 Borlase (p. 268). Sepia.

1762 Stroem (p. 137), Sepia.

1772 Scopoli (p. 127), Sepia officinalis.

1772 Pennant (Vol. IV, p. 55), Sepia officinalis.

1784 Schneider (p. 108), Sepia officinalis.

1784 Gronovius (p. 244), Sepia officinalis.

1791 Wolfen (p. 379), Sepia officinalis.

1801 Lamarck (Vol. 2, p. 7), Sepia officinalis.

1802 Bose (Vol. 2, p. 45), Sepia officinalis.

1805 Montfort (Vol. I, p. 171), Seiche commune.

1817 Cuvier (Plate 3), Sepia officinalis.

1822 Bowditch (Plate I, Fig. I), Sepia rugosa.

1822 Lamarck (Vol. II, p. 371), Sepia officinalis.

1824 Cams (p. 317, Plate 28), Sepia officinalis.

1824 Martens (Vol. 2, p. 436), Sepia officinalis.

1826 Payraudeau (p. 172), Sepia officinalis.

1826 Blainville (p. 84, Plate XI), Sepia officinalis.

1826 Risso (Vol. 4, p. 8), Sepia officinalis.

1827 Brughiere (Plate 76), Sepia officinalis.

1828 Delle Chiaje (Vol. 4, p, 51), Sepia officinalis.

1832 Deshayes (p. 944, Plate 3), Sepia officinalis.

1835 Ferussac and d'Orbigny (p, 260; Seiches; Plates 1, 2, 3), Sepia officinalis.

1837 Oken (Vol. 5, p. 534), Sepia officinalis.

1837 Rang (p. 90), Sepia officinalis.

1838 Potiez and Michaud (p. 8, Plate I), S e p i a officinalis.

1838 d'Orbigny (p. 20), Sepia officinalis.

1841 Cantraine (p. 14), Sepia officinalis = S. savigny Blainv.

1844 Philippi (Vol. I, p. 203), Sepia officinalis.

1851 Verany (p. 65, Plates 24, 25), Sepia officinalis. ‘i

1855 d'Orbigny (p. 272), Sepia officinalis.

1863 Aucapitaine (p. 368), Sepia officinalis.

1869 Targioni-Tozzetti (p. 57), Sepia officinalis.

1869 Lafont, (p. 11), Sepia officinalis.

Lafont, Sepia filliouxi (new species, identical with the typical Naples variety with a long
siphonal zone (Fig. 306 on p. 552).

1871 Lafont, Sepia fischeri (after Fischer, 1875, p. 20; new species, identical with the adult form
cf. p. 550).

1874 Fischer,?, (p. 368), Sepia officinalis.

1874 Fischer,?, (p. 369), Sepia filliouxi.

1875 Fischer,?, (p. 20), Sepia officinalis = fischeri Laf., filliouxi Laf.

1879 Tryon (p. 188, Plates 86, 87), Sepia officinalis.

1880 Tiberi (p. 25), Sepia officinalis.

1880 Stossich (p. 160), Sepia officinalis, j
1884 Rochebrune (p. 74), Sepia officinalis.

1884 Giard (pp. 309, 310), Sepia officinalis and filliouxi .

1885 Ninni (p. 159), Sepia officinalis, mediterranea.

1890 Cams (p. 453), Sepia officinalis, filliouxi Laf., fischeri Laf.

1894 Pelseneer (p. 206), Sepia officinalis. *

1896 Jatta (p. 149, Plates 2, 3, 7, 15, 16). Sepia officinalis. ' -

1908 Pfeffer (p. 57, Figs. 63, 64), Sepia officinalis.

1909 Bauer (Plate 10, many places). Sepia officinalis. .

1913 Meyer (Fig. I, different places). Sepia officinalis.

1916 Naef (Syst.) (p. 16), Sepia officinalis. ’ '

1921 Naef (Syst.) (p. 538), Sepiaofficinalis. 'i - : ■ ■ ^

1921 Grimpe (p. 299), Sepia officinalis (North Sea).

525

548 3. STRUCTURE OF THE ADULT ANIMAL

The habitus of the plump, dorsoventrally flattened species is well known,
although there are no good illustrations of live specimens (cf. Part 3, which
will appear later). The drawing in Jatta (1896, Plate 3, Figure 3) is based on
Verany (1851, Plate 24, Figure a) and has been reproduced repeatedly: the
outline is not exact, the coloration is that of a pale female, while the white
marginal line of the fins is a characteristic of the male. The exact outlines
of a live specimen are given in Figures 303, 304 and 307.

FIGURE 302. Sepia officinalis,
mature female, after a fresh, slightly con-
tracted specimen. Half natural size. Note
outline of mantle sac, fins, structure of
arms and tentacle clubs, especially the
arrangement of suckers, swimming and pro-
tective margins. Only the outer rows of
suckers on the protective margins of the
arms are visible. The spots on the fins are
in fact white. Note also the form of the
funnel and of the small skin folds on the
ventral side of the head.

FIGURE 303. Sepia officinalis, mature male, after a
preserved specimen. Half natural size, a) Ventral. Compare
with Figure 302, especially the arrangement of suckers, which
are arranged in 4 rows on the inner side of the arm, because of
the contraction of the protective margin. Note the contracted
buccal funnel, swimming margins of ventral arms, attachment
of funnel, form of fins and mantle, structure of club (arrangement
of suckers, protective and swimming margins), b) Dorsal.

Note the (white) longitudinal stripes at the margin of the fins
(Rl) which are characteristic for the male. Also arrangement of
skin warts (Wa); anterior mantle margin, funnel pockets (Tt);
eye region with cornea (Co), lid fold, pupil (Pu); interbrachial
membranes; swimming margins of ventral arms (tentacles
retracted).

549 Sepia officinalis appears drab because of its mainly grayish brown
coloration. Closer examination, however, shows striking effects of colora-
tion and pattern, caused by the action of iridocytes, chromatophores and the

526

opalescent, translucent consistency of the flesh. These effects are
intensified by the changing structure of the surface caused by the complex
musculature and innervation of the skin.

FIGURE 304. Sepia officinalis, male, after a live animal. Half
natural size. Anterior part of specimen in Figure 303, in 2 typical positions:
a) sitting on the bottom, with retracted head and arms; note the widespread
lateral margins of the ventral arms; b) swimming, about to project the
tentacles, the distal part of which is already extended between the arms;
note their position and length.

There are 3 types of chromatophores: dark, or more or less
reddish; yellowish brown; orange-yellow. The iridocytes give mainly
yellow green and reddish reflexes; some groups of iridocytes on the fins
and arms form white to whitish green spots (Figure 303) combined with
groups of brown chromatophores which may cover the iridocytes so that
they are replaced by dark spots, especially after preservation. A distinct
characteristic of the male is the white seam of more or less confluent white
spots near the margin of the fins. During the mating period, the males
compete with each other by displaying their colors, particularly by a diffuse
zebra striping of the dorsal surface (dark and pale in the female) and
spreading of the fins and the lateral margins of the 4th arms, which are
striped like the fins. The skin warts which appear in the excited animal
have a constant position. They form a longitudinal row on each side of the
550 mantle above and below the base of the fin, ventrally, some distance from
the base and lateral to the lateral lines (p. 527). Larger warts are present
on the dorsal side of head and mantle. The small warts are dense on the
dorsal side and usually not recognizable iii' preserved specimens. These
variable structures will be described in greater detail in Part III.

The shell shows a mixture of typical characters, resembling Belo -
sepia and specific characters. The dorsal plate is of normal form,
especially in the broadly flattening (Figure 309a) but not completely absent
wings. Proostracum absent anteriorly (p. 520), ventral process curved
downward posteriorly along the spine, as in Belosepia. Thin lines
which cross the growth lines radiate from the posterior end of the fork

527

and indicate the displacement of different points of the matrix during
development. The marginal incrassations are typical; the fork is folded
over (see p. 545) and becomes flatter on the posterior part of the dorsal
shield (Figures 283, 309c: iig).

Dorsal surface rugose, tuberculate, white in the middle; from the spine
radiate the shining, yellowish brown, noncalcified outer layers (p. 531), and
the similar marginal stripes of the otherwise calcified dorsal shield, which
correspond to the insertion of the muscular mantle (cf. Jatta, 1896, Plate 15,
Figures 42, 43; Appellof, 1893, Plate 1, Figures 1 — 2; examine material,
which is available everywhere).

The shell varies markedly, but my studies did not show any characters
which would justify the establishment of subspecies or varieties, because
the differences observed could not be correlated with other differences.

I shall mention only 3 points of divergence (see p. 551 on sexual differences).
1. The siphonal depression, i. e., the siphonal part of the marginal
incrassation may occupy V3 to almost % of the length of the shell. The
species can be divided in 2 sharply defined forms by this character: one
with a short siphonal part, the other with a long part. In the first group
(Figure 309), the length is about half of the whole phragmocone or less,
in the second group (Figure 306) it is distinctly longer. 2. The shell and
the whole animal grow further after sexual maturity. The siphonal
depression has a sharply angular anterior boundary in profile in the young
animal (Figure 309a) but it disappears later because the last formed
lamellae of the incrassation become increasingly shorter and incomplete.
Such a senile condition cannot be used as a systematic character. The
greater or smaller length of the siphuncle is already distinct in the juvenile
shell (Figure 308). 3. The lamellae of the incrassation are much more
densely arranged, i. e., far more numerous in a form of shell which shows
also other special characters. The size of adult animals varies widely,
from about 8 to 30 cm.

The fins occupy almost the whole sides of the mantle. Only a few
millimeters at the anterior margin (about ^35 of the length) remain free in
551 a normally contracted animal (Figure 303). Their width in the adult is

usually about a third of the width of the mantle in the live animal. Preserved
specimens rarely show this proportion. The mantle margin, olfactory
tubercle and eye region are typical. The pupil (see p. 533) is closed to a
narrow, m;- shaped slit; it resembles a bent sickle if opened. The iris flap
forms 2 rounded lobes.

The arms bear 4 regular rows of suckers, the tentacles about 125 suckers,
of which 5 — 6 in the third row are markedly enlarged (Figure 303). The
longitudinal rows differ in many other species of Sepia (p. 535).

The dorsal marginal row contains only slightly enlarged suckers. The
suckers of the 2nd row are at least twice the normal size. The suckers
of the 3rd row are enlarged 4 — 6 times. The 4th row resembles the 2nd
and the 5th resembles the first. The 3 marginal rows are pressed into
each other, forming an irregular marginal row which therefore contains
3 times the number of suckers in the corresponding parts. The club of
this group (p. 544) thus bears 6 rows. There is an inward curved swelling
at the apex, and 2 — 3 slightly larger, stalkless, flat suckers before it
forms a terminal part as in the Teuthoidea (p. 229).

528

The polymorphism of the arm suckers was described on p. 538 (see also
Plate XIII). To summarize: 1. The suckers at the base of the arms bear
more or less fused teeth which form a sharp edge. 2. Distally follow

suckers with long, crenelated teeth at the distal margin
and small, low teeth at the proximal margin. 3, The
terminal part bears small suckers with a widened
adhesive ring. 4. The small suckers of the club have
small, blunt teeth all around, especially on the distal
margin, and also show a widened adhesive ring. 5. The
large suckers of the club have relatively narrow
adhesive rings and more or less fused teeth, and
they are inflated.

The buccal funnel is usually shrunken beyond
recognition in preserved animals (but see Figure 2 of
Plate VI). Seven supports, points and attachments are
distinct in live specimens, but suckers are absent.

The typical conditions of the mantle cavity (p. 489)
are adapted to the compact form of the mantle sac
(Plate VII). The funnel bond is about ovoid, with the
apex posteriorly, the median margin is truncate and
the lateral margin more curved. The jaws (Plate XVIII,
Figure 1) and radula (Plate XV, Figure 5) show no
special characters.

There is a distinct sexual dimorphism. The sexes differ in the propor-
tions of the body, at least in the adult. The posterior part of the body of the
smaller female is relatively broader, especially posteriorly. Exact
measurements cannot be given, because the proportions depend markedly
on the state of contraction (Figures 302, 303). The large development of
52 the ovary (Plate VII), which affects the proportions of the body, affects also
the shell (Figure 306). The free marginal zones in the posterior part of the

FIGURE 305. Club of
Sepia officinalis,
natural size. Compare
with Figure 296, p. 536.

FIGURE 306. Shell of male (a) and female (b) of
Sepia officinalis. (after the variety
common in Naples). Note the wider shell of the
female, especially the markedly widened lateral
edges (Fr) of the shield in the posterior part, and
the slight widening of the fork.

529

shell of the mature female are markedly widened, more distinctly than in
other species of Sepia. These changes later affect also the fork, and the
whole shell assumes its characteristic form.

On the mating coloration of the male see p. 549; the special characters
of the suckers are described on p. 538, The larger distal part of the
hectocotylus is normal. It bears a few normal suckers at the base. There
is a coarsely wrinkled part of the inner surface with very small suckers in
which longitudinal rows are characteristically displaced: the 3 upper rows
are widely separated and the 4th row passes close to the 3rd.

4. POSTEMBRYONIC DEVELOPMENT

Freshly hatched animals show the characteristic habitus of these stages
(Figure 307), The arms are of about equal length; the ventral arms are not
markedly longer than the others in live and dead animals. The juvenile shell
is of particular interest (Figure 288). It is oval and very wide, and contains
7—8 air chambers. The ventral process is narrow, tuberculate, calcified,
distinctly separated from the widened lateral plates and without a soft
margin of conchiolin. The marginal incrassations and the narrow fork are
simply but distinctly differentiated, at least on the later septa, but their
connection is still distinct. The outer side of the first chamber bears a
small, blunt, conical, terminal spine (Vol. II, Plate XX).

553 About a month after hatching, the shell develops to the stage shown in
Figure 308. The outline is still simply ovoid, and the posterior parts of the
shield project little, especially in the region of the spine, where the
embryonic structure is still recognizable. The spine has already the form
of a large, slender cone which is apparently situated free on the outside of
the shield. The fork projects posteriorly although it is still very low. The
septa on the incrassations formed after hatching differ distinctly from those
formed before hatching: they are denser and widen rapidly anteriorly.

Further growth does not show distinct changes in principle, and the shell
gradually attains its definitive form. Figure 309 shows a juvenile sheU.

The fork still projects, the anterior end is again parabolic ally pointed and
the posterior part of the shield is widened (lateral plates). This part shows
a radial striation toward the embryonic shell; the lateral view of this part
shows the beginning of wings (Figure 310). The spine is large and projecting,
only little covered by the posterior margin of the shield. The whole shell
is still very wide.

The shell of the adult differs markedly from the above (Figure 309c).

The free margins of the shield become relatively much wider posteriorly,
but the whole shell becomes relatively narrower by its growth anteriorly.

In the region of the spine, the plate grows posteriorly and partly covers the
spine; the lamellae of the fork extend at first to the outer side of the fork
and disappear further on the shield. These changes are connected with the
fact that the posterior part of the shell is displaced anteriorly on the dorsal

554 side so that the spine does not project externally (Figure 291), because it
has become enveloped by noncalcified strengthening layers which are
deposited on the posterior part of the shield and contrast sharply in
coloration and sheen to the other parts (’'Dornhulle'* of Appellof).

530

(553)

FIGURE 307. Freshly hatched S e p i a offici-
nalis, drawn after a live animal. 4X. Dorsal
aspect of a swimming animal, with the tentacle
clubs ready to catch a My sis. Such prey can
be caught with great accuracy at a distance of
1-2 body lengths. The sudden extension and
retraction of the club brings the prey to the
mouth held by the suckers. It is then passed to
the arms and eaten. The habitus resembles that
of Spirula (Figure 277); the resemblance is
distinct in some parts, but other parts show the
specific characters of Sepia. The corneal fold
of the eye has been closed; the slit- shaped
pupil is of characteristic form. The lateral
margins of the ventral arms are distinct and,
like the lateral lines, pass at rest ventrolaterally
on the mantle sac, which is quite flat and
adheres to the bottom. The fins show the typi-
cal form posteriorly (usually incorrectly shown);
the growth at the anterior end is not completed.
There is an anchor- shaped glandular stripe in
the posterior dorsal part of the mantle sac. This
is Hoyle’s organ, which becomes active at
hatching and soon disappears.

FIGURE 308. Shell of a young Sepia officinalis, about one month
after hatching. 6 x. Note the general form, the development of the
siphonal depression (Si) and fork (Ga). Note posteriorly the perpendicular
ventral wall (Vw) and its free margin (Fr). Compare with Figures 288
and 282;

Rd — lateral edge; Hr — ventral process; Do — spine; Si^ and Si^g — 1st
and 16th septal neck ("siphonal stripe"). Beyond the 7th or 8th, the
septa are postembryonic; they become suddenly denser, indicating the
time of hatching, which can also be determined in older or fossil shells."

* This applies not only to Sepia but also to other types, e. g., of Nautilidae. Th. Staub (Verb. Schweiz,
naturf. Ges. Schaffhausen, 1921, p. 151) reports (according to my data) that one of the first air chambers
(the 7th in Nautilis pompilius)is markedly smaller than the preceding chambers, and the following
chambers increase only gradually in size. The small chamber is apparently the first formed after
hatching, at a time when the animal lived in difficult conditions. This is also the case with Sepia.

He also states that in stranded shells of adults, in contrast to young animals, the last chamber or
sometimes the last two chambers are markedly smaller (such shells also have a black, thickened margin);
specimens caught alive usually show a constant increase of the distance between the septas in the later
formed parts.

531

FIGURE 309. Shells of
Sepia officinalis,
a) Lateral; b) ventral view
of the shell of an almost
half-grown animal. 2 X ;
c) shell of an adult male
(% X) with an especially
short siphonal part (Si)
(variety 2).* Only half
of the lamellae of the
incrassation are shown.

Wu— incrassation; Hr —
posterior margin of shield;
Vw— ventral wall of
phragmocone; Rd — free
margin of ventral wall;

Ga — fork; Sr — siphonal
margin or last siphonal
stripe; Sw — last septum;
Ch — chitinous margin of
the shield; Kr — calcified
margin of shield; ug —
folded over fork(cf.
Figure 283 on p. 521).

555 e. SEPIA (PARASEPIA) ORBIGNYANA
FERUSSAC, 1826

1. DIAGNOSIS

Arms with 4 rows of suckers. Tentacle clubs with about 110 suckers,
and 3 particularly enlarged suckers in the 2nd row. Shell with a very large
posterior spine; margin of shield projecting freely from the spine and
(Figure 112 on p. 235) enveloping the posterior end of the mantle sac like
a cone. A process supported by the spine situated between the fins at the
posterior end. Hectocotylus with 2 zigzag rows of small suckers,
separated in its greater part by a folded area of skin.

2. LITERATURE

1826 Ferussac (d’Orb. 1835, p. 156), Sepia orbignyana.

1835 d'Orbigny (p. 273, Seiches, Plate 5), Sepia orbignyana.
1847 Delle Chiaje (Vol. I, p. 12, Plate 15), Sepia orbignyana.
1849 Gray (p. 109), Sepia orbignyana.

* Cf. p. 550. There are numerous such shells in the Zoological Station without indication of locality,
probably from the vicinity of Naples.

532

1851 Verany (p. 70. Plate 26), Sepia elegans.

1855 d’Orbigny (p. 274, Plate 13), Sepia orbignyana.

1863 Aucapitaine (p. 379), Sepia orbignyana.

1869 Targioni-Tozzetti (1) (p. 23), Sepia elegans.

1869 Targioni-Tozzetti (2) (p. 60), Sepia orbignyana.

1875 Fischer (p. 14), Sepia orbignyana.

1879 Tryon (p. 198), Sepia orbignyana.

1880 Tiberi (p. 25), Sepia elegans.

1884 Rochebrune (p. 104), Acanthosepion orbignyanum.
1886 Hoyle (p. 26), Sepia orbignyana.

1890 Cams (p. 454), Sepia orbignyana.

1890 Norman (p. 484), Sepia elegans.

1896 Jatta (p. 156, Plates 4, 7, 16), Sepia orbignyana.

1908 Pfeffer (p. 59, Figs. 65, 66), Sepia orbignyana.

1916 Naef (Syst.) (p. 16), Sepia orbignyana.

1921 Naef (p. 538), Sepia orbignyana (System).

1921 Grimpe (p. 300), Sepia orbignyana (?) (North Sea).

3. STRUCTURE OF THE ADULT ANIMAL

Figure 311 shows a preserved specimen of S. orbignyana. The
coloration is caused mainly by light (yellow) and dark (brownish red)
chromatophores and is dirty yellowish brown which may pass into pink
and reddish brown. The warts form two constant longitudinal rows in the
ventrolateral region of the mantle (Figure 311). Lateral lines are absent
or indistinct.

The shell (Figure 310) is narrowly oval with parabolical, pointed anterior
margin; there are distinctly delimited small wings posteriorly. The ventral
process projects from the spine. The posterior end of the shell forms a
spoon which surrounds the end of the mantle sac like the cone of the
Teuthoidea. This markedly convex, juvenile part of the shell in the middle
(Figure 314) is connected later by increasingly flatter parts. The fork
remains low but does not disappear as in S. officinalis. The very
characteristic siphonal stripes are transverse. They begin posteriorly
as flat, simple curves which gradually straighten in the middle and become
curved further on posteriorly. Still more characteristic is the later
development of posteriorly directed angles in the lateral part. The regular
arrangement and increase in size of these angles is a special character of
the subgenus (p. 545).

556 An important, distinctly specific character is the pointed process between
the fins, caused by the marked development of the rostrum which is not
covered by the recurved posterior margin of the shell (Figure 311) as it is
inS. officinalis. The rostrum is, of course, not visible in the intact
animal, although it projects in damaged specimens and has often been so
illustrated (1912, Ceph., p. 248). Because of this form of the rostrum, the
fins are widely separated posteriorly, but they are otherwise of typical
form. The fins are relatively shorter than in Sepia officinalis —
they leave about V12 of the mantle free anteriorly.

The mantle sac is narrow, corresponding to the slender shell, and is
only slightly flattened. The head is also relatively narrower than in Sepia
officinalis. Olfactory organ, funnel, eye region and arms are typical.

533

a

FIGURE 310. Shell of an adult female of S. orbignyana, natural
size, right side and ventral, a) Note the decreased curvature in
comparison with the juvenile shell (Figure 314) which extends hereto
the point marked with an x. Note also spine, wings, and incrassa-
tions. b) Note the differentiation of the shield; the noncalcified
(dark) marginal zone; wings; fork; the varying transverse striation
of the incrassation. Only half of the lamellae (siphonal stripes) are
shown.

The arms bear 4 rows of suckers which pass into 2 zigzag rows at the
base. The suckers are more sparse and less numerous than in S. offici-
nalis but similar in general. The tentacle clubs have also fewer suckers
(about 110) and are relatively shorter. The protective margin projects
proximally beyond the part with suckers, while the protective margins before
this zone are almost united and then disappear rapidly down the stalk.

About 3 suckers are particularly large.

Buccal funnel and funnel resemble those of S. officinalis (p. 551).
The funnel bond (Figure 312), however, is more elliptical, not narrowed
posteriorly.

557 The mantle cavity shows the typical conditions; jaws and radula show no
special characters (Plate XV, Figure 8; Figure 31 on p. 100).

Sexual dimorphism is markedly less developed than in S. officinalis:
the shell of the female is only slightly widened, and the nidamental glands
remain small, which is connected with the care for the offspring. Jatta
(1896, Plate 8, Figures 7, 8) confused the eggs of this species with those of
S. elegans and vice versa. The eggs of S. orbignyana are deposited
in a sponge, and complete gelatinous envelopes are not formed.

The greater part of the hectocotylus is modified: there are a few normal
suckers at the base and the distal quarter is also occupied by normally
arranged suckers. Between these zones is the folded inner surface,
bordered by 2 zigzag rows of small suckers (Jatta, Plate 16, Figure 12).

558 4. POSTEMBRYONIC DEVELOPMENT

Freshly hatched animals resemble the following stages so closely that
they are difficult to identify before preservation. The rostrum is still too

534

(557)

535

FIGURE 311. Mature female of Sepia or bigny a na , natural size. Note apex of body; short fins (cf. Figures 302, 303); slender
form; dorsal margin of mantle; arrangement of suckers on the arms; form of club (arrangement of suckers, length of swimming
margins).

small to be visible from the outside, and the outer spine is little marked
in much further developed stages (Figure 313). On the other hand, the
suckers form 4 regular rows on the arms, whereas they form only
2 zigzag rows in S. elegans.

FIGURE 312. Sepia or bigny a na , young female with
opened mantle cavity. The middle ventral part of the
muscular mantle is cut off and the mantle bonds are longi-
tudinally slit. The mantle cavity does not reach to the
posterior end (Plate VII): r

1 — accessory gland (invaginated glandular mass); 2 —
pores of the latter (invaginated zone of opening); 3
growing grooves for the "anlage" of funher parts of the
gland (cf. p. 539); 4 — adductor infundibuli lateralis.

The next stage already has the coloration of the adult, but it retains many
juvenile characters typical for the species. Fins, mantle margin, head and
arms resemble those of the young forms of other species of Sepia; on
the other hand, the marked projection of the rostrum at the posterior end
is a characteristic for this species., The arms are very short, particularly
their free part, because the interbrachial membranes occupy more than
half of the length of the arms. The mantle sac is wide because of the form } ,
of the shell which resembles that ofS. officinalis.

Larger stages than that shown in Figure 313 still show the above
characters. However, they are specific for subgenus and species as shown
in Figure 314.

The shield is broadly ovoid, with a blunt anterior end, as in offici-
nalis, and has distinct wings posteriorly. The spine is laterally
compressed, and the fork is typical and projects upwards posteriorly.
Particularly important is the course of the posterior boundary of the
559 lamellae of the lateral incrassations which is typical for Parasepia;

the first lamellae form a very shallow curve, in contrast to S. officinalis
(Figure 301 on p. 541); the later lamellae form a sharply broken line,
which forms a sharp corner near the fork that projects posteriorly and
separates a marginal zone. These corners form together a thin longitudinal
ridge on each side which extends on the siphonal part of the incrassation
near the lateral margin. The later lamellae also form a rather shallow

536

curve, in contrast to S . elegans (Figure 315). A common character of
S. orbignyana and S . elegans is that the posterior margin of the
shield bears a tuberculate deposit which is directed posteriorly and
corresponds to the beak ofS. elegans, but this gradually disappears.

The shell of the adult (Figure 310) attains the typical slender form, with
a parabolic anterior end and a markedly enlarged wing part.

FIGURE 313. Young Sepia orbignyana (2 x , drawn after life), 4— 6 weeks
old. The habitus resembles that of all young forms of Sepia (Figure 307),
but there is a distinct posterior point which contains the spine. Note the rela-
tively short fins; the short arms are connected most of their length; the iris
fold with pointed corners. The mantle margin is juvenile; it projects less
sharply anteriorly than later (Figure 311). The shell resembles that of similar
stages ofS. officinalis. The following characters are typical; juvenile
form of fins; form of mantle margin; funnel pockets (visible before the
mantle margin); convex cornea; large lens (black in the drawing); protruding
eyes.

FIGURE 314. Shell of a young S e p i a orbignyana. 4X.
Lateral and ventral. The lateral view shows the strong curva-
ture and gradual rotation of the lamellae of the incrassation,
which resembles the primary ventral curving of the phragmo-
cone in the Sepioidea. Note the free margin of the shield of
the wings; the posteriorly curved spine; the rough thickening
on the posterior margin of the shield which is directed toward
the spine (cf. Figure 283 on p. 521).

60 f. SEPIA (PARASEPIA) ELEGANS D’ORB.,

1 8 3 5 (= BISSERIALIS MONTFORT?)

1 . DIAGNOSIS

Arms with 2 simple or zigzag rows of suckers. Tentacle clubs with
60 —70 suckers, of which 2 (or 3) are markedly enlarged. Spine replaced
by a small, rough, longitudinal comb. Posterior margin of shield forming
a rough, beaklike median process which is curved posteriorly.

537

2. LITERATURE

1802 (1808) Montfort, S . bisserialis (identity uncertain).

1835 d’Orbigny (F^rrusac and Orb. p. 280; seiches, Plates 8, 27), Sepia elegans.

1835 d*Orbigny (p. 275, Plate 3), Sepia ruppellaria.

1844 Philippi (p. 203), Sepia rubens.

1849 Gray (p. 105), Sepia r u p pe 11 a r i a , — elegans .

1851 Verany (p. 73, Plate 26), Sepia bisserialis M.

1855 d'Orbigny (p. 285), Sepia elegans.

1869 Targioni-Tozzetti, (p. 62), Sepia bisserialis M.

1869 Fischer (p. 125), Sepia ruppellaria (including also S . bisserialis).

1875 Steenstrup, Sepia elegans (S. bisserialis).

1879 Tryon, Sepia ruppellaria (S. bisserialis).

1880 Tiberi, Sepia bisserialis (including S. rubens).

1884 Rochebrune, Rh om b os e p i on rupellarium (including also S . bisserialis — elegans).
1886 Hoyle (p. 24), Sepia elegans and S . ruppellaria (including also S . bisserialis).

1890 Norman (p. 484), Sepia ruppellaria.

1890 Carus (p. 454), Sepia elegans and ruppellaria (including also S . bisserialis. rubens).
1896 Jatta (p. 160, Plates 5, 7, 8, 16), Sepia elegans.

1908 Pfeffer (p. 60, Figures 67 — 69), Sepia elegans.

1916 Naef (p. 16), Sepia elegans (System).

1921 Naef (p. 538), Sepia elegans (System).

1921 Grimpe (pp. 299, 303), Sepia elegans (North Sea).

3. STRUCTURE OF THE ADULT ANIMAL

Figures 316 and 317 show the typical form of a live and a preserved
specimen. The bright coloration is caused mainly by light (yellow) and dark
(brownish red) chromatophores and varies between pink, yellowish brown
and brownish red with greenish reflexes. The ventral side is lighter and
shows distinct pale or raised lateral lines, and near them typically
distributed warts in a longitudinal row (Figure 317a).

The shell (Figure 315) is very characteristic. Its form closely resembles
that ofS. orbignyana. The posterior end of the dorsal shield bears
small, sharply delimited, lateral wings. The presence of a small beak,
directed posteriorly, on the ventral process of the posterior margin is
characteristic (cf. Figure 314). The fork extends as a narrow stripe on
the incrassation, the curvature of the siphonal stripes is more marked and
is shallow only in the middle. The shell is very fragile. Figure 316,
drawn after a live animal, shows the outline of the translucent anterior part
of the shell; preserved specimens are more or less deformed.

561 The fins are of normal form and slightly shorter than in S. orbignyana,
they leave anteriorly Vio~ Vii of the side of the mantle free. The mantle
projects sharply dorsally in the middle, but less than in S. orbignyana.
The head is normal. The cornea protrudes occasionally, suggesting a
specific character which is shown in a less pronounced degree also in
Figure 317.

The formula of the arms is normal (p. 533), but the differences in the
adult are small, in contrast to the youngest forms (Figure 319). The
position of the suckers is important. They are stated to be always biserial,
but the arrangement usually varies between 2 and 4 rows. All arms bear
2 regular zigzag rows in many specimens which become simple rows at the

538

base; a distinct quadriserial pattern is hardly ever found.'-' On the other
hand, there are all transitional stages to a simple biserial condition, which
is especially frequent in the male. However, zigzag rows are usually
present at least in the middle of some or all arms. The arms may differ
in this respect: a regular pattern of 2 rows is found most often on the
dorsal arms, least frequently on the ventral arms. The tentacle clubs
show the typical characters of the subgenus, but are relatively small and

bear only 60 — 70 suckers, i. e., much

FIGURE 315. Shell of an adult S e p i a elegans,
female, natural size. The dotted line indicates a
noncalcified marginal zone (1) which is particularly
distinct in the wings (2). Note at the posterior end;
the "beak" (3) and comb of the rostrum (4). The
fork forms a ridge. The development of the curved
siphonal lines which delimit the posterior boundaries
of the lamellae of the incrassation.

fewer than in S. orbignyana.

FIGURE 316. Adult, live female of Se pi a elegans,
natural size. Note form of fins and mantle; distribu-
tion of skin papillae; anterior mantle margin and
position of the translucent shell; cornea, lid fold, iris
flap, pupil (black); dorsal arms with weakly developed
swimming margin; ventral arms with strongly de-
veloped lateral margins. The small drawing shows
cornea, lid fold (3), pupil (4), iris flap (2) with
corners (1).

The buccal funnel bears no suckers. The funnel bond (Figure 318)
resembles that of S. orbignyana (p. 558). The mantle cavity shows the
typical characters of the genus (Figure 318), but the ink sac does not extend
562 as far posteriorly as in the other species. The genitalia are typical; the
nidamental glands are relatively small, but the accessory glands are
particularly large (Plate VII). Jaws and radula show no special characters.

* D’Orbigny (1839, Plate 27, Figure 5) shows the arms with 2 proximal and 4 distal rows, apparently diagrammatic.

539

FIGURE 317. Sepia elegans. Preserved, half-grown female, natural size. Compare with Figure 316,
which shows a slightly older, more slender animal with retracted tentacles. Note particularly form and
length of fins; mantle margin; warts and lateral lines on the mantle; slightly protruding cornea, pupils
and glandular lines; the rudimentary swimming margins on the 3 dorsal pairs of arms; the arrangement of
the suckers on the arms, the twisted tentacle stalks with 3 edges, the long swimming margins and the clubs
with two large suckers; buccal funnel and inner lip.

FIGURE 318. Opened mantle cavity of mature female of S epia
elegans (natural size). Note form of fins, mantle and thick-
ness of mantle. The eyes (cornea) are not protruding. Note;
funnel, funnel pockets and their connection with the head by the
lateral adductor muscle, funnel cartilage, gills and their attach-
ment, anal papilla, renal papillae, vena cava, accessory and
nidamental glands, ink sac, posterior aorta with branches,
posterior boundary of mantle cavity, translucent ovary; distal
part of oviduct ("genital process"), which projects from the left
branchial pocket. Note particularly the relation of the openings
of the nidamental glands to the zones of opening of the accessory
glands.

»

540

The hectocotylus bears a few normal suckers at the base. The distal
quarter or third is also normal, usually with two rows. There is a wrinkled
zone between them with very small suckers arranged in two, usually
simple, rows. The whole inner surface is turned upward.

Mature males are always slightly smaller than females. The maximal
dorsal length of the mantle is 7 cm for males and 8.1 cm for females.

4. POSTEMBRYONIC DEVELOPMENT

The youngest stages of this species closely resemble those of

5. orbignyana. However, closer examination of the arrangement of the
suckers on the arms identifies the species asS. elegans. A striking

563 character of the fully developed embryo is the large development of the

ventral arms (Figure 319). The interbrachial membrane is less developed,
and the shell is broadly oval, which is typical for the young forms.

FIGURE 319. Fully developed embryos of Sepia
elegans, 5x. Dorsal view; note the still short
fins; the translucent shell (3); the anchor-shaped
Hoyle's organ (4); the mantle margin projects
little, or not at all; an interbrachial membrane
connects the dorsal arms; the ventral arms are very
long. Ventral view; note form of the fins, ventral
arms with their lateral margins (1), and the small
yolk sac projecting from the buccal funnel; 2 —
glandular lines.

FIGURE 320. Shell of a young S. elegans. 4x.
Compare with Figure 314 and the above data. Note
the absence of a spine, the deposit at the posterior
margin (5), wings (4), narrow fork (3), the narrow
lateral edges, the posterior boundaries of the lamellae
of the incrassation (siphonal stripes), the general
curvature. The form is slightly rhomboidal.

Figure 320 shows the shell of the young form. It resembles that of
S. orbignyana (Figure 314), but the shell can be easily distinguished: the
rostrum is replaced by a small rugosity; but the tuberculate deposit on the
posterior margin of the shield is distinct and curved posteriorly and
dorsally (Figure 320a). The shield has a narrow, free, posterior margin,
the incrassations diverge gradually anteriorly. The fork forms a narrow
ridge which projects only slightly posteriorly and is not markedly widened.
The later lamellae of the incrassation are more strongly curved than in

541

S. orbignyana, and the outline resembles a rhombus. The other
characters are those of the subgenus (p. 545). Both the shell and the
animal become increasingly slender with age. The outer side of the shield
develops a rough longitudinal comb which replaces the spine and there are
only gradual changes later. The free margins of the posterior part of the
shield and the wings are more widened in the female; the deposit at the
posterior end persists as a beaklike process.

542

564 Chapter 42

FAMILY SEPIOLIDAE

Keferstein, 1866

Contents: a. Diagnosis. - b. Typical structure of adult Sepiolidae. - c. Juvenile forms of Sepiolidae (p. 571). -
d . Variation of the type of Sepiolidae (p. 572). — 1. Characteristics of the subfamilies (p. 572). - 2. Subfamily
Rossiinae (p. 574). - 3. Subfamily Heteroteuthinae (p. 576). •- 4. Subfamily Sepiolinae (p. 578). - 5. Eusepio-
linae (p. 579). - 6. Sepiola-like Eusepiolinae (p. 584).

a. DIAGNOSIS

Shell completely rudimentary, persisting at most as a noncalcified,
gladiuslike rudiment in the form of a delicate prooslracum. Primary
eyelid closed into a small pore on the eye chamber, forming a transparent
"cornea'* bordered from below by a secondary lid fold that does not
enclose the pore, which is situated further anteriorly. Mantle sac short,
bag -shaped, with large rounded fins dorsally. Septum of mantle cavity
strengthened by the retractor pallii medianus. A prominent lateral funnel
adductor present on each side between funnel bond and head. Arms without
protective margins; suckers of arms spherical.

b. TYPICAL STRUCTURE OF ADULT SEPIOLIDAE

The ancestral form of the family is named Protosepiola (Figure 330)
and is easy to describe. Its general appearance resembles that of a young
Ros sia (Figure 338), particularly in the form of the mantle, fins and head
and in the formation of the arm apparatus.

The shell of Protosepiola also resembles that of Rossia — a narrow
plate of conchiolin comparable to the free rhachis of the shell of Teuthoidea.
It bears posteriorly a delicate, leaf-shaped, pointed flag. This structure
565 is not a gladius, despite the superficial resemblance, which is accidental and
developed from a distinctly different basis. As explained on p. 503, the
rudimentary shell of the Sepiolidae developed from the shell of Sepioidea
and has to be considered as the proostracum, which alone persisted
(see Figure 321; cf. Jatta, 1896, Plate 14, Figure 31 ),

During later development, the rudimentary shell assumes a similar
relationship to the mantle as the anterior part of the gladius of the Loligi-
nidae (Figure 71 on p. 170), it becomes completely surrounded by the
muscular mantle and is situated on its inner side, covered only with a thin
layer which, as far as it borders on the mantle cavity, has to be considered

543

as a remnant of the primary mantle (Figure 323 Pr). This layer adheres
to the neck as a neck bond. The muscular mantle forms then a closed sac,
as in many Octopoda, which the Sepiolidae closely resemble in habitus.

FIGURE 321. Median sections through a younger and an older embryo of Sepiolidae (Sepietta oweni-
a n a), diagrammatic, about 30 x. a) The shell sac has a large posterior part (Hs) which corresponds to the
phragmocone (the X indicates the "anlage" of the shell, drawn only for explanation) and the muscular
mantle (Mm) is attached to it. b) The shell sac has already become retracted from the posterior part of the
body but has not been followed by the muscular mantle, so that shell sac and muscular mantle are connected
only by a thin plate of tissue which is later replaced by tissue of the mantle. Note the posterior apex of the
body (Sp) and Hoyle's organ (Ho). The attachment of the gonad (Go) is of interest. It has the typical
embryonic attachment in a), but the genital ligament in b) which connects the gonad (Go) to the dorsal
wall of the coelom (€61,062) has become very thin; it is later torn (cf. p. 486). Inner organs: mouth opening
(Al, Mu), jaws (Ok, Uk), subradular organ (Sr), duct of salivary gland (Gd), pocket of radula (Rd), dorsal (Ob)
and ventral (Ub) buccal ganglia, cerebral ganglia (Cg), pedal ganglia (Pg), visceral ganglia (Vg), statocyst
(St), esophagus (Oe), stomach (Ma), caecum (Bl),hind intestine (Ed), ink duct and gland (Td),vena cava (Vc),
gastric ganglion (Gg), kidneys (Ni), pericardium (C63), heart (Hz), gonad (Go), yolk organ (Do), septum of
mantle cavity (Ms), anus (Af), funnel organ (Dr), "anlage" of funnel valve (Tk), funnel (Tr), closed by a
membrane (Hy).

566 The fins resemble those of the Sepiidae,* extending far anteriorly along
the mantle and occupying a large part of the sides of the mantle. It is
certainly secondary and characteristic for the family that they are distant

* There is also a physiological resemblance, as Verany (1851, p. 58) described for a live Sepiola:

"elle nage avec beaucoup de grace, a I’aide de ses nageoires, qu'elle emploie comme des palettes,
leur donnant a volonte le movement progressif ou retrograde; mais jamais elle ne se sert de ses
bras pour aller en avant comme fait la seche^. Quand elle nage tranquillement, les bras tentaculaires
sont entierement contractes, et la tete est en grande partie refoulee dans le sac,dont elle ferme
Touverture."

^ Sepia climbs or creeps only in captivity. This type of movement is not natural but a vain
attempt to treat the walls as if they were objects to be removed or enemies.

544

from the posterior end. Figure 323 shows the typical form of the fins,
relatively long and narrow, especially in the Sepiadariinae, more strongly
rounded in most other Sepiolidae. The fin forms typical "earlobes"
anteriorly. They are articulated with the muscular mantle. The fins of the
embryo are based on large lateral invaginations of the shell sac (cf. Be-
1 opt era, pp. 493,494) from which the articulation capsules later develop.

FIGURE 322. Young Rossia macro-
soma (cf. Figure 338), after removal of
the end of the funnel and of part of the
wall of the head to show the pockets (Tt)
and base of the tentacles. Note also the
habitus and form of mantle, arms and fins

Sh - membrane between the LV and V
arms; Bt - ventral buccal pillar.

(Natural size.)

FIGURE 323. Adult Rossia macrosoma (specimen
in Figure 339), dorsal, natural size. The mantle is
folded back to show the anterior part of the "gladius"
(Pr), the neck and its connection with the funnel pockets.
Note also the closed secondary eyelids (Ld) and the
orbital pores further anteriorly (Po). The nerves from
the stellate ganglia (St) pass on each side from the head
to the mantle, between mantle and margin of funnel
pocket,

Co “ cornea.

567 The mantle margin forms 3 low projections, one mediodorsal and two
ventral, near the shallow funnel indentation. The olfactory organ is a low
papilla as in Sepia, and its margins are more or less raised above the
epithelium (Figure 32 6).

Eye region. The primary lid fold is closed above the eyeball during
embryonic development so that only a very small anterior pore remains.
Only the part above and around the lens develops into a cornea, while the
rest, including the area of the pore remains opaque (Figure 327). The
secondary lid fold then rises from the ventral side of the margin of the
cornea (Figure 323) but, in contrast to the Sepiidae (p. 533), this fold does
not include the pore. This proves that the secondary lid has developed
independently in the two families, perhaps also the cornea. The eye is

545

situated in its large chamber, and the cornea is curved free above the eye
(Figure 326; Plate XIX, Figure 10). As in the Sepiidae, the pupil varies in
form (Figure 324) which developed from the typical relationships described

on p. 115. It is adapted for vision upward, in
accordance with the mode of life of the animal,
which buries in the sand, so that only the eyes
and a respiratory slit are exposed, i. e. the
dorsal mantle margin and the funnel opening,
which is directed asymmetrically upward.

The arms differ little in length; the 1st pair
is slightly shorter, the 3rd slightly longer than
the others. The swimming margins are re-
presented by thin ridges of skin which continue
proximally in the "glandular lines" and are
absent on the ventral arms. On the other hand,
there are well developed membranes between
the bases of the arms; those between the 3rd
and 4th arms are especially large, larger than
is typical for the Decapoda (Figure 322). The
inner side of the arms bears 2 rows of suckers
with markedly projecting stalks which pass
rapidly into a quadriserial pattern near the
middle of the arms. True protective margins
are absent, but rudiments of their supports
persist as small, pointed tubercles on the outer side of the stalks of the
marginal suckers.

The suckers on the arms are very characteristic, but resemble the
suckers of the Sepiidae. They closely resemble the modified suckers of the
mature male of Sepia (p. 538) and are nearly spherical, with a strong
circular muscle, a narrowed horny ring, an edge without teeth and a deep
stalk pocket. If the marked accentuation of these changes in the Sepiolidae
is considered (Figure 325), it may be assumed that the modification of the
568 suckers in the mature male gradually passed to the female and was thus
transmitted to all Sepiolidae. Details of the morphology of the suckers,
which are characteristic for many genera and species, will not be described.
Distinctive characters as in the Teuthoidea (dentition) are not present and
I could not study the minor characters.

The extended tentacle stalks are relatively long, with an indistinct,
rounded outer edge and two more distinct edges which delimit the inner
surface. The stalk is inserted in a tubular pocket in which the contracted,
thickened basal part can be contained without coiling. The rest of the stalk
is situated in a kind of pocket formed by the widened membrane between
the 3rd and 4th arms, the tentacle becoming curved (Plate VIII). This
condition is typical for the Sepioidea (p.488) and differs from the primary
conditions in the Decapoda only in the deeper invagination of the base of the
tentacle.

The club is also typical for the Sepioidea (p.488): it probably had
8—16 primary rows of suckers, but the ventral rows were pressed into each
other so that only 12—13 rows of an original pattern of 16 remained
(Figure 339). The protective margins are distinct and united at the base

FIGURE 324. Form of iris margin
(Ir) in the Sepiolidae in different
stages of contraction (cf. Plate 19,
Figures 8, 11-14)

546

of the club. The ventral margin is widened and connected with the stalks
of the adjacent suckers so that they are apparently situated on the margin.

On the other hand, the dorsal margin is narrow and becomes indistinct
distally; The largest suckers are situated in the proximal part of the
dorsal rows. They resemble the smaller suckers of the club of the
569 Sepiidae (Plate XIIl); their margin is uniformly covered with blunt, sparse
teeth, and the wide adhesive ring has a papillated inner and a radially
striated outer zone. In contrast to the very short stalks of the arm suckers,
the suckers of the tentacles have long, thread-like stalks and the suckers
are round and cuplike. A swimming margin passes along the whole club, •
and is widened only in the proximal part, where it forms a rounded lobe
(as in Figure 331 ); but it forms only an edge in the distal part. (This is
the case in many species of Rossia, in Stolot euthi s, and all juvenile
forms of Rossiinae, Sepiolinae and Hetercteuthinae with a swimming margin.)
(Figure 331).

(568)

FIGURE 325. Morphology of the arm suckers of Sepiolidae. a) Normal sucker of S e pi a officinalis,
b) Arm sucker of Sepiola rondeleti (Trieste specimen), c) Arm sucker of Rossi a macrosoma,
view of opening, d) Same sucker in median section. Note the structure of the horny ring (Hr), especially
the free edge (Zr); adhesive ring (Hf); circular muscle (Rm); suction pad (Sp); insertion of stalk in c)
and d). The pit (Gr) is situated before the insertion of the stalk.

Sk - suction chamber; Tr - stalk; Rp - marginal papillae.

The buccal funnel consists of 7 parts, as in the Sepiidae. However,
the ventral supports, points and attachments are situated close together in
the middle and their fusion is imminent. The 6 typical buccal pockets are
only indicated, the funnel is also low, and the points have no suckers. The
funnel is typical, except for the presence of the lateral funnel adductor.

547

The neck bond (p. 566) is elliptical, with slightly raised margin and 2 flat,
closely approximated ridges in the middle. The funnel bonds form oblong
cups with nearly parallel lateral margins, rounded anteriorly and posteriorly.
They occupy slightly more than the posterior half of the funnel pocket. A
large, superficial muscle, the lateral funnel adductor, connects the funnel
bonds with the ventral side of the head. This muscle is homologous with
the lateral funnel adductor of the Sepiidae (p. 537) but not with the external
funnel adductor of the Ommatostrephidae (p.414). Externally visible
570 adductors of the funnel are absent here also. The inner opening of the

funnel and the funnel pockets are unusually wide, corresponding to the form
of the mantle.

(569)

FIGURE 326. Young Sepietta obscura.male, with opened mantle cavity, contracted, 5x. The ventral
mantle plate is cut off and the posterior part of the ventral wall of the funnel is also removed. Note the
short mantle sac with rounded fins (FI); the large, rounded head; the arms and especially the suckers (Na)
and their stalks (Nt); the membrane connecting the arms; the tentacles (Tn) which are partly retracted into
the pockets (Gt); cornea (Co), lens (Li) and small pupil (Po),and the pore before it; the olfactory organ
behind the eye (Go); the funnel with the lateral adductors (Ta). In the mantle cavity: anal papilla (A)
and padlike funnel gland before it (Td); near it, the forked origin of the median mantle adductor (Ap), which
is only the anterior support of the mantle septum (Ms), which is cut off along its attachment to the
mantle (x); the strong funnel retractors, the stellate ganglia (St) next to them, and behind them the gills
and their suspensory ligaments (Kb) attached to the branchial gland (Km). Branchial hearts (Kh) and their
appendages (Pd) and spermatophore gland (on the left) (Sp) visible through the skin.

K1 — attachment of a single branchial lamella; Bt - buccal pillar; Ga — base of ventral arm; x - reduced
supports of protective margin, attached to the stalk of the sucker.

548

FIGURE 327. Half- grown embryo of S e p i e 1 1 a oweniana,19x.
The typical habitus of Sepiolidae is already recognizable but the
posterior end is still pointed, with a small spine which later pierces
the chorion of the egg (Figure 39, Vol.II). The arms are retarded,
the tentacle club has a typical, lobe- shaped swimming margin (6).

8 - orbital pore, definitively contracted; 7 - pupil, visible through
the cornea (24); 9 — olfactory organ; 25 - pocket of tentacle.

The mantle cavity shows very characteristic conditions, particularly the
presence of a median mantle adductor, the function of which is not under-
stood. This muscle probably developed from the musculus rectus abdo-
minis (p. 105) and from the anterior margin of the median mantle septum,
along which the muscle appears far too early for the establishment of such
an origin (Figure 32 1).

The lateral displacement of the stellate ganglia is striking and can only
be interpreted as an adaptation to the modification of the complex of shell
and mantle because the narrow, rudimentary shell would cause the ganglia
to be situated close to each other (cf. Figure 323, p. 566). The actual
condition is perhaps a reminiscence of an ancient wide shell (cf. p. 565).

The funnel retractors are strong, connected only by a longitudinal skin
fold with the body (Figure 339). The gills are typical but their ligament is
relatively short and wide. The branchial hearts are typically situated.

The vena cava is displaced to the right; the fin vessels (posterior mantle
artery and vein) originate fah anteriorly and pass laterally close behind
the branchial hearts (Figure 335). The renal papillae resemble those of
the Sepiidae, but they are slightly more widely separated from each other
and from the anus and are less projecting.

Of particular morphological interest are the accessory nidamental
glands. Their form and function take a new course. In addition to the
normal nidamental glands, which resemble those of the other Sepioidea,
there are accessory nidamental glands the body and opening of which are
situated median to the renal papillae (Figure 329a) and lengthened anteriorly
(p. 5742). This structure may be assumed for the ancestral form but there
are no reasons to assume further changes for the derivation of the Sepia-
dariinae. There is, however, the question whether this apparently typical
gland could produce a luminous secretion. The point is whether accessory
glands have such a function at all. The projecting part of the oviduct, i.e.,
571 the genital process, has a fold around the base, particularly on the outer
side. The spermatophores are attached to this "foreskin" or to the
wrinkled surface in its vicinity.

The jaws (Plate XVIII, Figures 2—3) and radula (Plate XV, Figures 6—7)
of the Sepiolidae closely resemble those of the Sepiidae. The jaws are

549

slightly more delicate, and at least the inner marginal zones are light and
transparent. The radula of Rossia still has traces of secondary teeth in
the 3 median rows; the teeth of the second and especially of the third row
have lateral process at the base. Traces of marginal platelets are present
on the margin of the radula.

Hectocotylization affects mainly the left ventral arm. As in the Sepiidae,
some of the suckers of this arm are smaller, and there is a modification of
tile inner surface which bears the suckers. There is also a marked
enlargement of the suckers in the middle of the lateral arms, which is less
marked in the marginal rows of the other arms.

There may be warts on the skin, as in the Sepiidae, especially on the
dorsal side of head and mantle.

c. YOUNG FORMS OF SEPIOLIDAE

The Sepiolidae have large eggs (to 7 mm) like all Sepiodea, which hatch
in a markedly advanced stage. But they are less advanced than in the
Sepiidae, in which all or most of the suckers begin to develop in the egg
and attain their definitive form only later. The ends of the arms and of
the tentacle club grow further during postembryonic development and the
youngest forms have only a part of the later developing suckers.

An important character is the development of the swimming margin of
the juvenile club of the tentacle. In the embryo (Figure 327) this margin
is a rounded lobe at the base of the club which persists for some time after
hatching (Figures 347, 349) and remains more or less unchanged in some
Rossiinae and Heteroteuthinae.

There are usually only a few chromatophores. The young Sepiolidae
otherwise resemble the adults also in their generic characters.

A remnant of Hoyle's organ is still present at hatching in the form of
thin glandular lines which extend from the posterior end mediodorsally and
laterally toward the fins (Figure 345). There is a small spine behind the
fork, at the posterior end of the mantle, which is thought to facilitate
hatching. It is probably homologous with the conical terminal elevation in
the young Spirula (p.516) and like Hoyle's organ, disappears soon after
hatching.

572 d. VARIATION OF THE TYPE OF SEPIOLIDAE

It is very difficult to determine the relationships within the Sepiolidae and
to give them a clear systematic formulation, especially because 5 of the
14 genera were not available, and the published data are inadequate and
unreliable. I had doubts for some time that these 14 types could be arranged
in a genealogical tree and attempted to arrange the relationships in the form
of a network of lines. However, after completing my observations and on
the basis of new data by Wulker (l910) and Berry (1910—1913) on the exotic
species, I constructed the diagram shown in Figure 328.

550

The most modified forms (Rondeletiola and Sepietta) probably
developed in a series of stages to which the other forms remained more
or less similar. A systematic arrangement must therefore be based on a
close sequence of stages as described on p. 18. I proposed (1912) a
division into 4 subfamilies which I maintain as graded stages in which each
type is connected with the preceding as a special case; in other words, a
part of phylogeny. However, also in a genealogical tree of the morpholo-
gical relationships, the systematic boundaries (Figure 328) are difficult
to define because the stages are not defined. I have followed the ecological
character of the Heteroteuthinae and have placed Sepiolina in the
Sepiolinae, not in the Heteroteuthinae. Sepiolina resembles the Hetero-
teuthinae in the typical primary formation of the luminous organs and in
the arrangement of chromatophores and iridocytes on the ventral side of
the mantle. On the other hand, its habitus and many other characters
resemble those of the Sepiolinae.

SYSTEMATIC REVIEW

Family Sepiolidae

Subfamily 1.

Subfamily 2.
Subfamily 3.

Subfamily 4.

Sepiadariinae. Genera: Sepiadarium, Sepio -
1 o i d e a.

Rossiinae. Genera: Rossi a, Semirossia.
Sepiolinae. Genera: Sepiolina, Euprymma, and
the especially S e p i o 1 a-like forms Sepiola, Inio-
teuthis, Rondeletiola, Sepietta.
Heteroteuthinae. Genera: Heteroteuthis, Necto
teuthis, I r i d o t eu t h i s, St ol ot eu th i s.

1. CHARACTERISTICS OF THE SUBFAMILIES

a) Left ventral arm hectocotylized. Fins more or less Sepia-like, i. e.
longer than wide. Arms with 4 rows of suckers, at least in the distal part.

573 Shell absent. Dorsal margin of mantle fused with the skin of the head.

Arms distinctly connected at the base. Subfamily 1. Sepiadariinae.

Species of this subfamily do not occur in the European seas. Hectocoty-
lization and the form of the fins resemble those of the Sepiidae, and also the
quadriserial arrangement of the suckers, which occurs also in other Sepio-
lidae. The formation of a neck band is an interesting convergence to the
Sepiolinae and Heteroteuthinae, as is the loss of the shell.

b) One or both dorsal arms or one laterodorsal arm (p. 599) hecto-
cotylized. Subfamilies 2—4.

bi) Benthic Sepiolidae without an atypical interbrachial membrane which
reaches far anteriorly. Arms not markedly different in length. Tentacle
club widened, with distinct protective margins and a half-developed to
completely developed swimming margin (p. 569). Orbital pore permanently
open. Mantle cavity without luminous glands. Dorsal mantle margin free.
Neck bond well developed. Subfamily 2. Rossiinae.

551

b2) As bi but with a "neck band," a concrescence between the head and
the dorsal mantle margin. Neck bond absent. Usually with luminous glands,
which are embedded in the ink sac. Subfamily 3. Sepiolinae.

574 ba) Pelagic Sepiolidae. Coloration rich, with metallic sheen. Inter -
brachial membranes well developed. Dorsal arms always much shorter
than lateroventral arms. Tentacle clubs only slightly widened, with a rudi-
mentary swimming margin only at the base. Orbital pore (always?) closed.
Dorsal margin of mantle free or fused with head. Luminous glands large,
deeply embedded in the ink sac near its middle. Arms with 2 rows of
suckers, some of them markedly enlarged on 3rd arms of male. Subfamily 4.
Heteroteuthinae.

FIGURE 328. Diagram of the systematic- morphological relationships of the genera
of Sepiolidae ("genealogical tree"*), on the basis of the typical relationships and
under consideration of the morphological data. Note the mainly linear arrangement
of the main types: Sepiadariinae-Rossiinae- Sepiolinae — Heteroteuthinae. (Read
Rondeletiola forRondeletia; Protorondeletiola for Protorondel etra;
Metarossia forMitarossia.)

* It must be admitted that we consider the phylogeny on the basis of this scheme. There can be no question
of a certain knowledge of the genealogy, and we maintain in principle that typical similarities must be
considered regardless of phylogenetical interpretations (p. 43).

552

2. SUBFAMILY ROSSIINAE Naef, 1912

Diagnosis. Benthic species. Male with one or both dorsal arms hecto-
cotylized; suckers in middle of other arms of male markedly enlarged.
Typical protective membrane restricted to basal parts of arms. Dorsal
mantle margin free; neck cartilage well developed. Tentacle clubs
distinctly widened and with typical swimming margin.

FIGURE 329. Organs of mantle cavity of mature females of the hypothetical types Protosepiola (a)
and Protorossia (b). Note particularly the differentiation of the accessory glands (4). a) Each
gland with a process (2) anteriorly between the renal papillae (3). b) This process is differentiated as
a separate gland (2) with different structure and function (luminous gland).

1 ■“ mantle adductor; 5 -- zone of openings of accessory glands; 6 - nidamental glands; 7 — genital
process. Approximately natural size.

The ancestral form of the Rossiinae resembles the type of the family so
closely that only a few minor details have to be mentioned. A rudimentary
shell is present, and so is the neck cartilage, and the mantle margin is
completely free. The fins are still markedly longer than wide. The arms
and tentacle clubs resemble those of Ro s s i a, but not the buccal funnel,
which has to be assumed to consist still of 7 parts.

Important changes are found only in the mantle cavity. The anterior
part of the accessory nidamental glands has become histologically
differentiated and separated from the other part of the gland, which has
retained the typical structure. It is apparently phosphorescent, i. e. it has
become a "luminous gland” (about emission of light and its significance
see pp. 583, 633). As such a gland is important for both sexes and a
575 "transfer" from male to female is possible, it is not surprising that

"anlagen" of accessory glands develop also in the male, but the functionally
unimportant normal part of the gland becomes more or less reduced. Such
a "transfer" of characters from one sex to the other has been observed also
in Loligo forbesi (p. 205). Most Rossiinae do not show such a
differentiation of the accessory glands so that this process probably began

553

in the Heteroteuthinae and Sepiolinae and has to be assumed for their
ancestral forms. However, Chun (1915) described a Rossia (Figure 330)
in which both sexes have rudimentary homologues of the luminous glands
of the two following subfamilies, and the male has in addition rudiments
of the normal accessory glands.

FIGURE 330. Organs of mantle cacity of Ro s s i a mastigophora Chun, a) Female.
The luminous gland (a) is rudimentary, the true accessory gland (2) normally developed,
b) Male. The true accessory gland is also rudimentary. Diagrammatic, after Chun
(1915, p. 63). The organ which 1 consider as a rudimentary luminous organ is, according
to Chun, a small gland the structure of which closely resembles that of the accessory
gland. This applies to both rudimentary glands of the male which are apparently of the
same character.

These small glands have not been found in any other species of Rossia.
Chun only describes them without attempting a morphological interpretation.
My interpretation seems the only one possible, regardless of the need to
find a connection with the conditions in the higher Sepiolidae (Heteroteu-
thinae and Sepiolinae). The posterior glands of the male R. mastigo-
phora are obviously homologous with the accessory glands. On the other
hand, they closely resemble the anterior glands (Figure 330b), which must
therefore have the same morphological character. Two new developments
should not be assumed without sufficient grounds. The anterior glands
must then be considered as derived from the accessory glands and this
permits the proposed solution. The organ of Rossia mastigophora
is therefore a rudimentary luminous gland and it is not likely that it is a
new development because of its small size and weak differentiation. It is
more reasonable to assume that species of Rossia with well developed
576 luminous organs (as in Figure 329b), like those of the Heteroteuthinae,
lived at greater depths. The littoral species lost these organs, as did
later the species of Sepietta (Chapter 47). R. mastigophora is
thus apparently a transitional form. However, the subfamily is only in-
completely known. A study of Semirossia would be interesting, because
of the numerous points of resemblance between this form and the Sepio-
linae. Only the left dorsal arm of Semirossia is hectocotylized and
similarly as in Euprymma, its closest relative among the Sepiolinae.

554

3. SUBFAMILY HETEROTEUTHINAE Naef, 1912

Diagnosis. Nektonic Sepiolidae. Coloration rich, with metallic sheen.
Some suckers in middle of LV arms of male markedly enlarged. Arm
suckers almost completely biserial. Interbrachial membrane occupying
a large part of the arms. Neck cartilage normally developed only in the
anterior part, or completely replaced by the fusion between head and mantle.
Swimming margin of tentacle club rudimentary or widened only at the base;
club atypically narrow, with very small suckers. Both sexes with large
luminous glands, embedded in the ink sac.

FIGURE 331. Stoloteuthis leucoptera Vemll,2x. Reconstructed after a slightly deformed
specimen from the collection of Prof. G. Pfeffer, Hamburg. General characters of Heteroteuthinae:
coloration bright red (orange to brownish red), with a metallic sheen; fins very large, of typical
form; interbrachial membrane well developed; 3rd arm of male with 3 enlarged suckers, one of
them particularly large; luminous organs as in Heteroteuthis; ventral shield; mantle margin
typical; swimming margin of club widened and lobe-shaped only at the base. Specific characters;
formation of neck band as in the Sepiolinae, suppression of neck cartilage; short ventral arms; rela-
tively strong developed dorsal arms (2X).

FIGURE 332. Iridoteuthis iris (Berry) Naef, 1912. Diagrammatic, after Berry (1914). Specific
characters; size of head; small mantle sac; wing-shaped fins; fusion of entire dorsal mantle margin
with the head (but the head remains clearly recognizable); 8 rows of suckers on the club, (According
to Berry, 1914, p. 316, Figure 27, there is a swimming margin along the whole club; Berry's drawing is
perhaps incorrect in this point.) Coloration: sienna brown with variegated reflexes. Luminous glands
weakly developed. This species resembles Heteroteuthis in the proportions of the arms and the
form of the ventral part of the mantle. Cf. Plate VIII, Figure 3.

555

This subfamily is transitional between the Rossiinae and Sepiolinae in
many characters. I place in it Heteroteuthis, Iridoteuthis, Nec-
577 toteuthis and St o 1 ot eu th i s. The uniform habitus of these genera
shows that the subfamily is a natural transitional group, although the
differences from the Sepiolinae are not very sharp. The Heteroteuthinae
undoubtedly developed from the Sepiolinae; Sepiolina nipponensis
resembles Stoloteuthis leucoptera so closely that Berry (191 1 )
placed them in the same genus, but this resemblance is only an expression
of the close relationship between the two subfamilies, and if they are to be
distinguished, Stoloteuthis must be placed in the Heteroteuthinae and
Sepiolina in the Sepiolinae.

FIGURE 333. Nectoteuthis pourtalesi Verrill.
2x. Diagrammatic, after Verrill (1883). Specific
characters: length of ventral mantle margin, which
covers the funnel in the preserved animal: the charac-
teristic modification of the suckers and their stalks in
the distal part of the arm (note the small drawing x).
The following characters resemble Heteroteuthis:
proportions of arms; structure of interbrachial mem-
brane; the habitus as a whole, which may not be ade-
quately represented in Verrills' drawings. The dorsal
mantle margin is distinctly recognizable, but a neck
bond is absent. It probably resembles that of Irido-
te uthis.

There is a transition from R o s s i a-like to S e p i o 1 a-like conditions in
the series Heteroteuthis — Iridoteuthis — Stoloteuthis but there
is no direct connection between these genera (Figure 328), The ancestral
form of the subfamily (p. 19) must have had all "primary" characters of the
group by definition, i. e. the characters of Rossia and also the characters
of accessory nidamental glands and their derivates described on p. 574. We
then obtain the following picture. The ancestral form "Protohetero-
teuthis" resembles Stoloteuthis (p. 331 ) in habitus, i, e. Protohe-
teroteuthis differs from Rossia in size and form, and from Sepiola
in its stronger sheen, the large fins and the bright red coloration. It differs
from Rossia and Sepiola in its nektonic mode of life. The fins also
show another specific character; their margin is not simply rounded but
forms a posterior corner from which it is straight to the posterior part of
the base (Plate VIII, Figures 3, 4). This is very marked in young Hetero-
teuthis and Iridoteuthis (Figure 331); in Stoloteuthis and
Nectoteuthis it is apparently at least indicated (Figures 331—333).

The ventral mantle margin projects markedly anteriorly and has a shallow
funnel indentation between its rounded projections. The mantle margin

556

resembles in principle that of the Sepiolinae (Figure 363). The eye (cornea)
is especially large and protruding. A large interbrachial membrane extends
578 to the middle of the arms. The arms, except the ends of the ventral arms,
bear 2 rows of suckers; the dorsal arms are always more weakly developed
than the others. The apical suckers of the ventral arms become suddenly
very small and form more than 2 rows. The tentacle clubs are very narrow
and little or not widened near the stalk. Their swimming margin forms
a rounded lobe at the base (Figure 332); the ventral protective margin is not
markedly widened. The suckers are very small and uniform and are
arranged in numerous rows. The buccal funnel consists of 7 parts. The
neck bond is normal only in its anterior part; its posterior part has become
fused with the mantle and no longer has the characteristic structure. The
funnel is otherwise typical. The mantle cavity shows the primary conditions
of the Rossiinae (Figure 329b) but the development of the accessory glands
and the luminous glands derived from them is of particular interest. Highly
differentiated and effective luminous glands are present in both sexes
(cf. Heteroteuthis). On the other hand, the male lacks (probably
constantly) rudiments of the true accessory glands of the female; Hecto-
cotylization affects particularly one or both dorsal arms of the male
(but cf. Heteroteuthis). A few suckers in the middle of the LV arms
are markedly enlarged as an adaptation for holding the female.

Heteroteuthinae are nektonic animals, i. e. permanent swimmers
according to my observations and by comparison with live Heteroteuthis
with the data of Verrill (l88l)on Stoloteuthis. They have a bright
metallic sheen and a luminous, mainly red coloration. There is a mat,
sharply delimited area of different coloration on the ventral side (Figures 331,
333, 341). Expansion of the chromatophores in this area conceals the effect
of the luminous glands ('Ventral shield").

Of the 4 genera of the subfamily, Stoloteuthis is closest to the type.
Although Stoloteuthis differs most markedly from Heteroteuthis,
the two genera have a number of such important characters in common that
a close relationship between them is evident (coloration, sheen, interbrachial
membrane, enlargement of arm suckers of male, luminous organs). Irido -
teuthis and Nectoteuthis are already much more closely related to
the Mediterranean Heteroteuthis. The 3 Pacific genera complete the
picture and provide the frame for H et e r ot e u t h i s, which will be
described in greater detail (p. 595).

4. SUBFAMILY SEPIOLINAE Naef, 1912

Diagnosis. Benthic Sepiolidae. One or both dorsal arms hectocotylized,
the other arms, at least the 2nd and 4th pairs, bearing markedly enlarged
suckers in the middle. Interbrachial membrane very weakly developed
between 3 dorsal pairs of arms. Neck bond replaced by fusion between
head and mantle; skin of head continuing directly into mantle margin
(neck band). Typical swimming margins present along whole tentacle club.
Arms with 3—4 rows of suckers; apex of ventral arms with to 8 rows.

557

579 This subfamily, as defined here, includes also Sepiolina (Naef, 1912,
Teuthol. Notizen, No. 2, p. 248; cf. Berry, 1911, Z. Anz., Vol. 37; Proc. Ac.
Nat. Sc. Philadelphia, 1912, "Stolot euthi s n i pp o n e n s i s"). Berry
(1914, Bull. Bur. Fish., Vol. 32, p. 318) accepted this classification. How-
ever, Sepiolina is a transitional stage to the Heteroteuthinae before the
other Sepiolinae, and the precise boundary between the two subfamilies is
a matter of a feeling of form (p.572). The strictly Sepiola-like habitus
of S e p i o 1 i n a, which is apparently a benthic animal was decisive for me.
The hectocotylization is without distinct characters and does not permit
a decision. The mantle cavity of the female, however, shows conditions
resembling those in the Eusepiolinae. There is a kind of bursa copulatrix
on the left side, perhaps represented by an area of wrinkled glandular
epithelium (l could only make a cursory examination of a specimen in the
Zoological Collection in Munich). A weakly developed homonomous
structure is present also on the right side. The luminous organs resemble
those of the Heteroteuthinae, i. e. they are fused in the middle into a trans-
verse-oval body, above which passes the branched median mantle adductor
(Plate VIII, Figures 1, 2). Small pits indicate the zone of opening of the
glandular tubes (Figures 329, 375). This is apparently a primary form and
it gives therefore no indication about relationships.

All genera of Sepiolinae except Sepiolina can be placed together in
the Eusepiolinae, if a single transitional stage is assumed; only the Euse-
piolinae will be described.

5. EUSEPIOLINAE nov.

Diagnosis. Left dorsal arm of male markedly modified; a few suckers
near the base are lost but their stalks are modified and enlarged. A "bursa
copulatrix" in the form of a folded skin pocket develops correspondingly in
the female before and median to the base of the left gill. Luminous glands
sometimes absent, but usually separated in the middle and attached as ear-
shaped appendages on lateral lobes of the ink sac.

The general form of the body is typical for all Sepiolidae (p. 5 66). The
arms of the female also have no specific characters. The female does not
show the external characteristics of the group, because the fusion of the
dorsal mantle margin with the skin of the head ("neck band") occurs also
in some Heteroteuthinae (particularly Stoloteuthis) and in Sepiolina.
There are 3 characters: l) hectocotylization phenomena in the male;

2) corresponding changes in the mantle cavity of the female; 3) the difference
of the luminous glands from the typical form in the Heteroteuthinae and
Sepiolina.

1 . Hectocotylization of the arms of the male consists of 3 phenomena:
a) the arms become stronger with sexual maturity, although the tentacles
are markedly retarded (p. 126); b) the middle of the 3 ventral pairs of arms
580 show the usual enlargement of suckers; c) there is a marked modification
of the left dorsal arm, particularly of some of the stalks of the suckers.
Several stalks on the outer side of the arm discard the suckers and are
lengthened to strong spine- or toothlike formations which apparently have

558

a special function during copulation when the arm is introduced into the
mantle cavity of the female for the transfer of the spermatophores. (The
modified stalks are probably introduced into the "bursa copulatrix" and
widen it to allow the attachment of the spermatophores.)

(581)

FIGURE 334. Older embryo (a) and freshly hatched specimen (b) of S epiola sp. (S. ligulata ?)
a: 12 X; b: 8x). Note particularly the luminous glands (6), embedded in the laterally produced ink
sac (12). a) The luminous glands are oval also in younger embryos, with an oblong papilla which
bears the pores (5). b) They already closely resemble the definitive form (ear- shaped). A muscular
cord connects the luminous glands. This organ is characteristic for older embryos of S epiola, and
differs from that of similar stages of Sepietta and Rondeletiola. Note also: mantle sac, fins,
funnel pockets, funnel, funnel gland (8), funnel bonds (9) and the still weak lateral funnel adductors.

In die mantle cavity; the forked median mantle adductor (1); next to it, the visceral nerves and anal
papilla. Eyes, olfactory tubercle (10), base of tentacle (translucent) , the arms with their characteristic
suckers and stalks, clubs, rudimentary interbrachial membranes, buccal funnel. Figure b) shows also
the enlargement of the funnel gland and the displacement of eyes and olfactory tubercles. In the
mantle cavity: funnel retractors (7), gills, gill ligaments (still very short) , branchial gland (3) (trans-
lucent),gill lamellae, branchial hearts with translucent appendage, renal papillae; but mainly the
"anlagen" of the nidamental glands (2) between the branchial hearts, and before them, behind the
renal papillae (11), the epithelial zone from which the accessory gland (4) later develops (this gland
is not yet distinct). The yolk sac in a) has become detached; it would be much larger than the head
in this view.

2. The wrinkled area near the female genital opening present in Rossia
(SemiroBsia? Heteroteuthinae? ) and Sepiolina for the attachment of
the spermatophores is more differentiated in the Eusepiolinae and forms the
"bursa copulatrix." The systematic and morphological character of the
"bursa copulatrix" has never been recognized, although Racovitza noted that
the spermatophores are attached to a differentiated area. Its typical
development and variation in the different species are described on p. 581.

After the characteristic phenomena of hectocotylization had been
clarified and used for the definition of new species and genera (cf. Naef,
1912, Notes Nos. 2, 3 and 7), it could be assumed that corresponding
specific changes are present in the female. In fact, each species shows a
close correlation between the specific form of the hectocotylus and the
"bursa copulatrix," but we can only indicate this relationship in general at
this point. The characteristic structure of the bursa is naturally not
caused by that of the hectocotylus; nor does the bursa induce unilaterally

559

a change of the hectocotylus. Both structures developed together from the
conditions observed in the Rossiinae and have to be understood in connection
with the act of copulation. Except in the primary Sepiadariinae, copulation
in the Sepiolidae differs from the typical behavior of the Decapoda. The
displacement of the hectocotylus from the ventral to the dorsal side is
connected with the fact that the female of Sepiolidae is fertilized in a supine
position or from below.

Racovitza (C. R. Acad, Paris, v. 118, 1894, p. 722) described the copulation
of the Sepio la-like Eusepiolinae, probably after observations on Sepiola
atlantica d'Orb. (S. rondeleti does not occur at Roscoff). He writes:
"En Juin 1892 j'ai pu observer a Roscoff I'accouplement dela Sepiola
rondeleti i. Le male, quoique plus petit que la femelle, la saisit brusque-
ment et la retourne la face ventrale en haut. II introduit sa premiere paire
de bras dans la cavite palleale (le bras gauche est hectocotylise — Steen-
strup), la second paire est etendue le long du corps, la troisieme entoure la
nuque et la quatrieme est passee entre les bras de la femelle. L'accouple-
ment dure huit minutes. Pendant tout ce temps, le male fait des efforts
continuels pour tenir la femelle eloignee de tout point d'appui solide. Cette
derniere, ne pouvant respirer pendant I'accouplement, fait des efforts
ddsesp^res pour se debarasser de son compagnon dfes qu'elle arrive a fixer
ses bras quelque part."

581 Levy (Zool. Anz., Vol. 39, 1912, p. 285) described a different copulatory
position which is certainly atypical if it was correctly observed. Different
types and positions of copulation have also been observed in other Cephalo-
poda, including Sepia officinalis and Loligo vulgaris. I assume
that Levy confused the 3rd arms, which are specialized for holding the
female, with other arms; they appear in his drawing as a nonsensical
deformation, like that in preserved specimens. Cf. p. 586.

The earliest "anlage" of the "bursa copulatrix" is a crescent-shaped
invagination of skin posterior and lateral to the female genital process,
near the base of the gills (Figure 372b) and quite separate from the "fore-
skin" of the genital process. This is not always distinct in the youngest
stages (Figure 376a), because the crescent-shaped fold may cover the small
genital papilla to some extent, so that it might be confused with the "fore-
skin." In fact, the "foreskin" is situated nearer the papilla and can hardly
be recognized under a magnifying lens in the young forms. The whole
structure is later displaced toward the middle and the "foreskin" becomes
visible (Figure 376b). The bursa develops into a folded pocket which
extends dorsally to the muscular mantle. Cords of cutaneous muscles
radiate anteriorly, posteriorly and toward the middle from the opening
(Figure 374a). These muscles widen the opening and also spread a large
part of the inner side of the bursa (Figure 3 74b), This is a preparation for
copulation. A bursa ready to accept the spermatophores can easily be
confused with the genital opening, because the folded anterior margin of the

582 bursa covers the true genital opening and usually also the opening of the
nidamental and accessory glands of the left side. The bursa develops into
a large proliferation of skin which occupies one side of the complex of the
mantle cavity and makes its morphological interpretation difficult. After
the spermatophores have been attached to the edematous folds of the bursa,
usually in its deepest part, the bursa becomes contracted and later

560

(Figure 374c) reverts to its appearance before copulation because the
spermatophores are no longer visible. If the female then begins to deposit
eggs, sperm from the spermatophores trickles from the opening and
fertilizes the eggs.

FIGURE 335. Young male (b) and female (a) of S e p i o 1 a affinis. 4x. Note especially the enormous
development of the ear-shaped luminous glands (3); their general form and character are shown in Plate XIX,
Figures 6 and 17. Also note their position to the renal papillae (4) and the ink sac, which is visible shining
through in the interval, but the glands are complete. Figure a) shows also the typical "anlage" (5) of the
accessory gland and the form of the nidamental glands which is specific for the Sepiolinae (Figures 337,372)

(8); the genital papilla (6) which projects behind the left gill. In Figure b) the transversely passing posterior
pallial artery (18) and the posterior pallial vein (17) are visible (translucent) behind the branchial hearts.

Note in comparison with Figure 334: the further development of the olfactory tubercles (16), lateral funnel
adductors (14), funnel bonds (15), gill ligaments and branchial glands (10), renal papillae (4), the strong funnel
retractors. Also: stellate ganglia (13), pericardial gland (9), and dorsal part of funnel gland (12).

One of the folds which radiate from the bursa (Figure 376b) is particularly
important. This fold forms a curve around the base of the left gill and
extends to the inner side of the mantle some distance anteriorly. Its free
margin bears a fascicle of muscles the contraction of which during copu-
lation partly closes a space for the gills (Figure 377) and thus provides
protection for the gills; I have named it the left gill septum (on a right gill
septum cf. Figure 374)

3. Both sexes show a characteristic change of the form and structure
of the luminous glands. They begin to develop already in the embryo on
lateral lobes of the ink sac, which develop at the same time (Figure 334a)
and are displaced laterally soon after their appearance, becoming first oval
583 and later ear-shaped. Their openings, situated originally ventrally and in
the middle, are displaced to the outer anterior margin and together form a
small papilla.

The luminous organs usually develop before the accessory glands, so
that their derivation from them cannot be 'observed directly (but cf.

Figure 372). They are widely separated from the "anlagen" of the accessory
glands, except in Rondeletiola (Chapter 46). By the time the accessory
glands become distinctly developed, the luminous glands have already
reached their full size and maximal development (Figure 335); later they
become again smaller (Figure 360). I conclude, therefore, that the luminous
glands are connected with the search for food, protection and communication,
but not with reproduction, as might be assumed.

561

FIGURE 336. Habitus of a typical representative of Sepiolinae, S. oweniana,
resting on solid ground. On a sandy bottom the animal would burrow into the
ground with a few strikes of the fins and movements of the arms, leaving only
the eyes and part of the upper mantle margin with the curved funnel opening
on the surface (Plate XIX). Such a position at rest has been observed in all
Sepiolidae, including Rossi a. The appearance of the swimming animal is
shown in Figure 369c, but the tentacles are retracted (p. 568).

There are also some more secondary characters. The accessory glands
are fused in the middle. They are not sharply delimited, as their tubules
extend in all directions and may occupy a larger area. The right gland is
always much larger than the left and extends farther posteriorly (Plate XIX,
Figures 16, 17). A typical orbital pore is always present, in contrast to the
Heteroteuthinae. The fins are rounded. A swimming margin extends along
the whole of the tentacle club. The funnel and its bonds are moderately
long. The ventral arms are shorter than the lateroventral arms.

The skin is smooth, as in the Heteroteuthinae, but less shining (the sheen
is stronger on the posterior part of the ventral side and toward the flanks).

A less shining, raised part of the ventral surface of the mantle corresponding
to the "ventral shield" of the Heteroteuthinae is well defined on which the
chromatophores are darker and more distinct. This is most marked in
Sepiolina and Berry considers it correctly as an indication of the
relationship with S t o 1 ot e u t h i s, although he stresses its importance too
much. This definition and prominence of the middle of the mantle is not
always distinct in preserved specimens of other species, particularly in
species which have lost the luminous organs (Sepietta). As in the
Heteroteuthinae, this differentiation should be considered as a regulation
of light. The coloration varies between bright orange red, reddish brown and
dark brown.

Euprymna has a special position among the Eusepiolinae because of
the quadriserial arrangement of the arm suckers and the presence of more
than 32 rows on the tentacle clubs, the hectocotylization of the left dorsal
584 arm as in Rossia and the enlargement of the suckers in the outer rows
of the 3 ventral pairs of arms of the male; the 3rd pair of arms is therefore
not differentiated as in the following species. Euprymna is also larger
than the Sepiola-like Eusepiolinae and has denser and more uniformly
distributed chromatophores. Its habitus resembles that of Sepiola.
Euprymna has therefore to be considered as a branch of the Eusepio-
linae which has remained at a lower stage of development because its
special characters resemble those of the Rossiinae, particularly the two
little known species of Semirossia. All other Eusepiolinae belong to
a higher stage of uniform type; they are referred to as "Sepiola-like
Eusepiolinae. "

562

6. SEPIOLA-LIKE EUSEPIOLINAE nov.

1. Diagnosis. Arms with 2 rows of suckers, except on the apex of the
ventral arms. Left dorsal arm of male markedly modified, with 1—4 normal
suckers at the base, followed by a "copulatory apparatus" of modified stalks
of suckers and with a zone of more or less normal suckers in the distal part.
Enlarged suckers are present in the middle of the right dorsal arm and
especially on the dorsolateral and ventral arms of the male. Lateroventral
arms of male very strong, always curved like a hook after death and with
only small suckers in the adult. Ventral arms of male with a blunt tubercle
of varying size at the base.

I place S e p i o 1 a (including Inioteuthis Verrill), Rondeletiola
and Sepietta in this group. Sepiola is undoubtedly the genus from
which the other forms have to be derived; Sepietta and Rondeletiola
have become changed together from the type. Sepiola therefore deter-
mines the characteristics of the group.

2. LITERATURE

The data in the following records cannot be determined specifically but apply to the S e pi o la-like forms
in general.

1554 Rondelet, Sepiola (lib. XVII, Cap. X,p.519).

1555 Gesner, Sepiola rondeleti (lib. IV, p. 855).

1558 Bossuet, Sepiola rondeleti (p. 204).

1644 Aldrovandi, Sepiola rondeletii (lib. V,p. 63).

1755 Johnston, Sepiola rondeletii (tom.I,lib. I,Cap. III,p. 8,Fig.).

1755 Ruysch, Sepiola rondeletii (Plate I, Fig. I).

1767 Linne, Sepia sepiola (tom. I, pars VI, p. 3151).

1772 Scopoli, Sepia sepiola (p. 128).

1789 Herbst, Sepia sepiola (p. 80).

1791 Walfen, Sepi a sepiola (p. 225).

1799 Lamarck, Loligo sepiola (p. 16).

1817 Cuvier, Sepia sepiola (p. 10).

1817 Leach, Sepiola rondeletii (Vol. III.p. 140).

1822 Lamarck, Loligo sepiola (Vol. XI, p. 368, Ed. 12).

1823 Blainville, Loligo sepiola (Vol. XXVII, p. 134).

1824 Carus, Loligo sepiola (p. 318, Plate 29).

1824 Martens, Sepia sepiola (Vol.II, p. 436).

1826 Payraudeau, Loligo sepiola (p. 173).

1826 Rlsso, Sepiola rondeletii (Vol. IV, p. 7).

1826 D'Orbigny, Sepiola rondeletii (p. 149).

1827 Brughiere, Loligo sepiola (Plate 82, p. 136).

1831 Delle Chiaje, Loligo s e pi o 1 a ("C a 1 a m a i o sepietta, " "seccetella," Vol. IV, p. 59, Plate 58,

Fig. 30).

1833 Grant, Sepiola vulgaris (p. 78).

1835 Ferussac, Sepiola grantiana (p. 66).

585 1837 Rang, Sepiola rondeletii (p. 88).

1837 Oken, Sepia sepiola (Vol. V, part I, p. 535).

1838 Gervais and van Beneden, S. rondeletii (p. 421).

S. vulgaris (=^. grantiana).

S. deswigiana nov.

563

1938 Potiez and Michaud, S. rondeletii (tom. I,p. 7).

1839 Ferussac and D'Orbigny, S. rondeletii (also S. vulgaris and S. deswigiana) (p. 330,
Sepioles: Plate I).

1841 Delle Chiaje, S. rondeletiana (tom. I, p. 10).

1841 Cantraine, S. rondeletii (p. 15).

1844 Philippi, S. rondeletii (p. 203).

1849 Gray, S. rondeletii, also S. vulgaris, grantiana and deswigiana (p. 92).

1851 Verany, S. rondeletii (p. 55, Plate 22).

1853 Forbes and Hanley, S. rondeletii (Vol. IV, p. 220).

1855 D'Orbigny, S. rondeletii (p, 160,249).

1858 Adams, S. rondeletii (Vol. I, p. 40).

1863 Aucapitaine, S. rondeletii and S. vulgaris (p. 368).

1869 Jeffreys, S. rondeletii (also female S. atlantica) also S. vulgaris (Vol. V, p. 136).
1869 Targioni-Tozzetti, S. rondeletii and S. major nov., S. v u Ig ar is (p. 44).

1879 Tryon, S. sepiola ( = S. rondeletii = S. major) (p. 155, Plate 65).

1880 Tiberi, S. rondeletii (S. vulgaris, S. major?) (p. 23).

1880 Stossich, S. rondeletii and S. vulgaris (p. 159).

1884 Giard, S. rondeletii (p. 311).

1885 Ninni, S. rondeletii and S. vulgaris (p. 160).

1886 Hoyle, S. rondeletii ( = S. oceanica d’Orb.) p. 16).

1887 Steenstrup, S. rondeletii and S. petersii nov. (p. 19).

1889 Posselt, Sepiola rondeletii and S. petersi (p. 140).

1889 Giard, Sepiola rondeletii and S. petersi (p. 173).

1890 Girard, Sepiola rondeleti and S. petersi (p. 247).

1890 Cams, Sepiola rondeletii and S. petersi (p. 452).

1893 Joubin, Sepiola rondeleti (p. 2).

1894 Joubin, Sepiola rondeleti and S. petersi (p. 2).

1894 Pelseneer, Sepiola rondeleti (p. 205).

1896 Jatta, Sepiola rondeleti (p. 124, Plates 4, 5, 7, 8, 14).

3. Typical Morphology of the Adult of the
Sepiola-like Species

The hectocotylus has a uniform character in this group, also partly in
the aberrant species S. aurantiaca and S. ligulata. The only
exception is’ the little known Inioteuthis japonica Verrill (= Sepiola
(iniot euthi s) inioteuthis (Verrill) Naef, 1912). The following
characters may be considered as typical: there are 3 (4) normal or partly
enlarged suckers on the base of the genital arm (Figure 344), one of which
belong to the median and two to the lateral row. The stalks of the 2nd and
3rd sucker of the median row and the 3rd and 4th stalk of the lateral row
form the whole or the greater part of the "copulatory apparatus" (Figure 378).
This consists of 4 tubercles separated by incisions which correspond to or
develop from stalks of the suckers. The copulatory apparatus develops
during postembryonic development; the proximal stalks in each row are
displaced closer to each other so that the 4 tubercles form a transverse
row in which the 4th stalk of the lateral row forms the sublateral main tooth.
The suckers following the copulatory apparatus are lost together with their
stalks, one or several in each row. The distal part of the arm is always
slightly thickened and the stalks of the two rows become more or less
separated, forming a kind of basket on the inner surface of the arm, and a
number of suckers become enlarged, especially in the median row. This

564

part of the arm becomes curved outward during preservation, so that the
hectocotylus is very prominent in preserved specimens. A slightly
strengthened rudimentary interbrachial membrane usually connects the bases
of the dorsal arms of the male (Figure 373c).

586 There are numerous differences in the structure of the hectocotylus
which seem unimportant at first glance. However, there is a definite
morphological correlation between the hectocotylus and the "bursa copu-
latrix." This connection is quite distinct in some species (S. ligulata).

I therefore assume that each structural detail has a distinct functional
significance and systematic value as a specific character (cf. p. 12).
Successful copulation between individuals of different species, however
closely related, probably becomes often impossible or difficult mechanically.

Typical phenomena of hectocolyzation are also present in other parts
of the arms of the male. 1. The tentacles are markedly retarded in compar-
ison with those of the female (p. 126). 2. The LV arms are strongly

muscular and always become curved toward the mouth during preservation
(Naef, 1912, Not, 3, p. 263) so that they are difficult to examine. The suckers
of these arms develop normally at first but are later retarded and are
finally markedly smaller than those of the other arms. 3. A typical
character is present on the ventral arms. There is a blunt tubercle at the
base, directed toward the mouth; this is an outgrowth of the muscular arm
axis and is not formed by the stalks of the suckers. These two adaptations
are probably connected with the holding of the female during copulation
(p. 580). The second permits recognition of the male, but only in dead
material. The marked thickening of the head, of the arms and of some
suckers are also helpful.

In live and well preserved females the two lateral pairs of arms are
always much longer than the two median pairs (formula: 3=2, 4 = 1 ).

All Sepiola-like Sepiolinae are small, less than 9 cm long without the
tentacles, usually less than 6 cm.* Their habitus is typical for the Sepio-
lidae, with relatively large, but few and sparse chromatophores (Figure 373).
The identification of the species is particularly difficult, as I have shown
(Teutholog. Notizen Nos. 3, 7, 1912), especially if the material is scant and
not well preserved (see also Naef, 1916). Live animals (Plate XIX) are easy
to distinguish. The important systematic characters were difficult to define
i. e. to determine the constancy of correlation between the different
characters (loc. cit.). The following characters can be used in practice:
l) form of absence of a "gladius"; 2) form of fins; 3) form of ventral
margin of mantle; 4) formation of suckers of arms and tentacles; 5) forma-
tion or (secondary) absence of luminous glands; 6) coloration, caused by

587 the distribution and color of the chromatophores, and the presence of iri-
docytes, and particularly 7) the specific structure of the hectocotylus and
of the suckers of other arms of the male; 8) structure of the "bursa
copulatrix" in the female. The width of the neck band may also be used,

it is always less than one -third of the width of the mantle in the Mediterranean
species.

* The size is fairly characteristic for a species in a certain locality and often permits rapid identification.
On the other hand, individuals of different origin are not uniform. S. ow eniana from Bergen and
S. obscura and S. rondeleti from Trieste are much larger than the same species in Naples. These
forms have to be considered as different varieties.

565

1 . The gladius may be a distinct, flattened rod which is visible as a ,dark
line when the mantle is detached from the neck (cf. Figure 323 on p. 566);
it may form only a fine thread, or be absent (inioteuthis, Rondele-
tiola). 2. The fins may have a slightly indicated lateral corner (Figure 373),
they may be broader posteriorly and have a round margin (Figure 365), or
they may be nearly circular (Figure 369). These differences are recog-
nizable only in well preserved specimens. 3. The ventral margin of the
mantle may form a wide, very shallow funnel indentation with slightly
projecting corners (Figure 373), or else the funnel indentation is narrower,
more sinuate and forms an undulate line between two rounded projections
(Figure 331 ). The two projections with the funnel indentation between them
may extend anteriorly like a shovel (Figure 358), as in the Heteroteuthinae
(Figure 341). All these characters require adequate preservation. 4. The
stalks of the suckers of the arms may be more or less markedly lengthened
(Figures 354 and 361 ), the suckers may be multiserial at the apex of the
ventral arms (Figure 354) and some suckers in the maturing male may be
enlarged. 5. The luminous glands may be of typical form but may vary in
size (Figures 355 and 364), and they may also be absent (Sepietta) or
atypical (Rondeletiola). The latter variations are not connected with the
degree of maturity or similar factors, as Jatta assumed, but the proportions
are correlated with these phenomena (p. 583). 6. The distribution of chroma-

tophores differs markedly in the different species, . e. g their density on the
dorsal side of the head (Figures 365 and 369) and on the ventral side of the
mantle (Figures 363 and 367). The density increases markedly during
postembryonic development (Figures 346 and 351; 361 and 367). The sheen,
especially on the ventral side of the mantle, is very weak in forms without
luminous organs (Sepietta), stronger in forms with slightly developed
luminous organs (Rondeletiola) and always very distinct in forms with
large, typical luminous organs. This is always distinct in live animals
but not in preserved material. 7. The modified stalks which form the
copulatory apparatus are of specific and constant form in each species.

This also applies to the enlargement of some suckers (Figures 344 and 378).
8. The form of the bursa in certain conditions, e. g. in a fully extended state,
is also characteristic and constant for each species, as are changes in the
mantle cavity of the female which are connected with the form of the bursa.
The analysis of these characters was very difficult in the varied and in
part badly preserved material from Naples, because these structures are
physiologically variable (Figure 374) and their form had to be correlated
with factors like maturity and copulation, etc.

4. Juvenile Forms of the Sepiola-like Eusepiolinae

588

As in all Sepiolidae, the eggs of the Sepiola-like species are very large
and the animals hatch at an advanced stage of development. The length of
the freshly hatched animal may be about one-quarter of that of the female.

A phylogenetic reminiscence is the characteristic juvenile form of the
swimming margin (p. 569, Figure 327). In contrast to the Sepiidae, the num-
ber of suckers on arms and clubs is always much smaller than in the adult;
this applies also to the number of chromatophores (Figure 369; cf. young
Sepia, Figure 307). The neck band is already well developed; I never found
rudiments of the neck bond. Hectocotylization is absent in the young male;
the two sexes are identical externally.

566

589 Chapter 43

590

GENUS ROSSIA

Owen, 1834

a. DIAGNOSIS

Arms with 2—4 rows of suckers which gradually decrease in size toward
the apex. Both dorsal arms hectocotylized. Buccal funnel consisting of
6 parts because of the fusion of the ventral supports and points.

b. TYPICAL STRUCTURE OF ROSSIA

The only distinct character of the genus is that the buccal funnel consists
of 6 parts. All known species of Rossia have such a buccal funnel. This
character is distinctly atypical; all other characteristics of the subfamily
apply to Rossia. This character, however, defines Rossia as a more
specialized branch of the general stem (Figure 228). The relationships
caused by the differentiation of the accessory glands and the luminous
organs (p. 575) are not considered at this stage because of their hypothetical
character. Since luminous glands homologous with those of the Hetero-
teuthinae and Sepiolinae are absent (except in R. mastigophora) and
there are no accessory glands in the male (with the same exception), these
structures have apparently become reduced or have been secondarily lost,
in the sense that a differentiation of luminous glands from the accessory
glands and a transfer to the male does not take place (p. 575). The develop-
ment of the female genitalia is therefore of particular interest also in the
species of Rossia without rudimentary luminous glands. It has there-
fore to be assumed that some resemblances to previous morphological
conditions, particularly of the accessory glands will be found.

Some species of Rossia are incompletely known. I intend to publish
some contributions to the systematics of the genus. The species can be
identified by the following characters:

1. Biserial or quadriserial arrangement of suckers on the arms.

2. Presence or absence of constant papillae on the dorsal surface.

3. Number of suckers and rows of suckers on the club, and their size on the

club. 4. Form of the swimming margin of the club, with either a broad lobe
at the base, or a uniform margin along the whole club which tapers toward
the apex. 5. Presence of absence of rudimentary luminous glands and
accessory glands in the male (Figure 330). 6. General proportions of

mantle sac, fins, arms and tentacles. 7. Coloration and distribution of
chromatophor es .

The freshly hatched animal resembles the adult, except for the sexual
characters. The small terminal spine (p. 571) disappears rapidly. Only
^ "^acrosoma will be described.

567

FIGURE 337. Development of the female genitalia, b) Situs of an older embryo of Rossi a glaucopis
from Bergen. Compare with Figures 326 and 376. Nidamental glands (2) normal, but accessory glands (1)
extending far anteriorly between the renal papillae (4) (cf. Figure 334). a) Young R. glaucopis. The
accessory glands still project far beyond the renal papillae, but the differentiation of the anterior part is much
less marHed than in the earlier stage; the glands are undivided (higher magnification), c) Renal papillae,
accessory glands and nidamental glands of a young Rossia macrosoma. Accessory glands situated close
together and slightly folded. They do not extend so far anteriorly; they are attached anteriorly by a cutaneous
muscle w'hich can perhaps move them. lx. d) Situs of mantle cavity of a half-grown Rossia macrosoma.
The accessory glands are spread and situated completely behind the renal openings. Note also the genital
process (6), its attachment (5) to the body, and the "foreskin" at the base (7).

3 - mantle adductor; 8 - base of mantle adductor; 9 - anal papilla; 10 - pericardial gland; 11 - branchial
heart (translucent).

c. ROSSIA MACROSOMA (Delle Chiaje, 1829) Owen, 1834
1. DIAGNOSIS

Arms with 2 rows of suckers at the base and 4 rows in the middle and on
distal part. Papillae on dorsal side of mantle and head absent. Tentacle
club with more than 8 rows of suckers which are much smaller than on the
arms and show no marked differences of size. Swimming margin extending
along whole club and gently tapering only at the apex. Both sexes without
rudimentary luminous glands. Accessory glands absent in male.

ggj 2. LITERATURE

1829 Delle Chiaje, Sepiola macrosoma (Plates 71,72; cf. Philipiil)

1834 Owen, Rossia (p. 93).

1835 Ferussac and D’Orb., Rossia macrosoma (p.245, Sepioles Plate 4).
1839 Gervais and van Beneden, Sepiola macrosoma.

1841 Delle Chiaje, Sepiola macrosoma (V.I, p. 10).

1842 Ball, Rossia oweni and R. jacobii (male and female) (p. 193 and 349).

1844 Philippi, Sepiola macrosoma (p. 203; also, Delle Chiaje, Plate 72).

1845 D'Orbigny, Rossia macrosoma (p. 257, Plate 11).

1851 Verany, Rossia macrosoma (p. 60, Plate 23).

1853 Forbes and Hanley, Rossia macrosoma (Vol. 4, p. 221).

568

1853 Forbes and Hanley, Rossia owenii (also R. j a cobii).

1855 D'Orbigny (as 1845).

1856 Steenstrup, Rossia macrosoma (probably including R. owenii and R. j acobii) (p. 199, German:
p. 230).

1858 Adams, H. and A., Rossia macrosoma (Vol. 1, p. 39, Plate 4).

1869 Targioni-Tozzetti, Rossia macrosoma (1. p. 39, 2. p. 46).

1869 Targioni-Tozzetti, Rossia panceri (male of R. m a cr osom a).

1869 Jeffreys, Rossia panceri (Vol. 5, p. 133).

1879 Tryon, Rossia panceri (p. 159)

(also R. owenii (p.l59)).

1879 Tryon, Rossia panceri (p. 159).

1880 Tiberi, Rossia macrosoma (p. 23).

1880 Tiberi, Rossia panceri (?) (p. 23).

1886 Hoyle, Rossia macrosoma (p. 18) synon. R. panceri).

1886 Hoyle, Rossia owenii (?).

1889 Posselt, Rossia owenii synon. R. j acobii.

1889 Posselt, Rossia macrosoma (p. 141).

1890 Norman, Rossia macrosoma (also R. owenii as variety), (p. 469).

1890 Carus, Rossia macrosoma (also synon. R. panceri).

1894 Joubin, Rossia macrosoma (p. 3).

1896 Jatta, Rossia macrosoma (p. 134, Plates 2, 5, 15).

1896 Jatta, Rossia palpebrosa (p. 139, Plate 15).

1908 Pfeffer, Rossia macrosoma (p. 40, Figs. 38-42).

1916 Naef, Rossia macrosoma (p. 16).

1921 Naef, Rossia macrosoma (p. 538).

1921 Grimpe, Rossia macrosoma (p. 299).

3. TYPICAL STRUCTURE OF THE SPECIES

Only the club of the tentacle needs further description; the other
characters were described above. The club shows a slight modification
of the typical conditions in the Sepiolidae. The number of suckers is very
large (about 1,000), and their arrangement is difficult to describe, but it is
essentially as in Sepia. The dorsal marginal row contains only about
half the number of suckers in the ventral rows (about 60 : 130); they are
much larger dorsally at least in the proximal part, 3 times as wide as the
ventral marginal suckers. The suckers of the next 2—3 longitudinal rows
are smaller. The oblique, curved transverse rows contain more than
12 suckers in the middle and fewer toward base and apex. The longitudinal
rows are pressed into each other and the transverse rows are divided near
the ventral margin. The primary number of suckers is therefore apparently
much greater, possibly 32 (analogous to Sepia). The protective margins
are typical and connected at the base.

The sexual dimorphism is interesting; it affects the arms of the male
and the mantle cavity of the female. The suckers of the marginal rows of
the 2nd, 3rd, and 4th arm pairs in the mature male (Figure 339) are
markedly enlarged, those of the inner rows smaller than in the female. On
the other hand, the suckers of the middle and marginal rows of the female
are of uniform size. The suckers at base and apex of the arm of the male
592 are normal, as in the female; the suckers are largest in the middle of the
arm. A special hectocotylization is present on the dorsal arms, which place

569

the spermatophores into the mantle cavity of the female (p. 580). The
suckers of these arms are largest at the base, where they form only 2 rows;
they decrease gradually in size toward the apex. The marginal and median
rows are uniform, as in the female. The most important modifications affect
the arm itself: the 2 inner rows in the middle of the inner surface diverge
slightly, and 2 alternating rows of deep incisions (Po) separate the stalks, so
that a direct succession of the stalks of the two rows on each side is recog-
nizable and proves that the zigzag rows developed from simple rows on each
side (p. 118).

FIGURE 338. Young Rossia macrosoma,
natural size. The habitus of the family is typical:
form of fins, mantle, mantle margin, head, eyelids,
orbital pores (on small papillae), arms (showing
form and arrangement of suckers, interbrachial
membranes, but no protective margins), glandular
lines and base of tentacles. Characteristic for the
genus: the single ventral support of the buccal
funnel. Specific characters: club of tentacle
(swimming margin, large number of small suckers).

Near the base of the arm, where the biserial pattern becomes quadri-
serial, begins another and possibly more important change on the outer
side: a ridge of skin (Ls) extends from the marginal suckers, beyond the
middle of the arm, and then disappears; it is bordered on the inner side
by a longitudinal groove (Kr). Between this groove and the marginal
suckers the skin is folded and the stalks of the suckers are separated by
depressions.

The free distal part of the genital duct is typically situated and the
conditions in the mantle cavity are characteristic for the family, but
luminous glands are absent (p. 575).

The same applies to the mature female (Figure 340). However, the
vicinity of the opening of the oviduct shows an interesting differentiation
which will be discussed below. The skin of this area forms folded,
wrinkled proliferations between which the spermatophores are attached.

593 A stripe or crest of warts develops dorsal to the genital process, and from
there toward the base of the gills and anus. This is present only in mature
specimens and is not fully developed in the half-grown specimen in

Figure 340. The typical opening of the oviduct into the mantle cavity forms

594 pointed papilla which grows during postembryonic development from the
pocket of the gill base gradually toward the anus and to near the renal
papilla (Figure 340b). The opening forms a slit and is directed outward;

570

(593)

FIGURE 339. Mature male of Rossia macrosoma with opened mantle cavity and spread arms. Natural
size. The smaller drawing shows the inner surface of the left dorsal arm with the characteristic incisions (Po).
Note the arrangement and proportions of the suckers; modification of lateral surface of dorsal arms; buccal
funnel with medioventral support (Bt); tentacle stalk and club with suckers, protective (Ss.Sd) and swimming
margins (Sc); head and funnel, and their connection; situs of mantle cavity, particularly the forked median
mantle adductor (Ad); mantle bond (Mh), funnel retractors (with a septum which connects them with the body
and attaches them to the mantle); genital process (Pn) ("penis"). The posterior part of the genital system is
visible behind the left gill; renal papillae (Np), gills and branchial hearts (Kh). Behind the branchial hearts
pass the markedly displaced (cf. Figure 335) posterior mantle veins and arteries (Ap) which extend toward the
fins. These vessels are situated close to the lateral mantle veins (Vp) near the gills.

Ts - testis; Nt -■ pocket of spermatophores; Sp ~ spermatophore gland; Ka - branchial artery; Kv - branchial
vein; Ta -- lateral funnel adductor; Go ~ olfactory organ; Ls,Kr ~ see text on p. 592; Jr,Jd - ventral and
dorsal inner edges of tentacle stalk; Ax — muscular axis of club.

FIGURE 340. Mantle situs of a maturing female of Rossia macrosoma, natural size (cf. Figure 337):

1 - wrinkled area near female genital opening (homologous to bursa, p. 581); 2 -"foreskin" at base of
genital process; 3 - adductor of accessory nidamental glands (5); 4 - renal papilla; Mu - branchial
retractors; Vp — posterior mantle vein; VI — lateral mantle vein; x — median mantle adductor.

571

the process is attached on the inside by a skin fold which ends near the
anus. On the outside develops a kind of "foreskin," into which the process
can partly retract. A shallow pocket is formed between the "foreskin"
and the papilla.

The coloration varies from light yellowish brown with greenish reflexes
to dark reddish brown. The surface is always smooth. As in Sepia,
there are a large number of yellow and brown chromatophores, in contrast
to Sepiola, which has a similar habitus. The small number and loose
distribution of chromatophores are therefore not characteristic for the
family.

572

595 Chapter 44

GENUS HETEROTEUTHIS

Gray, 1849

a. DIAGNOSIS

Fins moderately large, inserted behind middle of mantle sac. Dorsal
margin of mantle free; anterior end of neck cartilage preserved. Ventral
mantle margin projecting like a shovel and covering greater part of ventral
side of head. Swimming margin of tentacle stalk forming a small lobe at the
base.

In addition to the Mediterranean species, this genus contains H. w e b e r i
Joubin from the Dutch East Indies and H. h a w a i i e n s i s Berry.* These
two species closely resemble H. di spar, but they are only incompletely
known; their special characters will be mentioned, but the Mediterranean
species will be used for the characterization of the genus.

b. HETEROTEUTHIS DISPAR (Ruppell, 1845) Gray, 1849
1. DIAGNOSIS

Both dorsal pairs of arms of about equal length and markedly more
delicate and shorter than the equally long ventral arms. Tentacles very
long; if extended, reaching more than the length of the body beyond the
longest” arms. Tentacles retracted in a spiral into a pocket formed by the
interbrachial membrane between 3rd and 4th arms.

596 2. LITERATURE

1845 Ruppell, Sepiola dispar (fide Veraoy, 1851, p, 64).

1849 Gray, Rossi a (Heteroteuthis) dispar (p. 90).

1851 Verany, Rossia dispar (p. 63, Plate 23).

1856 Steenstrup, Rossi a (Heteroteuthis) dispar (p. 230).

* "S tepha noteuthis hawaiiensis" Berry has a shorter funnel and shorter arms and tentacles, to judge
from preserved specimens. Its habitus resembles that of H. di spar, but the arms are less different (Berry,
1910, Figures 29,30) and the contrast between the two dorsal and ventral pairs is not marked. The 3rd arms
of the male are much longer, and the closer connection between the 1st and 2nd arms is on the left. The
other differences given by Berry have no systematic value (folds on the ventral side of the head are also
present in H. dispar; the fiinnel is covered only because of contraction during preservation, and a "ventral
shield" is present also in H. dispar).

573

1857 Troschel, Heteroteuthis dispar (p. 62).

1858 Claus, Rossia dispar (p. 259).

1858 Adams, H. and A., Rossi a (Heteroteuthis) dispa r (p. 40).

1879 Tryon, Rossia (Heteroteuthis)dispar (p. 162).

1880 Tiberi, Rossi a (Heteroteuthis) dispar (p. 23).

1886 Hoyle, Heteroteuthis dispar (p. 19).

1890 Carus, Heteroteuthis dispar (p. 453).

1896 Jatta, Heteroteuthis dispar (p. 142, Plates 7, 15).

1903 Lo Bianco, Cirroteuthis meangensis (p. 173).

1909 Lo Bianco, Cirroteuthis meangensis (p. 645).

1916 Naef, Heteroteuthis dispar (p. 16).

1921 Naef, Heteroteuthis dispar (p. 538).

3. STRUCTURE OF THE ADULT ANIMAL

The half -grown and adult Heteroteuthis differs markedly from all
other Mediterranean Sepiolidae in the form of fins, mantle sac, head and
arms, and particularly in its strong metallic sheen and bright, mainly
orange red to brownish red coloration ( Figure 341; Plate XIX, Figure 10;
Plate VIII; Plate XIX, Figure 20 ). Only the dorsal side of the eye is a dark,
sepia- brown; and a few darker chromatophores are present on the dorsal
side of the head. Fins and sides of mantle and arms are almost without
chromatophores and with a bright silvery sheen. A strong metallic sheen
is present also on other parts of head and mantle, and on the outer sides
of the arms; iris and outer surface of eyeball bluish. Of particular
interest is the ventral shield (p. 583), i. e. the anterior median part of the
ventral side of the mantle which is densely covered with chromatophores.
Each main chromatophore is surrounded by a ring of smaller chromato-
phores. The ventral shield regulates the luminous effect. The shield is
surrounded by a particularly shining zone which reflects incident light. The
layer of iridocytes in this zone covers the chromatophores. The live
animal shows a beautiful play of colors which the figure renders only in-
completely. Despite its plump form, the animal is of very handsome
appearance, especially because of its constant and vivid movements. The
skin is quite smooth, as in all Heteroteuthinae.

A "gladius" is apparently absent. In contrast to all other genera, the
fins are situated close to the posterior end and can change their position so
that their posterior base becomes almost contiguous; however, they never
occupy the normal position in the middle of the mantle. This position is
not primary, because the earlier subfamilies Rossiinae and Sepiadariinae
have typically situated fins. The corner near the posterior end of the base
of the fins (p. 577) is less distinct and appears more like a posterior
"earlobe." The fins are large, but moderately developed in comparison
with those of Iridoteuthis and Stoloteuthis (p. 576). The mantle
597 sac is short, parabolically rounded posteriorly, widened anteriorly. Its
margin projects dorsally in a shallow curve; it forms ventrally a shovel-
like process which covers the greater part of head and funnel.^' The funnel
indentation is shallow.

We consider this as an adaptation to nektonic life (p. 576). A similar variation between a relatively
straight margin of the mantle (shell) and a margin produced like a boat is present in the Nautilidae (p. 55)
(cf. Naef, Vierteljahrschr., Zurich). There are more benthic (planorboid) types and more nektonic
(nautiloid) types also in the Nautiloidea.

574

(598)

FIGURE 341. Almost mature male of He teroteuthis dispar, 2x. Note the distribution of
chromatophores on both sides and form of fins and mantle. The eyes with lens and iris are visible
through the cornea. Note the proportions of arms and suckers. The interbrachial membrane is
wrinkled on the ventral side, where the tentacles are placed (Plate VIII, Figure 2). The tentacle
club has a rudimentary swimming margin.

The head shows the characteristic form and differentiation of the family,
except that the orbital pores are completely closed (l could not find them in
the young stages either, Plate VIII). The cornea is also dorsally displaced
and the pupil circular (Plate XIX).

The arms differ markedly in length. The two dorsal pairs are of nearly
equal length and distinctly shorter than the ventral arms, which are of
almost the same length (formula: 3,4; 2, l). The relative size of the
suckers corresponds to the length of the arms. The interbrachial mem-
branes reach to half the length of the dorsal arms and to a third on the
ventral arms. The suckers are biserial and become rapidly very small
at the apex. The suckers on the apex of the ventral arms show a special
arrangement and form (Figure 342): they become much smaller than on
the other arms and form a more or less distinct pattern of 6 rows, each of
which is divided into oblique groups of three.

The tentacles are very long, like a lasso. If extended, they reach more
than the length of the body beyond the end of the arms. They are nearly
round in cross section, and taper gradually from basis to apex. The club
is slightly widened at the end, with more than 8 rows of uniform, very small
suckers and only slightly indicated protective margins. On the rudiment
of the swimming margin a special organ, perhaps a gland, apparently
developed: it forms a trough toward the inner side of the club (Plate VIII,
Figure l) and can therefore not have the function of a normal swimming
margin. It may be an "adhesion apparatus" to connect the two clubs
during forward swimming, as in the Oegopsida, or a luminous gland. The
retracted tentacle is coiled into spiral, more than in other Sepiolidae
(Plate VIII, Figure 2), and is then situated in a pocket formed by the
membrane between the 3rd and 4th pairs of arms so that only a small part
is situated in the normal pocket of the tentacles.

575

The buccal funnel consists of 7 parts. The neck bond is typical for the
subfamily, while it is usually more modified. Its posterior part is already
reduced and replaced by a fusion, but the anterior part (Figure 323 on p. 5 66)
has remained normal. The funnel is particularly long, and so is the funnel
bond, which is markedly widened in the anterior part but very narrow on the
whole.

598 The conditions in the mantle cavity are more or less typical for the
subfamily, but the ink sac (Plate VIII) is very large, so that the large
luminous organs do not cover it completely, and its posterior part remains
free. If the tears in my material are not artifacts, the pockets of the gill
base are perforated posteriorly, as in many Sepiolinae (Figure 380). The
luminous glands form together an oval body; the secretion of each gland'!'
is released through pores situated on a papilla directed anteriorly.

The female genitalia show some special characters. The nidamental
glands are situated far anteriorly (Plate VIII, Figure 2), as in Ros sia
mastigophora (Figure 330) and are relatively weakly developed (eggs
are unknown). The accessory glands are also weakly developed. There
is an area of glandular skin on each side between them and the luminous
glands, partly situated on the ink sac. I consider this as a separated part
of the general "anlage" of the accessory gland from which the luminous
glands also develop. The condition of my material did not permit a more
detailed examination. The projecting part of the oviduct has a characteristic
form and opens ventrally. The typical fold of the "foreskin" is situated at
its base and the oviduct gland is visible through the skin behind it.

(599)

FIGUI^ 342. Arms of a mature male ofHeteroteuthis dispar from Messina, cut ventrally.

The two dorsal arms of the right side are modified and connected. There are enlarged suckers on
the 3rd and 4th arms. The apex of the ventral arms is modified. Note also the insertion of the
buccal funnel. The tentacle stalks (5) are cut near the base; note their cross section (Jatta (1896,

Plate 15, Figure 36, cf. also Figure 29) gives a rather inaccurate drawing).

3 — connecting muscle; 6 — lost sucker. The female of this nektonic form is apparently not
turned on the back during copulation (p. 580), and the hectocotylus is introduced more laterally.

The hectocotylization is characteristic (Figure 342). It causes marked
changes of the entire arm apparatus. Only the two dorsal arms on the left
side are normal; the arms of the right side, the actual copulatory arms,

* The compressed tubules release a pulpy secretion in form of sausages which spread like fireworks from the
funnel and have a striking effect. The luminous organ at rest emits a soft light, as in the Sepiolinae.

A strong stimulus produces the luminous spray.

576

599 are connected by a strong muscular membrane to more than half their
length. Almost the proximal half of these arms is without suckers. The
distal part of the right dorsal arm bears normal suckers, except that the
first suckers are situated on the wrinkled, swollen skin which covers the
whole inner surface (4). On the other hand, the suckers of the 2nd arm are
smaller, and roughly the proximal half are situated in a groove (2) which
forms the inner surface of the arm and is directed toward the 2nd arm.
Wrinkles of swollen skin (l) form the margins of this groove. The LV
arms bear 9 normal suckers at the hasp, followed by 2 enormously enlarged
suckers; about 3 large suckers of decreasing size are situated more
distally, and beyond them 7 almost normal suckers which decrease rapidly
in size. Only the middle of the ventral arms bears moderately enlarged
suckers.

4. POSTEMBRYONIC DEVELOPMENT

The embryonic stages of H. dispar are unknown. The eggs probably
resemble those of other Sepiolidae (Plate XIX) but have thinner membranes.

The youngest known stage is shown in Figures 5 and 6 of Plate VIII.

This is a planktonic animal and its habitus differs markedly from that of
other Sepiolidae. The transverse widening of the fins is characteristic;
the base of the fins forms a kind of stalk of the widened other part of the
fins. This is only slightly marked in other Sepiolidae (Figure 346). The
fins have a distinct, more transparent and delicate marginal zone. The
mantle margin is still simple, almost transverse, the olfactory organ nearly
embryonic in structure and position (p. 570). The eyes are still relatively
small, with a sharply defined cornea but the lid fold is only indicated. The
arms are very short, but they can probably be markedly extended in life;
a strongly developed interbrachial membrane connects the whole arms
except at the apex. The proportions of the arms are already definitive
(3, 4; 2, l). Each arm bears only a few suckers in 2 rows. As in the

600 adult, the tentacles can be coiled (Plate VIII, Figure 2), but they are shorter,
and the club bears fewer suckers. Buccal funnel, funnel and the organs in
the mantle cavity resemble those of the adult, except for the genitalia.

The luminous glands are well developed. The sheen is weaker and the
coloration less bright than in the following stages; the living specimen was
quite transparent, and had only a few regularly arranged, orange -red chro-
matophores.

**C i r r o t e u thi s meangensis" Lo Bianco ("Puritan" 1903, 1909) is
a young stage of H. di spar. I examined the original specimen, which is
badly preserved. It is not always easy to identify the youngest stages of
Heteroteuthis, especially if they are badly preserved or without
tentacles; the cutaneous margin of the fins is easily lost, so that determi-
nation is even more difficult.

The older stages still differ markedly from the adult, but are easily
identified. Such a stage is shown in Figures 3 and 4 of Plate VIII. The
habitus is that of a young S e p i o 1 a but the fins are attached posteriorly,
there is a free dorsal mantle margin and an interbrachial membrane is

577

present. The fins show the typical form of the subfamily, they a re rounded,
with distinct "earlobes" and with a corner which is directed posteriorly and
inward and the margin is almost straight from the corner to the posterior
end of the base (Figure 332). The mantle sac is very short and very blunt
posteriorly. The mantle margin resembles that of the adult, but its ventral
projection is still less marked. The head is very broad, because of the size
of the eyes (Figure 332). Its differentiation resembles that of the adult.

The same is true of the funnel and the mantle cavity. Compared with the
adult, the arms are still short, with a large membrane and small differences
in length. The suckers are more numerous. The club of the tentacle also
bears a greater number of suckers and shows the characteristic structure
of the Heteroteuthinae (p. 578): it is hardly widened, the swimming margin
forms a rounded lobule at the base, and there are more than 8 rows of very
small suckers of uniform size. Sheen and coloration resemble those of
the adult, and the chromatophores are more numerous, especially on the
"ventral shield" (Figure 341). Such stages already show the typical reaction
of the luminous glands.

5110/2

578

601 Chapter 45

GENUS SEPIOLA

(Leach, 1817) s. restr. Naef, 1912

Contents; a. Diagnosis. — b. Typical characters (p. 602). — c. Variation of the type of Sepiola (p. 602).
d — Juvenile forms of the species in Naples (p.605). - e. Sepiola (Eusepiola)steenstrupiana
(p. 610). — f. Sepiola (Eusepiola) aurantiaca (p. 612). — g. Sepiola (Eusepiola)
ligulata (p.615). - h. Sepiola (Eusepiola) rondeleti (p. 618). — i. Sepiola (Eusepiola)
intermedia (p. 621). — k. Sepiola (Eusepiola) affinis (p. 624). — 1. Sepiola (Eusepiola)
robust a (p. 626).

a. DIAGNOSIS

Luminous glands typical, ear- shaped, well developed, attached laterally
to the ink sac.

Leach (1817) described S. rondeleti on the basis of the original
description by Rondelet. Rondelet's figure is thus decisive for the name
of this genus. We therefore give a reproduction of the original drawing
(Figure 343), which cannot be determined to the species. "S. rondeleti -
ana" d’Orb. 1839 (Sepioles, Plate 2, Figure 14) certainly belongs to
Sepiola because of the luminous glands, but the species is not certain
(see also pp. 618 and 640).

602 This genus contains the Mediterranean species described below, the

closely related S. atlantica d'Orb. (1893) and Sepiola ( I ni ot euth i s)
japonica Verrill (1881), as a representative of a separate subgenus.
Numerous species have probably not been described. '■=

b. TYPICAL CHARACTERS

The genus Sepiola is the type of the Sepiola -like Eusepiolinae, in
contrast to Sepietta and Rondeletiola and is characterized by the
absence of special characters. The luminous glands of Sepiola have the
form typical for all Eusepiolinae (p. 582), the corresponding modification
of the ink sac, the accessory nidamental glands and the whole mantle situs
(Figure 360). The typical structure of the hectocotylus is included in the
general description of the Sepiola -like species (p. 585). The

S. japonica (Tilesius, resp. d'Orb., 1839) is problematic, S. maculosa Goodrich, 1896 is also
inadequately described and belongs perhaps to Sepietta. The same applies toS. rossiaeformis
Pfeffer, 1884; which is probably a Sepietta; its diagnosis could be given after the original species in
the Hamburg Museum of Natural History.

579

hectocotylization of the other arms was described on p. 579 in the
characterization of the type of the Eusepiolinae. All known species of
Sepiola are small, little longer than 6 cm (without the tentacles).

(601)

FIGURE 343. Rondelet's original figure of a "Sepiola" (after Rondelet, De Piscibus, Liber XVII, Sepiola,
p. 519). The drawing does not permit identification of the species. This is a true Sepiola of my
classification for the following reasons: the distinct but inexactly drawn luminous organ, which even shows
by points the papillae of the opening; the form of the mantle margin which is folded back as is often the case
in dead, damaged specimens; the mantle margin differs distinctly from that of Rondeletiola or
S e p i e 1 1 a . The species is probably S. ligulata orS. aurantiaca, i. e. a species with markedly
projecting ventral mantle margin.

c. VARIATION OF THE TYPE OF SEPIOLA

The species are difficult to identify from the usually badly preserved
material, unless the structure of the hectocotylus of the mature or half-
grown male is used. The differences in the structure of the hectocotylus
are so marked that it is sufficient for determination of males. It is
particularly important because corresponding changes in the female (p. 580)
prove that this is a natural reproductive association. The sexual conditions
in Sepiola are important for the determination of the species. On the
other hand, it must be admitted that neither the types of hectocotylization
nor the characters of the bursa are constant. In addition to the well defined
major types (Figure 604), there are variants of uncertain systematic
importance. There may be varieties and hybrids, but we have no data by
which to recognize them. We consider the types which occur constantly at
certain localities as good species if they show certain characters of
hectocotylization that are correlated with a characteristic general
appearance (Figures 11 and 14 of Plate XIX) of adults of both sexes.

580

603 SUBGENUS EUSEPIOLA- nov.

Diagnosis. Tentacle club with 8 or fewer rows of suckers. Gladius
in the form of a distinct rudiment. Ventral mantle margin with narrow,
distinct funnel indentation and rounded projections which form an undulate
line lateral to it (p. 587). More or less developed suckers at the base and
on greater distal half of the hectocotylus.

The European species of Sepiola have some characters which are not
distinctly primary and are absent in the other species. The common
ancestral form of all Sepiolinae has to be assumed to have had a larger
number of suckers on the the tentacle club, which is the case in
Euprymna on one hand and in R o nd e 1 e t i o 1 a and Sepietta on the
other. The reduction to 8 (or less) rows is therefore secondary. Eight rows
on the hand part of Eu sepiola are the primary condition. The club of
Eu sepiola has a characteristic form; the hand part is relatively short and
broad because of the development of the ventral protective margin (p. 568),
on which a large part of the suckers is situated. The club tapers rapidly
to a narrow, distal part which bears a more or less regular pattern of
4 rows. The distal part is usually curved dorsally toward the swimming
margin in preserved specimens.

The typical structure of the hectocotylus is the same as in Sepiola and
the Sepiola -like species (p. 579). However, the type varies in the different
species, which are illustrated in Figure 344.

The mantle cavity and especially the "bursa copulatrix" show variable
conditions which are characteristic for the species and will be described
there. The proliferation of the bursa over the left renal papillae (Figure 370)
and the genital opening, and the displacement to the right side (Figure 362)
are remarkable. I have never observed a contraction of the bursa in
Eusepiola as in Figure 374. Another character is the following; a
small circular fold is formed around the opening of the nidamental and
accessory glands (near 1 in Figure 360) the contraction of which creates a
single opening for both organs. This fold encloses the opening of the oviduct
on the left and thus facilitates oviposition (near 3 in Figure 360). This
modification is of particular importance because it proves the close
604 relationship between accessory glands and oviposition which is still
problematical (see ecological part),

Sepiola atlantica inhabits the North Sea and the Atlantic coasts.

It resembles a number of other species of Sepiola, each in some
characters. The Mediterranean species resemble each other; but they
differ from S. atlantica in the absence of a heteromorphism of the apex
of the ventral arms which probably developed by secondary simplification
(polyphyletic). I assume that these species descended from forms
resembling atlantica which migrated from the Atlantic to the
Mediterranean and underwent secondary changes. The resemblance to the
atlantica group is especially marked in the structure of the hectocotylus.

The subgenus Eusepiola contains the European species, which resemble each other in a number of
secondary characters. The subgenus Inioteuthis (established as a genus by Verrill in 1881) differs
from Eusepiola in the absence of the gladius and in the characteristic form of the hectocotylus. On
Inioteuthis see Berry (1912, Cat, Jap. Ceph,, p. 405 "Inioteuthis japonica Verrill").

The ventral marginal rows become later pressed into each other so that there are only 6 distinct rows.

In some cases (S. steenstrupiana) the number of rows decreases still more, or the "anlagen" of 8 rows
do not develop also on the widest part of the club.

581

Except for rare abnormalities, the base of the hectocotylus bears 3 normal
suckers, followed by a typical copulatory apparatus (p. 585) which develops
from the stalks of 4 suckers, the 2nd and 3rd sucker of the median row and
the 3rd and 4th of the lateral row (3, 4, 2, 1). The last 2 stalks lose the
suckers and grow into toothlike bodies which become more or less fused and
curved inward. The outer tooth formed by the fourth stalk is always much
larger than the inner, which is often indistinct. The 2nd and 3rd sucker of
the median row form together a diffuse tubercle which is fused with the
former two formations into an ear- shaped organ. The tubercle forms a
605 wrinkled, rounded, flasklike process which is curved toward the median

side. The suckers following the copulatory apparatus (2 median and 1 lateral
inS. affinis) are lost. The distal part of the arm is slightly swollen; it
always curved outward in dead specimens and the suckers of the median
rows are, at least in part, larger than those of the lateral rows. There is
a more or less distinct dorsal edge (k in Figure 344) which begins from the fusion
of the two dorsal arms and ends on the dorsally directed lateral surfac e.

(604)

FIGURE 344. Dorsal arms of males of species of S e pi o la (Eusepiola). 2X. a) S. aurantiaca.

Note that the modification has extended to the right dorsal arm and note the form of the copulatory apparatus
on the left arm and the enlargement of the suckers on the distal part, b) S. atlantica. This species is
most closely related to the type, c) = g), left side. d)S. rondeleti. e)S. intermedia. OS. steen-
strupiana. g)S. ligulata. The shovel extends to the right, and the copulatory apparatus is divided
into a lobe directed anteriorly and a tooth (outer tooth) which is curved upward and outward (cf. c).
h) S. robusta . i) = e) seen from above; note the. groove (r) on the inner side of the arm. k) = 1) seen
from above, showing the edge (k) which borders on the groove. 1) S. affinis. Further explanations in
the text.

582

This group includes all the Mediterranean species, except S. aurantia-
ca, which is not strictly Mediterranean (Naef, 1912), and S. ligulata,
both of which show a different type of hectocotylization. As the common
characters of the atlantica group are typical for Eusepiola, the group
is not a separate systematic unit but only stresses the difference between
the type and the two above species. I do not consider these two species
as subgenera for certain reasons. Among the Mediterranean species of
the atlantica group, S. rondeleti, intermedia and affinis form
together the rondeleti group which is characterized by a dark brown
coloration (Plate XIX), and similar hectocotylization. S. rondeleti and
S. intermedia can easily be derived from S. af f i n i s (p. 642). I
considered S. affinis at first as a hybrid of rondeleti and inter-
media, because S. af f ini s is intermediate between these two forms.
However, S. affinis is much more common in Naples than the other two
species, especially S. rondeleti, with which it occurs together.

d. JUVENILE FORMS OF THE SPECIES IN NAPLES

The young forms collected or obtained from eggs can be easily dis-
tinguished in life from those of Sepietta and Rondeletiola because
of their completely developed, characteristic luminous glands (Figure 334),
which are visible through the mantle. They are absent in Sepietta and
markedly atypical in Rondeletiola (Figure 372; Plate XIX, Figures4,
16). They are small at hatching (p. 581), and develop gradually. They are
disproportionately large in half-grown specimens (p. 5 82) and later become
markedly smaller during the sexual functions, particularly in the female
(Figure 370).

Fully developed embryos and the youngest forms cannot be determined
with certainty, although they are apparently different. They will therefore
be described separately, and the species will be characterized by the older
stages, which can be identified, although the characters are difficult to
recognize. The young animals or mature embryos which I can distinguish
probably belong to some of the species described below. All these young
forms show at first the arm formula: 3, 2, 4, 1,

606 Species 1 (probably S. ligulata). This form was obtained from eggs
which are regularly found at the Amontatura. Circumstances did not permit
to obtain complete development so that identification was not possible.
However, eggs of Sepiola from the same locality almost always produce
animals of the same form, and S. ligulata is the only species that occurs
regularly in the locality in which the eggs were found. However, the rarity
of S. ligulata does not agree with the frequent occurrence of the eggs, so
that this argument of determination becomes doubtful.* The coloration
corresponds more or less to that of S, ligulata, but the young forms are
darker, more reddish brown, and do not show the orange -red or orange -
yellow tones of S. ligulata. These are the only embryos and young stages
of Sepiola which are common in Naples (other species are rare, and
occur accidentally) and they are particularly large. They may also belong

The habitat does not necessarily coincide with the area of oviposition. Cf. the ecological part.

583

toS. robusta. Figure 346 shows a young stage of Species 1, well-
preserved after 14 days in the aquarium; it had fed on My si s for a Week.
It did not show any marked changes in form, although it grew markedly
and its terminal spine, Hoyle's organ and the yolk sac were resorbed
(cf. also Figures 347 and 348).

FIGURE 345. Species 1, young Sepiola (s. ligulata?), freshly hatched, with a remnant of the yolk sac.

6x. Characteristic for this species are its size, the thick,, fleshy body, reddish brown coloration, distribution of
the dark chromatophores (the yellow chromatophores are suppressed) ; form of fins (Figure 352). Note also the structure
of the suckers (smaller drawing), which differs from that of the adult, the typical form of the stalks of the
suckers (4) with the distinct rudiment (2) of a support of the protective margin, the minute terminal spine (8);
Hoyle's organ (6; dotted lines); the well defined cutaneous margin of the fins (5), the typical mantle margin,
which becomes folded over during preservation, the glandular lines on the head (7). The lid is contracted
and dorsally open, the olfactory tubercle is still almost embryonic. The specimen is not much deformed by
preservation but the mantle is maximally contracted, so that the head is very prominent (Figure 353).

The natural form is better preserved in Figure 346.

Species 2. I have only a few fully developed embryos of this species.

Its origin is not exactly known, but it is certainly from the Bay of Naples or
its vicinity. I obtained them when I knew only "Sepiola rondeleti"

607 (p. 584). They probably belong to S. steenstrupiana or S. aurantiaca
because all other species are apparently excluded. S. robusta,

S. intermedia and S. affinis need not be considered, while S. ligulata
and S. rondeleti are placed in species 1 and 3. Characteristic for these
embryos is their slender, delicate body and their small, anteriorly situated
eyes. The fins are small, with a broad margin. There are dense, dark
(reddish brown) chromatophores. The form of the head is especially
characteristic (Figure 349).

Species 3 (probably S. rondeleti). Embryos of this type were found
rarely near the Zoological Station and along the shore, in localities where
S. affinis and S. r o n d e 1 e t i o ccur. S. affinis can be excluded because

608 of the density of chromatophores. This species resembles the preceding
species in its delicate body and in the distribution of the chromatophores,
but it has a distinctly different head with relatively large, laterally
protruding eyes. The mantle sac is also more rounded (Figure 350).

Species 4 (probably S. affinis) (Figure 351). I obtained these young
stages from the plankton near the shore, and only in small numbers. The
specimen could not have hatched long before. It differs from the forms

584

described above especially in the distribution of the chromatophores,
from species 1 (Figure 345) also in the fins with a broad margin and the
color (sepia brown) of the chromatophores, from species 2 in the form of
head and mantle and the size of the eyes, and from species 3 in its more
robust body and large eyes, but this may be at least partly due to its more
advanced development.

:607)

FIGURE 346. Young stage of Species 1, 14 days after hatching. The
body has a more or less natural form, although the mantle is slightly
contracted. Compare with Figures 345 and 347.

FIGURE 347. Young stage of Species 1, 25 days after
hatching. The form of the body is less well preserved
than in Figure 346. 6x. The juvenile characters of
the club and especially the swimming margin begin to
disappear (Figure 349). Note the differentiation of the
buccal funnel, which is contracted above the mouth;

FIGURE 348. Young stage of species 1, 14 days
old, but shrunken and badly preserved. This
figure shows the importance of careful preserva-
tion for systematic study. Only a specialist can
determine the form of the living animal from
such badly preserved material.

1 — swimming margin; 2, 3 — ventral protective
margin.

585

FIGURE 349. Species 2, young Sepiola ,
immediately before hatching. 6x. Note (in
comparison with Figures 345 and 350) the
form of mantle and head. The fins are small,
with a broad margin. The structure of the
club and the proportions of the arms are
typical (formula: 3, 2, 4, 1). There is a
small terminal spine; remnants of Hoyle's
organ and the yolk sac are also present.

FIGURE 350. Species 3, young Sepiola
(S. rondeleti?), immediately before
hatching. 6x. Compare with Figures 345
and 349. Size of body and form of head
are different. Note the parts mentioned
in Figure 349. The lens is distinct be-
cause the lids are wide open. Collected
on 27 June 1913.

FIGURE 351. Species 4, young Sepiola
(affinis?), (about 14 days old; collected
on 30 November 1911 near the Zoological
Station). Note form of fins (cf. Figure 345),
form of mantle, head, tentacle club
(swimming margin), and especially the
characteristic arrangement of the very
few, large chromatophores. 6x.

586

This young form apparently belongs to S. affinis, the most common
species in the coastal zone, but it may also belong to S. intermedia,
which also has few chromatophores. The specimen in Figure 352 probably
belongs to S. intermedia, because its whole character resembles this
species. At any rate, these two closely related species can only be
determined by direct breeding, because the youngest stages are more or
less similar. Another specimen which probably belongs to this species is
the young Sepiola caught near the Zoological Station (Figure 353),
although this specimen has only traces of chromatophores.

609

FIGURE 352. Young Sepiola of the same (or closely
related) species as in Figure 351 (affinis or inter-
media). 6X. Collected on 12 March 1912 at the
Amontatura. Note the compact body and the sparse
chromatophores, which are light reddish brown.

FIGURE 353. Y oung Sepiola of the same or a closely related species (S. affinis or intermedia). 6x.
Collected on 25 March 1913 near the shore. Good preservation shows the natural form of the head and
mantle. The mantle margin is distinct and shows the characteristic form of these young stages (see p. 587).

After examination of thousands of specimens, I can determine the older
stages of all species of Sepiola in Naples without difficulty. The
inexperienced may find it difficult to identify young females even according
to my data, but this is not very important and does not justify more
illustrations. Certain determination of badly preserved young females is
quite impossible; However, there are characters which make determination
of half-grown females possible; e. g. the apex of the ventral arms and the
swimming margin of the club in S. steenstrupiana (Figure 354) and the
mantle margin in S. aurantiaca (Figure 357). Half-grown females of
ligulata and robusta, ofrondeleti and affinis and of inter-
media and affinis are difficult to distinguish. On the other hand, half-
grown males can be easily identified by the structure of the hectocotylus.

587

even if this is not fully developed. It is practically impossible to confuse
adult males after careful examination of the hectocotylus.

Mature females can always be identified by examination of the "bursa
copulatrix." However, dissection of the mantle cavity and examination of
the bursa is not necessary if males of the same species are available.

Nor is it necessary for determination of the genus, i. e. the distinction of
Sepietta and Rondeletiola, if the species is European and the mantle
margin and tentacle clubs are well preserved (p. 587). On the other hand,
in exotic species (which do not belong to the subgenus Eusepiola)
examination of the luminous glands is essential.

610 e. SEPIOLA (EUSEPIOLA) S T E E N S T R U P I AN A
Levy, 1912

1. DIAGNOSIS

Suckers on the apex of the ventral arms becoming abruptly very small
and forming 4 rows. Tentacle club with markedly larger suckers in 2 dorsal
rows. Proximal part of swimming margin of club separated by a kind of
frenulum which connects the edge of the swimming margin with the
protective margin. Dorsal arms of male (Figure 344f) and mantle cavity
of female (Figure 355) modified as described below.

2. LITERATURE

1912 (February) Levy (Arch. Z. Exp., p. LVIII), Sepiola st e e n st r u pi a n a .

1912 (12 March) Naef (Zool. Anz., Vol. 39, p. 269, Figs. 1, 2), Sepiola tenera.

1912 (26 March) Levy (Zool. Anz., Vol. 39, p. 290), Sepiola steensirupiana,

1912 (20 August) Naef (Zool. Anz. , Vol. 40, p. 83), Sepiola steenstrupiana.

1912 Levy (Ibid., No. 2), Sepiola steenstrupiana.

1916 Naef (p. 16), Sepiola steenstrupiana.

1921 Naef (p. 538), Sepiola steenstrupiana.

3. DESCRIPTION

This species apparently occurs in Positano (Gulf of Salerno), near
Villefranche and perhaps along the coast of southern France. It has not
been found in the Bay of Naples, except for 2 specimens at Capo Posilipo
(Figures 354 and 355). I have not observed this species alive.

Closer examination shows marked differences between S. steen-
strupiana and all other species of Sepiola, but its habitus shows
no special characters, and the slightly atypical form of the fins in the young
(Figure 356) and adult male may be due, at least in part, to preservation.

The following parts are specific: 1) apex of ventral arms; 2) tentacle clubs;
3) structure of hectocotylus; 4) structure of female genitalia. Characters
of less importance are the lengthening of the stalks of the suckers on the

588

arms (Figure 354, but cf. Figure 361) and the distinct shining areas on the
posterior end and sides of the mantle sac, also in preserved animals.

FIGURE 354. Mature male of S. s t e e n s t r u p i a n a from Posilipo,
natural size. The smaller drawing shows the apex of the right
ventral arm (6) of another specimen. Note also the arrangement
and proportions of the suckers of the club, the characteristic
structure of the swimming margin of the club (1, 2), the muscular
median process on the left dorsal arm (corresponding to the comb
k in Figure 3440, the long bases of the arms, and the shining area
in the posterior part of the mantle sac. 5 — Orbital pore.

611 1. The ends of the ventral arms show a character which we assumed

for the preceding stage of Heteroteuthinae and Sepiolinae (p. 573) and which
is also present in H e t e r o t e u t hi s (Figure 342 on p. 599): the suckers
(smaller drawing in Figure 354) become suddenly very small toward the
apex and form several rows. Irregularities do occur, but the apex normally
bears 4 regular rows, in a short transitional zone with a transverse row
of only 3 suckers. The quadriserial arrangement develops by alternation
of pairs. This character is directly related to the condition in S. atlantica,
in which the heteromorphous apex of the arm is longer, wider, like an
appendage, and bears 4 — 8 rows instead of the usual 4 rows.

FIGURE 355. Situs of mantle cavity of a mature female of
S. steenstrupiana from Posilipo. 2x. Note the typical
form of head, mantle, fins, funnel, lateral funnel adductors,
funnel cartilages, olfactory organ, gills, ovary, nidamental
and accessory glands (2), the openings of which are covered
on the left but open on the right in a pore (1). Note
particularly the moderate size of the bursa (3) and the left
branchial septum (p. 582).

FIGURE 356. Young S. st e e n st r u pi -
ana from Positano. 6x. This stage
should be compared with the more ad-
vanced stages from the same locality.

It already shows indications of the
characteristic multiserial arrangement
at the growing apex of the ventral
arms; it is also recognizable by the
form of the swimming margin of the
club.

589

2. The tentacle club (Figure 354) differs markedly from that of the
other species: it bears 4 — 5 rows in its widest part and this did not develop
from an arrangement of 8 rows, because in this case the 4 — 5 ventral
rows would be pressed into each other. The two dorsal rows contain a
number of markedly enlarged suckers. Of particular interest is the
swimming margin: its proximal part, the only part present in embryos,
is sharply differentiated from the distal part, because a frenulum develops
between the two parts which extends from the protective margin to the edge
of the swimming margin. The proximal part of the margin thus forms a
trough like that in Heteroteuthis (Plate VIII), in which the distal part
does not develop at all. This development may be a reminiscence or a
beginning of differentiation which is constant in the Heteroteuthinae but
is suppressed in the other Sepiolinae. This is interesting as a species
character.

612 3. The dorsal arms of the male (Figure 354) are more distinctly

connected at the base than the other arms; on the left arm (i. e. the
hectocotylus) there is also a muscular process near this connection on the
median edge. This process forms a strengthening of the edge of the arm
as in S. atlantica (Figure 344k), and is also recognizable on the inner
side of the arm (Figure 344f). The copulatory apparatus is weakly differ-
entiated and is coiled anteriorly like the scroll of a violin, and is widened
from a stalklike base. The distal part is almost normal, except that the
two rows of suckers often more or less diverge. It is usually not as blunt
as in Figure 344 on p. 604. However, it is always thicker and slightly longer
than the right dorsal arm, which remains normal.

4. The mantle cavity of the mature female shows the typical conditions
of Sepiola. There is a small bursa copulatrix (3) which, however,
covers the left renal papilla, the genital papilla and the opening of the left
nidamental and accessory gland. On the right side, the opening of the
nidamental and accessory gland is surrounded by a distinct circular fold
which forms a pore.

This species is small. Adults are 3.5 cm long without the tentacles.

I could not observe the coloration of live animals. To judge from the dark
chromatophores of preserved specimens (mainly dark reddish brown,
expanded chromatophores are lighter and more reddish), the normal colora-
tion is probably orange red to reddish brown, if the chromatophores are
fully expanded. The translucent eyeball shows the usual yellow-, blue- to
brown greenish reflexes, and the ventral part of the mantle is surrounded
by a strongly shining, yellowish silvery zone which persists in preserved
specimens.

Material and localities: Villefranche (8 specimens), Canada (2),

Positano (4), Naples (Posilipo) (2).

f. SEPIOLA AURANTIACA Jatta, 1896
1. DIAGNOSIS

Tentacle club with about 8 rows of very small suckers. Ventral mantle
margin strongly produced, with a deep median incision. Dorsal arms of
mature male (Figure 344a) and mantle cavity of mature female (Figure 360)
with the modifications described below.

590

2. LITERATURE

S. a u r a n t i a c a Jatta, 1896 (Plate 5, Figure 4 (?); Plate 14, Figures 34, 35, 36, 38, 40 — p. 130). Pfeffer,
1908 (p. 49). Russell, 1905 (p. 455). Naef, 1912 (Z. Anz., Vol. 39. p. 271, Figure Id; Vol. 40, p. 85,
Figure Id). Ibid., 1916 (p. l6); 1921 (p. 538). S. pfe fieri Grimpe, 1921 (Zool. Anz., Vol. 52, p. 299;
Vol. 53, p. 4, Figures 1—5) is only a variety ofS. aurantiaca, which differs from the Mediterranean form
(var. neapolitana) in gradual differences.*'

613 3. DESCRIPTION

S. aurantiaca was described in 1896 by Jatta, who only accidentally
gave a definite diagnosis of the species by his description of the very
characteristic hectocotylus. Jatta did not know the species well; in
addition to the specimen to which the hectocotylus belongs, his material
contained specimens of S. ligulata and Rondeletiola minor named
S. aurantiaca. However, Jatta's color illustration (Plate 5) probably
represents S. aurantiaca, although this is usually much darker
(Plate XIX, Figure 13). A definite determination, however, is impossible
as the drawing is too sketchy. I have seen only a few preserved specimens
of Sepiola aurantiaca, but no live animals . S. aurantiaca
probably closely resembles S. ligulata in coloration and habitus, but
it is perhaps slightly darker, more reddish brown, at any rate with orange -
red tones, and perhaps also with yellow-orange tones. S. aurantiaca
probably lives in deeper water (200 — 400 m). Its typical form is shown
in Figures 357 and 358.

FIGURE 357. Sepiola aurantiaca. Mature
female, natural size. Note the general habitus and
especially the distribution of the chromatophores,
the form of the fins and of the ventral mantle margin.

FIGURE 358. S. aurantiaca. Mature male,
natural size. Compare with Figure 361. Note
the form of fins, mantle margin, and especially
the phenomena of hectocotylization on the left
dorsal arm, the curving of the 3rd arms and the
enlargement of the suckers on the other arms.

* The funnel indentation is still deeper, almost pointed. The suckers of the male are more enlarged. There
are apparently no changes on the base of the right dorsal arm. The tentacle clubs are larger and resemble
the type because of the distinct 6 rows and the slightly larger suckers. Grimpe *s controversy with Jatta is
therefore of no importance. Grimpe never distinguished S. pfefferi from the others, so that one does not
know which form is meant. Grimpe*s figures are obviously not very exact, descriptively or morphologically.
The suckers, for example, appear in opposite pairs, although they alternate also here.

591

The fins closely resemble those of S. ligulata. They have a
transverse posterior margin. The margin of the fins is tough, not
membranous. The mantle margin also resembles that of S. ligulata,
but it projects more markedly ventrally, like a shovel; the funnel
indentation is still deeper. The tentacles are rather short and delicate
and bear 8 longitudinal rows of rather uniform, very small suckers on the
widest part of the club.

614

FIGURE 359. Dorsal pairs of arms of mature male of
S. aurantiaca. 2x. Note the enlargement of the
suckers (1) on the outer arms (2nd pair) and also on the
middle part of the inner arms (1st pair), the attachments
of the buccal funnel (4) and particularly the typical
structure of some of the basal suckers on both dorsal
arms.i.e. the first two (2,3) of the right arm and the
suckers following the first two (normal) suckers on the
left arm (7,10). Further explanations in the text.

FIGURE 360. Situs of mantle cavity of
mature female of S. aurantiaca. 2x,
Note the generally typical structure of
all parts. The bursa displaces the left
renal papilla and also covers the genital
opening.

The dorsal arms of the male are shown in Figure 359, those of another
specimen in Figure 344a. Figure 358 shows the dorsal side. Two basal
suckers on the right dorsal arm have been lost and their stalks are more
or less modified. At their extreme development, the stalks form large,
slightly flattened processes, which are usually directed toward the left
arm (they project free in Figure 359). The right arm is otherwise normal,
except that the suckers in the middle are moderately enlarged. The left
dorsal arm is the true hectocotylus. It is very characteristic and permits
determination already in young specimens. It bears only 2 small suckers
at the base; the third, which belongs to the inner side, is lost, although
traces of its stalk (6) may be preserved. The copulatory apparatus is
characteristic: the 3rd and 4th stalks of the outer row form large processes
(7, 10) which are directed inward. They are connected at the base with a
toothlike structure (9) which is situated at the end of a muscular ridge (8)
and begins lateral to the basal suckers; it is also visible from the dorsal
side (Figure 358). This muscular ridge belongs to the outer side of the arm.
The part (12) which corresponds to the rugose tubercle and develops from
the 2nd and 3rd (or only from the 2nd) stalk of the median row resembles

* The sucker is restored in Figure Id inZool.Anz., Vol. 39, p. 269; Vol. 40, p. 83.

592

the corresponding structure in S. ligulata, but it is smaller ^nd less
flattened. This structure extends anteriorly toward the inner surface of
the arm in the form of an oval pad between the first suckers of the distal
part. The "shovel” of S. ligulata (p. 617) also has a distinct homologue
(11) in the form of a few (3) papillae which are probably the rudiments of
615 stalks of the median suckers, including certainly stalks 4 and 5 and possibly
stalk 3, which thus appears independently of the "wrinkled tubercle" (p. 604).
The distal part bears normal suckers, beginning from the 6th or 7th of the
lateral row. About the first 3 normal suckers of each row are markedly
enlarged, while the others become rapidly smaller toward the apex.

The left dorsal arm is shorter than the right, as in S. 1 i gu 1 a t a , at
least in preserved specimens. The dorsal arms are distinctly connected
at the base; a more or less rough ridge runs along the side of the arm from
the point where the two arms become separated to a low tubercle before
the middle of the arm (Figure 358a). The suckers of the 2nd and 4th arms
are markedly enlarged.

The mantle cavity of the female is less modified than would be expected
with such a hectocotylization (Figure 360). The bursa extends anteriorly
and covers the left renal opening and the genital opening anteriorly and
medially. The bursa extends little posteriorly and does not cover the
branchial septum.

To judge from preserved specimens, '■= the live animal is bright orange -
red with orange-yellow and brownish red tones. In specimens well
preserved in alcohol, the dark chromatophores are brownish red, often with
a carmine tone. Mature specimens are about 4 cm long without the
tentacles.

Material: Naples (12 specimens), Roscoff (1), Firth of Forth (1),

Bergen (4), North Sea (3).

g. SEPIOLA (EUSEPIOLA) LIGULATA

Naef, 1912

1. DIAGNOSIS

Tentacle club delicate, with about 8 rows of very small, uniform suckers.

Ventral mantle margin markedly produced, with a deep median indentation.

Dorsal arms of mature male (Figure 344 on p. 604) and mantle cavity of

mature female (Figure 361) with the modifications described below.

2. LITERATURE

1912 Naef (Z. Anz., Vol. 39, p. 271, Fig. 1, 2c), S. ligulata.

1912 Naef (Z. Anz., Vol. 40, p. 83, Fig. 1, 3c). S. ligulata.

1916 Naef (p. 16). S. ligulata.

1921 Naef (p. 538), S. ligulata.

* A point which is always overlooked is that the coloration of the live animal is not that of the chromato-
phores of preserved specimens. Partly contracted chromatophores are darker, more mat and less distinct
than in live specimens. After preservation of abundant material, one learns to reconstruct the coloration
of the live animal. The inexperienced person should confine his description to forms rather than colors
and should remember the physiological variability (e. g. irritation during preservation).

593

3. DESCRIPTION

This species is common, but never numerous, on mud bottoms at a
depth of about 200 m in the bay, including the " Amontatura, " where it always
616 occurs together with the much more common Rondeletiola minor

and Sepietta oweniana. The habitus of S. 1 i gu 1 a t a probably closely
resembles that of S. r o b u s t a and S. au r a n t i a c a , although I have never
seen S. aurantiaca alive. Figure 361 shows the typical form of the
species. The coloration of live specimens is shown in Figure 11 of
Plate XIX, but not all nuances of the coloration are exact: the animal is
almost golden-yellow, with yellowish orange to brownish red tones. The
periphery of the ventral side of the mantle and especially the posterior end
have a stronger silvery sheen than most other species of the genus. The
dark chromatophores are reddish brown in preserved animals, as in
S. robusta.

FIGURE 361. Sepiola ligulata. Mature female ( a - b ) and male (c -d ), preserved, natural size.
Note the typical habitus, the characteristic fins, the delicate tentacles, the distribution of chromatophores,
the specific modification of the left dorsal arm of the male and the slight enlargement of some suckers.

The fins differ in form from those of most other species (p. 587): they
form a rounded lateral corner directed partly posteriorly (cf. Figures 363
and 373). The posterior margin, however, is more transverse than in other
species. The fins are also relatively small. The ventral mantle margin
resembles that of S. aurantiaca, forming a markedly undulate line
(Figures 363, 365), not as shallow as in the typical species. There is thus
a deep funnel indentation and two large processes. However, these
characters are much less marked than in S. aurantiaca. The tentacles
are short and delicate. The widest part of the club bears about 8 dense
rows of uniformly arranged suckers of about the same size.

617 The dorsal arms of the male differ sharply from all forms of hecto-

cotylization observed so far (Figure 344g), particularly the left dorsal arm.
The 3 typical suckers at the base are followed by a copulatory apparatus
which does not resemble the basic form but is certainly related to it. The
main tooth, which is formed by the 4th stalk of the lateral row is curved

594

upward and is best recognized in lateral view (Figure 344c). The adjacent
secondary tooth, which developed from the 3rd stalk, forms only an
appendage or tubercle on the main tooth (cf. S. oweniana. Figure 37 8).
The adjacent formation, which developed from the 2nd and 3rd stalk of
the median row, and forms a wrinkled tubercle in other species (p. 604),
is here a thick, spatulalike lobe folded toward the distal part of the
arm and there is next to it a characteristic appendage which extends toward
the right arm. This appendage probably developed from the following
2 — 3 stalks of the median row; it forms a stalked plate or shovel-like,
slender structure and has 2 distal tubercles on the stalk part which are
apparently rudimentary stalks. This part may be analogous to the structure
which functions together with the wrinkled tubercle in S. rondeleti
(Figure 344d4) and to the rudiments observed in S. a u r a n t i a c a (p. 615),
possibly also the basal and median processes of the hectocotylus in
S. steenstrupiana (Figure 344f4).

FIGURE 362. Mantle situs of a mature female of
Sepiola ligulata. 2X. Note the large bursa,
which has broken through the mantle septum and
extends to the right side (y), leaving an open

passage between the two parts (x). The sperma-
tophores are attached on the whole organ or in the
skin. (Cf. Figure 364.)

The distal part of the arm is widened like a shovel or spoon, as in the
species of Sepietta. The widening is caused by the fusion of the
strongly lengthened stalks, which do not find space in a single row and are
curved irregularly inward and outward (at least in the preserved animal)
so that the original biserial pattern of the suckers is replaced by an
irregular arrangement. The first suckers of the lateral row are enlarged;
they have long, strong stalks so that the outer tooth of the copulatory
apparatus becomes situated in a kind of depression (Figure 344c). The
618 right dorsal arm shows only a slight enlargement of the median suckers,
also on the 2nd and 4th arms.

To this characteristic hectocotylus corresponds an equally characteristic
formation of the mantle cavity of the female, especially of the bursa
(Figure 362). The relationship between hectocotylus and bursa is quite
distinct and very instructive. The bursa proliferates anteriorly and covers
the renal papilla and the genital opening. However, the bursa breaks through
the mantle septum and occupies a large part of the right half of the mantle

595

cavity. This is unique in the genus and is a further development of the
conditions in S. rondeleti, in which the bursa forms a small caecum to
the right side (Figure 364). This caecum has here broken through. This
is of particular interest if we consider it in connection with the hecto-
cotylization. If the hectocotylus is introduced into the left half of the mantle
cavity, the shovel mentioned above (Figure 344g4) reaches the right side
of the mantle through the hole in the septum and can cover the part of the
bursa in that part. It is certainly not accidental that this shovel has a
homologue in S. rondeleti (Figure 344d4), which the right part of the
bursa is represented by a small caecum. However, the relationships in
S. rondeleti are not as clear as in this species, in which an extreme
character of the hectocotylus corresponds to the conditions in the mantle
cavity of the female.

S. ligulata is a small^species — about 4 cm long without the tentacles.

Material: about 200 specimens from the Bay of Naples and its vicinity,
mainly from the Amontatura.

h. SEPIOLA (EUSEPIOLA) RONDELETI
Steenstrup, 1856

1. DIAGNOSIS

Tentacle club with 8 rows; some dorsal suckers of hand part markedly
enlarged; 3 ventral rows pressed into each other, forming about 6 rows.
Ventral mantle margin markedly produced and with a dense row of
chromatophores. Dorsal arms of mature male and mantle cavity of mature
female modified as shown in Figures 344d and 364.

2. LITERATURE

1839 Fer. and d'Orb., Sepioles, Plate 1, Figures 1 — 2. The species shown under the name "S. rondeletii"
is probably Sepiola (Eusepiola) rondeleti. However, the male illustrated in Figure 5 is a Sepietta,
as shown by the drawing of the special form of hectocotylus in the Sepiolinae. Other species are listed under
the same name. The hectocotylus of this species was drawn by Steenstrup (1856, Plate 1, Figure 9), but he
619 did not realize that he was dealing with one of the numerous forms (species) of the presumably well known
S. rondeleti.* However, I have attributed this species (1912, 1916, 1921) to Steenstrup. Also Verany .
(1851, Plate 22, Figures c, d) apparently examined this species; at least, his drawings do not agree better
with any other species. According to Verany, this is a variety from the Posidonia area in Paestum. The text
confirms that the species in question is in fact S. rondeleti: "La variete c.d. se rencontre plus communement
sur les fonds sabloneux, aupres des rochers couverts d'algues; quand la mer est tres houleuse, on les prend
meme dans le port de Genes; elle parait sedentaire et ne pas voyager pas bandes, car jamais on n'en peche
des quantites, et on la trouve dans toutes les saisons de Tannee. Elle vit assez longtemps en captivite. "

Also: Naef, 1912 (Z. Anz., Vol. 39, p. 270, Figure Ic; Vol. 40, p. 85, Figure Ic: S. rondeleti);

Ibid., 1916 (p. 16: S. rondeleti), also 1921 (p. 538: S. rondeleti).

596

3. DESCRIPTION

Figure 363 shows the typical form of this species. The coloration of
healthy, fresh specimens is shown in Figure 14 of Plate XIX, but it should
be brighter and more coffee brown. The chromatophores are dark violet
brown in preserved specimens. The largest, well preserved females can be
identified without difficulty. They are much larger than the other species
of the rondeleti group (p. 605) and have much denser chromatophores;
this is less marked in smaller specimens, but is always recognizable: the
outer side of the arms of S. af f i n i s and i n t e r m e d i a always bears
a single distinct row of very large chromatophores (Figures 365 and 367),
which are present in the young forms of all species; this part is more
densely covered with numerous, irregularly arranged chromatophores in
several rows inS. rondeleti. The chromatophores which are at the base
of each stalk on the sides of the arms of all species and also form a
regular row are not included.

FIGURE 363. Male and female of S. rondeleti, from Naples, natural size. Mature specimens, preserved.
Note the specific characters: form of fins, chromatophores, mantle margin, large clubs of female, left
dorsal arm of male. Note also the enlargement and small size of the suckers on the arms of the male
and the orbital pore. The LV arms of the male were prevented during preservation to become curved in
the manner characteristic for all Sepiola-like species (p. 586, Figure 361).

620 The fins form a large curve which extends laterally and slightly
posteriorly, as in all species of the rondeleti group, and also in
S. robusta and Sepietta obscura. Also the ventral side of the fins
bears a few chromatophores, and numerous chromatophores are present
on the dorsal side (Figures 363 and 365). The mantle margin is of typical
form, as in the whole rondeleti group: distinctly, but not markedly
produced ventrally; the neck bond is fairly broad.

The tentacles are well developed, particularly in the female (Figure 363),
especially the clubs, and they bear accordingly stronger suckers and denser

597

chromatophores on the outer side. The suckers of the dorsal rows are
markedly enlarged in the proximal part. The broadest part bears about
6 rows, but the ventral row consists of several rows. A pattern
of 8 rows may be considered as basic. The apex of the ventral arms is
normally biserial; and an irregular compression of rows as a result of
contraction cannot be compared with the regular arrangement described
for S. steenstrupiana.

FIGURE 364. Situs of mantle cavity of a mature female of S. rondeleti, before copulation. Note the
form and size of the bursa copulatrix (4). The funnel is opened to show funnel gland (7) and funnel valve
(8; see Figure 374). On the inner side of the mantle, note the attachment of the mantle septum (5) and the
linear mantle bonds (6), which are much longer than the funnel bonds;

1 — luminous gland; 2 — accessory gland; 3 — evagination of the bursa through the mantle septum; 4 —
extension of the bursa to the inner side of the mantle.

The dorsal arms of the male (Figure 344d) also differ from those of all
other species. The right arm is normal and resembles that of the female.
The left arm is always curved outward in preserved animals; it is blunt,
laterally compressed but widened dorsoventrally. Its narrow inner surface
is directed toward the middle and bears 3 slightly enlarged suckers at the
base, a median row of larger suckers and a lateral row of small suckers
in the distal part. The copulatory apparatus consists of an inward curved
formation like an eyelet which developed from 4 stalks (p. 585). The
elements which develop from the outer row can be distinguished laterally,
especially the lateral tooth. The wrinkled, strongly inward curved tubercle
belongs to the inner row. A process opposite, which also develops from the
321 inner row, forms a groove which opens toward the middle together with
the wrinkled tubercle. This is followed distally by 1 -- 2 smaller suckers.

The mantle cavity of the mature female (Figure 364) shows marked
special characters. The bursa is very large and extends not only far
posteriorly but also behind the gill along the gill septum and toward the

598

inner side of the mantle (4). The bursa also forms a caecum into the
right half of the mantle cavity, pushing the soft mantle septum between
body and median mantle adductor before it. The bursa reaches anteriorly
to the renal papilla and luminous gland. The renal papilla, however, is
intact, though often covered by the bursa.

This species is apparently widely distribiated along the Mediterranean
coasts. On the other hand, specimens from other localities are not quite
identical with the variety in Naples. All mature specimens from Naples
are of the size shown in Figure 363a; females are always markedly larger
than males. The animals are 5.5 cm long without tentacles in Trieste,
Material: Naples (24 specimens), Trieste (9), Villefranche (9),

Cancale (1). All specimens from Naples were found isolated on the sandy
bottom in the coastal zone (Mergellina).

i. SEPIOLA (EUSEPIOLA) INTERMEDIA
Naef, 1912

1. DIAGNOSIS

This species belongs to the rondeleti group (p, 605). Tentacle
club with about 6 rows of very small suckers. Mantle margin typical,
with very sparse, large chromatophores. Dorsal arms of mature male and
mantle cavity of mature female showing the modifications described below
(Figure 344: e, i; Figure 366).

2. LITERATURE

1908 Pfeffer (p. 52, Figs. 53— 57); "S. rondeletii."

1912 Naef (Z. Anz., Vol. 39, p. 270, Fig. Ih; Vol. 40, p. 83, Fig. Ih).

1921 Naef (p. 538).

1921 Grimpe (Z. Anz., Vol. 52, pp. 299, 302; Vol. 53, p. 4).

"S. intermed ia " belongs to Sepiett a (species?), not to S. inter m ed ia .

3. DESCRIPTION

Figure 365 shows the characteristic appearance of this species. The
outlines are characteristic among the forms in Naples, especially of the
males, but there is a marked resemblance to S. af finis in form and
coloration which is dark coffee brown, with a strong shade of reddish
brown, not dull gray as in Figure 14 of Plate XIX. This species is
markedly smaller than S. rondeleti and easily distinguished from it
without detailed examination, if the specimens are well preserved. The
loose distribution of the large chromatophores is characteristic. They are
absent on the ventral side of the fins, which otherwise resemble those of
S. rondeleti. The mantle margin is of typical form.

622 The tentacles are very delicate, especially in the male, and have the
normal arrangement of small, almost uniform suckers. Figure 344e, i.

599

shows the dorsal arms of the male. The right arm is normal, except for
the slight enlargement of the suckers toward the middle, especially in the
median row. The left arm bears at the base the 3 typical suckers, which
are not markedly enlarged. These are followed by the copulatory apparatus,
which does not show the characters observed inS. rondeleti: the
wrinkled tubercle curved toward the middle projects free and does not form
a groove (or spout) together with an accessory elevation of the inner margin,
although distal to the apparatus the stalks of the inner row have prolifera-
tions which correspond to such an elevation. The suckers in the distal part
of the arm vary in size, particularly in the inner row. The basal apparatus
is followed either by 2 markedly enlarged suckers, of which the proximal
is slightly larger, or by 3 enlarged suckers, of which the median is the
largest, or by a very small sucker, then a very large and then a moderately
large sucker. The last condition is apparently a transition between the first
two; it also resembles the arrangement in the closely related Sepiola
affinis. The distal part is swollen; it is also laterally compressed and
curved outward, as in S. rondeleti. The apex is blunt. The arm is
much longer than the right but relatively shorter than in S. affinis.

FIGURE 365. Dorsal and ventral view of preserved male and female of S. intermedia. Natural size.
Note form of fins, distribution of chromatophores, the different contraction of the mantle in the two sexes,
the marked enlargement of the suckers and the structure of the left dorsal arm in the male.

The dorsal view shows further characteristic details (Figures 344i and
365a). The dorsal arms are connected at the base; the edge extending from
this connection forms a curve distal to the 3 basal suckers and reaches the
side of the arm, and then gradually disappears. Together with the
longitudinal swelling formed by the wrinkled proliferations of the first stalks
of the distal part, this edge borders a very characteristic groove. A less
developed groove of this form is also present inS. atlantica and
S. affinis (Figure 344b and k). and there are weak indications in
623 S. robusta (Figure 369a). This structure becomes particularly prominent
by the sharp constriction on the side of the arm between the 3 basal suckers
and the copulatory apparatus and forms the entrance from the inner surface

600

of the arm into the above groove. The suckers of the 2nd and 4th arms of
the male are particularly enlarged in this species (Figure 365b).

FIGURE 366. Situs of mantle cavity of a mature female of S. in ter media. 2x. Compare with Figures 364
and 368. Note the small bursa copulatrix (6) which does not cover the left genital opening. A small
circular fold (3) connects the left genital opening with that of the left nidamental and accessory gland.

Note the mantle constrictor (4), which extends from the mantle septum to the base of the left gill and in
fact runs along the mantle, which has been removed. Behind this muscle, along the septum, is situated
the broad concrescence between body and mantle. Further laterally is the caecum (7), which represents
the reduced left half of the posterior mantle cavity. (The skin which lines the mantle is preserved.)
Otherwise, the parts are typical (Figure 360 on p. 614).

1 — right accessory gland; 2 — circular fold connecting the opening of the accessory gland with that of the
nidamental gland.

The conditions in the mantle cavity of the mature female are very
characteristic (Figure 366). The bursa copulatrix is very small; its
wrinkles reach the gill septum, but they are not connected with the mantle
(Figure 364). The genital opening is exposed and connected with the left
nidamental and accessory glands by a circular fold. Especially charac-
teristic is the posterior part of the left half of the mantle cavity because
of the formation of the mantle constrictor, a cutaneous muscle which extends
from the insertion of the median mantle adductor on the mantle to the base
of the left gill (Figure 366). This muscle is situated on the mantle, not on
the body as it appears from examination of the figure; it has become
situated on the body only after removal of the mantle. Contraction of this
muscle causes a more or less complete constriction of the posterior part
of the mantle cavity, and at least its temporary closing and narrowing.

This effect is accentuated by the concrescences between mantle and body
in an area along the insertion of the mantle cavity septum on the left side
and along the muscle toward the base of the left gill. Thus, all that remains
of the left half of the mantle cavity is a narrow caecum which opens into the

601

rest of the mantle cavity near the left gill (7). A similar narrowing of the
left posterior part of the mantle cavity is observed in the maturing female
of S. atlantica: the attachment of the median mantle septum widens
624 toward the bursa copulatrix, but this widening extends more or less
uniformly to the posterior end so that this part of the mantle cavity is
narrowed along its whole length and the posterior part is not constricted as in
S. intermedia.

These changes are associated with the changes of the hectocotylus and
with copulation, but the relations are still not clear. The relatively short
distal part of the hectocotylus may be connected with the closure of the
posterior part of the mantle cavity.

This species is not rare in the deeper parts of the coastal zone and is
often caught by fishermen. Good specimens are occasionally collected
near the shore.

Material: over 100 specimens from Naples and vicinity.

k. SEPIOLA AFFINIS Naef, 1912

l. DIAGNOSIS

Species of the rondeleti group (p. 605). Tentacle club with 6 rows
of very small suckers, including slightly enlarged suckers in the 3 dorsal
rows near the base. Mantle margin typical. Chromatophores sparse.
Dorsal arms of mature male (Figure 344k, 1) and mantle cavity of mature
female (Figure 36 8) showing the modifications described below.

2. LITERATURE

Naef, 1912 (Z. Anz., Vol. 40, p. 84, Fig. 2b), 1916 (p. 16), 1921 (p. 538). Pierantoni, 1918 (p, 128).

3, DESCRIPTION

Figure 367 shows the appearance of this species. Figure 18 of Plate XIX
the coloration of a well-preserved specimen. This species of the
rondeleti group closely resembles S. i nt e r m e d i a and younger
females of the two species are difficult to distinguish. However, the
chromatophores of S. affinis are more densely arranged and S. affinis
resembles S. rondeleti also in other characters, but the whole habitus
is similar to that of S. i n t e r m e d i a . It is not impossible that this species
developed by hybridization of the other 2 species and it probably hybridizes
with them even at present. This will be discussed elsewhere.*

* The validity of this species is proved by the fact that it occurs alone and has a distinct distribution, where
it is much more common than any assumed parent species; these occur only rarely in this area, while
S. affinis is absent in other areas.

602

The tentacles (Figure 367) are very delicate, like those of S. inter-
media, particularly in the male. They are of typical formation, with
about 6 rows of small suckers, of which those near the base of the dorsal
625 rows are slightly enlarged. The dorsal arms of the male are characteristic
(Figure 344k, 1). They are intermediate between S. rondel eti and
S. intermedia, but they also resemble the dorsal arms of S. atlantica
(Figure 344b). The 3 small, normal suckers at the base are followed by
the copulatory apparatus which, resembles that ofS. intermedia.

The apparatus contains, from the outside to the inside, a main tooth which
developed from the fourth stalk of the outer row, a smaller tooth formed
by the third stalk of the same row, then a wrinkled tubercle which
represents the 2nd and 3rd stalk of the median row. The stalks of the
following suckers of the median row show wrinkled proliferations which
are less marked than in S. intermedia. Dorsal examination shows
(Figure 344k) that they partly border a small, indistinct groove, together
with the thick edge (k) from the point of fusion of the two arms. In contrast
to S. intermedia (Figure 344i), this edge is simple and there is no
sharp constriction between the basal suckers and copulatory apparatus. The
.suckers of the lateral row are uniformly small in the distal part. In the
median row 4 — 6 small suckers are followed by 3— 4 very large suckers
with a characteristically lengthened, thick basal pad; the row consists
further on of small suckers to the blunt apex. This pattern varies to
some extent. The median row may begin with 4 very small suckers,
in rare cases with 5 — 6 suckers, of which the first two are slightly larger,
or with 5 — 6 uniformly small suckers. 4 small suckers are the rule
for specimens caught near the Zoological Station (Figure 344k).

FIGURE 367. Dorsal and ventral view of female and dorsal view of male of S, affinis, natural size.
Preserved specimens. Note distribution of chromatophores, form of fins and mantle; in the male; the
slightly tapering posterior end, the thick head (as in Figure 365), enlargement of arm suckers, modification
of left dorsal arm, the curved 3rd arm and the delicate tentacles.

The conditions in mantle cavity of the mature female resemble those of
S. rondeleti, but they are less marked (Figure 368). The bursa

603

copulatrix is small; it has a process toward the gill septum, but does not
reach the mantle. Both renal papillae remain free. The bursa does not
reach into the posterior part of the mantle cavity, or into the right side
626 (Figures 364, 366). A rudimentary mantle constrictor (p. 623) is

apparently indicated in some specimens, but changes as in S. intermedia
do not occur.

The coloration of live specimens is as in S. intermedia and
S. rondeleti. However, warmer reddish brown tones are rare in the
specimens in Naples. Preserved animals show violet brownish chromato-
phores (together with yellow ones, which become rapidly bleached). The
length is usually distinctly less than 4 cm (without tentacles); larger
specimens have not been found in Naples.

Material; Naples (about 100 specimens); Cancale (southern France)

(1 male); Venice (1).

FIGURE 368. Situs of mantle cavity of a mature female of S. affinis, soon after copulation. 2x.
Compare with Figures 366 and 364. The bursa is moderately large and without particular characters (2, 1).
The organs are typical (funnel, luminous glands, anus, median mantle adductor, median mantle septum,
gills, ovary, nidamental and accessory glands). The common opening of the nidamental glands and the
accessory glands (4) is visible on the left; there is a renal papilla (3) on each side.

This is the most common species of Sepiola around Naples.

S. affinis lives at a depth of a few meters on sandy bottom, usually
together with Sepietta obscura, especially near the harbor of
Mergellina, but also near the Station to Cape Posilipo.

604

1.

SEPIOLA ROBUSTA Naef, 1912

1. DIAGNOSIS

The species belongs to the rond eleti group. Tentacle club large, with
about 8 rows of suckers, those in the dorsal rows in the widest part of the
club markedly enlarged. Mantle margin typical. Chromatophores dense.
Dorsal arms of mature male (Figure 344) and mantle cavity of mature
female (Figure 370) showing the modifications described below.

2. LITERATURE

Naef, 1912 (Zool. Anz., Vol. 39, p. 271, Figs. 1. 2; Vol. 40, p. 84, Figs. 1, 3), 1916 (p. 16), 1921 (p. 538).

627 3. DESCRIPTION

Figure 369 shows the typical appearance of the animal. Its coloration
resembles that of S. ligulata, but the dorsal side is slightly darker and
more reddish brown. The dark chromatophores of preserved animals
closely resemble those of S. ligulata in distribution and tone (dirty
reddish brown). In the middle of the dorsal side, however, they are very
dense and darker, almost violet brown. In the live animal, these colors are
orange red and reddish brown (p. 615). This species resembles S. ronde-
leti in size (about 5 cm without tentacles). However, there is no close
relationship between S. robusta and any other species. It is clearly an
isolated type. Its name refers to the strong development of the whole body
and especially of the tentacles. S. robusta apparently occurs on sand and
mud bottoms. It has never been found close to the shore or in deep water
catches of the Zoological Station. I have not seen it alive.

The fins form a broad curve laterally and slightly posteriorly. They
bear moderately dense chromatophores on the dorsal side. The ventral side
is without chromatophores. The tentacles are very strong, with broad
clubs, especially in the female. The enlargement of the suckers in the
widest part of the club and in the dorsal rows is more marked than in the
other species (S. rondeleti resembles it in this respect).

628 The head of the mature male is markedly thickened, but the suckers are
only slightly enlarged in comparison with S. intermedia (Figure 365).

The left dorsal arm of the male (Figure 344h) bears the 3 typical suckers

at the base, of which the distal sucker of the outer row is markedly enlarged.
Then follows the copulatory apparatus, and then a distal part in which the
suckers at first diverge markedly and then again converge. The distal
half of this part resembles that ofS. rondeleti: the median row contains
slightly enlarged suckers, the adjacent lateral row consists of small
suckers; the arm is always curved and laterally compressed. There is
dorsally (Figure 369a) a trace of the groove on the median lateral surface
of the arm which was described for S. intermedia (p. 622), The right
dorsal arm bears markedly enlarged suckers in the middle of both rows.

605

FIGURE 369. Sepiola robusta. Male and female, preserved, dorsal and ventral, natural size. Note
particularly the thick, compact body; the strong clubs with very large suckers; chromatophores; hectocotyli-
zatiOD of arms of male (left dorsal arm, enlargement of many suckers, curvature of LV arms).

FIGURE 370, Situs of mantle cavity of a mature female of S. robusta. 2X. Note the extent of the
bursa copulatrix (3) which proliferates anteriorly over the renal papillae but leaves the genital opening
free or only partly covered (2). A few very large eggs are shining through in the posterior part:

1 — common opening of accessory and nidamental glands of right side; 4 — part of bursa which extends
to the lateral mantle septum.

The mantle cavity of the mature female shows no special characters.
As in S. steenstrupiana, aurantiaca and ligulata, the bursa
copulatrix proliferates anteriorly over the renal papilla, which I could

606

not find; a single papilla would be sufficient because the two renal sacs
are connected in the Decapoda. The genital opening is sometimes covered,
sometimes free. The bursa does not extend posteriorly as far as in
S. rondeleti and it does not reach the mantle along the gill septum.

Material: about 150 specimens from the Bay of Naples.

607

629 Chapter 46

GENUS RONDELETIOLA
Naef, 1912

DIAGNOSIS

Ink sac pear-shaped, with luminous glands in both sexes, embedded in
the middle; luminous glands of female connected with accessory nidamental
glands. "Gladius" rudimentary or absent. Tentacle club with more than
8 rows of uniform, small suckers. Ventral arms with normal biserial
suckers to the apex. DL arms of male with markedly enlarged suckers only
in the ventral row, beginning from the 3rd or 4th sucker; one or two
proximal enlarged suckers with markedly widened openings. Funnel
indentation wide, shallow and with weakly indicated, blunt corners laterally.
Only one species:

RONDELETIOLA MINOR Naef(l912), 1921
a. DIAGNOSIS

Luminous glands fused, forming a lens -shaped organ embedded in the
middle of the ink sac (Plate XIX, Figures 4, 16). Accessory nidamental
glands of female closely connected with the above organ. Luminous glands
opening in 2 small pits near the middle, opening together with the
accessory glands in the female. Rudiment of shell absent already in half-
grown animals. Tentacle club with about 16 rows of suckers. Left dorsal
arm of male with 3 small suckers at the base, followed by the copulatory
apparatus (p. 585), which forms a transverse swelling with a very strong,
inward curved lateral tooth. Distal part of arm not widened like a spoon;
suckers of distal part enlarged to the apex in the inner row but only
in the proximal part of the lateral row (Figure 374d). Base of right gill of
female separated from median and posterior part of mantle cavity by a
septum (for protection during copulation. Figure 374a).

630 b. LITERATURE

1912 Naef (Z. Anz., Vol. 39, pp. 248, 267, Fig. li, 2d; Vol. 40, p. 82, Figs, li, 3d), "Sepietta" minor.
1914 Chun (p. 17, Atlantic), "Sepiola rondeleti." Examination of one of Chun’s specimens showed that
it does not differ from the Mediterranean form.

608

1914 Schkaff (p. 592), "Sepietta" minor.

1916 Naef (p. 3, Fig. Id), "Ro nd e let i a" m i n or .

1918 Pierantoni (p. 122, Plates 6— 8), "Rond e le t i a" m i n or .

The generic name is preoccupied by a fish; therefore:

1921 Naef (p. 538), (System) Rondeletiola minor. See also p. 148.

c. SYSTEMATIC POSITION OF RONDELETIOLA
MINOR AND DEVELOPMENT OF THE
LUMINOUS ORGANS

Differentiation and characterization of this species, one of the numerous
forms known under the name of Sepiola rondeleti, was rather
difficult. It may be surprising that a species which is so difficult to
recognize should be given generic rank (C. Chun made amiable sceptical
remarks). A detailed study, however, showed a number of very distinct
characters which are not only important systematically but are also of
great biological interest. This refers above all to the formation of the
luminous glands and the genital organs, which made it necessary to establish
, a new genus (1916).

I placed this species at first (1912) in Sepietta, but the study was not yet
complete, and I saw only the common contrast to the species of Sepiola
which proves the special relationship between the species originally placed
in Sepietta. We must therefore establish a category of a higher rank,
namely the Sepietta group, within the S e p i o 1 a -like Eusepiolinae.

Diagnosis. The Sepietta group consists of Sepiola- like
Eusepiolinae (p. 584) with the following characters: Ink sac pear-shaped.
Luminous glands not ear-shaped (as in the true species of Sepiola).
Tentacle club with more than 8 rows of small suckers. Funnel indentation
very wide, with indistinct projections of mantle margin laterally. Apex of
ventral arms without increased number of rows of suckers. Dorsal arms of
male always connected at the base for a short distance. Laterodorsal arms
with enlarged suckers only in the ventral row, of which 1—2 proximal
suckers have large, characteristically widened openings. Rudiment of shell
very weakly developed, if present at all.

The characteristics of this group can be completed by description of a
young female. Its stage of development is still indifferent, so that it does
not show definite characters. The habitus and the whole external
organization of such young stages closely resemble those of Sepiola.

These forms differ from Sepiola only in secondary specific characters,
not in any general characters. The opened mantle cavity, however, shows
631 distinct general characters of the Sepietta group, at least in the female
(the male could be easily recognized as a Sepietta), i. e. the character-
istic "anlagen" of the accessory nidamental glands on the broad, pear-
shaped ink sac. In the typical Eusepiolinae (Figure 334), these glands are
situated behind the renal papillae and are not connected with the ink sac,
while the "anlagen” of the luminous organs are situated more anteriorly
and are embedded in the ink sac. We considered these luminous organs as
derived from the accessory nidamental glands. The two structures are here
apparently again united at a higher stage in the middle between the renal

609

papillae, but maintaining the general relationship to the ink sac. We
consider this as a case of atavism, because it seems impossible that
heterogeneous organs become fused in this manner. These "anlagen"
replace both the accessory and the luminous glands. They also differ in
form and structure from those of the typical species of Sepiola: they
show characteristic, coarse, radial ridges (Figure 48 on p. 125), some of
which reach the ridgelike, sharply defined margin of the "anlage," while the
others, which alternate more or less regularly with the former, do not
reach the margin. This condition is characteristic for Sepietta and
Rondeletiola. The two rounded "anlagen" are contiguous in the middle
and are slightly pressed into the ink sac, while in other cases they are
situated loosely on it.

FIGURE 371. Main drawing; female of a Sepiett a -like form (advanced young stage of Rondeletiola
minor or Sepietta obscura). 6x. Collected in plankton, in shallow water near Naples. Note the
general habitus, and the whole external organization, which resemble a species of Sepiola, but the
chromatophores are very sparse. Note also form of mantle and head, the arms, margin of fins, boundary of
cornea, the suckers with their stalks, club (7) with the larval swimming margin (6). Ventral view; typical
conditions of the Sepiolinae in the funnel pockets, funnel, lateral funnel adductors, olfactory organ with
circular wall (4), cornea (5), funnel cartilage and funnel gland (3). Mantle cavity: funnel retractors, gills,
branchial hearts, nidamental glands (2), mantle septum, median mantle adductor. Note particularly; the
broad, pear-shaped ink sac with the characteristic oval, ridged "anlagen" of the accessory glands (1) between
the renal papillae. Small drawing; freshly hatched Rondeletiola minor, 6x. About 1 day old, obtained
from eggs collected in the Amontatura (Figure 15 of Plate XIX). Note the small size, the short dorsal arms,
the otherwise typical habitus of the subfamily, but particularly the absence of luminous glands (cf. Figure 334).

632 The "anlagen" have exactly the same form in females of Sepietta.

However, they are completely absent in males of Sepietta; I did not find
them after examination of hundreds of older and younger specimens (p. 641).
Rondeletiola minor differs in this respect. These "anlagen" are
present in both sexes (Plate XIX), like the luminous glands of the typical
Eusepiolinae (Figure 334) and the rudimentary luminous glands of Rossi a
mastigophora (Figure 330). They are at first uniformly developed in
both sexes (as in Figure 371b) but they grow more rapidly in the female and

5110/2

610

become larger than in the male (Figure 372a), but without recognizable
differences in structure. They then become differentiated into a typical
accessory gland, as the tubules form at first a strictly radial pattern
between (or on) the ridges which radiate from the center.

FIGURE 372. Situes of mantle cavity of young Ron d e 1 e t i ol a minor, a) male, 4.5 X; b) female, 4x.
Note the typical conditions of head, olfactory organ, funnel, mantle, gills, branchial hearts, anus, and renal
papillae (3), origin of median mantle adductor. Note in the male, the "anlage" of the spermatophore glands
(4) behind the left gill. The "anlagen" of the nidamental glands (5, dotted) are visible in the female. The
median mantle septum has been cut off close to the body. "Anlagen" of accessory glands (1) are present in
both sexes. They are much smaller in the male, although they have already reached the peak of development.
(The female is also slightly older.) 6 — genital process; 7 — "anlage" of bursa copulatrix; 8 — median mantle
septum.

The internal differentiation cannot be discussed here (see Part II). It
results in both sexes in the formation of a structure with the character of
an accessory gland (Figures 374, 375) but is situated slightly more
anteriorly than usual. There is a separate rounded pit on each side in
which the pores are concentrated (Figure 300), while the bodies of the
glands form a more or less unified organ.

In fresh females two elements can be distinguished in the glands without
microscopic examination (Plate XIX, Figure 16): a mat, transparent lens
is situated in the center slightly more anteriorly, surrounded laterally and
posteriorly by the typical accessory gland. Each of these two differentiated
zones includes part of the developing tubules. This shows that the lens
633 developed from a typical "anlage" of an accessory gland (cf. also Pierantoni,
1918, Plate 8, Figure 21). (Dnly the lens is present in the male and is
therefore apparently the product of the "anlage" of a true, whole accessory
gland, although it is not fully developed (Figure 4 of Plate XIX). The lens
is undoubtedly a luminous organ, which produces a luminous secretion, as
I established in many living specimens.-' It is important morphologically,
what was only a hypothesis until now, that this luminous gland belongs to
the accessory gland or develops from it. This luminous gland is certainly
homologous to that of other Eusepiolinae, and has to be derived by
modification from the typical form. It seems unnecessary to assume that

* The emitted light is not very strong, and the eyes have to be adapted by spending some time in darkness.
Not only live but also recently dead animals and also the removed glands produce a slimy secretion which
becomes dissolved or emulsified in seawater and produces a milky phosphorescence.

611

the luminous gland is a new development after disappearance of the typical
structure. This has to be excluded, because of the close relationship
between Rondeletiola and the other Sepiola-like Eusepiolinae.

We showed above that the "anlagen" of the accessory glands of Rondeleti-
ola have a characteristic position; they replace topographically both the
true accessory glands and the luminous glands of other Eusepiolinae and
enter into special relationships with the ink sac which develops further.
Preparation of the "lens" of the adult animal shows distinctly that it is
embedded in the ink sac and fills a pit in the form of a segment of a
sphere. One might consider this organ as a rudiment of the typical
luminous gland of Eusepiolinae; it is possible that Rondeletiola
developed from a form in which these organs were more or less rudi-
mentary, especially because the closely related Sepietta shows no trace
of them, except perhaps the special position of the accessory gland
(Figure 371). This is a functional organ which differs so markedly from
the typical structure fp. 581) that, instead of rudimentation, this has to be
considered as a new, very characteristic aberration of ancient develop-
mental potentialities and norms of a highly problematic phylogenetic
character.

The differentiation of the "accessory gland complex" of Rondeletiola
differs in the following points from the primary conditions which have to
be assumed for the typical Eusepiolinae.

1. The luminous glands and the accessory glands develop in fact from
a single "anlage"; this confirms what was previously only a hypothesis
(p. 575, Figure 329a). 2. The luminous and accessory glands of the

female remain unified in position and retain a common zone of opening.

3. The luminous glands of both sides remain united in the middle in a
single median glandular body of characteristic form which cannot have
resulted from rudimentation of the typical organ. These characters
634 induced me to establish a new genus in contrast to my early views, which
were based on superficial knowledge. I thought at first that Rondele-
tiola minor is a transitional stage to Sepietta, in which the luminous
glands have become quite rudimentary without disappearing completely.

I had to correct this with the above discussion. It is not certain that
Sepietta ever had luminous glands like those described above. On the
other hand, the common ancestral form probably showed similar conditions.

d. YOUNG FORMS OF RONDELETIOLA MINOR

Young females are practically impossible to distinguish from species of
Sepietta in mixed material. Males can be identified only relatively late,
after clubs and hectocotylus have developed their characters. After
dissection, males can be recognized by the absence of rudimentary
luminous glands, females only after the "lens" of the luminous glands has
become distinct (this can only be determined by examination of the mantle
cavity of fresh material (Plate XIX)). A well-developed but very delicate
gladius is characteristic for Sepietta. Despite these difficulties, I have
definitely determined the youngest stage of Rondeletiola (Figure 371,
below) because I have bred them from eggs (Figure 15 of Plate XIX).

612

They can be recognized by their small size which differs from that of the
youngest stages of related species (cf. Figure 18 of Plate XIX and
Figure 381). The freshly hatched forms have very small dorsal arms
(cf. Vol. II) and also have a specific form.

e. TYPICAL STRUCTURE OF THE HALF-CROWN
AND ADULT ANIMAL

The general form of Rondeletiola shows no specific characters;
only careful preparation reveals its characteristic form, e. g. the slender
form of the female. The coloration of the live, healthy animal (Figure 13
of Plate XIX) probably closely resembles that of Sepietta neglecta.
However, its coloration differs from that of the other species of Sepiolinae
which occur in Naples; it varies between orange -yellow, orange-red and
brownish red, but it varies also according to the locality. Many specimens
are bright orange- red, but the chromatophores of these preserved animals
are reddish brown. Rondeletiola minor differs from the otherwise
similar Sepiola ligulata in a much looser distribution of chromato-
phores and the absence of strongly shining areas on the posterior end and
sides. However, traces of such shining areas are present and more
distinct than in Sepietta. Rondeletiola is also intermediate between
Sepiola and Sepietta in the consistency of the flesh. I can usually
recognize the genus by this character, even when the specimen is discolored
and shows no other specific characters. The mantle is more transparent
in Sepietta, more milky and fleshy in Sepiola.

635 A "gladius" is absent in the half-grown animal, perhaps since hatching;
nor is there a trace of the shell sac after this stage. The fins are rather
small; they resemble those of Sepiola ligulata, S. aurantiaca,
Sepietta oweniana and S. neglecta and have an indication of blunt
lateral angles. The mantle margin forms a very wide, shallow funnel
indentation and slight, blunt lateral angles. The mantle margin of badly
preserved specimens is often simply transverse, if it is not mechanically
pushed in by the funnel.

All arms, including the ends of the ventral arms, bear simple biserial
suckers. The buccal funnel consists of 7 parts. The tentacle clubs are
typical, only slightly widened, with about 16 rows of very small, uniform
suckers. The ventral marginal rows are more or less transversely pressed
into each other. As in Sepietta and Euprymna, the club appears
velvety because of the large number and small size of the suckers.

The sexual dimorphism of this species is very characteristic.

Figure 374d shows the dorsal arms of the male, unfortunately of a badly
preserved specimen. However, all important characters are distinct.

The 3 typical small suckers at the base are followed by a characteristic
copulatory apparatus. This shows a very strong, hornlike lateral tooth
which is curved outward anteriorly and inward and is formed by the stalk
of the 4th sucker. This tooth is so large that it is visible dorsally in the

636 preserved animal (Figure 373). Next to the large tooth is a smaller tooth
which develops from the stalk of the 3rd sucker of this row. Then comes
a transverse, edged swelling which is raised slightly in the middle of the

613

arm and ends in a sharp angle on the median side. This swelling develops
from the 2nd and 3rd stalk of the median row. Its middle elevation
corresponds to the second stalk, the marginal one to the 3rd; they form at
first a more diffuse tubercle which gradually develops into a transverse
crest. In the distal part of the arm, the 5th and 6th suckers of the outer row
are lost, providing space for the large main tooth. Then follow 1 — 3 small
suckers on thin, long stalks and further distally several (4) larger suckers,
followed by gradually smaller suckers to the apex. The median row of the
distal part begins with markedly enlarged suckers which become gradually
smaller toward the apex, but from the 6th sucker onward the suckers are
637 larger than the opposite suckers in the lateral row. The arm is longer than
the right dorsal arm, thickened and usually curved dorsally and laterally.
The first suckers of the 2 rows of the distal part diverge slightly, but they
do not form a spoon-shaped widening as in Sepietta (Figure 37 8).

The dorsal arms are connected at the base by a low, swollen membrane
(see p. 630). The right arm is normal and shows only a slight enlargement
of the suckers in the middle.

(635)

FIGURE 373. Rondeletiola minor, ventral and dorsal view of female and male, natural size. The
head of the female is extended, showing the funnel pockets, lateral funnel adductors and olfactory organ.

The head of the male is more retracted. Observe the specific form of the fins, structure of club, distribution
of chromatophores. Note particularly in the male the presence of enlarged suckers on the 2nd and 4th arm.s,
the modification of the left dorsal arm, the thickening of the head, and the bell-shaped, tapering mantle sac.

The enlargement and modification of the suckers of the 2nd arms in
the mature male is important. As in Sepietta, it is restricted to the
ventral row. There are 3 (2 — 3) small suckers at the base, followed by
2 (or 1) large suckers with characteristically widened openings. Then
follow normal, enlarged suckers, which become gradually smaller distally.
Both rows have enlarged, normal suckers in the middle of the ventral arms.

The mantle cavity of the maturing female shows interesting relationships
(p. 634). The luminous glands are united with the accessory glands.-' Their
openings are situated in rounded pits near the openings of the nidamental
glands. They are bordered posteriorly by sickle-shaped folds which reach

* The two components cannot be distinguished without preparation (Figure 374; cf. Figure 16 of Plate XIX).

614

the median mantle adductor and can extend more or less over it, so that '
the opening of the nidamental glands becomes closer to that of the accessory
638 glands. The folds correspond to the folds which unite the openings of the
two organs in Eusepiola (Figure 370 on p. 628), but the opening of the
luminous gland and that of the accessory glands must remain free if
our interpretation is correct. Cf. p. 639.

FIGURE 374. Situs of mantle cavity of female and dorsal arms of male of Rondeletiola minor. 2x.
a) half-grown female; b) just fertilized female; c) female ready for oviposition. Note the varying condition
of the bursa (Bu), contracted in a), widened in b); c), the bursa has become contracted again over the sperma-
tophores (Sp). Figures a) and c) show the typical form of the genital opening (Gp) with the fold of the foreskin.
The openings of the nidamental glands (Nd) are surrounded by a similar fold (Fa). The luminous glands (Lo)
are fused with the accessory nidamental glands. The two gill septa (Kr, Ks) are characteristic and asymmetrical.
The other parts (head, funnel indentation, anus, mantle muscles, gills, nidamental glands, etc.) are typical.

Lk — body of luminous gland; Lo — opening of luminous gland and accessory nidamental gland; Be — opening
of contracted bursa copulatrix; St — folds (muscular cords) radiating from opening of bursa; Tk — funnel valve;
Td — funnel gland; Az ~ main tooth of copulatory apparatus; Nz — lateral tooth; Ho — tubercle on median
side at end of transverse crest, formed by 3rd stalk of longitudinal row. The two arms are shortened because
of preservation in formol.

The genital opening of the female is a ventrally directed slit on the
genital papilla which projects from the foreskin, i. e. slightly lateral and
posterior to the opening of the left nidamental gland (Figure 374c). The
opening of the bursa copulatrix is situated further posteriorly and laterally.
The whole organ is contracted before maturity (Figure 374a); fibers

615

radiate from the narrow opening in all directions but mainly toward the
median mantle muscle. They form folds which indicate the area over
which the whole bursa is extended before copulation (Figure 374b).

(637)

FIGURE 375. Situs of mantle cavity of mature male of R o n d e 1 e t i o 1 a minor. 2x. a) viscera after
removal of mantle and opening of funnel to show the funnel gland (1), the genital process ("penis," 6) before
the anus, and at base of left gill, and posterior part of funnel retractors. Median mantle septum and median
mantle adductor cut off close to the base. The following parts are visible (translucent, dotted): branches of
veins (5) with transverse connection; branchial hearts; behind them, transversely, posterior mantle artery;
parts of the genital organs (7); 8 — perforating posterior end of pocket of gill base, b) Parts in situ, mantle
removed. Median mantle septum and median mantle adductor cut off near attachment on mantle. Note the
typical topography, form of head, olfactory organ, funnel, lateral funnel adductors, funnel bonds, funnel
pockets, mantle sac, gills, branchial hearts, posterior mantle artery (behind branchial hearts), genitalia,
mantle septum and muscle, anal region, funnel organ, and renal papillae. Between the renal papillae, the
luminous glands, fused into a rounded body, and the pit of their opening. Genital process covered.

There is a branchial septum at the base of each gill. Both septa are,
however, differently developed. The generalized left septum, formed by a
simple muscle, extends posteriorly from the outer margin of the opening
of the bursa, median to the gill, and is attached on the mantle behind the
base of the gill, but without covering it (p. 582). The specific right septum
originates medially before the base of the gill and extends to the mantle
slightly lateral and anterior to its origin. It thus covers the posterior
part of the gill and separates and protects its base.

In the female ready for copulation or in the mated female the bursa
expands (Figure 374b) so that its folded inner surface is turned outside
and covers the renal, genital and nidamental openings of the left side. It
also extends markedly posteriorly, this extension being effected by the
radial muscles (Figure 374a), mainly by the muscle which forms the left
branchial septum. The spermatophores are attached to the mucosa of the
widened bursa. The bursa later becomes contracted again to the condition
shown in Figure 374a, and it is necessary to open the closed bursa to find
the spermatophores (Figure 374c).

The mantle cavity of the male shows the typical conditions of the
Sepiola-like species, except for the luminous glands. The luminous
glands with their characteristic opening pit are united into a rounded

616

body between the renal papillae. According to preservation they are
surrounded by a more or less distinct dark margin which indicates the ink
sac (Plate XIX, Figure 4; cf. Figure 375).

Branchial heart and gill are connected posteriorly by the posterior
mantle artery, which passes transversely and forms a distinct edge where
it reaches the mantle. There is apparently a slit between this edge and
the body leading to the pocket of the gill base, which is thus apparently
already perforated, as it certainly is in Sepietta and probably also in
Sepiola. This perforation is comparable but not homologous to that in
the Loliginidae and Ommatostrephidae (Figure 95 on p. 205 and Figure 207
639 on p. 419), because the topographical correlations there are quite different.
The opening in these two groups is situated before, not behind the posterior
mantle artery. However, the two arrangements are equivalent for the
circulation of water for respiration (Figure 32 on p. 100).

Rondeletiola minor occurs in the Bay of Naples at a depth of 150 —
200 m on mud and fine sand. It appears in masses (about 5000 specimens
were caught on one occasion) and is the most common species of Sepiolidae
in the area. It occurs also in the Atlantic as proved by a specimen which
Chun (1914, p. 17) named "Sepiola rondeleti." R. minor is small,
usually less than 4 cm long without tentacles, rarely slightly longer.

Material: about 10,000 specimens from the Bay of Naples.

Addendum

f. DERIVATION OF RONDELETIOLA

To understand this type, its systematic position has to be defined.
Rondeletiola is a variant of the Sepietta group (p. 360), which is
a special case of the Sepiola -like Eusepiolinae (p. 584). The
Eusepiolinae (p. 579) are specialized Sepiolinae (p. 578), which are derived
from the Rossiinae (p. 574). The variation of the luminous glands is a
peculiar, discontinuous process. The following characteristics are typical
for the Rossiinae (Figure 329): a) the transfer of the accessory nidamental
glands to the male; b) the displacement of these glands anteriorly; c) the
differentiation of the anterior part into a luminous gland; d) the separation
of the anterior part from the normal part. This condition is accentuated
in Eusepiola (p. 603) by the following: e) the luminous glands develop
before the accessory glands and therefore appear independent (Figure 334);
f) the openings of the accessory gland become connected with those of the
nidamental glands and the genital openings (p. 603; Figure 370 on p. 628).
This condition cannot be attributed to the common preceding stage of
Eusepiola and Rondeletiola (Figure 328), but the last two
specializations (e, f) must have been absent in Sepiola if a retrogressive
differentiation of the preceding step (d) is to be assumed to have taken place.
The skin folds involved in step f (Figure 374fa; cf. also p. 643,1^2)
open anteriorly in Rondeletiola, and (detached?) cutaneous glands are
present between the luminous and accessory glands of Heteroteuthis
(Plate VIII, Figure 2), also in Sepiolina, and perhaps also in
Euprymna and Inioteuthis. This may be a gradual formation
and reduction of the type within a closely related group, partly within the

617

old "Sepiola rondeleti." It must be based historically on a
corresponding change of the potencies of formation, i. e. an enrichment
and a reduction of the genotype. Stages a — f may correspond to specific
factors of formation. We thus approach an analysis of the hereditary
mass which should be continued with the exact methods of genetics.

640 Chapter 47

GENUS SEPIETTA

Naef, 1912

a. DIAGNOSIS

Luminous glands absent. "Anlagen" of accessory nidamental glands
apparently absent in males. Ink sac narrow, pear-shaped. Shell rudiment
weakly developed. Tentacle club with many more than 8 longitudinal rows
of suckers. Left dorsal arm of male with 3—4 small suckers at the base,
followed by copulatory apparatus in the form of a transverse crest with
3—4 distinct tubercles; distal part widened into a spoonlike form, tapering
to a pointed apex and with several enlarged suckers in the median row and
markedly lengthened (often fingerlike) stalks of suckers in the lateral row.
Dorsal arms connected at the base, especially in the male. Dorsolateral
arms of male: 3 — 6 markedly enlarged suckers in the ventral row, the
first 1—3 suckers with characteristic, widened openings. Suckers of
dorsal row small. Apex of ventral arms with normal, biserial arrangement
of suckers.

b. LITERATURE

Many of the early authors certainly knew this genus. Sepietta appears under various names in addition
to Sepiola. For example, Figure 5 of Plate 1 (Sepioles) in Ferussac and d'Orbigny (1839) certainly shows a
Sepietta . One might be tempted to consider the genus as characterized by Figure 5, which shows a male in
dorsal view, with a widened, spoon-shaped hectocotylus which is characteristic for the genus. However, the
species cannot be determined, the form is not that of Sepietta (Figure 379a), and the enlargement of the
suckers on the arms does not show the characteristic form. This change was interpreted to have been caused by
disease of a "Sepiola rondeleti," the suckers of which had become larger and harder (loc. cit., p. 233).
The funnel indentation of another female considered as the same species (loc. cit.. Figure 2), is narrow,
distinctly like in Sepiola . The club bears 4 rows of suckers (Figure 3). It is impossible to identify the
species (the author placed the form at the beginning of the Sepiolinae, next to Sepiola atlantica).

The generic name Sepiola, however, has been defined by Leach (Rondelet) (p. 601).

Peters (1842, Sepiola, part) distinguishes between species of Sepiola with a pear-shaped ink sac (i. e.
without luminous glands) and those with a trilobed ink sac (i.e. with luminous glands). He did not
make a generic distinction between the two groups because he was not convinced that the characters
are constant and he did not realize their significance.

641 Steenstrup (1887) grouped at least the species of his genus Sepiola according to the form of the ink sac,

which he considered as a constant character. He did not distinguish between the species, and his data
were therefore not accepted.

Jatta (1896) confused Sepiola, Sepietta and Rondeletiola. He noticed the differences between the
luminous glands, but considered them only as "una variazione organica non constituente carattere
specifico." Jatta denied a correlation between these differences and the characters of the gladius, form

619

of the fins, clubs and suckers of the tentacles. His rejection of a correlation tp. 129) is excusable
because his "Sepiola rondeleti" included not less than 10 species the characteristics of which
are extremely difficult to determine.

Levy (1912) reinstated the distinction of Peters and proposed to separate the genus Sepiola, but he had
insufficient knowledge of the characteristics of the species. He established the new genus,
"Sepidium": "Ink sac pear-shaped, without appendages" and included in it the species of
Sepietta from Naples and Trieste (also Se pie tt a oweniana and S. ob sc ur a), without
proving that they are similar except in the form of the ink sac; he named them Sepidium
owenianum. Levy arrived at this distinction mainly because his Mediterranean material consisted
almost entirely of Sepietta (some specimens sent by myself, with a note to the effect that they belong
to a genus different from Sepiola), while at Roscoff he only obtained Sepiola of a constant
character (S. atlantica), except for a single male of S. aurantiaca Jatta, which he did not
recognize and determined asS. vulgaris Gerv. and v. Beneden. Levy probably came to the
conclusion that the differences of the ink sac are constant systematic, not physiological characters.

I maintain my view on this subject (Z. Anz., Vol. 40, pp. 78 — 81) and Levy’s impolite answer (loc.
cit., Vol. 41) does not need further discussion. The name Sepidium is preoccupied by a beetle
(Fabricius).

Naef (1912, Nos. 2, 3, 7). I established the correlation between the formation of ear-shaped appendages

(luminous glands) on the ink sac and certain forms of hectocotylization, structure of the club and other
characters, and I proved their systematic significance. This resulted in the establishment of the new
genus Sepietta, which is identical with Sepidium Levy after my removal of Sepietta minor
Naef, 1912.

Naef (1916) Syst. Ubers., p. 16. New Sepiolidae, p. 3, Figures 1 — 2. Cf. also Grimpe (1921, Teuthol.

M. VII, p. 299; VIII, p. 2).

c. DERIVATION AND DEFINITION OF THE
TYPE OF THE GENUS

The genus Sepietta differs sharply from Sepiola in the absence of
luminous glands and the resulting simplification of the ink sac. This
difference can be explained phylogenetic ally as a loss of the glands during
adaptation to certain ecological factors. From a systematic -morphological
point of view, Sepietta is closely and specifically related to the true
species of Sepiola (cf. "S ep i o 1 a - like Eusepiolinae," p. 584),

Sepietta forms with them a group for which ear- shaped luminous glands
are typical and resembles in this character another preceding stage of
the classification. Still more specific is the relationship with Rondele-
tiola (p. 630). As stated above (p. 573), there are rather indifferent
young forms which suggest that the two genera developed almost directly
from a common ancestral form (Figure 328) which we name Proto -
rondeletiola. We cannot add much to the diagnosis (p. 630) of this
ancestral form of the "Sepietta group" (p. 622). At any rate, it must
have had luminous glands of uncertain form (p. 633); the loss of these
glands in Rondeletiola must have resulted directly in conditions like
those in Sepietta.

642 Sepietta can obviously not be derived directly from R o nd e 1 e t i o 1 a
minor because R. minor is more specialized than Sepietta in many
characters: the gladius is delicate in Sepietta, but is is always present,
whereas R o n d e 1 e t i o 1 a minor does not have a gladius; the branchial
septum on the right side of Rondeletiola (p. 638) is a special formation,
while it is absent in Sepietta. This is the definition of the genus.
However, there are other special resemblances of relative diagnostic value
between the species of Sepietta which stress the monotypic character
of the group.

620

d. TYPICAL STRUCTURE OF THE ADULT ANIMAL

The habitus of the species of Sepietta does not differ from that of
the Sepiola-like Eusepiolinae. This similarity has led to mistakes and
confusions (p. 584), even by renowned specialists. I can determine at a
glance dead or alive, more or less well preserved specimens of
Sepietta, without determining any special characters by the absence
of an intensive sheen on the ventral part of the mantle and at the posterior
end, the coloration of the live animal and the consistency of the flesh,

643 especially in recently dead specimens. These characters are difficult to
describe. However, only morphological characters should be used for
determination.

FIGURE 376. Situs of mantle cavity of two young female Sepietta oweniana; a) 4X; b) 3X. Note
the general characters of the family (Sepiolidae) and subfamily (Sepiolinae); head, funnel, mantle, fins,
gills, septum of mantle cavity, median mantle adductor (2), renal papillae (10). Further: posterior opening
of pockets of gill base (9) (cf. p. 638); translucent anal region (b). Note especially: development of the
genitalia, nidamental glands (8), folds of opening (13), accessory gland (1) and its gradual displacement
posteriorly (cf. Figure 371), genital process (5) with fold of foreskin (a, the fold is covered by the "anlage"
of the bursa and the papilla becomes visible after removal of the gill). Bursa copulatrix (4, b— 6); b shows
the muscles which radiate from the opening of the bursa and the adjacent branchial septum (7), which encloses
the gill base; 11 — branchial gland; 12 - vena cava.

A "gladius” is always present in species of Sepietta, but it is a very
delicate, colorless thread in the anterior dorsal part of the mantle which is
difficult to find. The fins vary markedly and none of the existing forms can
be regarded beyond doubt as the basic form. The R on d e 1 e t i o 1 a - like
fins of S. oweniana and neglecta are more likely to be the basic form
than those of S. obscura. Both forms are within the range of variation
of Sepiola (p. 587), and they do not differ much from the type of

621

Sepiolinae. The free mantle margin resembles that ofRondeletiola,
but the funnel indentation is wider and shallower (Figure 382b, d), and
the blunt, projecting corners lateral to the indentation are still less
distinct and unrecognizable in badly preserved specimens.

FIGURE 377. Situs of mantle cavity of a half mature female of S e p i e 1 1 a oweniana, 3x. The genitalia
are typical; the later striking form of the bursa copulatrix (6, Figure 380) is at this stage indicated only by
radiating muscles (5). On the right, the openings of the nidamental and accessory glands (3) are connected by
small skin folds (1) (cf. S e pi ola , Figure 370 on p. 628 and Rondeletiola, Figure 374 on p. 636). These
openings are included in that of the oviduct on the left (4).

The tentacle club of Sepietta bears always more than 8 rows of
suckers (as many as 32). The suckers form a velvety surface on the club
because they are uniformly very small. The ends of the ventral arms are
also not modified, i. e. they do not bear suckers which become suddently
smaller and form 4 or more rows.

The conditions in the mantle cavity of the male are as in Rondeletiola
(Figure 375) except for the absence of luminous glands. They differ only
little from those in Ro s s i a macrosoma (Figure 339). The conditions
in the female are more complicated and are best considered ontogenetically.

644 There is also a resemblance to Rondeletiola because the pockets of
the gill base a.re perforated posteriorly (p. 638, Figure 376b, g). The
indifferent conditions in Figure 371 soon develop into conditions charac-
teristic for the genus (Figure 376): the "anlagen" of the secondary
nidamental glands are displaced farther posteriorly, at first behind the
renal papillae (a), later even further (b). They form together a pad which
projects at first markedly but later sinks into the skin and gradually assumes

622

the typical form (Figure 377). The tubules of the gland spread finally more
or less irregularly beneath the skin, as in other Sepiolinae, while the
openings become concentrated in a shallow pit before the opening of the
nidamental gland (Figure 377). At the same time they become differentiated
and reach their full size, and the opening of the oviduct comes to be
situated closer to the opening of the left gland. Small skin folds formed
by circular muscles connect the outlet of the accessory and nidamental
glands on the left, and they become also connected with the opening of the
oviduct but do not form a common pore as in Sepiola (Figure 370 on
p. 628). The disturbance caused by the retrogressive differentiation of
the luminous glands is apparently replaced by a functional equilibrium
through the intermediate stage observed in Rondeletiola (p. 639).

FIGURE 378. Dorsal arms of mature males of Sepietta, 2x. Note the general similarity in the widening
of the hectocotylus, the structure of the copulatory apparatus, the connection between the two dorsal arms and
the enlargement of the suckers in the middle of the right arm and on the distal part of the left arm, especially
in the median row. Note the difference in the formation of 3 or 4 suckers at the base, the enlargement of
certain suckers in the median row of the hectocotylus, and the form of the median part of the copulatory
apparatus (1 — 4 are the parts of the apparatus described on p. 584); 5 — attachment of dorsal buccal pillar (6);
7 — enlarged sucker of median row; 8 — lengthened stalks of lateral row in distal part of hectocotylus.

The bursa copulatrix develops as a crescent- shaped depression
adjacent to the genital papilla (Figure 376a) which becomes deeper and
folded like a funnel (Figure 376b). It also forms a strong branchial septum
and separates the posterior part of the gills. In the mature animal the form
of the bursa differs in the various species but not markedly from the general
type of the Eusepiolinae in the characters of the genus. The left branchial
645 septum is always distinct. A right branchial septum is absent, as in

Rondeletiola (Figure 374). There are thus no essential differences
from Sepiola in the mantle cavity of the female or from the type of the
Sepiola -like forms, except for the reduction of the luminous glands.

On the other hand, the hectocotylus is of so uniform structure that at first
(Naef, 1912) I neglected the differences and placed all specimens in
S. oweniana. Further examination and study of the correlation between
hectocotylus and other characters led me to a new view (Naef, 1916).

At any rate, hectocotylization is fairly uniform. The copulatory apparatus
(p. 584) is normal. It consists of the following parts, beginning from the
outside: 1) a hooklike, inward curved main tooth which develops from the

623

fourth stalk of the lateral row; 2) a smaller lateral tooth, adjacent to the
main tooth which is formed by the third stalk; 3) a flask-shaped rugose
tubercle; 4) a median elevation, which is often indistinct and completes the
transverse crest formed by the above parts on the inner surface. The distal
part of the arm always shows a distinct, spoon- shaped widening, formed
by the flat axis of the arm and the lengthened stalks of the suckers. The
first stalks of the lateral row are especially long and form the high margin
of the spoon. The lengthening of the stalks is accentuated by a large
interval near the main tooth because at least one sucker and its stalk have
been lost. The apex tapers but it is normal at the end. There are a number
of enlarged suckers in the middle, especially several in the median row
near the copulatory apparatus.

The right dorsal arm is normal, except for some enlargement of the
suckers in the middle. The dorsal arms are connected at the base by a
short, thickened membrane.

The coloration of the species of Sepietta is yellowish brown to reddish
brown; bright orange -red and dark brown chromatophores, as in Sepiola,
are absent. The species are larger than Sepiola, 7 — 9 cm long without
tentacles.

d. SEPIETTA OWENIANA (Pfeffer, 1908)

Naef, 1912

1. DIAGNOSIS

Tentacles long, well developed; clubs with about 32 rows of very small
suckers. Fins with a slightly indicated, blunt corner. Hectocotylus with
4 suckers at the base; median row of distal part with 2 large suckers,
then 2 — 4 smaller and then again 2 larger suckers. Mantle cavity of
mature female with the characters described below.

646 2. LITERATURE

1839 d'Orbigny, Sepiola oweniana (part). The description applies also to S. obs cur a .

1896 Jatta, Sepiola r onde let ii (part). The description covers all Sepiolinae in Naples, except
S. aurantiaca. Only Figures 26 and 27 of Plate XIV are characteristic for the species.

1908 Pfeffer, Sepiola oweniana. Pfeffer gave a characteristic drawing of the hectocotylus, without
distinguishing the species from its closest relatives.* The name is therefore valid.

1912 Levy, "Sepidium owenianum." (The material consisted of Sepietta oweniana and
S. obscura. — Z. Anz., Vol. 39, p. 289.)

1912 Naef (ibid., p. 266, Figure le; Vol. 40. p. 82, Figure le), Sepietta oweniana.

1916 Naef (p. 7, Figures Ic, 2d), Sepietta oweniana.

Lozano y Rey (1905) published (Plate 4, Figure 62) the drawing of a hectocotylus of "Sepiola
rondeletii" which probably belongs to this species. However, there are 6 small suckers at the base
and the suckers of the median row of the distal part are enlarged according to the formula i,2 , 3, 4, 5, 6, 7. .
The first detail is certainly not exact, the second may be a variant of the above form. Cf. p. 647.

624

3. DESCRIPTION

The species of Sepietta differ little from each other in external
characters. The general form resembles that of Rondeletiola (p. 635);
Sepietta is slightly slenderer than the type of Sepiola which is not
a practical character in view of the physiological variation of the
proportions. S. oweniana is characterized by its size (Figure 379);
adult specimens from Naples are 7 cm long without tentacles; specimens
from Bergen are 8 — 9 cm long, (preserved or dead specimens); fresh
animals are Viq longer. Exact measurements are impossible because of
the extensibility of the long arms. The coloration (Figures 8 and 12 of
Plate XIX) of well preserved specimens is yellowish brown to dirty
647 orange-red or reddish brown, distinctly brighter than inS. obscura,
which lives near the shore.

(646)

FIGURE 379. Dorsal view of male and female ofS. oweniana, natural size, from Naples. Note the
typical habitus of Sepiola. Special characters; form of fins, form of dorsal arms and their connection,
enlargement of suckers in the male, especially the two cuplike suckers on the 3rd arm. Note also the size
(cf. Figure 383).

The fins resemble in form those of Sepiola ligulata and aur anti -
aca, Rondeletiola minor and Sepietta neglecta. They have
a lateral angle which is very blunt but recognizable in well preserved
specimens. This angle is directed slightly posteriorly, in contrast to
S. obscura (Figure 379). The tentacle clubs bear about 32 rows of very
small, uniform suckers in the oblique rows so that they appear velvety.

The hectocotylus bears 4 normal small suckers at the base, but
it does not differ otherwise from the hectocotylus of other species of
Sepietta. The copulatory apparatus shows the 4 typical elevations:
a large, usually inward curved, lateral tooth, followed by a deep incision.

625

then a flasklike, rugose body which is bent toward the arm, and a strong
adjacent edge which ends on the inner side of the arm at the basal pad of the
second sucker of the distal part. The median row of the distal part begins
with 2 markedly enlarged suckers, followed by 3 (rarely 2 or 4) smaller and
then by 2 larger suckers and becomes gradually indistinct toward the apex.
The outer row of the distal part contains moderately enlarged suckers
to the apex.

FIGURE 380. Mantle cavity of a mature female of S. oweniana, ready for mating. 2x. (Cf. Figure 384.)

The only specific character is the large bursa copulatrix (9), which covers the renal papilla and genital
opening (7) and pushes the median mantle adductor aside. Typical for all the species of Sepietta are the
posterior openings of the pockets of the gill base, the large branchial septum (10), and the absence of luminous
glands. The other characters are those of the Eusepiolinae. Compare cornea, olfactory pits, funnel, gills,
nidamental (11) and accessory glands (5), ovary (12), median mantle adductor (8), renal papilla (4); formation
of retractors of funnel (1) and cephalopodium (2), which are distinct because of stretching of the fresh specimen;
3 — lateral pocket; 6 — pit of opening of accessory glands; 13 — lateral funnel adductor; 15 — olfactory organ;
16 — lateral mantle vein; 17 — posterior mantle vein.

The presence of 4 suckers at the base is interesting, in contrast to the
3 suckers in most species of Sepiola, inRondeletiola minor and
Sepietta obscura. It has to be assumed that the 2nd sucker of the
648 inner row has become detached from the copulatory apparatus without
causing any marked modification. This change can be explained by an
abnormal case observed in S. obscura (Figure 378a), i. e. the presence
of a small, rudimentary sucker between parts 3 and 4 of the copulatory
apparatus. I assume that the second stalk of the median row is not an
essential part of the copulatory apparatus, and that the parts medial to the
lateral tooth are formed mainly by the third stalk of the row.

626

The structure of the hectocotylus has also here a relation to the formation
of the bursa copulatrix, but the exact relationship is not clear. The
difference is small, but the presence of 3 or 4 suckers at the base changes
the structure of the copulatory apparatus, to which a marked difference
of the female organs corresponds (see below). The 2nd arms are also
characteristic: the ventral row contains 3 small suckers (constantly,
in my experience), then 2 large, widened suckers and finally normal
suckers of gradually decreasing size.

FIGURE 381. Young, freshly hatched S. oweniana.
6x. The form is typical for Se pi ola . Note the
size and the sparse chromatophores. Cf. Figures 345,
349, 350, 371.

The bursa copulatrix of the mature or fertilized female is very large
and resembles that of Sepiola rondeleti (Figure 364). The bursa of
S. oweniana proliferates anteriorly over the renal papilla and extends
laterally to the branchial septum. It apparently does not become contracted
after mating. I examined a large number of specimens, some of which were
certainly fertilized, without finding a contracted bursa.

S. oweniana lives in the Bay of Naples at a depth of 80 — 400 m on a
mud bottom, usually at depths of 120 — 200 m.

Material: about 3000 specimens from Naples, 1 from Palermo, 1 from
Villefranches, 1 from Liverpool, 1 from Plymouth, 30 from Bergen.

349 4. JUVENILE STAGES

The youngest specimens can be distinguished from S. obscura and
S. neglecta by the coloration of the chromatophores. However, this
difference does not permit a reliable determination. The distinction from
Rondeletiola is also very difficult, except in males, in which the
"anlagen" of the luminous glands are either present (in Rondeletiola)
or absent. Figure 381 shows a specimen of S. oweniana hatched in the
aquarium. The species lays eggs in the aquarium just as well as in nature
(Amontatura).

e. SEPIETTA NEGLECTA Naef, 1916
1. DIAGNOSIS

Habitus as in S. oweniana, but always markedly smaller (Figure 382).
Tentacles very delicate; clubs with very small, uniform suckers in more

627

than 16 oblique, transverse rows. Hectocotylus with 4 suckers at the base;
median row of distal part at first with 4 markedly enlarged suckers.

Mantle cavity of mature female as in S. oweniana. Fins with slight,
blunt lateral angle, as in S. oweniana.

2. LITERATURE

1916 Naef, a) (p. 9, Figs, lb, 2b), Sepietta neglecta.
1921 Naef (p. 538), ibid.

3. DESCRIPTION

This new species could be considered as a variety or subspecies of
S. oweniana. The only difference between the two forms is in the
structure of the hectocotylus; all other differences are relative. This does
not mean that S. neglecta is indistinctly defined, but only that it is closely
related to S. o w e n i a n a . It is very difficult to distinguish between
immature, badly preserved females.

The coloration of preserved and probably of live animals does not differ
from that of S. obscura. Live S. neglecta would then be reddish brown.
The chromatophores are more loosely distributed than inS. obscura.

This species is much smaller than S. oweniana; mature specimens are
about 5 cm long without tentacles.

The fins resemble those of S. oweniana (Figure 379) but are more

650 distinctly heart-shaped. The tentacles, however, are smaller than in
S. oweniana and bear much smaller suckers.

The hectocotylus (Figure 378c) is characteristic: it bears 4 small,
normally situated suckers at the base. These are followed by a copulatory
apparatus which at first glance does not differ from that of S. oweniana.
However, there are some specific characters. The median elevation is
less distinct and forms a blunt, diffuse tubercle which passes on one side
into the fused basal pads of the first 2 large suckers of the distal part
and on the other side reaches the second elevation, which forms a flasklike
body. The third elevation is like a tooth and characteristically connected
with the large, usually inward curved outer tooth. The distal part of the
arm is widened like a spoon. The inner row of suckers begins with
4 uniform, markedly enlarged suckers, then follows a markedly smaller
sucker and then about 15 suckers which decrease gradually in size. The
second sucker of the outer row is little larger than the first, and the
other suckers decrease gradually toward the apex.

651 The mantle cavity of the mature female resembles that of S. oweniana.

I did not have fresh specimens for detailed examination.

The copulatory apparatus closely resembles that of S. o w e n i an a ; it
develops from the 3rd sucker of the median row and from the 3rd and 4th
sucker of the lateral row. The 5th sucker of the lateral row is lost. The
ventral row of the 2nd arm of the male has 2 — 3 small suckers at the base,
then follow 2 very large suckers with widened openings and then normal
suckers of gradually decreasing size to the apex. It differs from the equally

628

large S. oweniana in the very narrow head, but this cannot be exactly
measured because of the marked variation. The head is less narrow in
the mature male (Figure 382c). Younger females can be distinguished from
S. oweniana by their darker coloration. This is, however, only reliable
when the specimens are well preserved. Older animals can be recognized
by the delicate tentacle clubs; the hectocotylus gives distinct characters
for identification. The distribution of this species is little known.

S. neglecta is much less common in the Bay of Naples than the two
other species; it lives in shallow coastal waters (according to coloration;
deep water forms are usually more orange red).

Material: About 100 specimens from the Bay of Naples (without data on
the exact locality and usually in bad condition).

FIGURE 382. Sepietta ne gleet a , dorsal and ventral view of female and male, natural size. Note the
characteristic enlargement of the suckers on the left dorsal arm of the male and on the dorsolateral arms and
the characteristic form of fins and delicate tentacles. Mantle markedly contracted by preservation.

f. SEPIETTA OBSCURA Naef, 1916
1. DIAGNOSIS

Tentacles long, strong; clubs with about 12 rows of small suckers;
dorsal rows with markedly enlarged suckers in widest part of club. Fins
with broadly rounded margin (Figure 383). Left dorsal arm of mature
male with only 3 normal suckers at the base; median row of distal part
beginning with a very large sucker, followed by suckers of gradually
decreasing size toward the apex. Mantle cavity of mature female with
moderately large bursa copulatrix which becomes contracted after
copulation to a narrow opening in the form of a papilla (Figure 384).

629

2. LITERATURE

1839 d'Orbigny, Sepiola oweniana (part).

1851 Verany (p. 56, PI. 22, Figs. 1, b) (cf. p. 566), Sepiola rondeletii.
1896 Jatta, Sepiola rondeletii (part) (posthumous material).

1908 Pfeffer (p. 49, Figs. 48 — 52), Sepiola oweniana (? part).

1912 Levy, Sepidium owenianum (part) (specimens from Trieste).

1912 Naef, Sepietta oweniana (part) (specimens from Trieste).

1916 Naef (p. 4, Fig. la, 2a), (a) Sepietta obscura.

1921 Naef (p. 538), (a) Sepietta obscura.

652 3. DESCRIPTION

This species is certainly not a variety or subspecies of S. oweniana,
despite its close similarity. It is difficult to distinguish the two species
without the following instruction. I knew that this species has special
characters already in 1912, but I did not know the characters of the bursa
copulatrix, which are very significant and also increase the systematic
value of the hectocotylus. I did not consider the characters of the
hectocotylus important enough for the establishment of a new species,
but thought that they were variations of that of S. o w e n i an a . It later
became clear that the structure of the hectocotylus is constantly
connected with a distinct form of fins and club, and with a distinct form
of the bursa copulatrix.

FIGURE 383. Sepietta obscura, dorsal and ventral view of male and female, natural size. Specific
characters: form of fins (cf. Figure 382), relatively small suckers, distribution of chromatophores.

In addition to the structure of the hectocotylus and the corresponding
characters of the mantle cavity of the female, the species is easily
recognized by two characters: 1) The fins are almost circular, with
broadly curved lateral margin which does not have the parabolic lateral
corner that is present in the preceding species. 2) The tentacle club bears
moderately small suckers which are much less numerous than in other

630

species of Sepietta and arranged in about 12 rows in the widest part of
the club. The suckers of the dorsal rows are nnarkedly enlarged. Tentacles
and clubs otherwise resemble those of S. oweniana; they are robust,
in contrast to S. neglecta. Live, healthy animals are reddish brown to
dark brown dorsally, distinctly darker than S. oweniana. The chromato-
653 phores of alcohol-preserved specimens are accordingly darker, dirty

reddish brown, often with violet tones. The chromatophores on the ventral
side have the same color, while those of S. oweniana are always lighter
reddish brown.

The species is not as large as S. oweniana in Naples, and probably
in other localities where the two forms occur together. However, the size
of specimens from different localities is not exactly commensurable
(cf. pp. 646 and 586). The largest specimens from Naples are 5 cm long
without tentacles, those from Trieste are much larger (cf. p. 621,

S. rondeleti from the same locality), 7.5 cm long, like S. o w e n i an a
in Naples. Verany (Figures a and b of his Plate 22) probably had such
large specimens; at any rate, preserved specimens often have such an
appearance.

FIGURE 384. Situs of mantle cavity of a fertilized
female of Sepietta obscura. 2x. Note the
characteristic contracted bursa copulatrix. From
the papilla-like opening of the bursa (1) radiate
folds formed by muscles. Note the strong branchial
septum (4), its suture with the mantle has remained
after removal of the mantle; openings (2) of ovi-
duct, accessory (3) and nidamental glands which
are united on the left. The conditions are otherwise
typical for the subfamily and genus; luminous
glands absent, and pockets of gill base with a
posterior opening.

The hectocotylus (Figure 378a) is connected at the base with the right
dorsal arm by a short swollen fold. The right dorsal arm differs from that
of the female only in a slight enlargement of the median suckers. The
hectocotylus begins with 3 small, normal suckers; there is also rarely a
very small, rudimentary sucker near the crest (Figure 378a), between the
first and second elevation. The apparatus forms a transverse crest with
4 elevations, the outer elevation usually forming a more or less detached,
inward curved tooth. The second elevation is less developed than the others.
The third is rounded and more projecting. The inner elevation is a simple
edge beginning on the inner side, behind the first of the distal suckers.

The inner row of the distal part of the arm consists of normal suckers,
but the first suckers are more enlarged than on the right arm. The first
sucker is the largest, rarely the second (more frequently in specimens
from Trieste). The suckers decrease uniformly in size toward the apex.

631

The distal part is widened and spoon- shaped, as in S. oweniana,

S. neglecta and Sepiola ligulata. The stalks of the first suckers
of the outer row are lengthened and often connected with each other,

654 forming a high margin of the spoon. The stalks are often curved inward,
like the fingers of a violinist, and the widened axis of the arm is
comparable to the palm of the hand, the inner row taking the place of the
thumb. The outer row has uniformly decreasing suckers toward the apex;
the maximum is approximately at the 3rd sucker. As already the first
sucker has a more or less long stalk, the margin of the spoon has a deep
gap which may be closed by the tooth of the copulatory apparatus. The
apparatus is formed by the stalks and basal pads of 4 suckers. Relatively
late, when the other parts are already distinct, the enlarged fourth sucker
of the median row is lost, while its stalk forms the crestlike inner part
of the apparatus.

The suckers on the other arms of the male are relatively less enlarged
than in the other species of Sepietta (Figures 382c and 383c). The
laterodorsal arms have 3 — 5 suckers at the base; then follow 2 largest
cuplike suckers and then normal suckers which gradually decrease in size
toward the apex.

The bursa copulatrix of the mature female is much smaller than in
S. oweniana and neglecta, as shown by the radiating muscles in
Figure 384. It does not extend as far posteriorly or toward the branchial
septum. The bursa does not become so strongly contracted after copulation
that its opening is reduced to a narrow slit which projects outward like a
papilla. The left renal papilla is unchanged.

The species occurs along the coast of Naples from Castel dell'Ovo
to Posilipo at a depth of 3 — 12 m on sandy bottom. S. obscura is
therefore easy to collect.

Material: Naples (500 specimens), Trieste (300); several specimens
from the French coast.

632

655 Chapter 48

ORDER OCTOPODA

Leach, 1818

Contents: a. Diagnosis. — b. Typical structure of the adult animal (p. 656). — c. Typical postembryonic
development (p. 668). — d. Variation of the type of Octopoda (p. 669).

a. DIAGNOSIS

Dibranchiata (p. 90) with 8 prehensile arms, of which the 3rd arms are
homologous with the tentacles of Decapoda (p. 115). Suckers arranged in
one or two rows, without a horny ring around the opening, and therefore
unable of being transformed into hooks. Suckers sometimes with lengthened
stalks which are not separated from the sucker by deep constrictions.

Buccal arms ("buccal funnel") completely absent. Funnel without valve.

Renal opening situated near base of gills, i. e. near base of efferent vessel.
Dorsal mantle margin fused with neck on a wide area; neck plate absent.
Mediodorsal part of mantle cavity extending posteriorly between stellate
ganglia in the form of a wide sac which is secondarily connected with the
ventral mantle sac further posteriorly in the branchial region. Axis of
gills with a wide longitudinal canal between the afferent and efferent vessels
and connected with the mantle cavity between the branchial lamellae. Median
mantle septum open posteriorly, while the anterior part forms a strong
muscle the insertion of which extends anteriorly to the anal region. Female
gonoducts symmetrical, male gonoduct present only on the left side.

Internal shell completely reduced, without indication of differentiation.

As this long diagnosis shows, the Octopoda are a sharply defined,
uniform group which differs more markedly from the general type of the
Dibranchiata than the Decapoda. It is therefore not difficult to establish
a general picture on the basis of the diagnostic characters and to define
the type of the group which will be named "Protoctopus."

656 b. TYPICAL STRUCTURE OF THE ADULT ANIMAL

The typical Octopoda differ markedly from the Decapoda already in their
general form, although some forms of the Decapoda (Histioteuthidae,
Sepiolidae) resemble the form of the Octopoda. The octopod body consists
of an ovoid to sac -shaped part and strong arms connected by a membrane.

The shell is a cartilaginous plate in which a distinction of parts
comparable to the parts of the typical shell of Dibranchiata is not possible.

657 There is no trace of a phragmocone or other morphological elements.

633

It has to be assumed, however, that the shell consists mainly of a remnant
of the proostracum with a flattened cone. At any rate, the retractors of
funnel and cephalopodium are inserted also here on the shell, and the fins
are attached on its sides.

(656)

FIGURE 385. Lateral view and sagittal section of Protoctopus. The general habitus shows the conditions
of young Octopoda (cf. Plate IX). The shell is still visible in the posterior part of the body in the form of a

transverse clasp, at the sides of which the fins are inserted. Note the anterior mantle margin, funnel pockets,

olfactory organ, funnel; the transverse, rectangular pupil (Figure 389) and iris, primary slit of lid and

secondary lid; also the parts of the arm apparatus and the mouth with outer and inner lip, etc. In the

mantle cavity: the muscular median mantle septum (Ms) which reaches far anteriorly but is perforated
posteriorly, i.e. the median mantle adductor. Note the large dorsal mantle cavity (Md). The shell
(Sch) is rudimentary and replaced in its greater part by the muscular mantle (Mm).** C — coelom; G —
gonad; Si — venous sinus; Ao — anterior aorta; M — stomach; B — caecum; Vn — genital vein; Ag —
genital artery; N — renal sac; H — heart chamber; Ao — posterior aorta; Vc — vena cava (its bifurcation
posteriorly); Pk — pancreas part of liver (Lb); Ed — hind intestine; Vd — fore intestine (ingluvium);

Tb — ink sac; Gd — poison gland; Si — venous sinus; St — statocyst; Vg — visceral ganglion; Pg — pedal
ganglion; Cg — cerebral ganglion; Bo, Bu — dorsal and ventral buccal ganglion; Vb — interbrachial vein;

Z — tongue; S — subradular organ.

If the muscular mantle is moderately contracted, the posterior part of
the body is narrowly oval with a parabolic, pointed apex. However, its form

* As in the Teuthoidea, the shell of Octopoda apparently lost first its hydrostatic function, i. e. the

phragmocone, and then the supporting function, i.e. the proostracum, when the body became less compact.
The recent forms do not make rapid movements like the typical Decapoda. They float rather than swim,
like pelagic or planktonic animals.

** The shell of the preceding stages, in which the metamorphosis is less advanced, can be visualized by
extending the anterior and posterior margin at the expense of the mantle, until a longitudinal oval plate
with a spoon-shaped terminal cone is obtained (cf. Figure 87, also p. 657).

634

varies markedly since there is no solid support. It can be extended like
a cigar or shortened into a baglike or nearly spherical form.

The muscular mantle of the Dibranchiata (Sepiolidae, p. 565) reaches its
maximal development in the Octopoda. It suppresses the shell completely,
apparently through stages, as in the Teuthoidea. The shell is reduced
not only at the end of the cone, where the "anlage*' disappears from the
ventral side and leaves the posterior end free, but also from the anterior
dorsal margin. The two halves of the muscular mantle of all Octopoda are
contiguous anteriorly and the proostracum is reduced as in Spirula
(p. 516). However, this process cannot be traced any more ontogenetically
because the embryos of the Cirroteuthidae, i. e. Octopoda with a relatively
well developed shell, are unknown, while the shell is too far reduced
already in the "anlage" in the Polypodoidea (Vol. II, Plate XXV). The
conditions shown in Figure 385 are apparently not primary morphologically,
as a better developed shell (in the sense of the general type of Dibranchiata) does
occur: e. g. in the Vampyroteuthoidea, which resemble the general type of
Dibranchiata most closely, the shell is stated to form a round, dishlike plate
in the dorsal part of the posterior mantle sac (Sasaki, 1920, Proc. U. S.

Mus., Vol. 58, p. 23). Such a structure would be at least a resemblance
to the more normal conditions in the Teuthoidea with a flattened cone
(p. 146), although a typical differentiation into cone, proostracum, etc.
is absent (Naef, Fossile Tintenfische, 1922, pp. 285 and 25).

The muscular mantle is attached around the whole shell, also at the
anterior margin, so that the complete inclusion of the shell rudiment, which
is present in most species, is prepared. The muscular mantle has reached
here the maximum of its possible development (compare the Sepiolidae,
p. 565), while the shell, which was a protective structure, becomes a mere
remnant. Its supporting function is restricted to points of greater
muscular stress, while the posterior part of the body is not supported.

The reduction of the shell indicates the increase of active against passive
658 adaptations (p. 138; see also Fossile Tintenfische, pp. 33 and 163). This
does not refer to the speed of movement but to an increased variety of
aggressive and defensive muscular actions (cf. Part III),

A number of other structural changes can be explained by the development
of the muscular mantle: A nuchal concrescence develops instead of the
missing connection with the proostracum, while the supporting role of the
latter is replaced by the muscular development of the septum of the mantle
cavity. As in the Sepiolidae (p. 570), the median mantle septum has probably
extended anteriorly to the anal region and become connected with the
musculus rectus abdominis. This extended fibers to the mantle through
the septum, and strengthened its origin on each side of the rectum. Thus
developed the strong median mantle adductor between the region of the hind
intestine and the muscular mantle which obliterated the primary character
of the septum that was at first only a fold of skin by the entry of the median
mantle artery into the mantle cavity (Figures 81, 82 on p. 183). A secondary
median connection (Figure 385) developed posteriorly between the two
halves of the mantle cavity so that the adductor apparently passes free
through the cavity. This formula obviously does not correspond to the
ontogeny of the recent forms, nor to their phylogenetic stages.

The development and fusion of the mediodorsal part of the muscular
mantle is connected, as in Spirula (Figure 252 on p. 475), with the

635

formation of a large dorsal part of the mantle cavity, which extends
posteriorly to the region of the rudimentary shell. This "dorsal mantle
sac" corresponds to the part of the mantle cavity of the typical Decapoda
which forms a shallow caecum between the stellate ganglia, behind the
neck plate (Figure 37 on p. 110). The dorsal mantle sac also forms a
caecum in the young Octopus, which is divided from the ventral part of
the mantle cavity on each side by a septum in which the stellate nerves
pass (Plate X, Figures 5, 6); it later becomes perforated and is connected
secondarily with the lateroventral mantle sac in the branchial region
(Figure 392).

The posterior part of the mantle sac bears dorsally on each side a well
developed, rounded, winglike fin as a rudder which is attached on the lateral
margin of the shell. It is doubtful whether an articulation of the fins has to
be assumed in view of the reduction of the shell and the character of the
fins (cf. p. 95). The fins may have been attached to the rudimentary shell
by the shell sac (still or again?).*

659 The anterior mantle margin is fused with the head on almost the whole
dorsal side, so that little more than the ventral half remains free. The
olfactory organ is situated in the corner of the open slit, sometimes free,
sometimes situated beneath the contracted mantle margin. The other part
of the mantle slit is occupied by the funnel, which consists only of funnel
pockets and funnel as a neck bond is absent; both are visible laterally
(Figure 394). The olfactory organ is a low, oval papilla which may be
embedded more or less deeply in the swollen skin, so that an olfactory pit
is formed. However, this disappears when the subcutaneous tissue shrinks

FIGURE 386. Cross section through the head of E led one
moschata, diagrammatic. The cavities are shown dark:

Or — orbit; Si — venous sinus around optic ganglion (Op) and
white body (Wk), connected on the median side by the
periesophageal sinus. Note also esophagus, duct of poison
gland and buccal arteries. Cg — cerebral ganglion; Pg —
pedal ganglion; Kk — head cartilage; Vc — cephalic vein;

Vo — ophthalmic vein; Vn — circumorbital vein; Tr — funnel.
The accessory parts of the eye (cf. p. 96) are complicated by
differentiation of the lid, the lower margin of which (Pu) has
moved below the upper margin (Po) and functions as a kind
of cornea. The swollen secondary lid (SI) is situated near the
swollen primary lid. Ir — iris; Li — lens.

or when the skin is stretched.

The eye is formed mainly as in P r o t o d i b r a n c hu s (p. 96), but the
surrounding area shows a number of characteristic conditions. As in
Protodibranchus, the ventral and especially the dorsal margin of
the iris projects and covers parts of the lens so that the pupil is narrowed

“ The "support of the fins" of the Octopoda (Cirroteuthoidea) could also be interpreted as a fin cartilage,
which is suggested by the presence of isolated cells of cartilage (AppelloO. However, a fin cartilage
is present also in parts of the true shell (Sepia), and the embryology of the homologous structure in the
Octopodoidea proves the shell nature of the "cartilaginous rods" (Vol. II). An anatomical-embryological
study is desirable (cf. Vols. II and III).

636

to a longitudinal, open, almost rectangular opening which forms a slit if
contracted.'* **' The longitudinal oval opening of the primary lid is situated
dorsal to the pupil in the young forms, as in the Decapoda (Figure 394).

The dorsal and ventral margin of this opening become independent during
postembryonic development, so that the dorsal margin can be drawn over
the ventral margin at the corners, and the ventral margin can be drawn
beneath the dorsal. This is effected by an independent continuation of the
musculature of the two margins, that of the dorsal margin on the outer side,
that of the ventral margin on the inner side of the primary lid. Contraction
of these muscle fibers causes a diaphragmlike closure of the lid opening
above the pupil. Each margin can also produce the same effect separately.

660 The two lid margins leave at first a slit between them which leads directly
to the lens. The primary parts of the lid situated directly above the eye are
completely transparent. Around this zone rises an opaque, circular fold,

as in the Sepiolidae (Figure 369 on p. 627) and Sepiidae (Figure 303 on
p. 548). This secondary lid may form another cover over the transparent
zone, giving effective protection but also a restriction of the field of vision
or even a complete interruption, depending on whether or not a small
opening remains.

The Octopoda have only 4 pairs of arms. It is not certain which arms
of the Decapoda are absent. At any rate, the absent arms are certainly
not the 4th or 5th, but one of the 3 dorsal pairs, as the embryology shows
(Vol. II, Plates IV, XXVII, XXXIII, etc.). At any rate, it is not correct to
consider the Octopoda as Decapoda which have lost the tentacles. The
absent arms, of which no trace is left in the embryo either, are probably
the dorsal arms of the Protodibranchiata, i. e. the arms homologous with
the dorsal arms of the Decapoda. The dorsal arms are the most weakly
developed in many Decapoda already in the "anlage" (Vol. II, Plates V,

XXIII, XXVII, etc.) and their loss is thus most probable. The 8 arms are
longer than in the typical Decapoda but show a typical rectangular cross
section with outer, inner and lateral surfaces. The strong, very extensible

661 membrane which connects the outer edges of all arms is not present in
the Decapoda in this form, at any rate not between the ventral arms. This
membrane is well developed between the ventral arms of the Octopoda
(but see Figure 452). The inner surface of all arms is characteristic: it
bears a row of rounded suckers which increase in size from the base to
about the first quarter and then gradually decrease in size to the apex,
where they become invisible to the naked eye. The suckers are accompanied
on each side by a row of cirri which alternate with them, as was assumed
already for Protodibranchus. In contrast to the Decapoda, however,
the cirri stand free on the surface without connection by skin folds
(protective margin. Figure 43 on p. 116).

* The pupil of the recent Octopoda always forms a transverse slit in the light which maintains its position to
the horizontal plane by rotation of the eyeball; the slit is widened at both ends, because its dorsal margin
is curved, with the open side upward. In twilight or after accommodation the slit widens to a narrow
rectangle. The pupil of preserved specimens is usually oval.

** Such a protection of the eye is obviously more necessary in benthic than in nektonic forms. It is therefore
less developed in the Argonautidae than in the Octopodidae. It shows definite regressions in the Bolitaenidae
and Amphitretidae because of their adaptation to pelagic life; this is probably simply an inhibition of the
typical development described above. The only known lid apparatus in the Cirroteuthoidea is that of
Opisthoteuth is depressa (Meyer, 1906, Plate 16, Figure 30), a benthic species with a well-developed
diaphragm.

637

FIGURE 387. Morphology of the lid apparatus in the Octopoda. a) Condition in the larva with almond-
shaped opening of primary lid. b) The dorsal margin extends over the corner of the lid, the ventral margin
below it. c) The modified margins become contracted, and narrow the slit like a diaphragm, d) The
contraction is complete and permanent and the slit is closed, e) The previously ventral margin of the
primary lid has been displaced dorsally on the inner side of the orbit, forming on the pupil a "pseudocornea"
which covers the lens, except in a complete or abnormal state of relaxation, f) The displacement has
increased further, so that a spontaneous opening of the primary lid is excluded. Around the primary eyelid
develops (already in a) a secondary, circular lid fold which is swollen, muscular and opaque so that the
contraction of this fold restricts or inhibits vision. The parts of the primary lid inside the secondary lid fold
are transparent.

Pp — pupil; PI — corner of primary lid; SI — secondary lid; Kr — line of displacement of the two corners;

Po — primary upper lid; Pu — primary lower lid.

FIGURE 388. Mouth region ofProtoctopus.
Arms spread. Note the structure of the mouth:
The free biting edge of the lower jaw (Uk)
which covers that of the upper jaw (Ok), the
papillated inner lip (II), the smooth-edged
outer lip (Al), and the absence of rudiments
of mouth arms. The 8 inner suckers which
replace the mouth arms functionally belong
to the prehensile arms. Sh — interbrachial
membrane; Ci — cirrus.

The suckers of Protoctopus and of the Octopoda (Figure 389) show
mainly the conditions of P r o t o d i b r an c h u s (Figure 29 on p. 98). They
are situated not directly on the surface of the arm but on a muscular,
movable, pillarlike stalk which can be markedly lengthened in life, but
can also be shortened to such an extent that the suckers of preserved animals

638

are often apparently sessile, especially if the swollen subcutaneous tissue
encloses the contracted stalk. The suckers are rounded, with a narrow
marginal ring which consists in young forms of almost separated papillae
(Plate XIII, Figures 1 —3). The adhesive ring is marked with thin radial
grooves and projects slightly against the wall of the chamber in the form of
rounded papillae. The suction chamber widens considerably from the
margin toward the bottom. It also has a well developed muscular wall which
consists of predominantly radial musculature, the contraction of which
widens the chamber and creates the suction. The adhesive ring of a sucker
adheres completely to the substrate during suction; smaller objects (like
the leg of a crustacean) may also be enveloped in a bivalvular folding of the
margins so that the margin of the chamber comes into direct contact with
the prey. Detachment is by relaxation of the radial muscles in the wall
of the suction chamber and by contraction of the weaker systems of circular
662 muscles near the chamber and between the radial muscles. Figure 389a

shows the typical structure of the sucker, in comparison with the Decapoda.

As had to be assumed for Nautilus, Orthoceras and Protodi-
branchus, the ends of the arms of all Octopoda grow continuously (the
same applies to the Decapoda, but to a lesser degree), so that the absolute
number of suckers cannot be given. Because of the new formation of
suckers at the apex, their development can be followed not only in the half-
grown but in some cases also in the adult animal (Plate IX, Figure 394;

Vol. II, Plates XXVIII and XXXI).

FIGURE 389. Typical suckers of Octopoda (a) and Decapoda (b). Diagrammatic longitudinal sections through
a sucker and its base. There is a differentiated subcutaneous connective tissue the musculature of which is
connected with the muscular axis of the arm (ax). From the central nerve cord (nv) extends a nerve which
forms a small ganglion (gg) behind the suction chamber (sk). The inner wall of the sucker contains a marginal
ring (rr), an adhesive ring (hr), a wall ring (wr) and a bottom part (sp) which forms a muscular suction pad.
The adhesive ring also has a muscular wall (hm). On the other hand, the wall ring is muscular only in the
Octopoda; it is hardened by a thick cuticle in the Decapoda (x) and contains only epithelium. The suction
chamber of the Octopoda (x) is lined with a delicate, continuously renewed cuticle. The adhesive ring of the
Decapoda is also formed by cuticle; however, this consists of particles also in the Octopoda which are
movable with respect to each other as they are connected only by a thin layer. The stalk of the sucker is
differentiated into a stalk and a basal pad (bp) in the Decapoda and has an inner network of crossed muscle
fibers, its outer surface consists of a layer of circular muscles (ri) and beneath it a layer of longitudinal
fibers (Ig); rg — circular musculature of sucker (particularly differentiated in the Decapoda); z — teeth on
free margin of wall ring; y — supporting edge; v — depression in suction pad; ab — muscle for detachment
of sucker, i. e. to retract the marginal ring in the dorsomedian zone. Compare this description with that
of Niemiec (1885). Note the general agreement and also the almost entirely muscular structure of the sucker
of Octopoda which excludes a mechanized operation as that in the Decapoda (p. 120). The dead suckers
obviously have no adhesive function, and a transition to a claw-shaped or hook-shaped sucker is excluded.

Cf. Fossile Tintenfische (1922, pp. 26 — 27).

639

FIGURE 390. Situs of mantle cavity of
Prot octopus. Note the fusion of the mantle
margin with the head, position of olfactory
organ (16) at this point; insertion of funnel
pockets (13) on the mantle; the formation of
"funnel corners" (14) and their relation to the
small indentations of the muscular mantle,
into which they project. Note the course of
the funnel retractors (9) toward the rudi-
mentary shell, which still serves for their
insertion (Figure 392 on p. 664). The median
mantle adductor (i. e. the modified musculus
rectus abdominis) surrounds the anus (12).

Note the line of attachment of the septum of
the mantle cavity (3) on the muscular mantle
(cut); form and attachment (17) of gills;
topographical relationship of gills to kidneys
(1) and genital papillae (2); their support by
the strong, superficial gill retractors (7), which
radiate posteriorly to the muscular mantle;
position of branchial hearts (6) and renal sacs
with the translucent venous appendages (5).
Entrance to dorsal mantle sac (11) anterior to
stellate ganglia (10). The fins are terminal,
as in the fossil Palaeoctopus. This position
must be considered as primary, although it is
no longer common in the recent Octopoda.

8 — Efferent branchial vessel; 4 — atrium of
heart; 15 — funnel.

FIGURE 391. Situs of mantle cavity of a female of
Octopus s a 1 u z z i i , natural size. The mantle is opened
at the insertion of the median mantle adductor and spread.
The funnel corners (13) are slightly folded over. Lateral
spreading of the mantle caused spreading of the gills,
showing the gill ligament and branchial gland (10) and
the attachment (15) of the lamellae, the vessels extending
to the branchial gland and the branchial adductors (14):

1 — genital process, retracted into a "foreskin"; 2 —
visceral nerve; 3 — venous appendages of venous branches;
4— heart, translucent; 5 — atrium; 6 — branchial vein;

7 — renal papilla; 8 — branchial heart; 9 — gill retractor;
10 — branchial gland: 11 ~ stellate ganglion; 12 —
anterior mantle adductor; 13 — funnel corner.

640

(664)

FIGURE 392. Situs of mantle cavity of a half-grown Oct opu s vulgaris, natural size, a) Ventral view.
The muscular mantle is cut without changing the natural topography. One gill is opened by removing the
outer branchial lamellae at the base (1, 2). Note the narrow base of the lamellae and the wide central canal
which opens in the lateral part of the mantle cavity between the ligaments of the median branchial lamellae
(Fe in b). Tr — funnel; Tt — funnel pocket; Mu — lateral funnel adductor; Te — funnel corner; Af — anus;

Rt — funnel retractor; Ad — median mantle adductor; Po — genital opening (papilla completely retracted);

Od — oviduct; Np — renal pore; Ms — mantle septum, extending posteriorly as a ridge on the mantle; Kr —
branchial retractor; Kh — branchial heart; Kv — branchial vein; Ka — branchial artery (bulbus); Km —
branchial gland; Mf — mantle groove, into which the funnel corner of the live animal hooks; Ax — remaining
part of gill axis; 3 — efferent vessel of branchial lamella (cut), b) Dorsal view, showing the whole dorsal
mantle sac, which extends far posteriorly and is connected again with the lateral part of the mantle cavity
behind the anterior mantle adductor (Va); the median branchial lamellae (Bl) are therefore visible, and also
the central canal (Fe) between the lamellae. Note also the extension of the dorsal mantle sac anteriorly
between the insertions (As) of the funnel pockets: if the neck bond had been lost, the insertions would
be on the body, not on the mantle. The actual condition can be understood only if the lost neck bond was
fused with the mantle and transferred the insertion of the funnel pockets to this (p. 663; see also Figure 323
on p. 566). Ha — posterior mantle adductor; Ap — left lateral mantle artery; Av — passage through the
muscular mantle near the canilaginous rod (Sh), which shines through; Mm — muscular mantle, cut; As —
funnel pocket (Tt), cut at its insertion on the muscular mantle; Te — funnel corner; St — stellate ganglion;

Dm — dorsal mantle margin, fused with head; PI — opening of secondary lid, with the primary upper lid inside.

There is no trace of an inner circle of arms (buccal funnel) probably
already in Protoctopus and the 8 basal suckers are therefore situated
directly around the outer lip.

The funnel shows some special characters, particularly the absence of
a neck bond. The funnel pockets are attached instead to the mantle, and the
anterior part of the dorsal mantle sac is situated between the two points of
attachment (Figure 392). This condition is remarkable. Fusion of the neck

641

bond with the mantle in the Dibranchiata (Sepiolidae, Cranchiidae) creates
at first a markedly different condition: although the funnel pockets are
attached on the mantle, they remain attached to the neck, and a mantle
cavity is absent (Figure 323 on p. 566). We therefore have to assume that
the mantle cavity of the Octopoda has extended secondarily anteriorly and

663 that the rest of the neck bond, if it is still distinct, would be situated on the
inner side of the dorsal mantle between the two lines of insertion of the
funnel pockets. Instead of connecting the mantle with the neck, these
insertions are displaced to the mantle by the extending mantle cavity. This
morphologically unexpected condition follows from the function of these
parts. As the funnel pockets must adhere to the mantle, which makes
extensive movements, unlike in the Decapoda, in which the dorsal part of

664 the mantle is supported by the proostracum and neck bond, the pockets must
be inserted on the mantle.

The corners on which the inner margin of the funnel and the pockets are
contiguous are rounded processes which are curved outward because of
their elasticity and the contraction of the musculature. They are pressed

665 into indentations on the inner side of the mantle and this strengthens the
closing of the mantle slit which is otherwise formed by adhesion. This
prevents the funnel from being pressed out of the mantle cavity during
strong contraction and the water from flowing out at the sides instead of
through the funnel. (The inner margin of the funnel often projects outward
in dying or damaged specimens. Compare the improvement of this
primitive closure of the funnel with the typical arrangement in the
Decapoda and Argonautidae, Figure 312 on p. 558 and Figure 435 on p. 727.)
There is a typical funnel gland, which resembles that of the Decapoda in
form and differentiation (Plate VII, Figure 2), but a funnel valve is absent
in all Octopoda. If water enters the mantle cavity, the dorsal and ventral
margin of the funnel are pressed together and close the funnel opening
(Figure 404). This is connected with the type of swimming. The movements
of Octopoda are never as definite and controlled as in the Decapoda; the
funnel of the Decapoda functions much more passively and mechanically
than in the Octopoda. Even the permanent swimmers among the Octopoda
are not able to swim long distances; Octopoda are planktonic rather than
nektonic forms (cf. also the Cranchiidae, p. 392).

The mantle cavity shows a number of specific characters, in addition to
the situs (Figure 390). The stellate ganglia are widely separated because
of the development of the dorsal mantle cavity. The gills are attached far
anteriorly, and the renal papillae are situated close to the entrance to the
atrium, i. e. near the efferent branchial vessel (Naef, 1912, Coelom, p. 331;
Naef, 1913, p. 441). The branchial hearts are situated laterally on the
abdominal complex and protrude into the mantle cavity. A thin cutaneous
muscle, the superficial retractor (Plate II, Figure 1), extends far
posteriorly from the efferent branchial vessel. The renal sacs cover
the greater part of the abdominal complex.

666 The typical picture of the gill is shown in Plate X and Figures 392 and
393. The gill does not appear as a product of the folding of a primary
lamella (p. 73, Plate IV), but as the result of alternating incisions on a
compact substrate: the incisions are sharp, the swellings between them
rounded. The branchial lamellae are thick disks which little resemble the
typical form (p. 188); they are attached to the apex by ligaments of the

5110/2

642

667 second order which are markedly shortened on the ventral side. The
lamellae of the 2nd order are similar and are incompletely divided by
incisions into lamellae of the 3rd and 4th order. Between the afferent and
efferent vessels (see pp. 73, 135, 489) passes a longitudinal canal which
opens between each two lamellae and permits fresh water to flow around
the organ. The detailed distribution of the vessels is markedly modified:
the afferent vessels pass on the outer side of the lamellae of the 1st, 2nd,
and 3rd order. Each branchial lamella is accompanied by a reduced or
intercalated lamella (Jn) on the proximal and distal side.

(665)

FIGURE 393. Typical structure of the gill of Octopoda. a) Branchial lamella, lateral. The lamella is
a thick disk with alternating incisions and rounded projections; thin afferent vessels (Ab) form zigzag-shaped
anastomoses (An) on the outer side, b) Cross section of a segment of the gill. Ka — branchial artery;

Kv ~ branchial vein; Ca — longitudinal canal; Km — branchial gland; Mv -■ branchial vein; Kb — gill
ligament; Mm — muscular mantle; Kn — branchial nerve. The arrows show the direction of the blood stream;
1 — afferent vessel of a lamella in the ligament; 2 — same in lamella of the second order (Bl), passing on
the outer edge (Ab); 3 — efferent vessel, passing in the interior and reaching the branchial vein; Jn —
intercalated branchial lamella; Aa — artery of gill axis; Bdj— attachment of lamella of the second order;

Bd — same of lamella of the first order.

The jaws of all Octopoda differ from those of the Decapoda in their
weakly developed biting processes (Plate XVIII), which are only indistinctly
delimited from the rest of the jaw. The radula is not characteristic; it
has the 9 typical rows of teeth, of which the marginal teeth are simple plates
(marginal platelets). The teeth of the median row have a secondary cusp on
each side, those of the submedian row only on the outer side; both lateral
rows consist of true brushlike teeth (cf. Plate XVI, Figure 8). Some sub-
groups show marked differences.

The biology ofProtoctopus, or the typical mode of life of all
Octopoda, is assumed to resemble that of a recent Octopus, but with
more active swimming for migration or search for prey. At any rate, the
recent Octopoda cannot be directly derived from a specific benthic ancestry
form.

643

(666)

FIGURE 394. Older embryo of Octopus vulgaris. 30 X. (If disturbed, such stages hatch, detach the
outer yolk sac and are viable.) This is the simplest form of a typical octopod, the fins of which have been
lost. The mantle margin is slightly shrunken, so that funnel and funnel pockets are open posteriorly;

1 — 4 — arms; 5 — sucker; 6 — apex of flagellum; 7 — first indication of interbrachial membrane; 8 —
outer yolk sac; 9 — pupil; 10 — primary lid; 11 — funnel; 12 — funnel pocket; 13 — its posterior margin;

14 — posterior margin of funnel; 15 — olfactory organ; 16 — fused mantle margin; 17 — free mantle margin;
18 — mantle sac.

(666)

FIGURE 395. Median section through an almost fully developed embryo ofO. vulgaris, slightly
diagrammatic. 34 X. The general topography is typical for the Octopoda. Note the relationship between
outer and inner yolk sac (Do) and between this and the liver. The inner yolk sac replaces topographically
the liver in the embryo. Note also the dorsal mantle cavity (Md) and the partition of the ventral cavity by
the still complete mantle septum (Ms, hatched), which extends almost to the anus. Go — rudimentary
gonad; Co — coelom; Hz — heart; Ap— posterior aorta; Ni — kidneys; Af — anus; Mv — mantle cavity;

Tb — ink sac; Lb — liver; Pa — pancreas; Gg — gastric ganglion; Ma — stomach; Sn — blood sinus; Vc —
vena cava; Jn — ingluvium; Ao — anterior aorta; Gd — poison gland; Oe — esophagus; Vg — visceral
ganglion; Pg — pedal ganglion; Cg — cerebral ganglion; Ub — infrabuccal ganglion; St — statocyst; Kv —
cephalic vein; Dr — funnel gland; Tr — funnel; Gs — sublingual ganglion; Sr — subradular organ; Rd — pocket
of radula; J1 — inner lip; A1 — outer lip; Ok — upper jaw; Mu — mouth; Do — yolk sac.

FIGURE 396. Apex of arm of a young E led one
moschata (cf. Plate IX, Figure 5). Note the trans-
verse ridges separated by deep incisions. This
continues on the apex also in the adult, while the
proximal "anlagen" form rounded papillae and suction
chambers by invagination of the skin (cf. Nautilus,
p. 63, Figure 11). 24 x.

644

The sexual dimorphism is assumed to be as in Protodibranchus
(p. 106): enlargement of some suckers in the mature male, the body of
which was probably more slender than in the female. The spermatophores
were transferred by one or both arms of the 3rd pair, without permanent
differentiation. The spermatophores are no longer attached in the mouth
region (probably only partly because of the absence of the mouth arms) but
in the mantle cavity, on or near the female genital opening (cf. Sepiolidae).

Nidamental and accessory nidamental glands are absent. The right
gonoduct is absent in the male. Genital processes are only weakly developed
668 and can be retracted so far that pocketlike depressions are formed instead
of papillae (Ocythoe). There is no entrance into a genital pocket at the
base of the male genital process (Figure 35 on p. 105). This pocket
becomes closed at an early stage during postembryonic development,
because the free margin of the opening becomes fused with the process
(Naef, 1913, Figure on p. 448).

c. TYPICAL POSTEMBRYONIC DEVELOPMENT

The early young stages of Octopoda already show distinctly the character
of the order in the uniform formation of 8 arms even if abnormal differ-
entiations appear later. Compare, for example, Tremoctopus (Vol. II,
Plate XXXI and Figure 437) and Argonauta (Vol. II, Plates XXXVI,
XXXVII, and Figure 455). However, there is no distinct larval character
followed by a metamorphosis (but see the Polypodoidea, p. 673).

c

FIGURE 397. Larva of O c t o pu s macropus from the
plankton of Messina. 10 x. The funnel should be longer
and extended anteriorly as in Figure 174 on p. 355. The
arms are markedly contracted and curved inward during
preservation. Note the brushes on the more highly
magnified part of skin (44 x). Specific characters are
the distribution of the chromatophores and the slender
body with the very narrow head.

The earliest stages of the Cirroteuthoidea are unknown and therefore we
do not know how far the characters of the larvae of Polypodoidea can be
generalized. Common to all subgroups is certainly the very peculiar
apparatus (probably for protection and floating): The whole body is covered
with small tubercles, spines or brushes which develop from large basal
cells in cuplike depressions of the epithelium (Figure 397c). These
tubercles often split later (especially in Octopus macropus —

Figure 397c) into a stellate or tuftlike bundle of thin bristles. They are

645

present in all young Octopoda known (Octopus, Scaeurgus,
Trematopus, Argonaut a, Ocythoe). There is no trace of them
in the Decapoda.''!=

669 The eyes of the youngest Octopoda resemble those of the Oegopsida;

they have a wide open primary lid and a more or less protruding eyeball.-"!'
The arms are very short and bear very few suckers, the number of which
increases later (Figure 396 on p. 667).

d. VARIATION OF THE TYPE OF OCTOPODA

The classification of the Octopoda has been discussed recently by
various authors, but no definite result has been obtained (Naef, 1912;

Thiele, 1915; Grimpe, 1916; all in Zool. Anz,). This is partly due to
methodical errors and partly to the difficulty to find a suitable position for
the incompletely known fossil Palaeoctopus newboldi Woodw.

(see also Naef, 1921, System.).

I agree with Grimpe (1917) on the classification of the recent species.

He placed the Cirrata opposite to the Incirrata and defined both groups.

On the other hand, I do not admit that the habitus of Palaeoctopus is
sufficient to place it in the Octopodidae because of the presence of muscular,
well-developed fins, and I do not agree with the names of the two subgroups.

I stressed in the introduction (p. 14) and in a previous work (Naef, 1919,
p. 22) that it is not permissible to determine the position in a certain group
by the presence or absence of certain characters, i, e. by a rigid diagnosis.
The following example will illustrate my views. Reinhardt and Prosch
(1844) divided the Octopoda into Apteri and Pteroti in order to place a rare
type (Cirroteuthis) and stress the contrast between this form and the
other genera of Octopoda. However, Octopoda with finlike structures (e, g.
Pinnoctopus d’Orb., 1845) but otherwise resembling the forms without
fins have since been described. The fins can thus not be used as a
diagnostic character. Lutken (1882) and Hoyle (1886) therefore replaced
the division into Apteri — Pteroti by the Trachyglossa and Lioglossa,
according to the presence or absence of a toothed radula. New difficulties
appeared, however, with the discovery of species (Vampyroteuthis)
which closely resemble Cirroteuthis (i. e. the Lioglossa) but have a
well-developed radula. Thiele (1914), who occasionally discussed the
"phylogenetic method" (Thiele, 1910 and 1913, Zool. Anz., Vols. 35 and 51;
see also Naef, 1912, Zool. Anz., Vol. 50, p. 336), placed Vampyro-
teuthis in the Trachyglossa as a consistent diagnostician. Grimpe (1916,
670 p. 353) rejected this view with good reasons, but he repeated (loc. cit.) the
mistake of Reinhardt, Prosch, Lutken and Hoyle: he considered the presence
or absence of cirri on the arms as a new character and maintained the
terminology of his predecessors (Cirrata — Incirrata). The same happened

* These formations apparently persist on branched skin warts in the growing animal, perhaps in their original
number (p, 705). Compare, for example, Octopus arborescens Hoyle, 1904.

** This condition is very variable physiologically. Preservation causes a retraction of the eyeball, also in
adults. Only careful cocainization can prevent the retraction of the eye (Figure 399 on p. 676 and
Figure 402 on p. 679). The natural condition is probably caused by accumulation of blood in the
ophthalmic sinus (Si in Figure 386 on p. 659),

646

again: Berry (1913, 1914, Figure on p. 275) described Laetmoteuthi s,
which has no cirri but otherwise resembles Cirroteuthis. The cirri
may be lost in badly preserved specimens, but this is not certain and does
not alter the fact that such determinations are wrong in principle,
especially when they are based on a single character, A specialized form
may lose the cirri but it should not be removed from the group to which it
belongs. It is particularly wrong to name such systematic units. If
Reinhardt and Prosch had used an indifferent name, it could have been
maintained. All three formulations are based on a morphological element
the presence of which is typical for the preceding systematic stages, i. e.
for the Dibranchiata and Octopoda in general (see also p. 19). The presence
of the character in question does not prove a closer relationship and neither
does its absence. Systematic groups have to be based on a type, i. e. from
a center which may be either an ideal type or an ancestral form. However,
uncertainty can be avoided in practice by choosing an existing species or
a small group (if possible by using the "morphological instinct," p. 18) as
the type of a newly established systematic category ("nominal type" in the
sense of Naef, 1919, p. 21; 1921, p. 534), as a certain specimen serves as
the type of a species. The name of these forms may be used for the whole
group (e. g. Sepia — Sepioidea, Teuthis==!= — Teuthoidea), as is usually
done in the case of families (Sepiola — Sepiolidae). I therefore divide
the recent Octopoda into Cirroteuthoidea (after Cirroteuthis Eschricht) and
Polypodoidea (after Polypos, Polypodes Aristotle ^ O c t o p u s , Octopo-
didae auct.).

It is doubtful whether P al a e o c t o p u s newboldi can be placed into
these two groups, as its position will remain uncertain in any case. Unless
the evidence determined otherwise, I have not placed fossil Decapoda in
well characterized recent groups, but kept them separate. In one doubtful case
(p. 145), I placed Pa 1 a e o lo li g o in the Mesoteuthoidea, i.e. in a fossil group
considered as a preceding stage, and not in the Metateuthoidea, to which this
transitional form is perhaps equally closely related. In any case, I could
671 not have placed Palaeololigo in the Myopsida or in the Oegopsida. Nor
canPalaeoctopus be placed in any of the two recent groups of Octopoda,
but it is a type which is more closely related to the ancestral form of all
Octopoda than the recent representatives of the order. This view is based
in part on the subterminal position of the fins which is typical for all
Dibranchiata (young Decapoda). The position of the rudimentary shell and
the course of the vessels show that the position on which remnants of the
base of the fins in the adult should be present is dorsal and not terminal
(Sh in Figure 392 on p. 664), also in species in which fins are absent. The
group of recent Octopoda stands thus in contrast to P ala e o c t o pu s . 'i''’'

If we wanted to place the genus in the recent groups, its place would be in
the Cirroteuthoidea because of the fins. However, the typical form of the
ink sac of Palaeoctopus differs from that of the recent fin- bearing
Octopoda in which this was studied. The presence or absence of cirri on
the arms of the fossil can obviously not be determined, although they were
probably present in ancient Octopoda. Palaeoctopus resembles the
Octopodidae in habitus but differs from them in the arrangement of the

* Cf. p. 208.

In other words, the "morphological position" of the fins has been displaced dorsally in the recent Octopoda

(see p. 94).

647

suckers, which are biserial in the Octopodidae. The uniserial arrangement
in the El ed one -like forms (q.v.) is undoubtedly secondary, as the
structure of the musculature shows.

FIGURE 398. Palaeoctopus newboldi H. Woodward
1896. 0.5 X. Reconstruction by L. Dollo, 1912 (p. 126).
From the Upper Cretaceous (Senonian) of Syria. The eyes
are drawn in unnaturally. Compare with Figure 397 on
p. 668 to obtain a better picture. The habitus resembles
that of a young O c t o pu s macro pus. The mantle
sac is probably deformed and more slender in life. Cf. also
1922, Fossile Tintenfische, Figure on p. 286.

Jatta stressed the systematic importance of the funnel gland (1896,
p. 22). This is particularly important because of the scarcity of definitely
672 formed structures in the soft body of the Octopoda. Comparison of the

known forms shows the following: 1. The basic form of the organ (p. 102)
can be expressed diagrammatically as M -shaped, e. g. in Argonauta
and Ocythoe. 2. This organ becomes W- shaped in many Octopoda
(A m p h i t r e t u s, O. vulgaris, macropus, defilippii, saluzzii,
(Scaeurgus) unicirrus (cf. p. 714), patagiatus, Alloposus
mollis, Tremoctopus). 3. VV -shaped organs are present in
O. (Scaeurgus) tetracirrus, partly also in unicirrus, and in
O.hoylei, c a 1 i f o r n i c u s, valdiviae, Opi s thot euthi s (Chun,
1915, p. 537). 4. The Bolitaenidae apparently have only the A-shaped
middle part. 5. Only 2 small, rounded pads (a) are present in the
Vampyroteuthidae. Cirroteuthidae (?).

CLASSIFICATION OF THE OCTOPODA

I. Fossil forms, which cannot be placed in suborders II or III and
occupy an intermediate position, like Palaeoctopus Woodward.

648

Suborder I: Palaeoctopoda Naef, 1921

Fins terminal, suckers uniserial, ink sac well developed, habitus
Octopus-like Family 1. Palaeoctopodidae Dollo, 1912

To this family belongs only "C ala i s" newboldi de Sowerby,

1846 =P al a e o c t o p u s newboldi (de Sow.) Woodward, 1896. Cf.
Figure 398 on p. 671 and Naef, 1922, Fossile Tintenfische, pp. 285 — 286,
Figure 97.

II. Pelagic and abyssal Octopoda, with usually wing- shaped, muscular
fins. Arms with 2 rows of cirri on the inner surface, which accompany
the uniserial suckers and alternate with them. Shell rudiment present,
single. Arms connected by a large interbrachial membrane to the apex.

Suborder II: Cirroteuthoidea Naef, 1921

A. Mantle slit wide. Funnel and radula well developed

Family 2: Vampyroteuthidae Thiele, 1914

V a m py r o t e u t h i s Chun, 1913, M e 1 an o t e u t h i s Joubin, 1912,

La et m ot euthi s Berry, 1913, Hy m e n ot e u th i s Thiele, 1916.

B. Mantle slit markedly narrowed. Radula rudimentary or absent.
Funnel small or rudimentary.

a) Head and body distinctly delimited from arm crown, though often

very short Family 3: Cirroteuthidae Keferstein, 1866

Cirroteuthis Eschricht, 1836, Cirrothauma Chun, 1911,

St au rot euthi s Verrill, 1879, Froekenia Hoyle, 1904, Chunio-
teuthis Grimpe, 1915, Cirroctopus n.g. (p. 675).

b) Head and body connected with the disklike arm crown as a central

elevation and not distinctly delimited from the arms

Family 4: Opisthoteuthidae Verrill, 1883

Opi s thot e uthi s Verrill, 1883.

673 III. Pelagic and benthic Octopoda without true fins, partly with a more
or less projecting ridge or fold of skin ("lateral line") which borders the
sides of the mantle. Arms with 1—2 rows of suckers. Cirri absent. Inter-
brachial membrane usually leaving greater part of arms free. Shell
rudiment consisting of 2 separate "cartilaginous rods" in the mantle
musculature, or absent.

Suborder III: Polypodoidea Naef, 1921

A. Pelagic Octopoda with gelatinous body and uniserial suckers. Eyes
small, widely separated, very prominent, at least in the young forms.

All 5 median rows of radula with multicuspid teeth. Series Ctenoglossa
Naef, 1921.

a) Ventral mantle margin broadly fused with funnel. Lateral margin

free. Interbrachial membrane large, umbrella-like. Eyes telescopic.
Habitus medusa-like Family 5: Amphitretidae Hoyle, 1886

Amphitretus pelagicus Hoyle, 1885; Cf. Ijima and Ikeda, 1902.

b) Ventral mantle margin free. Arms short. Eyes widely separated,
with long optic nerve. Olfactory tubercle stalked. Third arms the longest.

649

Hectocotylization consisting of enlargennent of all or only the distal suckers
on the right 3rd arna Family 6: Bolitaenidae Chun, 1911

Bolitaena (Steenstrup, 1859; Hoyle, 1886), Chun, 1904; Eledonella
Verrill, 1884; probably also Vitreledonella Joubin, 1900, which has
a habitus resembling that of Eledone.

B. Littoral or pelagic Cephalopoda of Octopus type. Pelagic forms
always with 2 alternating rows of suckers on the arms, most littoral
forms also. Eyes large, situated close together, moderately prominent
in early youth. Cartilaginous rods in mantle present. Multicuspid teeth
present at most in median row of radula. Series Heteroglossa Naef, 1921.

a) Littoral Octopoda. Male resembling female and of about the same

size. Hectocotylized arms ventrally with a longitudinal groove and an
about spoon- shaped, modified apex, otherwise normal. Adult male with
enlarged suckers on all arms Family 7; Octopodidae d’Orb., 1835

See Chapter 50.

b) Pelagic Octopoda. Male much smaller than female and often
markedly different. Hectocotylized arms abnormal on the whole, coiled
inside a pocket before use, lost in copulation and later regenerated.

True "hectocotylus." Cartilaginous rods in mantle not recognizable in

the adult, traces of rods present in the embryo

Family 4; Argonautidae Naef, 1912

See Chapter 53.

674 The suborder Cirroteuthoidea will not be described in detail as it

does not occur in the Mediterranean. The Cirroteuthoidea do not contain
any primary forms among the recent Octopoda, at least not in external
characters. However, they may be considered as predecessors of the
Polypodoidea because they show typical characters of the Octopoda;
cirri on the arms, a single shell rudiment and well developed fins.* On
the other hand, they show secondary adaptations which should not be
assumed for the other groups, i. e. the extremely well developed inter-
brachial membrane, the gelatinous body, fins of characteristic form and
a reduced funnel gland. All these special characters are connected with
the planktonic -pelagic** and abyssal mode of life of the group. The same
applies to the absence of the ink sac, which is apparently a general
character of the suborder.

* These are general characters of the Dibranchiata (p. 19).

*■* Some forms of this group are markedly like medusae, for example, Cirrothauma Chun, 1911 or

O pisthote uthis medusoides Chun, 1915. The movement by repulsion and the radial arrangement
of the arms suggest such adaptations.

650

675 Chapter 49

SUBORDER POLYPODOIDEA

Naef, 1921

a. DIAGNOSIS

True fins absent; delicate ridges of skin or fin margins sometimes
present on sides of mantle. Arms with 1—2 rows of suckers which are
never accompanied by cirri. Rudimentary shell consisting of 2
"cartilaginous rods" situated obliquely in muscular mantle on both sides
of median dorsal line but not in median contact, or rods absent.

b. TYPICAL STRUCTURE OF THE POLYPODOIDEA

The ancestral form of the Polypodoidea, which we name Protopoly-
pus, differs from the general ancestral form of the Octopoda in the
absence of fins. This is a simplification of the primary structure which
can be explained ecologically by a reduction of swimming movements.

Fins must be assumed to have been present (p. 656); they are present in
the embryo in the typical position (Vol. II, Plates XXV, XXXVII) and show
a normal correlation to the mantle sac, but they later disappear without
trace. Whether they persist or reappear in the form of "lateral lines" is
not clear. At any rate, the ridges which form the lateral lines are not
equivalent to fins^'= as they do not show the typical muscular structure or
the relationship to the rudimentary shell, although they may be widened into
margins and even make undulating movements (cf. Figures 425 and 431).

The rudimentary shell of the adult consists only of 2 "cartilaginous rods"
(Figure 392 on p. 664) situated obliquely on each side in the posterior
676 dorsal part of the muscular mantle but not in median contact. (The primary
connection was probably situated nearer to the posterior end.) The
epithelial tubes in which the rods are situated develop during the embryonic
stage as lateral evaginations of the shell sac and replace it at least partly.
As Figure 392 shows, the cartilaginous rods serve mainly as points of
attachment for the retractors of funnel and cephalopodium, that is the

* The recently described Stauroteuthis mawsoni Berry 1917 (Austr. p. 8 — 9) suggests, however,
that the lateral lines are reactivated rudiments of fins. This is a young form of Cirroteuthoidea with
large, well developed fins which occupy about the posterior half of the mantle sac but continue as large
folds toward the eye and the posterior end, where they are united. This animal should be considered as
the type of a new genus which I name Cirroctopus because it resembles the normal Octopodidae
although it undoubtedly belongs to the Cirroteuthoidea.

651

"posterior mantle adductors" which develop from the posterior part of the
retractors of the cephalopodium in the Octopoda (p. 664). Each cartilaginous
rod is a rounded, spindle-shaped body which contains isolated cells in its
mass (Appellof, 1899), like the cartilaginous deposits of other shells of
Cephalopoda; they have a chitinlike consistency (cf. Jatta, 1896, Plate 25).
The rods are usually slightly bent at the point of attachment of the muscles.

FIGURE 399. Freshly hatched E 1 e d on e moschata,
5x. The habitus is typical for young Polypodoidea,
but the eyes are retracted by preservation (p. 699) and
the arms are markedly shortened. The interbrachial
membrane is still weakly developed. Compare also
the mantle situs of such a stage (Plate X, Figure 1).

The arms are as assumed for Protoctopus (p. 660). However, a
typical element is absent: the cirri which accompany the suckers have
disappeared. This is probably connected with a change of food which makes
these fine tactile organs unnecessary. Typical Polypodoidea feed on larger
prey (Crustacea) than the plankton-feeding Cirroteuthoidea. The inter-
brachial membrane is also slightly less developed since it is not used as a
net for small fry but for the capture (poisoning) of larger prey. The
suckers are well developed and show the typical biserial, zigzag arrange-
ment which is simplified secondarily in some forms (E led one).

The funnel, mantle cavity and jaws of the Polypodoidea resemble those
of Protoctopus (pp. 662, 665). On the other hand, the radula shows a
number of special characters: at least the teeth of the median row have
accessory cusps, and the successive teeth differ in the arrangement of the
677 cusps, as recorded by Steenstrup (1859, MS; cf. Hoyle, 1886, Catalogue,
p. 21 ) for "Bolitaena microcotyla,"a species of Ctenoglossa. The
arrangement of the accessory cusps varies in the longitudinal row so that
about each 5th tooth resembles the first (Plate XVI, Figures 4, 5). This
apparently improves the efficiency of the radula.

The mode of life and diet probably resembled that of the recent species
of Octopus, i. e. a reduction of the swimming movements in comparison
with the ancestral form and the Cirroteuthoidea. The distribution of the

652

species is therefore due to the pelagic -nektonic juvenile forms. The sexual
dimorphism is of the general type of the Octopoda (p. 667). However,
comparison of Amphitretus (after Sasaki, 1917) with the Octopodidae
shows some special characters of the hectocotylus. The right LV-arm
bears a copulatory apparatus near the apex which resembles that of the
Sepiolinae (p. 585). It is situated transversely on the inner surface of the
arm, like the spout of a pot, with the apex distally. The terminal part of
the arm bears diverging, reduced suckers which indicate a spoon shape,
as in the Sepiolinae. The structural resemblance is obviously associated
with a similar function. As in the Sepiolidae, the spermatophores are
placed in the mantle cavity of the female. However, while the Sepiolidae
have a bursa copulatrix (p. 5 80) for the attachment of the spermatophores,
the spermatophores of the Polypodoidea (Cirroteuthoidea?) are placed directly
in the genital opening so that internal fertilization is possible, although this
does not always take place before oviposition (see Vol. II).

c. POSTEMBRYONIC DEVELOPMENT

The benthic character is distinct in the structure of Polypodoidea, but
the contrast between the adult animal and the planktonic young stages is
more marked and causes a kind of metamorphosis which cannot be assumed
for the general type of Octopoda. We can speak here of larvae as in the
Oegopsida and Loliginidae. Figure 397 on p. 66 8 and Figure 409 on p. 6 87
show the typical habitus of these larvae (but see p. 107).

The Polypodoidea have relatively small eggs (1—2 mm) which hatch in
a correspondingly undeveloped state. On the other hand, some species of
El e done and Octopus have large eggs (6 — 9 mm) with a shortened
metamorphosis in the egg. A freshly hatched animal of these species
already shows the habitus of the adult. This is apparently the rule in
El e done. The typical larval form does not differ much from the normal
form of Octopoda since neither mantle sac nor head show distinctive
characters. At any rate, the mantle sac never lacks the armature of spines
or bristles of the young Octopoda (p. 6 88), while the undeveloped lid gives
the head its special appearance (p. 666). More striking is the inhibited
condition of the arms: there are 8 short, uniform arms, with 3 well
developed larval suckers and a whiplike terminal part without suckers
678 (Plate IX, Figure 2).^'= The terminal part later develops further suckers,
after a distinct interval and not as a continuation of the first row of
"anlagen. " The apex grows further, like a vegetative center (cf. Oegopsida,
p. 234). The subsequent suckers are formed in a single line, also in the
biserial forms (Plate IX, Figure 6) so that a large part often remains
uniserial until it assumes a zigzag pattern, as in the Decapoda (Figure 245
on p. 462; Vol. II, Plate XXl).

* These stumpy larval arms, the whiplike ends of which are probably particularly sensitive, can be extended
suddenly for a short distance by contraction of transverse and circular fibers. 1 observed this in laboratory-
bred young Octopus vulgaris after offering them the much smaller larvae ofArgonauta as prey.
The arms thus behave like the tentacles of Decapoda (p. 119): the fingerlike ends of the arms at first and
then the suckers fasten onto the prey, which is then eaten. The small Argonauta do the same with
still smaller planktonic Crustacea.

653

The eyes become at first increasingly prominent and may temporarily
appear stalked, as in many Oegopsida (Figure 402). They are later again
retracted although some adult Polypodoidea have markedly bulging eyes
(Figure 413 on p. 699 and Figure 420 on p. 708). The musculature of the
dying animal is always contracted spasmodically, and preserved specimens
therefore almost never show a natural picture.

FIGURE 400. a) O. vulgaris, obtained from eggs;
freshly hatched larva (21 June 1912). c) Young
stage found on 19 June 1913 in the plankton and
closely resembling a), b) Slightly older young
stage from the plankton, with retracted head and
funnel and with contracted primary lid the margin
of which appears like a papilla. This specimen
might also belong to another species (O. s a 1 u z z i i ?).
Note the distribution of chromatophores (those
situated deep in the dorsal side are not visible;
see Plate X, Figure 6). Note also form of mantle
sac, mantle margin, position of olfactory papilla
with respect to the mantle margin, funnel and
funnel pockets, eye and lid opening. Each arm
bears 3 suckers and ends in a whiplike apex.

FIGURE 401. Advanced stage, probably ready to
begin its benthic life, a) Ventral view, b) Part
of dorsal surface, 44 x. c) A single spine, 200 x.
The species cannot be identified. The habitus is
typical for the Octopoda, the only difference being
in the short larval arms. The distal two of the
three larval suckers are recognizable on most arms.
Some "anlagen" are arranged in a zigzag (cf.
Figure 409 on p. 687). The spine has a fine
longitudinal striation which corresponds to its
splitting into bristles (this may take place later
or is suppressed here).

679 The olfactory organ forms a flat papilla on the outer corner of the
mantle slit. It is later retracted into the slit and covered with skin
(Plate IX, Figures 2, 3; Figure 409, on p. 687).

There are at first only a few chromatophores as in the youngest forms
of the Teuthoidea (Plate IX). They are more numerous on the head and
arms than on the mantle sac (Plate IX) and gradually become more
numerous toward the metamorphosis. The chromatophores are light
(yellow to orange) or dark (reddish brown to dark brown).

The occurrence or persistence of such nektonic -planktonic young forms by
inhibition of the juvenile characters beyond the larval period is apparently
realized in the Ctenoglossa and makes it possible to define the typical
character of the group (see p. 680).

654

FIGURE 402. Young Octopus vulgaris, drawn alive,
3x, in typical position, waiting for prey. Note the
spirally coiled arms, the extended protective membrane,
form of head (caused by the protruding eyes), asym-
metrical position of funnel, distribution of "main"
chromatophores on mantle, head and arms. 5 warts on
mantle sac and the wart (dotted) situated above each eye.
Such animals burrow in the sand and between small
stones, like adults between larger stones.* **

d. VARIATION OF THE POLYPODOIDEA

The Polypodoidea consist of 2 sharply defined groups (p. 673). The first
group is not represented in the Mediterranean, but it will be briefly defined
for a better understanding of the second group.

1. CTENOGLOSSA

Diagnosis. Planktonic -nektonic Polypodoidea. Suckers on arms uniserial
or in a zigzag pattern distally. Skin and partly also the musculature
gelatinous. All 5 median longitudinal rows of radula with comblike multi-
cuspid teeth. Hectocotylus without longitudinal groove (for transfer of
spermatophores) and without longitudinal canal derived from this groove.

680 The Ctenoglossa are smaller, strictly planktonic forms. Their pelagic
habits are evidently a secondary development (absence of fins). Many
characters are distinctly nektonic and probably developed from the
Polypodoidea by suppression of the transition to benthic life (p. 679).

This group consists of the Amphitretidae and Bolitaenidae, which I
described in 1912 (Zool. Anz., Vol. 40, p. 196). The relationship between
these two families is expressed mainly in the radula (cf. Thiele in Chun,
1915, pp. 492, 493, 532). Type genus: Amphitretus,

* If they are kept on sand in the aquarium, they often hide completely deep under the surface of the sand.

** A number of medusa-like forms belong to this group (see also p. 674). The most striking isAmphitretus
pelagic us (cf. Ijima and Ikeda, 1902), a species with thick, gelatinous, swollen skin. On the other hand,
Vitreledonella Joubin has the appearance of a transparent E led one, but with very small eyes and
long arms, weakly developed membrane and markedly retarded LV arms. It is perhaps only an atypical
species of Heteroglossa.

655

2. HETEROGLOSSA*

Diagnosis. Suckers usually biserial; if they are secondarily uniserial,
there are about 100 or more on each arm in the adult. Radula with multi-
cuspid teeth only in the median row; the teeth of two adjacent rows never
have more than 2 cusps. One 3rd arm markedly hectocotylized in the
male, at least at the end.

In contrast to the uniform habitus of Ctenoglossa, this group contains
forms of very different habitus (Argonaut a, ^ Tremoctopus,

El e done, etc.), but Oc t o p u s represents the type of all these forms.

Some Argonautidae (Alloposus, Ocythoe; cf*. Figure 448) do not
differ markedly from Octopus. Also the most extreme forms of
hectocotylus show a uniform basic type which can be identified with the
normal form in the Octopodidae. It is characterized by a longitudinal
groove on one of the 3rd arms of the male, and a modification of the distal
part in its continuation. The Octopodidae can be considered as the preceding
systematic stage of the Argonautidae in general habitus and in characters
of the hectocotylus, i. e. the type of Argonautidae developed from that of
the Octopodidae as a central group, and can be identified in most parts
with the actual conditions observed in species of Octopus (e. g.

O. vulgaris). The ancestral form of an Argonauta argo can be
defined thus.

* The names of the 2 groups are based on a single character (although this is not advisable in general,
p. 670). However, the names are so indifferent that they could be used for any form of radula.

656

681 Chapter 50

FAMILY OCTOPODIDAE
d'Orbigny^ 1835

Contents: a. Diagnosis. — b. Typical structure of the adult animal. — c. Typical development (p. 686). —
d. Variation of the Octopodidae (p. 688).

a. DIAGNOSIS

Suckers uniserial or biserial. Shell rudiment always present in mantle
in the form of "cartilaginous rods." Sexes almost identical, except for the
hectocotylus. One (rarely both) 3rd arms hectocotylized, i. e. shorter than
the opposite arm, with a more or less distinct spoon without suckers at
the apex and a spout- shaped copulatory apparatus on the inner side before
the apex; interbrachial membrane on ventral side of hectocotylus with a
gliding groove for the spermatophores at the margin which leads to the
apparatus.

b. TYPICAL STRUCTURE OF THE ADULT ANIMAL

The typical habitus of the family is represented by Octopus vulgaris
except for its size.

The Octopodidae are strictly benthic forms, and differ markedly from the
pelagic Polypodoidea (Ctenoglossa), especially when resting or crawling
(Figure 403). The body is sloping, the head much higher than the posterior
end of the mantle. The arms and the anterior part of the head are bent from
the base at almost a right angle to the longitudinal axis and the arms are
spread radially or coiled into a spiral (Figure 402 on p. 679). The mouth is
also directed downward. In this normal position, the funnel must be
displaced laterally, from the left to the right or vice versa. The respiratory
movements can then be easily observed (p. 682).

All Octopodidae have "cartilaginous rods" (Figure 392 on p. 664) in the
mantle and differ in this respect from the Argonautidae and perhaps also
from the Ctenoglossa, in which such rudiments of the shell have not been
found.

682 The mantle sac has the general form which varies markedly physiologi-
cally. (Figures 403 and 404 show the normal habitus of the living animal.)
The width of the mantle slit and the form of the free mantle margin also
depend entirely on the state of contraction. The mantle slit occupies

657

normally about the ventral side of the body, and the fusion with the head
about half of the dorsal side.

Olfactory organ and head are typical in form and position. The lid
apparatus is always complete, so that the dorsal and ventral margin of the
primary lid cannot open directly (Figure 387 on p. 660). The ventral margin
has been markedly displaced dorsally during postembryonic development
(p. 659); it covers the pupil and forms a "pseudocornea. " If a probe is
inserted into the pocket formed below the upper lid, it always reaches into
the eye chamber, which is thus only apparently closed. The secondary lid
is swollen and only its ventral margin is free and folded, as in the
Sepiolidae (p. 567). The opening is therefore dorsally displaced during
complete contraction.

FIGURE 403. Live Octopus (Scaeur-
gus) tetracirrus (cf. p. 681), which
shows the characteristic posture of the
Octopodidae: the bending of the arms
downward, the position of trunk and
funnel, the spiral curvature of the arms
(the 1st arm in the figure on the left is
regenerated, the 3rd arm is hecto-
cotylized). The eye shows the second-
ary lid, upper lid, pseudocornea and
pupil. Compare with Figure 402 on
p. 679.

FIGURE 404. Same animal, alive. 0.5x. Movement of mantle and funnel, a) Inhalation; b) exhalation.
Note change of mantle margin, funnel and funnel pockets, particularly of the funnel opening. Note the lid
apparatus and pupil, which is slightly more closed in a) and more open in b).

All arms bear biserial suckers and are connected proximally by a well
developed interbrachial membrane which also extends for some distance
683 along the outer edge toward the apex. Funnel and mantle cavity are typical
for the Octopoda. Luminous organs are absent.

The radula shows characters which distinctly resemble those in the
Ctenoglossa. This applies particularly to the form of the teeth in the

658

median row: they are mnlticuspid, with 2 — 3 accessory cusps lateral to the
main point, which is constant and uniformly developed. The accessory
cusps are markedly asymmetrical and never the same in successive teeth
of the same row; an identical or similar tooth only recurs after a number
of teeth. This seems at first completely irregular, but closer examination
shows that there is a pattern.

Hectocotylus of Octopodidae (see Figure 411).

FIGURE 405. a — d. End part of the hectocotylus of
the species in Naples; a) O. s a lu z z i i ; b) O.
macropus; c)0. vulgaris; d)0. defilippii.

1 — longitudinal groove; 2 — copulatory apparatus;

3 — spoon (lateral margin); 4 — transverse grooves
of spoon; 5 — longitudinal ridge on spoon; 6 —
rudimentary spoon.

FIGURE 406. Beginning of conducting groove of
O. vulgaris, a) Outer view; b) inner view.
The membrane is impressed longitudinally and
slightly obliquely, so that the beginning of the
groove passes from the outer to the inner side.

A number of papillae, probably sensory, form a
serrated profile on the inner margin. The papillae
apparently form the transition from the funnel or
from the genital process ("penis") which passes
through it.

However, this pattern is complicated and of no further interest at this
point (see Plate XVI, Figures 4, 5). The two adjacent rows contain bicuspid
teeth, of which those of the paramedian row are very small. The teeth of
the following rows are stronger and have laterally projecting basal plates
which (in contrast to the Ctenoglossa) do not have further cusps. The jaws
show no special characters.

The sexual dimorphism is small in the live animal, as the male often
hides the hectocotylus, which often resembles an injured arm (Figure 403
on p. 692). The right 3rd arm is usually hectocotylized, abnormally also
684 the left 3rd arm. The hectocotylus is distinctly shorter than the

corresponding normal arm. It has a slightly widened, spoon- shaped end
part without suckers. There is a light groove along the ventral edge which
is formed by the margin of the interbrachial membrane. It is transversely
striated, and may form a canal in the live animal, but it often disappears
in preserved specimens. The spermatophores are passed along the groove

659

FIGURE 407. Situs of mantle cavity of a female of O. vulgaris during oviposition. 0.5X. The mantle is
cut and spread. Note the general characters in comparison with Figure 392 on p. 664 and Figure 408 on p. 685.
The filled ovary (Ov) inflates the abdominal complex to some extent, but not as much as in Figure 408. The
oviduct glands (Ei) are enlarged. The spreading of the preparation shows the topographical relations of the
visceral nerves (Vn), pericardial funnels (Pt), renal papillae (Np) and venous appendages (Va).

Kr — retractor of gills; Ns — posterior border of a renal sac; Vb — branchial vein; Kh — branchial heart;

Km — branchial gland; Kb — ligament of gill; St — stellate ganglion; Ma — anterior mantle adductor;

Tr — funnel retractor; Gp — genital papilla; Ad — median mantle adductor; Tt — funnel pocket. Note that
the free "genital process" has not disappeared, but persists in the part of the gonoduct distal to the oviduct
glands; however, it is attached to the body (cf. Figure 423 on p. 712 and Figu^res 427 — 429 on p. 719).

to the end part. The groove begins proximally at the retracted margin of the
the membrane between the 3rd and 4th arms (Figures 405, 406 on p. 683).
There is a characteristic entrance at this point which accepts the sperma-
tophores from the funnel or genital process during copulation. The margin
of the membrane is slightly interrupted at this point; the part extending
from the 4th arm diverges to the inner side of the membrane, while the
continuation begins slightly outward and is rapidly closed into a groove.
Before the indentation, the free part of the margin bears papillae or
denticles probably of sensory character, which evidently function in the
conduction contact with the funnel or the genital process (Figure 428).

The exit of the groove in the end part also has a special structure: the
groove turns to the inner side of the arm and reaches the transverse

660

copulatory apparatus which resembles the spout of a milk pitcher. Only
685 this spout is probably introduced into the genital opening of the female,
while the spoon clasps the papilla which bears the opening.

FIGURE 408. Situs of mantle cavity of O c t o pu s vulgaris. 0.5 x.

Left: mature female before oviposition, with extremely enlarged ovary displacing the other organs. The
organs are drawn in their natural position as far as possible. However, the mantle sac is markedly shrunken,
so that the whole abdominal complex is pressed toward the exit. The mantle sac should be much wider (cf.
Figure 407). Te — funnel corner; Od — oviduct; Np— renal papilla; Va — venous appendage; Kr — gill
retractor; Ns — boundary of renal sac (pressed anteriorly); Ov — ovary; Mh — posterior margin of mantle cavity.

Right; Almost mature male, which at any rate already attempted copulation. (The genitalia appear relatively
small, cf. Figure 423.) 1 — visceral nerve; 2 — "penis diverticulum"; 3 — renal sac; 4 — position of testis;

5 — gill retractor; 6 — branchial ganglion; 7 — branchial gland; 8 — anterior mantle adductor; 9 — "penis,"
i. e. end part of genital process (this part forms only a papilla).

The opened mantle cavity of the female shows a number of important
conditions which vary markedly according to the physiological condition.

In comparison with the juvenile conditions (Figure 392 on p. 664), the
genital papillae of the mature animal are displaced toward the anus, often
to its level (Figures 40 8 and 407). The oviduct glands are often visible

* Compare p. 677. The longitudinal groove is characteristic for the Heteroglossa, but homologues of the

terminal formations of the hectocotylus are present also in the Ctenoglossa, e. g. in A m ph i t r e t u s (Sasaki,
1917, Figure on p. 363). The spoonlike terminal part of the Octopodidae is represented here by the whip-
like apex with a flat inner side bearing a row of papillae on each side which are probably reduced suckers.
Proximal to them is the part corresponding to the spout, which does not continue into a groove. A groove
could not function here since the membrane extends almost to this point.

661

under the skin before the renal papillae, but they may also be situated much
deeper. The glands can be recognized by their rounded form and radial
686 structure; they are situated near the distal oviduct, which is distinct under
the skin. The ovary is extremely large and contains about 100,000 eggs;
its size disturbs the topography of the mantle cavity (Figure 408). A genital
duct is present in the male only on the left; it ends in a more or less long,
free genital process (Figure 408 on p. 685 and Figures 427 — 429 on p. 729).
The duct bears a caecum which is directed inward and posteriorly ("penis
diverticulum" of Marchand, 1907) (Figure 412 on p. 712 and Figure 432
on p. 722).

The coloration of the dark chromatophores in the Octopodidae varies
between light orange red, yellowish brown, reddish brown and dark, almost
blackish brown or grayish black and permits varying tones and patterns in
combination with the yellow and yellowish brown chromatophores. The
patterns are accentuated by the variability of the skin, which forms wartlike
elevations oh the whole body but especially on the dorsal side. Some of the
warts are of complex form and branched. Some of these warts and "cirri"
are more or less constant, especially those above the eye (Figure 403 on
p. 682), which are always symmetrical.

c. TYPICAL DEVELOPMENT

Most Octopodidae pass through the typical larval development (p. 677)
which includes the stages shown in Figures 400, 401 and 409. Stages older
than that shown in Figure 401 do not occur in the plankton — they sink to the
bottom and begin a metamorphosis, but its stages could not be found in
Naples. The animals probably bury into the sand or detritus, like the older
stages (p. 679), and only appear later as small forms of Octopus
(Figure 410 on p. 6 88) with numerous suckers on all arms. They differ
from adults only in minor details.

Many Octopodidae (mainly species of Octopus and El e done) hatch
at this stage and are already quite large (Figure 399 on p. 676). This
development is associated with the size of the egg. Normal eggs are about
2 mm long, forms which direct development have eggs 6 mm long or more.

In O. digueti, in which care for the young is highly developed (cf.
Argonaut a), the eggs are 3X9 mm. The bristles or spines described
on p. 668 are present in all larvae. As I observed in Octopus vulgaris,
these structures have a function in hatching. The animal hatches with the
posterior end first, as is usual in the Dibranchiata, through a relatively
narrow opening at the thick end of the ovoid capsule where the spermatozoon
687 entered, i. e. at the micropyle. The animal must squeeze through this small
opening, with the posterior end first, using the spines as barbs. If the
embryo is situated wrongly in the egg, i. e. with the mouth toward the
opening, it cannot hatch and dies, unless assisted. It cannot turn around
in the capsule and the spines, which are directed outward, push against the
edge of the opening. I have no explanation for the role of the spines in
pelagic larvae. However, they certainly increase friction and help to keep
the animal afloat as its specific weight is greater than that of sea water.

The spines do not impede movement as they are directed anteriorly, while
the rapidly swimming animal moves backward. The stellate tufts (Plate IX,

662

Figure 1) do not always split but they often form brushes or later blunt
tubercles (Figures 678 and 705). The splitting of the spines into tufts of
bristles may be connected with the transition to benthic life: particles of
sand, detritus, etc. adhere to them and make the animal unrecognizable
to enemies. The small glands which produce the spines later probably form
the basis for the formation of skin warts. At any rate, the last visible
remnants of spines or bristles are situated on these transient warts
(Figure 410). The thin cutaneous muscles which form the warts probably
develop in connection with the glandular cups which produced the spines
(cf. p. 705).

FIGURE 409. Larva of O c t o pu s (v u 1 g a r i s ?) from the plankton in Naples, October 1913. lOx. Note the
increase in size and the further development of all parts in comparison with Figure 400 on p. 678. The arms
have apparently grown particularly; they show the beginning of a zigzag pattern of the distal suckers (5). as
the magnified (26 x) distal part of an arm shows (d).

1 — primary lid, surrounding the pupil; 2 — olfactory organ; 3 — funnel pocket (its posterior opening is visible
because of the shrunken mantle); 4 — larval sucker; 6 — growing terminal flagellum.

I observed the development of the hectocotylus only in O. vulgaris
which does not show the typical formations distinctly. The longitudinal
groove is already distinct in stages of only V4 of the definitive length when
688 the distal part has developed and the apex begins to develop. The formation
of suckers ceases and a wider part than on the other arms is formed instead
of the spoon. The surface of the arm further proximally forms a transverse
swelling, the "anlage" of the beak, which continues into the ventral lateral
edge toward the base. Further details could perhaps be found in a species
with a more strongly developed spoon and beak. It would be interesting to
know whether the beak is formed by "anlagen" of suckers, and whether
such "anlagen" could be identified in the spoon (cf. p. 685). This could
clarify the differentiation of this genital apparatus.

d. VARIATION OF THE OCTOPODIDAE

The Octopodidae contain a large number of species, the habitus of which
resembles that of Octopus vulgaris but shows numerous and
diversified differences. The mantle sac may be narrow and slightly

663

pointed or broadly rounded posteriorly, which is usually associated with
the relative width of the head. The arms may have very different propor-
tions: the 1st, 2nd or 3rd pairs may be very strong and long. The average
length of the arms also varies markedly; some species (P. macropus,
defilippii, etc.) have very long arms, other species have very short
arms. It is doubtful whether the uniserial or biserial arrangement of
suckers on the arms is typical. The arrangement of the musculature in
El e done suggests the second. Yet it is doubtful whether the uniserial
forms (El e done, etc.) are closely related. The very variable habitus,
hectocotylus and radula of these species suggest the contrary. On the other
hand, there are important resemblances between uniserial and biserial
species.

The form of the suckers varies markedly but they also differ with age and
689 according to their position on the arm. Figure 3 on Plate XIII shows a
juvenile sucker from the apex of the arm and Figure 2 an older, fully
developed sucker. I suggest therefore not to use the finer differences
for identification. The number of suckers on an arm varies with age; it
is characteristic for the species in the adult, but only in a wide range of
variation, between 70 and 300. Arms of different length bear about the
same number of suckers; longer arms have larger suckers, and shorter
arms smaller suckers.

(688)

FIGURE 410. YoungOctopus vulgaris, drawn after a well-preserved specimen. 4x. (cf. Plate IX,
Figure 3). It was killed in swimming position (more correctly, in the position when sinking to the bottom).

It is of about the same size as the specimen in Figure 402 on p. 679. Note the form of mantle sac, funnel,
head and arms and particularly: funnel pocket (Tt), olfactory organ (Ro, partly covered by the upper part of the
free mantle margin), secondary eyelid (Al, ring-shaped), primary upper lid (Ol), transverse pupil (Pu),
interbrachial membrane (Sh), ends of arms, suckers, deep chromatophores (Ch) of arms (the others are
suppressed), warts (Hh) on dorsal side of mantle, bases of head and arms. Tr — funnel.

The interbrachial membrane varies markedly and extends distally along
the posterior outer edge of the arm, sometimes in the form of a wide
margin. The membrane may be thickened by muscles at the margin and
instead of extending separately on the outer edge of an arm, the margins
may be contiguous on the dorsal side of the arm and form a dorsal margin.

The radula differs little from the type. The most markedly different
form is present in O. valdiviae Chun (1914) and some species of
El e done, in which the teeth are unicuspid. The jaws also show some
special characters, but these cannot be used as systematic characters
since the descriptions (cf. Plate XVIII) are not exact enough; they show
marked variations and postembryonic changes.

664

FIGURE 411. Young female of O ct opu s
saluzzii, natural size. Collected May 1915
at the Amontatura. The form is typical for
a dead, contracted animal with retracted eyes
(p. 678) and sac-shaped mantle (cf. Figure 413);
specific characters are the different length of
the arms and the margin of the interbrachial
membrane (cf. on this the chapter on this species).

which I have not examined in detail
markedly from the type. Vitrei

The funnel gland gives better
characters. There are two types:
the basic W form and two separate
Vs (see p. 672, Jatta, 1896, p. 22;
also Berry, 1917, p. 15).

Other systematic characters are
given by the coloration and sculpture
of the surface. Some species show a
characteristic coloration or pattern
when the chromatophores are extend-
ed, e. g. the orange color of
"Scaeurgus" tetracirrus, the
grayish black of some exotic species,
or some spotted patterns. Only the
specialist can use the varying
brownish tones, but there are distinct
differences between living individuals,
also if they are of similar size and
proportions. No fisherman has ever
confused O. saluzzii with O. vul-
garis as Ferussac and d'Orbigny
did, apparently because they had only
preserved material.

Sculpture and coloration are
variable, but the observed conditions
are characteristic for the species.
There may be completely smooth
forms, but there is usually a
typical distribution of tubercles,
at least temporarily, particularly
an almost constant warty process
("cirrus") above each eye. Other
tubercles are also constant in a
more or less characteristic form in
some species, sometimes only some
of them, or else the skin shows a
permanent pattern which corresponds
to the distribution of the tubercles.
The appearance of the surface of pre-
preserved specimens naturally
depends on the condition of the animal
and on the type of preservation and
these characters have therefore to
be used with care. The body may be
variously deformed in alcohol -
preserved specimens.

Some non- Mediterranean forms
show a general habitus which differs
edonella richardi Joubin

665

(cf. pp. 673, 680) may be mentioned as a bathypelagic aberrant form of
Octopodidae, resembling, but not closely related to the Bolitaenidae.

Another variant of the type of the family is apparently the specimen which
Michael Sars caught at a depth of 100 m at Station 58. The habitus of the
form resembles that of the Cirroteuthoidea='= — head and mantle sac are
abnormally broadly connected and are very short, and the eyes open
dorsally; the mantle slit is markedly narrowed, just large enough for the
large funnel; the arms are short, biserial, with well developed interbrachial
membranes.

Especially important for identification of the species is the hectocotylus.
The terminal part shows an obliteration of the typical differentiation into
spoon and beak and the whole structure is usually small in these cases
(El e done. Figures 427, 433). However, also a small apex of the
hectocotylus may show the normal form (O. defilippii. Figure 405 on
p. 6 83). If the differentiation is typical, the spoon may be long and narrow
or wide and short or it may reach unusual proportions of length and width.
The sculpture of the bottom of the spoon (Figure 405 on p. 6 83) may also
vary, showing transverse ridges and stripes, and a median longitudinal
stripe. The length of the normal, proximal part of the hectocotylus (except
the groove) may also vary. The number of suckers is not quite constant
691 also in the same species, but there are certain norms.'''''' The hectocotylus
ofVelodona is apparently contracted before the apex into a coil and
shows a shortening of the spermatophore groove (Chun, 1914).

The hectocotylus may be developed on the right or left side also in
closely related species. This has been used for the establishment of
"Scaeurgus." However, the species which Berry (1914) described and
illustrated as Octopus hoylei certainly closely resembles the
Mediterranean S c . tetracirrus and Sc. patagiatus of Hawaii,
although its hectocotylus is normally situated on the right side. On the
other hand, the two Mediterranean species of "Scaeurgus" differ
markedly from each other except that both have the hectocotylus on the
left side. Both forms probably form, together with other Octopodidae,
a natural group in which the hectocotylization has become variable.

Sc. tetracirrus shows a special relationship to O. hoylei in the
structure of the interbrachial membrane, the gelatinous consistency, the
sculpture of the skin and the form of hectocotylus and funnel gland. As to
whether Scaeurgus is a valid form, it has to be considered that Appellof
(1 893) found a E 1 e d o n e cirrosa with two symmetrical hectocotyli.

This may be a rare abnormality (cf. p. 654) but such a displacement of
the hectocotylus from one side to the other is probably not an important
change. On the other hand, the differences in the special formation of the
genital arm are far more significant.

Some species show problematic phenomena of hectocotylization also on
the ends of the other arms of the mature male. The suckers may become
transverse, often divided into two halves, and form papillae without suction
chambers (see El e done; Figures 430 and 433). These structures have

* I examined and drew this form in the laboratory of C. Chun, who probably described it (1914?), although
I have not seen a description.

** Such norms can only be established after examination of numerous specimens, since the hectocotylus may
be regenerated after injury, although this is much less frequent than with the other arms.

666

probably sensory functions during copulation and are also found in biserial
species of Octopus.

The variable proportions of the genitalia in the mantle cavity are shown
by comparison of Figure 408 on p. 685 (small spermatophores) with
Figure 423 (large spermatophores). The female genital papillae may also
differ markedly from large processes to minute papillae retracted in a
kind of foreskin; they may be thick and blunt or pointed, and their distance
from the anus is variable. Some species have no ink sac; Grimpe (Nordsee,
p. 300) uses this character for the establishment of new genera. (For
further details, see Vol. III.)

Only examination of the whole material would permit the establishment
of a classification. Introduction of new names is not important. The
correlation of the following characters would have to be examined for the
definition of natural genera; 1) size of eggs (p. 686); 2) form of head and
mantle in the live animal (p. 688); 3) relative length of arms (pp. 693, 703,
708); 4) formation of the interbrachial membrane (p. 699); 5) development
692 of a constant sculpture on the skin (p. 690); 6) development of the funnel
glands (p. 689); 7) occurrence of lateral lines or fin margins (p. 675);

8) narrowed mantle opening (p. 690); 9) form of the radula (p. 6 89); 10) ink
sac (p. 691); 11) gelatinous -transparent structure of the body (p. 6 80);

12) interbrachial pockets in the mouth area (Cistopus); 13) form of apex
of hectocotylus (spoon) (p. 690); 14) abnormal apices of other arms of the
mature male (except the hectocotylus).

The "genera" Eledone and Scaeurgus are obviously artificial
groups, which were established on the basis of striking characters. It
cannot be decided at present whether the new classification of Grimpe
(1921, Nordsee, p. 300) is valid. The validity of Pinnoctopus d’Orb.,
1845 is also doubtful, because the large. Sepia -like, posteriorly fused
fin margins of this form may be artifacts formed by the relaxed skin and
the drawing is certainly exaggerated. Another doubtful form, Cistopus
Gray, 1849, is based on a single specialization in an otherwise typical
species of Octopus (Tryon, 1879, p. 127, Plate 39, Figures 61, 62). The
same applies to Velodona Chun, 1915. At any rate, the variety of forms
in the family has not yet been worked out, and the numerous workers who
have dealt with the many species of the "genus" Octopus have not yet
done serious systematic -morphological work. We have indicated above
what should be done but we cannot solve the problem. We shall therefore
attempt to determine provisionally the position of the available species by
a precise diagnosis, as in the case of Sepia and Loligo (p. 193 and
545). There are evidently two groups, represented by the generic types
Octopus and Eledone, if only for practical purposes. Regardless of
the traditional meaning of these names, we consider the distinction between
forms with a direct and an indirect development (i. e. between small and
large eggs, p. 6 86) as more important than all other characters and would
use this distinction for the establishment of subfamilies if it were certain
that all species with uniserial suckers resemble the Mediterranean
Eledone. We would then distinguish Eledoninae and Octopodinae, but
not in the sense of Grimpe (1921, Nordsee, p. 300). We consider as related
to Eledone the Octopodidae represented by O. digueti de Rochebrune
1896, i. e. forms with regularly biserial suckers and with eggs which are
5 — 9 mm long (usually about 6 mm without capsule). (I have often examined

667

eggs of such Octopodidae with young forms and with half developed embryos
resembling those ofEledone in the Museum of Natural History in
Hamburg and in the State Zoological Collection in Munich. ) However, these
embryos differ from those of Eledone in their regularly biserial suckers,
which are present in large numbers already in the embryos. The eggs in
Munich are from Sagamibai (Japan), the origin of the material in Hamburg
is unknown. Both show isolated, club-shaped egg capsules and differ in this
respect not only from the Mediterranean Octopus but also from
Eledone. See also Tryon, 1879, Plate 19, Figure 3. I propose the name
Paroctopus, type species: P. digueti for these species.

693 Chapter 51

GENUS OCTOPUS
Lamarck, 1798

(= Polypus Schneider, 1784; Hoyle, 1901)

Contents; a. Diagnosis. — b. Typical character. — c. Variation. — d. Octopus vulgaris (p. 697). —
e. Octopus saluzzii(p. 699). — f. Octopus macropus (p. 702). — g. Octopus defilippii
(p. 707). — h. Octopus tetracirrus (p. 710). — i. Octopus unicirrus (p. 713).

a. DIAGNOSIS

General appearance almost typical. Eggs small (about 1.5 — 3 mm),
development indirect (p. 686). All arms with regularly biserial (zigzag)
suckers. Ink sac well developed. This genus also includes the species of
"Scaeurgus" Troschel 1857 (cf. p. 681) but the hectocotylus is on the
right side (if not stated otherwise).

b. TYPICAL CHARACTER

The genus cannot be exactly defined even with the restricted definition.
The descriptions of most species are too incomplete, because of either lack
of material or insufficient systematic -morphological knowledge, to permit
determination of their systematic position. The diagnosis applies to all
the Mediterranean Octopodidae except Eledone, i. e. a group containing
very different forms, although it is certainly not defined too narrowly.

The description of the family may be considered as the norm (p. 681),
also for the young stages, because Octopus is the central group (p. 16)
of the family, with a restricted diagnosis.

c. VARIATION

Octopus shows, however, marked variations from the type which
apply to the following parts and should be described for the various species;

1. Relative length of the arms and relative size of suckers on the arms.
2. Sculpture of suckers. 3. Absolute number of suckers on the arms of
694 more or less mature specimens. 4. Form of funnel gland (W or VV).

669

695

FIGURE 412. Half-grown male of Octopus
vulgaris. VaX- Drawn after a fresh specimen
(compare with Figure 41 1 on p. 689). Funnel
and mantle sac are not strongly contracted or
reduced in size but are as in the swimming ani-
mal (ventral side). Note the proportions of
arms, suckers, and interbrachial membrane;
the point of entrance at the hectocotylus (x),
the longitudinal groove (y), and the almost
rudimentary spoon (z).

5. Presence of lateral lines (fin
margins) on the mantle. 6. Occurrence
of constant skin warts, especially on
the dorsal and ventral sides of the
mantle sac. 7. Structure of the teeth
of the radula. 8. Form of jaws.

9. Coloration if the chromatophores
are extended. 10. Enlargement of
certain suckers, especially in the male.

11. Structure of apex of hectocotylus.

12. Hectocotylization on the right or
leftside. 13. Form of the egg. 14. Struc-
ture of spermatophores. 15. Form of
external genitalia in the mantle cavity.
16. Form, size and attachment of
interbrachial membrane.

A complete description of the
species based on a single dead speci-
men is usually only possible if the
specimen is a well preserved, mature
male and uncertainties are not ex-
cluded even then. The arms are very
contractile and exact measurements
are therefore useless. Only a great
difference in length may be considered
as characteristic and regenerated arms
cannot always be recognized. Of some
species, not a single intact speci-
men has been obtained and all arms
are often repeatedly regenerated. The
relative size of the larger suckers
usually corresponds more or less
closely to the length of the arm. The
absolute number should be used with
care, mainly because of the frequent
regenerations. It cannot be used at all
in young specimens because it has not
yet reached the normal value. The
norm also varies from one race or
variety to another in the same species
(O. macropus, p. 704). The funnel
gland also is not always constant in
form in the same species (O. uni-
cirrus, p. 714) but it usually is.

The lateral lines are very character-
istic for the different species. It is
not always easy to distinguish between
transient and constant skin warts, but
their characteristic form and
consistency are often of systematic
value. The teeth of the radula usually
differ little from the type (Plate XVI)
and have only little value as species

670

characters. The jaws also show only few distinct characters and the
coloration is only significant if it differs markedly from the usual dirty
brown, or if it shows a constant and characteristic pattern. The characters
of the hectocotylus are very distinct and the special form of its apex is
characteristic also for larger groups. The structure of the hectocotylus is
certainly more important than the side (right or left) of hectocotylization.
Eggs and spermatophores are little known, but they certainly differ from
one species to another. This is especially true for the spermatophores,
the characters of which, in connection with the type of hectocotylization,
would probably provide a basis for the natural systematic s of the whole
group. The form of mantle sac and head is very variable, but it is specific
in the live animal; it is worthless in preserved specimens because of the
various deformations.

d. OCTOPUS VULGARIS Lam., 1798
1. DIAGNOSIS

Dorsal skin more or less covered with warts and divided into areas.
Dorsal arms distinctly shorter, lateral arms (hectocotylus not included)
much longer than the others. Suckers of lateral arms of large specimens
(length over 20 cm without arms) markedly enlarged, beginning from the
15th~17th sucker (especially in the male, in which two suckers, usually
the 17th and 18th, may be more than 3 cm wide) and gradually decrease in
size toward the apex. Apex of hectocotylus forming only a small (about
7 mm), indistinct spoon (Figure 405 on p. 6 83). Coloration yellowish brown,
grayish brown or reddish brown in life. Skin with warts. Lateral lines on
mantle absent.

2. LITERATURE

1554 Rondelet (p. 813), Polypus prima species.

1558 Gesner (Vol. IV, p. 870), Polypus prima species.

1606 Aldrovandi (p. 15, Fig. 16), Polypus cum duplice acetabulorum ordine.

1757 Hasselquist (p. 33), Octopodia.

1758 Linne (1. Part VI, p. 3149), Sepia octopus,

1767 Johnston (p. 5), Polypus.

1792 Bose (p. 24, Plate V), Sepia rugosa.

1802 Bose (Vol. I, p. 47), Sepia octopus.

1805 Montfort (Vol. II, p. 113, Plates 23 — 25; Vol. Ill, p. 5, Plates 27, 28), Poulpe commun, Poulpe fraise.
1817 Cuvier (Vol. II, p. 7), Poulpe granuleux.

1817 Leach (p. 394), Polypus octopodia.

1822 Lamarck (12th edition, Vol. 11, p. 361), Octopus vulgaris.

1824 Carus (p. 319, Plate 31), Octopus vulgaris.

1825 Blainville (p. 465, Plate 2), Octopus vulgaris.

1825 Savigny and Audouin (Vol. I, p. 9, Plate II), Octopus vulgaris.

1826 Blainville (Vol. 43, p. 188), Octopus vulgaris.

1826 Blainville (Vol. 43, p. 188), Octopus brevitentaculatus.

1826 Payraudeau (p. 172), Octopus vulgaris.

1826 Risso (Vol. 4, p. 3), Octopus vulgaris.

671

1826

1827

1828
1829
1829
1829
1833
1835

1837

1838
696 1841

1841

1844

1849

1851

1851

1853

1855

1858

1863

1869

1869

1879

1880
1880
1882

1885

1886
1890
1890
1890
1894
1894
1896
1901
1908
1916
1918

1920

1921

1921

1922

d'Orbigny (p. 142), Octopus vulgaris.

Bruguiere (p. 136), Octopus vulgaris.

Ehrenberg, Octopus vulgaris.

Delle Chiaje (vol. 4, pp. 40 and 55). Octopus vulgaris.

Sangiovanni (p. 321), Octopus vulgaris.

Wagner (p. 225), Octopus vulgaris.

Oken (Vol. V, p. 536), Sepia octopodia.

Ferussac and d'Orbigny (pp. 26 and 38), Octopus vulgaris. (The authors place here also
O. saluzzii Ver.)

Rang (p. 62), Octopus vulgaris.

Potiez and Michaud (Vol. I, p. 6), Octopus vulgaris.

Delle Chiaje (Vol. I, p. 2; Vol. V, p. 68), Octopus vulgaris.

Cantraine (p. 18), Octopus vulgaris.

Philippi (p. 201), Octopus vulgaris.

Gray (p. 6), Octopus vulgaris. Gray places here also O. saluzzii Ver. and O. u n i c i r r h u s D.
Ch. (Scaeurgus).

Verany (p. 16, Plate 8), Octopus vulgaris.

Verany (p. 45, Plate 12), Octopus tuberculatus.

Forbes and Stanley (Vol. IV, p. 209), Octopus vulgaris.
d'Orbigny (p. 168), Octopus vulgaris (defined as by Gray).

Adams (Vol. I, p. 20), Octopus vulgaris.

Aucapitaine (p. 290), Octopus vulgaris.

Jeffreys (Vol. V, pp. 144, 145), Octopus vulgaris.

Targioni-Tozzetti (pp. 17, 19), Octopus vulgaris. T. distinguishes a large specimen as
O. troscheli.

Tryon (p. 113), Octopus vulgaris (- O. octopodia L.).

Tiberi (p. 10), Octopus vulgaris.

Stossich (p. 157), Octopus vulgaris, also O. troscheli .

Verrill (p. 72), Octopus vulgaris.

Ninni (p. 159), O c t opu s v u 1 g a r i s , also O. t r os c he li .

Hoyle (p. 7), Octopus vulgaris.

Colombatovic (p. 7), Octopus vulgaris.

Norman (p. 466), Octopus vulgaris.

Carus (p. 459), Octopus vulgaris.

Pelseneer (p. 206), Octopus vulgaris.

Joubin (p. 25), Octopus vulgaris.

Jatta (p. 212, Plates 4, 7, 8, 22, 23), Octopus vulgaris.

Hoyle, Polypus vulgaris.

Pfeffer (p. 20, Figs. 11 — 13), Polypus vulgaris.

Naef (System, p. 16), Polypus vulgaris.

Grimpe (Brehm, p. 593, Fig.), Polypus vulgaris.

Wiilker (p. 56), Polypus vulgaris (Red Sea).

Grimpe (pp. 298, 300), Octopus* vulgaris (North Sea).

Naef (p. 538), Octopus vulgaris (System).

Naef (Foss. Tintenf., p. 288, Fig.), Octopus vulgaris.

3. STRUCTURE OF THE ADULT ANIMAL

Adults in Naples are 20 — 25 cm long without arms and weigh 2 — 3 kg.
Exceptionally, animals weighing 5, 7 or even 12 kg are found, Jatta

* Grimpe reinstated the traditional name Octopus (1920, Zool. Anz., Vol. 51, p. 206). Hoyle (1901,
Manchester) unnecessarily revived the names which Schneider (1884) had proposed for Eledone
(M os ch it e s) and Octopus (Polypus); these names are invalid, according to the rules of nomenclature
but they confused the terminology. We therefore use the original names.

672

697

reported specimens of to 25 kg (1896, p. 216). I have never seen such
animals. They probably live at greater depths, like large Sepia
offic inalis.

The arms are more than 4 times as long as the body. The dorsal arms
are the shortest, followed by the ventral arms and then by the two lateral
pairs, which show very variable proportions (Figure 412). The size of the
suckers increases from the mouth to the 18th or 19th of the row, and then
gradually decreases toward the whiplike end of the arm. Each arm bears
about 240 suckers. Some suckers of both lateral arms, from the 15th — 17th
onward, are particularly large (usually the 17th and 18th sucker). Such
enlarged suckers occur in both sexes but the enlargement is less marked
in the female. The largest suckers on the dorsal and ventral arms are also
the 17th — 19th, usually the 18th.

The mantle cavity shows essentially the typical conditions (p. 665,

Figure 392 on p. 664 and Figure 407 on p. 6 84). Note also there the
topographical changes connected with the maturation of the female
(Figure 408 on p. 685).

The jaws (Plate XVIII, Figures 4, 5) show also the changes which take
place during growth: they become more strongly chitinized and pigmented.
This must be remembered during comparison of the jaws. The margins
of the jaws of young animals, especially of the upper jaw, may be so
transparent that they are overlooked and simulate characters which do not
exist. Most drawings of jaws of Octopus are without systematic value,
not only because the author has no information on the constancy or
variation of the characters but also because the illustrations are too inexact
to show specific characters. The size of the jaws must obviously be
considered. The lower jaw, measured in the midline, is about V12 of f^e
length of the body. The typical radula is shown on Plate XVI, Figure 5.

The mantle cavity of the female shows no special characters (Figure 390
on p. 663 and Figure 407 on p. 684).

The lateral arms of the male show a marked enlargement of the suckers.
The 17th and 18th sucker of each arm are the largest; they may be wider
than 3 cm. The hectocotylus develops on the right side. It is only
moderately shortened and bears about 170 normal suckers, followed by a
small, indistinctly differentiated but otherwise typical spoon which is rarely
longer than 7 mm from the last sucker to the apex of the arm in the mature
animal (Figure 405 on p. 683 and Figure 412 on p. 694).

The skin of young animals bears transient warts, of which the two largest
are absent only rarely and in very young specimens. The whole dorsal
surface of the mantle, head and arm bases of older animals is densely
covered with small papillae, with larger papillae of different size between
them. Relaxation of the skin musculature creates a characteristic division
of the surface into flat elevations separated by thin incisions. This is
absent in most other species.

Merculiano (Jatta, Plate 4, Figure 1) well reproduced the typical
coloration, a varying pattern of brownish yellow, reddish brown, dark
brown and gray tones in a large variety of combinations (cf. ecological
part). These tones combine with the skin warts into a highly characteristic
picture, especially in young animals, which may resemble a stone overgrown
with algae, a sea urchin, or a sponge.

* Cf.. for example, Chun (1915, Figure 52).

673

4. POSTEMBRYONIC DEVELOPMENT

O. vulgaris has a typical metamorphosis. The youngest postembryonic
stages are shown in Figures 400^ 401 on p. 67 8 and Figure 409 on p, 687,

The determination is certain for specimen a (p. 678), which was hatched
from the egg; the two other specimens, found in the plankton, are closely
similar. Such larvae are not rare in the plankton of the bay in summer
69 8 and autumn and those hatched in the aquarium can be kept alive for some
time, to about 12 days. The larvae show the typical characters (p. 66 8)
of the earliest pelagic stages of Octopoda. They do not differ markedly
from those of other species of Octopus, particularly O. saluzzii and
O. defilippii. On the other hand, O. macropus has a markedly
different juvenile form, at least in slightly older stages (Figures 417, 418
on p. 705, Figure 419 on p. 706 and Plate IX). Thin spines cover the
entire mantle, the head, the projecting funnel and the arms except the apex.
These spines are directed anteriorly and are important during hatching
(cf. p. 687); they may disappear during preservation.

A slightly more advanced larva is shown in Figure 409 on p, 687; mainly
the arms have developed further. The older larva shown in Figure 401 on
p. 67 8 probably also belongs here. I do not know the next stages which lead
to the definitive form (p. 6 86) and we return therefore to the young animal
in Figures 402 and 410 (see also Plate IX, Figure 3). This specimen
already resembles the adult in structure and mode of life. It shows only
small remnants of the early armature of spines or bristles and these
disappear rapidly. On the other hand, there are distinct juvenile characters,
especially in the live animal (Figure 402 on p. 679), particularly the
characteristic form of the head and the still relatively short arms, which,
however, already show the typical proportions of the adult. The arms of
the smallest specimens which are found in winter and spring between stones,
sand and detritus in the coastal zone bear about 40 — 50 suckers. The head
is characteristic because of the markedly prominent eyes, which may be
stalked when the musculature at the base of the eye is contracted and the
venous sinus of the eye is filled with blood (cf. Figure 386 on p. 659).

The following specific changes are present in comparison with the larval
form. The olfactory organ is retracted more or less under the mantle
margin. The typical lid apparatus is formed above the eye (p. 660):
secondary lid, primary upper lid, pseudocornea or primary lower lid. The
conditions in the mantle cavity resemble those in the adult. But mainly the
posture of the body is typical for Octopoda living on the bottom: the arms
are bent ventrally, the funnel laterally, and the arms are coiled into a spiral
(p. 6 82). The skin is also adapted to life on the bottom. In addition to the
large chromatophores, which are present already in the larvae and which
are now the main chromatophores in the deeper layers of the skin, there
are numerous small chromatophores, some of them yellow, others brown.
These small chromatophores provide a protective adaptation to life on the
bottom. The skin musculature produces numerous warts on the dorsal side,
some of them especially large and constant in all specimens. Particularly
constant (Figure 402 on p. 679 and Figure 410 on p. 688) are one wart above
each eye, 4 on the dorsal surface and one at the posterior end; they become
dendriform appendages or spines by contraction (p. 697).

5110/2

674

699 The older stages need no special description as they rapidly beconae

similar to the adult condition. Of particular interest is their rapid growth:
with good care and feeding the weight usually doubles within 10 days. During
longer captivity, however, the growth rate decreases. With only moderate
and irregular care, the weight of 2 specimens increased from 43 and 58 g
on 30 June to 174 and 180 g on 30 July. Lo Bianco (1909, p. 652) reports
the following increase in weight: 8 August: 65 g; 6 October: 600 g, i. e.
a tenfold increase within 2 months.

e, OCTOPUS S ALUZ ZII Verany (1837)

1. DIAGNOSIS

Dorsal arms only slightly shorter, lateroventral arms only slightly
longer than the others. Normal arms with about 190 suckers. Interbrachial
membrane continuing distally as a broad, very extensible margin along the
ventral outer edges. Skin of preserved specimens usually smooth and
700 slightly swollen and gelatinous. All arms of male with several strongly
enlarged suckers near the 10th --12th sucker; other suckers gradually
decreasing in size in both directions. Hectocotylus of mature male with
about 135 — 140 suckers, with a narrow spoon, more than 3 cm long, bearing
transverse ridges at the apex. Live animal yellowish brown to orange
yellow, with slightly warty skin.

(699)

FIGURE 413. Young female of O. saluzzii, 0.6 X. a) In a glass dish, viewed from above, arms extended.
Note the outline of head and body, form of eyes, particularly of pupil and funnel. The interbrachial membrane
extends along the ventral side of the arms in the form of broad margins, b) Same animal, swimming.

These observations make the assumption unnecessary that giant Cephalopoda are very old. Most species live
about one year, at any rate, probably not longer than 2 — 3 years. The same applies to the fossil forms
(cf. Naef, 1922, p. 207).

675

2. LITERATURE

1837 Verany (p. 93. Fig.), Octopus salutii.

1840 Verany (Fig. 5), Octopus saluzzi.*

1844 Philippi (p. 201), Octopus saluzzi.

1845 Ferussac and d'Orb., (pp. 27, 33), Octopus vulgaris. The authors do not consider the species as valid.
1849 Gray (p. 6; Gray agrees with Ferussac and d'Orbigny), Octopus vulgaris.

1851 Verany (p. 20, Plate 9), Octopus salutii.

1855 d'Orbigny (pp. 179, 191), Octopus vulgaris. He considers O. sa luti and O. unicirrus D. Ch.
as forms of O. vulgaris.

1869 Targioni-Tozzetti (p. 18), Octopus salutii (? ).

1879 Tryon (p. 114), Octopus salutii.

1886 Hoyle (p. 7), Octopus salutii.

1890 Carus (p. 459), Octopus salutii.

1896 Jatta (p. 224, Plates 4, 21, 22), Octopus salutii.

1916 Naef (p. 16), Polypus saluti.

1921 Naef (p. 538), Octopus salutii.

3. STRUCTURE OF THE ADULT ANIMAL

This species closely resembles a half-grown Octopus vulgaris,
in habitus, but the coloration and the consistency of the skin differ
considerably, in addition to specific characters of the hectocotylus,
interbrachial membrane, size and number of the suckers, etc. The body
is short and compact as in O. vulgaris.

Arm formula: 3; 2; 1 = 4. The 3rd pair is distinctly longer than the
2nd which is markedly longer than the other two pairs (Figure 411 on p. 689).
The suckers show the typical arrangement (p. 676, not as stated by Jatta,
p. 225) and there are about 190 on each arm. A marked enlargement of
some suckers is present only in the male (not as stated by Jatta) (Figure 411
on p. 689).

The interbrachial membrane is very characteristic. It is supported by
a muscular thread at its margin and extends to ^/g of the arms toward the
apex, but only at the ventral outer edge. It thus forms a very delicate and
elastic margin in the live animal which resembles a veil. Because of its
elasticity and marginal support, the interbrachial membrane forms pockets
and bulges in which prey may be captured. Chun (1915) published a similar
picture of Velodona. Preservation causes a more or less marked
contraction of the membrane and obliterates the most prominent and
distinctive character of the species (Figure 411 on p. 689), especially when
701 the arms are extended. In specimens with coiled arms, their margins are
always distinct (Jatta, Plate 22, Figure 1), but not as markedly as in the
live animal.

The mantle cavity (Figure 391 on p. 663) is typical in general and closely
resembles that of O. vulgaris. A minor character is shown by the
female genital openings: in the only mature female I have examined, they
form pointed papillae which are slightly retracted into a kind of foreskin.

The openings are also not situated close to the anus, as in O. vulgaris.

* This name must be derived from the name Saluzzi; the confusion .is caused by the use of different spellings
by the author.

676

The jaws (Plate XVIII, Figure 7) do not differ markedly from those of
other Octopodidae, but the biting process of the upper jaw is of specific
form and there is a deep median incision in the posterior part of the lower
jaw. The radula is typical (cf. the inexact Figure 8 of Plate 21 in Jatta).

FIGURE 414. Inner part of arms and mouth region of Octopus saluzzii, natural size. Note the
arrangement and relative size of the suckers; also the relation of the interbrachial membrane (3) to the
dorsal (5) and ventral (4) edges, along which the membrane forms broad margins. In the mouth, the
apex of the lower jaw, around it the fluted inner lip (2) and the smooth outer lip (1).

Hectocotylization consists in 1) an enlargement of some suckers near
the base of all arms, of which 1 — 2 are particularly large; the suckers
become gradually smaller toward the apex, so that many suckers are little
enlarged; 2) the 3rd arm of the right side forms the hectocotylus. I counted
136 and 139 normal suckers on the hectocotylus in two cases. The end part
of the hectocotylus (Figure 405 on p. 683) is characteristic. The spoon
is very long, narrow, with numerous sharp basal incisions between thin
transverse swellings. The spout of the spermatophore groove is very
distinct. The margins of the spoon are often spread so that its structure
becomes indistinct. The whole terminal part is over 3 cm long in mature
specimens; it is much smaller in half-grown specimens but is recognizable
by its structure.

702 Merculiano reproduced the coloration and habitus of the live animal very
well (Jatta, 1896, Plate 4, Figure 3). The animal is bright yellowish brown
to orange-yellow with brown and reddish brown cloudy and light spots. The
surface is smooth (frequently in life, but usually after preservation), except
for the warts above the eyes. The skin is always slightly gelatinous and
swollen. A length of only 8 cm (without arms) may be considered as normal
for mature specimens.

677

4. JUVENILE STAGES

I have some very young specimens, all of them preserved. They have
still shorter arms and fewer suckers but their coloration and interbrachial
membrane are already characteristic for the species and differ. distinctly
from those of O. vulgaris, which they closely resemble otherwise. The
coloration is more pink than yellowish brown in alcohol-preserved
specimens.

f. OCTOPUS MACROPUS Risso, 1826
(=0. RUBER Rafinesque, 1814?)

1. DIAGNOSIS

Arms of different length, very long (formula: 1, 2, 3, 4). Interbrachial
membrane short, restricted to the base of the arms. Dorsal arms very
strong and with markedly enlarged suckers in both sexes, especially in the
male. Hectocotylus with a long (2 cm or more), narrow spoon with a thin,
light, longitudinal stripe at the bottom and a narrow transversely striated
zone on each side of the stripe. Live animals with large, transient warts
and reddish brown (yellowish brown to brownish red) coloration with many
large whitish spots.

2. LITERATURE

1826 Risso (Vol. IV, p. 3), Octopus macropus, description.

1829 Wagner (p. 225), Octopus macropus.

1829 Delle Chiaje (Vol. IV, pp. 40, 56), Octopus macropus.

1829 Sangiovanni (p. 319), Octopus macropodus.

1835 Ferussac and d’Orbigny (p. 18), Octopus cuvieri. (t O. lechenaultii, macropodus,
longimanus. Poulpes, Plates 1, 24, 27.)

1837 Rang (p. 61, Plate 90), Octopus macropus.

1839 d’Orbigny (p. 16, Plates 3, 4), Octopus cuvieri.

1841 Cantraine, Octopus ruber Rafinesque 1814. C. considers this insufficiently characterized species
as a synonym ofO. macropus.

1841 Delle Chiaje (Vol. I, p. 3; Vol. 5, p. 65), Octopus macropus.

1844 Philippi (p. 201), Octopus ruber Raf.

1849 Gray (p. 13), Octopus cuvieri.

1851 Verany (p. 27, Plate 9), Octopus macropus.

1851 Verany (p. 32, Plate 7 bis. Fig. 3), Octopus alderii. juv..'

1855 d'Orbigny (p. 173), Octopus cuvieri.

1856 Steenstrup (p. 234), Octopus macropus.

1857 Troschel (p. 59), Octopus macropus.

1858 Adams, H. and A. (p. 19), Octopus cuvieri.

1869 Targioni-Tozzetti (pp. 23, 24), Octopus macropus and cuvieri.

1879 Tryon (p. 122), Octopus cuvieri.

1880 Tiberi (p. 10), O ctopus macro pus.

1880 Stossich (p. 57), Octopus macropus.

1885 Ninni (p.l). Octopus macropus.

1886 Hoyle (pp. 11, 95), Octopus macropus (= O. cuvieri?).

1886 Appellof (p. 6), Octopus cuvieri.

678

1887 Brock (p. 597), Octopus cuvieri.

1890 Colombatovic (p. 7), Octopus macropus.

703 1890 Carus (p. 460), Octopus macro pus.

1894 Joubin (Amb.) (p. 58), Octopus macropus.

1894 Joubin (Hirond.) (p. 212), Octopus macropus.

1896 Jatta (p. 217, Plates 6, 23, 24), Octopus macropus.
1916 Naef (p. 16), Polypus macropus.

1920 Wulker (pp. 50, 56), Polypus macropus.

1921 Naef (p. 538), Octopus macropus.

3. STRUCTURE OF THE ADULT ANIMAL

Adult or half-grown specimens of this species are easily recognized and
distinguished from all other forms. However, this species is possibly
heterogenous because the specimens differ markedly in some characters
(p. 705).

FIGURE 415. O. macropus. Vs^* male; b, b| — female; both mature. Note the resemblance.

Both have a narrow head and a slender mantle sac (cf. Figures 412, 413, 420). The sexual differences are
caused by the development of the ovary in the female. Drawing after live animals. The drawings are
designed to show that the form of the body of Octopodidae can be very specific, despite the variation by
different contraction of mantle and head.

The habitus is determined by the length of the arms, the form of the body
and the characteristic coloration which Merculiano naturally reproduced
in Jatta (1896, Plate 6, Figure 1, mature female). Figure 415 shows a
mature male and female. The mantle has a specific form already in the
young stages (Figure 419 on p. 706). However, the posterior end is more
blunt and the whole mantle sac is plumper and relatively larger. The free
704 ventral margin of the mantle is not a simple transverse line, but is slightly
but distinctly produced in the middle. The head is very narrow.

679

FIGURE 416. Almost mature male
of O. macropus. VsX- Ventral
view of a fresh specimen. Note
the characteristic form, the short
7Q5interbrachial membrane, the long
unequal arms with suckers of dif-
ferent size. Note particularly the
hectocotylus; the number of
suckers is reduced, and the distal
part is characteristic; x —
entrance to the spermatophore
groove (y); z — spoon.

The arms are very long, whiplike. If they
are completely relaxed, the body forms only
about V? of the total length. The arms differ
markedly in length; formula: 1, 2, 3, 4
(proportions: 8:7:5:4 or 9:8:6. 5:6); the
DL and LV arms differ most markedly in
length. The suckers are numerous, but differ
in size on the different arms; those of the 1st
pair are often twice as large as those on the
4th pair. The dorsal arms bear distinctly more
numerous suckers than the ventral arms;

I counted up to 300 on the dorsal arms, in
larger specimens usually about 270, but the
number of suckers never exceeded 240 on the
ventral arms. There is apparently a variety
with short arms, with only about 150 suckers
on each arm (correspondingly fewer on the
hectocotylus). However, a distinction is
difficult because young specimens have
generally fewer suckers. I consider therefore
the form with long arms as typical and do not
consider the form with short arms as a variety
or subspecies. A very short interbrachial
membrane connects the base of the arms and
extends only little along the arms, except in
the spermatophore groove.

The mantle cavity shows no special charac-
ters and jaws and radula (cf. Jatta, Plate 23,
Figure 13) show the typical conditions of the
family. The biting process of the upper jaw
is very short, and the lower jaw has a deep
incision in the posterior part (Plate XVIII,
Figure 6). Hectocotylization consists in a
slight enlargement of the suckers of the
proximal part of the arms and the formation
of a very characteristic hectocotylus on the
3rd arm of the right side; it is little more than
half as long as the opposite arm and bears only
about 110 — 115 suckers (80 — 90 in the form with
short arms). The end part consists of a narrow
spoon, over 2 cm long in large specimens.

A typical spout leads from the spermatophore
groove to the spoon. The bottom of the spoon is
transversely striated, with a light median line
in the whole spoon. Many other species with
a transversely striated bottom of the spoon do
not show such a line (see Figure 405 on p. 683).

The coloration of the live animal is usually
darker and lighter brownish red or reddish
brown, often more yellowish brown. The whole
dorsal side of the body, head and arms is
covered with large, whitish spots which may

680

form light, pointed warts. Only the warts above the eyes are constant,

3 or 4 on each side, one of which projects more than the others. Numerous
small warts may be present on the dorsal and ventral side, probably in
connection with the tufts of bristles of the larva. These warts apparently
show sometimes remnants of such bristles, but I did not make a micro-
scopic examination.

O. macropus is much smaller than O. vulgaris. The mature
animals examined are 10 — 15 cm long (without arms) and weigh to 1.5 kg.
This species is common on the market in Naples. It is valued as food and
is known under the name ’’Purpessa," i. e. the female of "Purpo verace"

(O. vulgaris). Today, the fishermen know the sexes well.

4. JUVENILE STAGES

I have a number of developmental stages of this species, which are all
characteristic. The youngest forms are shown in Figures 417 and 418, in
Figure 397 on p, 668, and on Plate IX, Figure 1. These are still planktonic
larvae, but the number of suckers is markedly greater than the primary
number of 3 suckers (the smallest larva shows 7 suckers). Special
characters: 1. Head and mantle sac are already slender, but otherwise
larval. 2. The skin is gelatinous, swollen (the larva was therefore first

706 considered as that of "S c a eu r gu s t e t r a c i r r u s"). 3. Tufts of bristles

are present in the largest specimen. 4. The chromatophores have a
characteristic distribution which permits identification of the larva by
comparison with older stages. The specimen from Messina (Figure 397 on
p. 668; Plate IX, Figure 1) is particularly well preserved. It is less
deformed than the others and has a wide, ovoid mantle sac. Each arm
bears 10 suckers, and the spines have already split into tuft- or star-shaped
structures (part of the periphery is shown in the smaller drawing). The
characteristic form and the habitus of the animal are well reproduced in the
drawing on the plate. The chromatophores are dirty reddish brown.

(705)

Larvae ofO. macropus from the plankton. 10 x .

Figure 417, from Porto d'Ischia, 20 August 1912; Figure 418, from the Bay of Naples, old material. The
small drawing c shows the typical spines of the skin, magnified about 40 x, from the periphery. The spines
have not yet split into bristles. Contraction has caused some deformation; compare with Figure 397 on p. 668
and note the number of suckers (7) and the distribution of chromatophores.

681

FIGURE 419. Young stages of Octopus macro-
pus, natural size, a) One of Jatta's specimens
of "O. a 1 d e r i i . ” The arms are relatively short,
the membrane is weakly developed, and the mantle
sac is slightly shortened by preservation. The eyes
are retracted; in life they are as shown in
Figures 397 on p. 668 or 400, 401 on p. 678.
b) and c) Older animal which shows distinctly the
character of the species (caught 1 May 1912 near
Posilipo). The natural form of the head and
mantle is well preserved because of careful cocain-
ization. Both stages show clearly the specific
distribution of the chromatophores.

Young but already benthic animals have been known for a long time;

Jatta (1896, Figure 19, Plate 7; Figures 19 — 26, Plate 20; p. 228) described
them as "Octopus alderii." The original description of this "species"
(Verany, 1851) does not permit identification. At any rate, the specimens
described by Jatta belong to O. macropus. The genital organs are just
beginning to develop. These stages are intermediate between the larvae
described above and the older forms which are easy to identify. They
resemble the larvae in the arrangement of the chromatophores, especially
on the mantle — the chromatophores are concentrated ventrally at the
posterior end and at the free margin of the mantle but they also form a
707 loose group in the ventral median line (Figure 397 on p. 668). This
arrangement remains distinct in the older stages, but the number of
chromatophores is increased.

Already the alderii stages closely resemble the adult O. macropus
in the form of mantle sac, head and bases of the arms. On the other hand,
the arms are still short, as in the juvenile forms, and do not differ
markedly in length, but the formula is already definitive (l, 2, 3, 4), The
mantle may Show various deformations, but it is usually pointed at the
posterior end, with a bellylike widening near the end and a slightly bell-
shaped constriction at the anterior margin. The markedly slender body is
distinct also in preserved specimens (Figure 419 on p, 706). Alcohol-
preserved specimens usually have a head of abnormal form because of the
retraction of the eyes, which protrude in the live animal as shown in the
Figures 397 on p. 668 or 400, 401 on p. 678. However, the head is much

682

narrower than in O. vulgaris and most other species (except O. defilip-
pii), probably as in the older stage of Figure 419c. The chromatophores
are rather dense and reddish brown on the dorsal side.

The arms of the specimen in Figure 419c, d, resemble those of the later
stages. The dorsal arms should be still larger than the others in this
stage; the dorsal arms are shorter in this specimen because regeneration
has certainly taken place.

g. OCTOPUS DEFILIPPII Verany, 1851
1. DIAGNOSIS

Small species, with markedly^ different, very long arms which tend to
autotomy. Formula: 3, 2, 4, 1. Suckers of arms correspondingly different
in size. Interbrachial membrane very short, but reaching beyond the
middle on the posterior edges of the arms. Hectocotylus with about 100
(70 — 104) suckers and a small, indistinct spoon. Coloration dirty, spotted,
grayish brown to yellowish brown. Length 4 — 5 cm without arms.

2. LITERATURE

1851 Verany (p. 30, Plate II), Octopus De Filippi.

1869 Targioni-Tozzetti (p. 20), Octopus De Filippi.

1879 Tryon (p. Ill), Octopus defilippi.

1880 Tiber! (p. 11), Octopus De Filippii.

1886 Hoyle (p. 8), Octopus defilippi.

1890 Carus (p. 460), Octopus De Filippii.

1896 Jatta (p. 221, Plates 4, 24, 25), Octopus defilippi.
1916 Naef (p. 17), Octopus defilippi.

1921 Naef (p. 538), Octopus defilippi!.

708 3. STRUCTURE OF THE ADULT ANIMAL
I

The general appearance of the live animal is shown in the colored
drawing by Merculiano in Jatta (1896, Plate 4, Figure 2). O. defilippii
has a slender body, like O. macropus, and this is accentuated by the
long arms. However, it has a different coloration and makes characteristic
sluggish movements with the arms. The mantle sac, which is shown in
Figure 420, is often contracted like a sausage or inflated into a sac or
an egg. The eyes of the living animal are very prominent, as observed
overwise only in juvenile forms (Figure 397 on p. 668).

The interbrachial membrane is very short at rest but is very elastic.

It extends markedly beyond the middle of the arm at the posterior outer edge
and it can be widened into broad margins temporarily (Jatta, Plate 4).

Arm formula: 3, 2, 4, 1. The 3rd pair is markedly longer (not including

709 the hectocotylus), the 1st pair markedly shorter than the others. If the
3rd arms are relaxed, they are about 5 times as long as the body without

683

arms, the 1st pair about 3 times as long. The suckers are correspondingly
unequal and there are about 140 on each arm.

(708)

FIGURE 420. Mature male of O c t op u s defilippii, natural size. Note the form of mantle, funnel and
eyes. A short membrane connects the long arms. The hectocotylus bears a relaxed, spread groove (y) and
a small, indistinctly differentiated terminal part (z). The smaller drawing on the right shows the animal
behind a stem of Myriozoum, to which it appears adapted. The figure on the left shows it "stilt-walking"
on the proximal parts of the arms. The stalked eyes are distinct. The main drawing is based on a freshly
killed animal, x — entrance to spermatophore groove.

The conditions in the mantle cavity are typical and resemble those in
O. vulgaris . The jaws are very delicate and small, and the radula (after
Jatta, Plate 24, Figure 7) is simplified by suppression of the accessory
cusps, also in the median rows. The hectocotylus shows a varying number
of suckers so that a norm cannot be given. There are usually 70 to 90, but
30 and 104 may also occur. A hectocotylus with less than 100 suckers is
probably regenerated, and this is certainly the case in those with only

684

30 suckers. There is apparently never a completely intact arm apparatus
because of the tendency to autotomy which leaves only stumps that are later
rapidly regenerated. However, the hectocotylus is usually not discarded;

I have not found a single case of loss of the hectocotylus. This important
arm shows evidence of regeneration only as a rare exception. Its apex
resembles that of O. vulgaris, i. e. it is typical and the spoon and
beak are very small and indistinctly differentiated (Figure 405 on p. 683);
it is usually less than 3 mm long, never longer than 4 mm.

The coloration resembles that of O. vulgaris, with which O. defilippii
is closely related; the variations of the coloration are also similar. There
are grayish, yellowish and reddish brown tones, often with a greenish
iridescence around the eyes and in other areas. The darker areas often
form a netlike pattern, dark transverse stripes on the arms, spots on the
dorsal surface, etc. The warts are transient; only those above the eyes
are constant. Skin warts are often present in large numbers and of
different size and dense on mantle, head and arms (Jatta, 1896, pp. 223 —
224). The skin is soft and especially loose on the body so that the surface
is very variable. This species is small, rarely longer than 5 cm without
arms, at least in Naples.

4. JUVENILE STAGES

I consider as belonging to this species a number of small, delicate larvae
from the plankton in Naples, which otherwise resemble those of O. vulgar-
is and do not require a special description. I also have some young stages,
about 1.5 cm long (without arms), which already resemble the adults except
for the presence of larval chromatophores as in Plate IX, Figure 3 and the
relative short arms with fewer suckers. If they are 2.5 cm long (without
arms) they resemble the adult in all characters.

710 g. SPECIES OF "SC AEURG US"

As stated above (p. 691), the presence of the hectocotylus on the left side
does not characterize a natural group of Octopodidae. However, we placed
the following species of Octopus in a provisional group, because these
forms are perhaps the nucleus of a natural subgenus which is diagnosed
provisionally as follows:

Hectocotylus usually on the left side, with short, wide, typically
differentiated end part with a distinct spoon. Skin warts constant in their
greater part, dense on mantle, head and bases of arms.

This group includes the two following species and a few exotic forms with
a left hectocotylus, and also Octopus hoylei Berry, 1914, with a
hectocotylus on the right side (p. 691). A more exact definition of the group
cannot be given and the group is only mentioned because the name
Scaeurgus (Troschel, 1857) has been used for the following species.

This is perhaps the central group of the genus and therefore of the family,
but current knowledge does not permit its being given a definite rank,
perhaps as a subgenus.

685

FIGURE 421. End part of hectocotylus of Mediterranean species of the Scaeurgus
group, natural size, a) Octopus (Sc.) tetracirrus; b) Octopus (Sc.)
unicirrus. 1— "spoon"; 2 — beak (copulatory apparatus); 3 — spermatophore groove.
The typical formation of these parts in the Octopodidae appears here in its simplest
form, but as a mirror image of the normal.

h. OCTOPUS ("SCAEURGUS") TETRACIRRUS
(Delle Chiaje, 1829)

1. DIAGNOSIS

Skin swollen and gelatinous. Funnel gland always divided into two
V-shaped parts. (Duter side of arms with a distally tapering margin,
strengthened by longitudinal muscles at the margin, which is forked at the
base into the margin of the interbrachial membrane on each side. Two
warty "cirri" behind each other always above each eye. Coloration bright
orange- yellow to orange-red in life. Length 3 — 9 cm without arms.

2. LITERATURE

1829 Delle Chiaje (Vol. IV, Plate 72), Octopus tetracirrhu s.

1835 Ferussac and d’Orbigny (p. 36), Octopus tetracirrhus.

1841 Delle Chiaje (Vol. I, p. 4, Vol. V, p. 65), Octopus tetracirrhus.
1844 Philippi (p. 202), Octopus tetracirrhus.

1849 Gray (p. 11), Octopus tetracirrhus.

1851 Verany (P.25, Plates 7 and 7bis), Octopus tetracirrhus.

1855 d'Orbigny (p. 175), Octopus tetracirrhus.

1858 Adams (p, 20), Octopus tetracirrhus.

1879 Tryon, (p. 119), Octopus tetracirrhus.

1880 Tiberi (p. 12), Scaeurgus tetracirrhus.

1886 Hoyle (p. 14), Scaeurgus tetracirrhus.

1887 Fischer (p. 334), Scaeurgus tetracirrhus.

1890 Carus (p. 461), Scaeurgus tetracirrhus.

1896 Jatta (p. 230, Plates 5, 25), Scaeurgus tetracirrus.

1915 Naef (p. 17), Scaeurgus tetracirrus.

1921 Naef (p. 538), Scaeurgus tetracirrus.

711 3. STRUCTURE OF ADULT ANIMAL

The habitus of this species is so characteristic that it cannot be confused
with any other species after seeing it alive (Figure 422 and p. 622). The
colored drawing of Merculiano (Jatta, 1896, Plate 3, Figure 2) reproduced the
natural color correctly, but the position of the arms is that of a dead
animal. My drawings are based on live animals and show a more natural
form.

The arms are never intact and always injured or regenerated; they are
of uniform structure and of about the same length. I cannot give further

686

details from the 3 fresh specimens at my disposal. The arms are very
short and thick, especially in the young specimen shown in Figure 424;
they are about 2.5 times as long as the body in the adult. The interbrachial
membrane is very characteristic and a similar structure is present only
in the related Octopus hoylei Berry.

FIGURE 422. O c t o pu s ( S c a e u r g u s) tetracirrus. Mature male, alive, dorsal, natural size. Note the
typical posture, the characteristic form of the soft body, which is covered with small warts. Note the convex
eyes with upper lid and pseudocornea, the 4 cirri above the eyes, the form of funnel and arms. The hecto-
cotylus is on the left. The interbrachial membrane is strengthened at the margin and is connected with the
margins on the outer side of the arms.

The very distinct marginal muscles of the interbrachial membrane do
not continue on each side on the outer edge of the arm but they are fused on
the dorsal surface. From their point of union extends a common muscular
cord which passes in the median line of the arm. Contraction of this
muscle raises a broad skin fold (Figure 422). This may be the same
margin which occupies the posterior outer edge of the arm in other species
and has become connected with the margin of the membrane which
otherwise occupies the anterior edge of the arm (cf. Figure 413 on p. 699).

The funnel glands are always divided into two V-shaped parts, as in
most species of the group (Jatta, 1896, Plate 25, Figure 12). Jaws and
712 radula are more or less typical (loc. cit.. Figures 6, 7). The opened

mantle cavity of the mature male shows the conditions shown in Figure 423.
The translucent venous appendages are finely grapelike in both sexes and
do not consist of large fragments. The genitalia are very large and displace
the viscera of the posterior part of the body. The projecting end part of
the genitalia with the "penis diverticulum" is also very large, which is

687

probably connected with the large size of the spermatophores. The hecto-
cotylus is typical and formed on the left side. The spoon has shallow
transverse grooves on the bottom (Figure 421).

FIGURE 423. Mantle cavity of a mature male of
Octopus (Scaeurgus) tetracirrus, natural
size. Compare with Figure 410 on p. 688 and
Figure 422 on p. 711. The large gonad fills most
of posterior part of the body, displacing the renal
sacs (3) anteriorly. The genitalia are very large
(1, 2, 4, 5, 6). The branchial hearts are displaced
laterally and anteriorly:

1 — "penis"; 2 — "penis diverticulum"; 3 — renal
sac; 4, 5 — spermatophore gland; 6 — accessory
sperm at ophore gland.

FIGURE 424. Young O. t e t r a c i r r u s with
opened mantle cavity. 6x. The characteristic
appearance is caused by the thick, swollen skin,
which makes the differentiation indistinct, and
by the very short (contracted) arms. Note the
projecting inner margin of the funnel, left
funnel pocket, funnel retractors, stellate ganglia,
gills (branchial glands, gill ligaments, branchial
hearts), renal papillae, median mantle adductor
(cut at the mantle), and the small indentations
on the inner side of the mantle, into which the
inner margin of the funnel fits.

Especially characteristic are the coloration and the texture of the skin.
Jatta (Plate 5, Figure 2) published a good reproduction of the coloration,
which is bright and uniformly orange-yellow to orange- red. A greenish
tone is present above the eyes, and the eye is light yellow inside the lid.

713 The skin is swollen, gelatinous and so loose that the slightest pressure
deforms it (the name in Naples is therefore ’’Purpo incammisato"). The
surface bears numerous small tubercles in a more or less regular

688

arrangement on mantle, head and bases of arms (Figure 422). Lateral
lines are absent. Two warty processes ("cirri") are always present on
the dorsal side of each eye. O. tetracirrus is not large; the largest
specimens are 8 — 10 cm long without arms.

4. JUVENILE STAGES

The young animals are typical for the family. However, the skin is '
swollen already in the smallest specimens. This and the characteristic
coloration make identification easy. The arms remain short for a longer
time than in the preceding forms (cf. Figures 401 and 424). The mantle
cavity of young animals is normal.

i. OCTOPUS ("SC AEURGUS") UNICIRRUS
(Delle Chiaje, MS) d'Orbigny, 1839

1. DIAGNOSIS

Skin tough, densely covered with small warts also on ventral side of
mantle. Mantle sac with distinct lateral lines. Interbrachial membrane
continuing on outer edges of arms. Only one distinct wart ("cirrus") above
each eye. Coloration in life reddish brown with greenish and orange tones.
Length about 7 cm without arms.

2. LITERATURE

1838 Delle Chiaje (MS. in.Ferussac), Octopus unicirrhus (cf. Jatta, p. 234).

1839 Ferussac and d’Orbigny, Octopus unicirrhus (cf. also O. c u v ie r i , Plates 3 — 4, partly
belonging here).

1849 Gray (pp. 7, 19), Octopus vulgaris (part.).

1851 Verany (pp. 22, 25, Plates 12 and 12 bis), Octopus cocco.

1855 d’Orbigny (pp, 169, 192), Octopus vulgaris (part.).

1857 Troschel (p. 57), Scaeurgus coccoi (= O. u n i c i rr hu s).

1858 Adams (p. 19), Octopus cocco.

1869 Targioni-Tozzetti (p. 21), Octopus unicirrhus.

1879 Tryon (p. 127) (?), Scaeurgus coccoi.

1880 Tiberi (p. 12), Scaeurgus unicirrhus.

1886 Hoyle (p. 14), Scaeurgus unicirrhus.

1890 Carus (p. 461), Scaeurgus unicirrhus.

1896 Jatta (p. 234, Plates 3, 25, 26), Scaeurgus unicirrhus.

1916 Naef (p. 17), Scaeurgus unicirrhus.

1921 Naef (p. 538), Octopus unicirrhus.

689

3. STRUCTURE OF THE ADULT ANIMAL

714

Except for the coloration, the general appearance of this species shows no
special characters (cf. Jatta, Plate 3, Figure 2). The habitus is typical for
the family but there is a special resemblance toEledone cirrosa.

FIGURE 425. Octopus (Scaeurgus)
uni cirrus, mature male, ventral, natural
size. Note the form of the mantle.

Lateral lines visible in anterior part of
mantle. Small warts cover the mantle.
Note also the relative development of the
arms, the attachment of the interbrachial
membrane and the structure of the hecto-
cotylus (cf. Figure 412 on p. 694).

The arms differ only little in length; the two lateral pairs are slightly
longer than the median pairs and bear more numerous suckers, 160 — 165
as against 150 — 155 (Figure 425). The interbrachial membrane is well
developed. It is deeply incised on the ventral side, and extends in the form
of fin margins far along the ventral outer edges of the arms, to about half
of the length of the arms distal to the interbrachial incisions. The funnel

690

gland is sometimes W-shaped; sometimes it is divided into two V-shaped
parts, as in O. tetracirrus. The mantle cavity shows the typical
conditions, but the translucent venous appendages resemble those of
O. tetracirrus. Radula and jaws are typical, but the jaws show a
characteristic distribution of the more strongly colored and chitinized
zones (Figure 10, Plate XVIII) and the biting process of the upper jaw is
markedly curved. The hectocotylus of the specimen illustrated bears 81
normal suckers, then a typically differentiated terminal part with a very
long and fully developed spout (Figure 421 on p. 710) and a strongly convex
spoon with a smooth floor.

715 The coloration of the live animal is bright yellowish brown to reddish
brown with greenish tones (Jatta, Plate 3, Figure 2). The colors are
irregularly cloudy and spotted, in contrast to O. tetracirrus. Light
lines pass along the outer and inner edges of the arms. The whole dorsal
surface of mantle, head and arms is densely covered with tough, rounded
warts. In contrast to the preceding species, the warts are constant and are
isolated on the plane surface. Similar warts are also present on the ventral
side of the mantle (Figure 425). There is only one "cirrus," which
corresponds to the posterior cirrus of O. tetracirrus, above each eye.
Larger but transient skin warts are also present on the dorsal side.

A typical lateral line passes along each side of the mantle, often irregularly
and interrupted, and it disappears posteriorly and toward the anterior
margin.

4. JUVENILE STAGES

I do not have or cannot identify early planktonic and benthic young stages
of this species, which is not surprising because of their typical character.
Young animals a few centimeters long already closely resemble the adults
except for the shorter arms.

691

716 Chapter 52

GENUS ELEDONE
Leach, 1817

(Moschites Schneider, 1784; Hoyle, 1901)
a. DIAGNOSIS

Suckers uniserial on all arms, forming regular zigzag rows in some
places temporarily. Eggs very large (about 6 mm). Development direct.
Hectocotylus always on the right side. Interbrachial membrane not forming
broad margins (as in Velodona Chun) on ventral outer edges of arms.

The Mediterranean species show also the following characters: 1. Apex
of arms of mature male without normal suckers but with atypical elevations
which developed from the suckers. 2. Hectocotylus without a typical
terminal part with a spout and a spoon. 3. Radula typical for the family.

One might consider including Velodona togata Chun, 1914 in the
genus and deleting the last sentence of the diagnosis. However, the special
character of this form cannot be overlooked despite its resemblance to
El e d o n e .

The Eledone group probably developed from a typical form of
Octopus. The uniserial arrangement of the spermatophores is certainly
secondary. As other distinct characters are not present, the group may not
be a natural monophyletic unit. (Indirect development connected with the
large size of the eggs and the heteromorphous ends of the arms of the male
occur also in species of Octopus; cf. p. 692.) However, the Mediterran-
ean species are certainly more closely related to each other than to any other
species of Octopus.

717 b. ELEDONE MOSCHATA (LAM., 1799) LEACH, 1 817
1 . DIAGNOSIS

Mantle sac without lateral ridges, with transient skin warts on the dorsal
side and with a rather smooth ventral side. Normal arms of female with

* This is suggested by the alternative arrangement of the musculature at the base of the suckers; it permits
a regular alternating displacement of the suckers, which is particularly distinct in E. cirrosa, in which
this IS observed frequently on many of the arms, in live and dead animals. This character is therefore not
very important, not even as a generic character, and still less for larger groups. Cf., for example, Joubin
(1918, pp. 39 — 40), who established the name "Eledonidae” for the species of Eledone of the European
ccasts, together with Graneledone n.g. (with warty skin, and large spoon on the hectocotylus:

E. verrucosa Verrill), Bolitaena, Eledonella and V i t r e 1 e d on e 1 1 a .

692

about 100 suckers. Apical suckers of arms of male replaced by 2 rows of
transverse, flattened papillae. Hectocotylus with 60 — 75 normal suckers,
followed by a short, rugose terminal part (5 — 7 mm) without a spoon
separated from the spout. Coloration dirty grayish brown with dark spots
on the dorsal side in life, with a musky odor. Length about 12 — 14 cm
without arms.

2. LITERATURE

— Aristoieles (lib. IV, cap. I), Eledw^i
~ Plinius (lib. IX, cap. XXX), O z a i n a .

1533 Belon (p. 333), Eledona,

1554 Rondelei (lib. XVII. cap. VIII, p. 516; cap, IX, p. 417), Polypus tertia spec.

1558 Gesner (lib. IV, Vol. IV, p, 871), Polypus tertia spec.

1558 Bossuet (lib. IV, pp. 740, 871), Polypus tertia spec.

1606 Aldrovandi (cap. Ill, pp. 42, 43), Eledona, Bolitaena, Ozolis (?).

1758 Seba (III, Plate 2), Polypus fern in a.

1798 Lamarck (Vol. I, pi. 22), Octopus moschatus.

1802 Montfm (III, p. 80, Plate 34), Poulpe musque.

1802 Bose (I, p. 48), Sepia moschata.

1814 Rafinesque (p. 29), Ozaena moschata.

1817 Cuvier (Vol. 2, p. 8, 3^ ed.), Poulpe musque.

1817 Leach (III, p. 138), Eledone moschatus.

1820 Ranzani (II, p. 80), Eledone moschata.

1822 Lamarck (Vol. II, p. 363, 12© ed.), Octopus moschatus.

1824 Carus (p. 326, Plate 32),. Octopus moschites.

1825 Blainville (Vol. Ill, p. 365), Octopus moschatus.

1826 Risso (Vol. IV, p. 2), Eledona moschata.

1826 d*Orbigny (p, 145), Eledon moschatus.

1826 Payrandeau (p. 172), Octopus moschatus.

1827 Blainville (Vol. 43, p. 190), Octopus moschatus.

1829 Deile Chiaje (Vol. 4, pp. 43, 56, Plate 57), Octopus moschatus.

718 1829 Sangiovanni (p. 317), Octopus moschatus.

1833 Oken (Vol. V, Section I, p. 538), Sepia moschata.

1835/48 Femssac and d'Orbigny (p. 72, E led on s, Plate I bis, III), Eledon moschatus pars.
Also E. a 1 d r o V a n d i D. Ch. and E. g e n e i VeT.

1837 Rang (p. 64, Plate 91), Octopus moschatus.

1838 Potiez and Michaud (p. 7), Eledone moschatus.

1841 Cantraine (p. 19), Eledone moschata.

1841 Deile Chiaje (Vol, V, p. 66), Eledone moschata.

1844 Philippi (p. 202), Eledone moschata.

1849 Gray (p, 21), Eledon moschatus. Also E. a Id r ov a nd i D. Ch.

1851 Verany (p. 7, Plates 4, 5, 6), Eledon moschatus.

1855 d*Orbigny (p. 195), Eledon moschatus. (Also E. aldrovandi andE. genei.)

1858 Adams (Vol. I, p. 21, Plate I), Eledon moschata.

1863 Aucapitaine (p. 366), Eledon moschatus.

1869 Targioni-Tozzetti (p, 28), Eledone moschata.

1871 Woodward (p. 165), Eledone moschata.

1879 Tryon (p. 128), Eledone moschata.

1880 Stossich (p. 2), Eledone moschata.

1880 Tibet! (p. 12), Eledone moschata.

1884 De Rochebruoe (p, 159), Eledone moschata.

1885 Ninni (p. 159), Eledone moschata.

1886 Hoyle (p. 16), Eledone moschata.

1887 Fischer (p. 331), Eledone moschata.

693

(717)

FIGURE 426. Half-grown fe-
male of E led one m os-
chat a. 0.5 X. Note the
typical form of mantle, funnel,
head and arms, the slight dif-
ferences in the length of the arms,
arrangement of suckers and
form of interbrachial mem-
brane. Drawing of a live
animal in swimming position.

1889 Posselt (p. 139), Eledone moschata.

1890 Colombatovic (p. 8), Eledone moschata.

1890 Cams (p. 462), Eledone moschata.

1894 Pelseneer (p. 206), Eledone moschata.

1896 Jatta (p. 239, Plates 3, 7, 26, 27), Eledone moschata.
1901 Hoyle (p. Ill), Moschites moschata.

1908 Pfeffer, Moschites moschata (partly T. c i r ro s a)

1912 Naef (Not. 6. p. 749), Moschites moschata.

1916 Naef (System, p. 17), Moschites moschata.

1921 Naef (p. 538), Eledone moschata.

3. STRUCTURE OF THE ADULT ANIMAL

The form of an adult Eledone moschata
is typical for the family but the suckers are
uniserial. It differs from E. cirrosa in the
absence of lateral ridges. The coloration of the
live adult (see the good drawing by Merculiano
in Jatta, 1896, Plate 3, Figure 4) is mainly gray,
grayish brown to reddish brown, always with dark
brown to blackish spots on the dorsal side. Along
the outer edges of the arms passes a sky blue
line caused by dense iridocytes which is most
distinct on the ventral side and continues more
or less distinctly on the margin of the inter-
brachial membrane. The species is not very
large, but about twice as large as recorded by
Jatta (p. 243): the largest specimens examined
are 10 — 12 cm long without arms, their total
length about 4 times as long. The surface is
often quite smooth, except for the warts near
the eyes, but there may be dense warts of
different size on the whole dorsal surface.

Lateral ridges on the mantle are absent.

The arms are of about equal length;
formula: 1, 2, 3, 4. The interbrachial membrane
is typical, moderately long, and disappears
rapidly on the outer edges. There are about
100 suckers on each arm in larger specimens,
with slight variations. Their size increases
rapidly from the 8th or 9th sucker of each row
and then gradually decreases toward the apex.

The funnel gland is W- shaped. The mantle
cavity shows the typical conditions of the
Octopodidae in the "anlage" (Plate X, Fig-
ures 427—429). Both jaws have weakly
developed biting processes; and the upper jaw
is slightly curved in the dorsomedian line, while
the lower jaw has a characteristic distribution
of pigmentation (Plate XVIII, Figure 9).

694

719

Situs of mantle cavity of Eledone moschata. The muscular mantle is opened and more or less

spread, so that the organs become clearly visible.

FIGURE 427. Almost mature female,
in natural form and position. Note:
funnel, funnel pockets, funnel retrac-
tors, anterior mantle adductors,
stellate ganglia, gills, branchial
glands, gill ligaments, gill retractors,
branchial hearts, renal papillae,
distal parts of oviduct with papillae
of the opening, mantle adductor (cut)
and anal papilla. The genital papil-
lae are situated distinctly more
posteriorly than in Figure 408 on
p. 685.

FIGURE 428. Almost mature male.
The left gonoduct (Gn) extends
much further anteriorly and ends in
a very long and thin genital pro-
cess (Pn) which can apparently be
introduced into the point of entry
of the spermatophore groove on
the hectocotylus (compare with
Figure 408 on p. 685 and Fig-
ure 423 on p. 712). A "penis
diverticulum" is apparently absent.
Ol, U1 — upper and lower lip of
anal slit; Si — wings on anal slit;
Ad — insertion of median mantle
adductor (cut at Sp); Np — renal
papilla; Kr — gill retractor; Kh —
branchial heart; Kv — branchial
vein; Km — branchial gland;

Kb — gill ligament; Tr — funnel
retractor.

FIGURE 429. Young male with
still undeveloped genital process
(3) at the end of the gonoduct (2)
which otherwise grows along the
body. 1 — superficial gill
retractor.

695

The radula (Plate XVI, Figure 4) is typical for the family. The mantle
cavity of the maturing animal shows some special characters in the
development of the genitalia (see Figures 427, 428 and 429). The oviducts
are relatively thick (cf. p. 684), which is connected with the large size of
the eggs, and the short free papillae of the opening are situated far behind
the anal opening (cf. p. 685). On the other hand, the male gonoduct is thin,
with a long, very narrow, terminal part (genital process) which can
apparently be introduced directly through the funnel opening into the point
of entry of the hectocotylus (p. 683). This is definitely impossible in many
other species (p. 685), so that copulation must take place through the funnel
opening.

Hectocotylization consists of: 1) formation of the hectocotylus;

2) modification of the apex of all arms; 3) enlargement of the largest
suckers on the arms.

720 1 . The hectocotylus bears about 60 — 75 (average 70) normal suckers; it has

a structure at the end, the typical differentiation of which is still indicated,
but only in well preserved specimens (Figure 405 on p. 6 83). The
spermatophore groove is directed normally to the middle of the inner side
of the arm and continues in a groovelike depression which apparently
corresponds to that of the typical "spout." The apex of the spout is raised
only slightly, forming a small tubercle at the apex so that no space is left
for a spoon. However, careful preparations show a possible remnant of a
spoon. It is rugose and therefore difficult to examine.

FIGURE 430. End of arms of a mature male of E led one
moschata. a) hectocotylus; b) normal arm. The
hectocotylus (a) still shows indistinctly the normal differen-
tiation at the apex, with a rudimentary spoon (1) and a spout
(beak) (2, 3) which almost reaches the apex. In (b) the
proximal suckers are transformed into flat transverse papillae
or lamellae (7), each equivalent to half a sucker. Transverse
ridges (6) connect these papillae. The lamellae pass (5) into
normal "anlagen" of suckers at the apex (4) (cf. Figure 396
on p. 667). 8 — spermatophore groove. 4x.

2. The arms of the half- grown male form no further normal suckers at
the apex, but the "anlagen" of the suckers become separated transversely
and the depression which probably represents the "anlage" of the opening

696

of the sucker becomes a thin transverse groove which comes to be divided
into 2 parts, together with the "anlage, " so that 2 rows of transverse
papillae or lamellae are formed which remain connected in pairs by a low
ridge. The papillae are more or less in the form of a shoe. They are
situated with their narrow base on the arm, widen slightly laterally and form
a narrow sole with a thin groove which is directed toward the inside of the
arm. The first formed (proximal) papillae still resemble suckers, or the
last formed suckers already resemble the papillae. Transitional forms
and irregularities are frequent.

3. All arms of the mature male have enlarged suckers near the base.
This late modification is especially distinct at about the 4th— 12th sucker.
This is least marked on the dorsal arms.

721 4. YOUNG STAGES

The freshly hatched animal (Figure 399 on p. 676) resembles a typical
young form of Octopodidae after the metamorphosis. This specimen is
deformed because of preservation in formol (cf. Figure 402 on p. 679). The
coloration is reddish brown, darker dorsally than on the ventral side.

Warts or lateral ridges are absent but remnants of the spines or bristles
are still present and are not completely absent also in older stages. The
young form gradually attains the definitive form by growth of the mantle sac
and increase of the relative length of the arms. At a length of a few
centimeters, the young animal already closely resembles the adult. The
mantle cavity of the young forms (Plate X, Figure 1) shows no special
characters: the gills are relatively smaller and less compact than in the
adult (p. 666), i. e. they resemble the gills of Decapoda. The base of the
gills covers the weakly indicated renal papilla.

c. ELEDONE CIRROSA (LAM., 1798)

D'ORBIGNY, 1839

1. DIAGNOSIS

Mantle with distinct lateral ridges (or narrow margins). Dorsal surface
densely covered with warts, which never disappear completely. Female with
more or less equally long normal arms with more than 120 suckers. Arms
of mature male with transversely compressed rudimentary suckers which
are more or less laterally produced, like cirri. Hectocotylus with about
70 suckers and a formation at the end which corresponds to the "spout" of a
typical hectocotylus but is not followed by a spoon.

2. LITERATURE

1606 Aldrovandi (p. 14, Plate 14), Polypus "in quo una tantum acetabulorum series exprimatur. "
1799 Lamarck, Octopus cirrhosus.

697

1802
1814
1820
1827
1827
1829
722 1829

1839

1839

1841

1841

1844

1849

1849

1851

1855

1855

1860

1869

1879

1880
1880

1884

1885

1886

1887

1889

1890
1890

1896

1897
1901
1908
1912
1916
1921
1921

Montfort (p. 55, Plate 32, p. 67, Plate 33), Ozaena aldrovandi.

Rafinesque (p. 29), Ozaena aldrovandi.

Ranzani (Dec. I, p. 81), Octopus aldrovandi.

Blainville (Vol. 43, p. 191) (?), Octopus aldrovandi.

Grant, Octopus ventricosus.

Sangiovanni (p. 315), Octopus leucoderma.

Delle Chiaje (Vol. 4, pp. 45, 57), Octopus aldrovandi. Assumes a closer relationship to
O. cirrosus.

Ferussac and d'Orbigny (pp. 73, 77, 78), Eledon moschatus (pars).

Ferussac and d’Orbigny, Eledon cirrhosus.

Delle Chiaje (Vol. 1, p. 5; Vol. 5, p. 66), Eledone aldrovandi.

Cantraine (p. 20), Eledone aldrovandi.

Philippi (p. 202), Eledone aldrovandi.

Gray (pp. 21, 22), Eledone a Id r o v a n d i . (pars).

Gray, Eledone octopodia.

Verany (pp. 12, 15, Plates 1 — 3), Eledon aldrovandi and E. genei.

(1845) d'Orbigny (pp. 195, 197), Eledon moschatus (pars).

(1845) d’Orbigny, Eledon cirrhosus.

Steenstrup (p. 232), Heledone aldrovandi.

Targioni-Tozzetti (p. 29), Eledona aldrovandi.

Tryon (p. 129), E led on a aldrovandi.

Stossich (p. 158), Eledona aldrovandi.

Tiberi (p. 13), Eledona aldrovandi.

Rochebrune (pp. 159, 160), Eledone aldrovandi, E. cirrosa and E. genei.

Ninni (p. 159), Eledone aldrovandi.

Hoyle (pp. 15, 102), Eledone cirrosa. Considers correctly E. a Id r ov a nd i , E. genei as
synonyms of E. cirrosa. Compare also: Notes, p. 278.

Fischer (p. 331), Eledone aldrovandi.

Posselt (pp. 139, 140, 240), Eledone a Id r o v a nd i and c i r r osa .

Norman (p. 468), Eledone aldrovandi and cirrosa. (?).

Cams, Eledone aldrovandi and cirrosa.

Jatta (p. 243, Plates 5, 27, 28, 29), Eledone a Id r o v a n d i and c i rr os a .

Hoyle, Eledone aldrovandi and cirrosa.

Hoyle, Moschites aldrovandi and cirrosa.

Pfeffer, Moschites cirrosa (pars) (Pfeffer confuses E. m osc ha t a with this species).

Naef (Not. 6, p. 750), Moschites cirrosa.

Naef (System, p. 17) , Moschites cirrosa.

Grimpe (pp. 298, 301), Eledone cirrosa (North Sea).

Naef (System, p. 538) , Eledone cirrosa.

3. STRUCTURE OF THE ADULT ANIMAL

Merculiano illustrated the live animal in natural colors (Jatta, 1896,
Plate 5, Figure 1). The dorsal side is darkest in the middle and the colors
vary between orange -yellow, orange -red and reddish brown. The ventral
side of the mantle has a greenish iridescence and is usually very light,
sometimes partly orange-yellow. The lateral lines form whitish ridges
or very narrow margins. The dorsal side is covered with numerous small
723 warts (which are also present on the ventral side) and some larger warts.
These are transient, except those above the eyes, which never disappear
completely. A distinct musky smell is not recognizable. The adult animal
grows to about 12' cm or more (without arms). Its form is typical for the
Octopodidae, without any special characters. The arms are of about equal

698

length, with a normal interbrachial membrane. Each arm of the adult
bears more than 120 suckers which rapidly increase in size to the 6th
and then just as rapidly decrease in size to about the 9th or 10th; they
become smaller very gradually toward the apex.

(721)

FIGURE 431. Half-grown male of Ele d one c irr os a , natural
size. Note the typical habitus of Octopodidae, the form of the
hectocotylus (y), the more or less equally long arms, uniserial
suckers and lateral ridges (slightly displaced ventrally) on the
mantle sac:

m — superficial outer funnel adductor; x — point of entrance to
spermatophore groove; y — spermatophore groove; z —
terminal part of hectocotylus.

The funnel gland is W- shaped. The situs of the mantle cavity
(Figure 432) shov/s normal conditions. The form of the jaws is shown on
Plate XVIII, Figure 8. The superficial gill retractors differ markedly
from the type (Figure 432 on p. 722); in contrast to all other Octopodidae,
they are united in the middle and inserted together on the mantle. The
radula is typical (compare the inexact drawing in Jatta, Plate 28, Figure 4).

699

i

(722)

FIGURE 432. Situs of mantle cavity of a half-grown male ofEledone cirrosa, natural size. Note the
connection of the funnel pockets and the anterior mantle adductors with the mantle (see Figures 428, 429 on
p. 719):

1 — branchial gland; 2 — branchial ganglion; 3 — branchial retractors, fused at (4), which is characteristic
for the species; 5 — penis diverticulum (cf. Figure 408 on p. 685, and Figures 423, 424 on p. 712); 6 — vena
cava; 7 — median mantle artery; 8 — lateral mantle vein.

FIGURE 433. Apex of arms of the adult male of Eledone cirrosa. 4x.

a) Hectocotylus, without spoon (cf. Figure 405 on p, 683 and Figure 430 on
p. 720), but with a well -developed beak (1). 2 — longitudinal groove.

b) Normal arm, on which the suckers form transversely compressed forma-
tions (6) which resemble the typical "anlagen" (p. 667) and the lateral parts
of which (5) project more or less like "cirri" which are distinct from the
suckers (4). Normal "anlagen" of suckers (3) are present at the apex.

Genitalia of the mature female: The projecting free parts of the oviducts
are rather long and pointed at the end (cf. E. moschata. Figure 427 on
p. 719). In contrast toE. moschata, a small penis diverticulum is
distinct on the male gonoduct. Hectocotylization is in general as in
E. mo sc hat a (p. 719): 1. The hectocotylus is situated on the left side,

(however, Appellof, (1893) observed also bilateral development). It bears
about 70 normal suckers; the terminal part resembles the spout of the
normal hectocotylus of Octopodidae but a spoon is absent, i. e. it has been

700

lost (Figure 433). This resembles the condition in E. mo s chat a in which
the spoon is still an indistinct rudiment (Figure 430 on p. 720), 2, The
normal arms bear reduced, characteristically modified suckers at the
apex which form again almost normal "anlagen" at the extreme end from
which they apparently developed. They are transversely compressed, with
724 a reduced opening and form characteristic lateral processes which are like
cirri in the largest specimens. These structures are only supposedly less
developed in Mediterranean specimens than in the northern material
because adult specimens from the Mediterranean are difficult to obtain.
Posselt (1898) therefore incorrectly distinguished the Mediterranean
E. aldrovandi from the northern E. cirrosa on the basis of these
characters; he even referred an especially large specimen from Nice to
the northern species. The "cirri" are more or less distinctly differentiated
from the sucker and are probably homologous to the lamellae or papillae
of E. m o s c h at a . 3. The proximal suckers are markedly enlarged on all

arms. They are least enlarged on the dorsal arms. The largest sucker is
about the 6th in the row, as in the female, and the suckers are enlarged to
about the 10th.

E. cirrosa is not as closely related to E. m o s c h a t a as might be
assumed. This is proved mainly by the structure of the spermatophores in
the two species (see description below).

4. YOUNG STAGES

I have no freshly hatched animals of this species but they probably
resemble those of E. moschata, except perhaps in the distribution of
the chromatophores and in coloration, which is probably yellowish brown
to orange-red, i. e. lighter than inE. moschata.

A young animal, 12 mm long without arms, was caught at a depth of
200 m at the Amontatura. This specimen differs little from that shown in
Figure 399 on p. 676. The chromatophores are lighter, reddish brown, and
much less numerous. In contrast to E. moschata, the arms bear a
distinct single row of large chromatophores. This row is still present
much later, when the animal is about twice as large. Lateral ridges are
still absent, as in E. moschata, but they soon appear (at a length of
2 cm), at least temporarily, and leave no doubt regarding identification.

At the same time the skin warts become constant (these are transient in
the very young stages), and the coloration assumes its definitive character.
The arms grow slowly, so that young forms have short arms for a long
time; only half-grown specimens show more or less the definitive
proportions.

701

725 Chapter 53

FAMILY ARGONAUTIDAE
Naef, 1912

(Philonexidae d'Orbigny, 1845)
a, DIAGNOSIS

Nektonic Heteroglossa. Arms with biserial suckers. Funnel connected
with the mantle by a solid bond, which may be replaced by fusion. Mature
male markedly smaller than female; one arm of 3rd pair markedly modified
into a hectocotylus. Hectocotylus, with one or several spermatophores,
detached during copulation. It survives in the mantle cavity of the female
and fertilizes the eggs actively.

b. DEFINITION AND POSITION

I extended and defined the family of d'Orbigny in 1912, and placed in it the
genera Alloposus, Tremoctopus, Ocythoe and Argonauta,
because of their typical resemblance. The rank of family given to this unit
is based on the structure of the system. A higher rarik would necessitate
placing it opposite the Octopodidae within the Heteroglossa (etc., see p. 673),
which would be an unjustified complication of the nomienclature. It might
be justified to give each genus the rank of family, but this would require
reformulation of the relationships between them, which are described below.
However, it seems more important to stress the resemblances within the
group than the differences between them, because the relationships are very
interesting and characteristic.

c. TYPICAL STRUCTURE

Only a few species of this very diverse group have survived, but its ideal
ancestral form (p. 680) has to be assumed to resemble Octopus.
Alloposus, the least aberrant genus, shows this resemblance to the
Octopodidae in its general habitus and also in some very special characters,
726 The Argonautidae have no trace of an inner shell, at least in the adult.

These remnants are quite small and are early reduced already in the group
from which they developed.

702

The arms are always distinctly different. The dorsal arms are markedly
longer than the laterodorsal arms ab origine and these are markedly longer
than the lateroventral arms. The relative length of the ventral arms varies.
All arms bear two rows of alternating suckers with very extensible stalks
which are often pillarlike. The suckers in each longitudinal row are
connected by folds which are attached on the outer side (Figure 449) and
form a structure which resembles the protective margins of the Decapoda,
but it is not equivalent to them morphologically. It may disappear and is
formed by muscles from one sucker to the other; contraction of these
muscles raises the elastic skin into folds.

The eyes are large, very prominent, round in lateral view. The iris
opening is also round, in contrast to the benthic Polypodidaev''

FIGURE 434. Diagrammatic cross section through the arm of
Ocythoe, female. 2x. Note the round muscular axis of the
arm (1); the connective tissue and skin around the axis; the
differentiation of the skin into a sucker, a stalk of a sucker,
and a connecting fold (6) at the position of the other row in
which the section passes in the interval between 2 suckers.
Note also the arrangement of the musculature of the sucker
(4, 5, 8) and that of the stalks. 2 — longitudinal fibers;

3 — transverse fibers; 7 — chromatophores; 9 — marginal ring.

The lid apparatus is less complete than in the Octopodidae. The
"pseudocornea" still has the character of a lower lid (Figure 449 on
p. 753); it can be opened so widely that seawater reaches the lens when
the upper lid is correspondingly relaxed. Such a condition is never found
in well preserved specimens of Octopodidae (Figure 387 on p, 660).

The funnel gland is W-shaped, as in most species of Octopus. This
basic form undergoes a partial modification within the family (p, 672), On
the other hand, the connection between funnel and mantle is markedly
strengthened. Tremoctopus shows the basic form of this connection
which is a direct further development of the conditions which begin to
develop in the Octopodidae. However, the funnel corners of the Octopodidae
enter only slightly into shallow grooves in the mantle and are attached on a
low swelling (Figure 392 on p. 664), while a more solid connection developed
in the Argonautidae: the groove is deepened into a pocket, and the funnel
corner is curved into a hook, firmly attached by adhesion. The various
modifications can be derived from this form. This also explains the lesser
development of the mantle adductor, the function of which is replaced by the
727 "apparatus of closure." The whole change has to be interpreted as an
adaptation to nektonic life, like the particularly strong muscular mantle.

The following characters of the mantle cavity may be considered as
typical for the Argonautidae: 1. The origin of the median mantle adductor
is restricted to its anterior part near the anus (Figure 408 on p. 685 and
Figure 450 on p. 754); there is therefore a remnant of the mantle septum
in the whole posterior part of the mantle cavity only in the form of the
posteriorly disappearing attachment of the median mantle adductor on the

* The benthic forms of the Decapoda tend to have a transverse, slitlike pupil, while the nektonic species of
the deeper water (Heteroteuthis in the Sepiolidae, the Oegopsida in the Teuthoidea) have a round pupil.
This is a direct adaptation to the environment (pp. 174, 533, 567).

703

mantle. This permits greater freedom of movement of the mantle
during swimming (I). 2. The superficial gill retractors radiate in the
ventral skin (Figures 450 and 466) and do not reach the posterior end as in
the Octopodidae (Figures 427 — 429 on p. 719). 3. The ventral skin which
borders on the mantle cavity bears isolated chromatophores, which are
absent in the Octopodidae (Figures 450 and 470). These chromatophores
can only be of use to the half- transparent young forms but they persist in
the adult. 4. The genital openings are situated more or less close to the
base of the gills (Figures 450 and 466), which covers the male openings
completely (Figure 470).

The jaws (Plate XVIII, Figuresll, 1 2) are relatively delicate, slightly
convex, with weakly differentiated biting processes. Their posterior parts
are unpigmented, transparent in fresh specimens and very short in the upper
jaw. The radula varies markedly (Plate XVI). Ocythoe (Figure 8)
resembles the typical conditions in the Dibranchiata most closely;
Alloposus also shows primary characters. Two regular, symmetrical
lateral cusps on the teeth of the median row are characteristic. Accessory
cusps are absent. The marginal plates are well developed.

FIGURE 435. Diagrammatic longitudinal section through the connection between funnel corner and mantle
in the Octopoda. a) Typical condition (Octopus); the funnel corner extends only slightly into an
indentation of the mantle, the margin of which is thickened, b) Alloposus; the funnel corner is fused
with the mantle, c) Trem octopus; the funnel corner forms a hook which extends into a pocket of the
mantle (cf. Plate X, Figure 4). d) Ocythoe; the funnel corner is coiled into a snail-like knob and is
firmly enclosed in a corresponding depression of the mantle, e) Argonauta; the funnel corner is hooked
into the mantle; it is differentiated like the funnel bond of the Decapoda and is curved around a tubercle of
the mantle like a snap button.

The sexual dimorphism is of particular importance for the characteriza-
tion of the group. Hectocotylization or external modification of the arms of
the male (p. 86) is a general phenomenon in the Cephalopoda and so are

728 differences in the proportions (habitus) of adults of both sexes. However,
nowhere have these changes attained such an extreme degree as in this
family, which are the true "Hectocotylifera." The sexes differ so markedly
in size that it is difficult to believe that they belong to the same species.
The different size of the sexes and the divergent structure of the whole arm
apparatus of Argonauta (Plate XI) are so marked that nobody at first
would place both sexes in the same family. Closer examination, however,
shows very characteristic resemblances on the basis of which the
morphologist can establish the relationship even without direct observation
of the actual sexual relationship.

704

Especially characteristic is the genital arm, which always belongs to the
3rd pair and is homologous to that of the Octopodidae. Its original form may
be assumed to be as follows: One VL arm became detached from the
interbrachial membrane and was situated free between the directly
connected DL and V arms; it formed a short but otherwise normal basal
stump with a few suckers. Between the stump and the arm is a constriction
which has to be considered as the predetermined site of autotomy
(Figure 436 on p. 732). This arm is longer than the others. As in the
Octopodidae, there is a groove along its ventral outer edge which may be
widened in some places and may form a reservoir for one or several
spermatophores. The arm, which bears 2 rows of more or less normal
suckers, ends in a cylindrical, hollow process, the "penis" (Figure 436 on
p. 732); its canal is a continuation of the spermatophore groove and is
probably formed by fusion of the margins of an adjacent groove. This arm
is not free but is situated in a pocket between the 2nd and 4th arms and is
coiled in it; the interbrachial membrane is contracted above it, forming a
sac which can expand posteriorly under the skin and protrude externally
as a swelling below the eye (Figures 446 and 46 8). Such a hectocotylus can
be derived directly from the hectocotylus of the Octopodidae (p. 683). The
"penis" is therefore homologous to the copulatory apparatus (beak), while
the spoon is displaced laterally and reduced (but see p. 749). This
characteristic structure, which is further modified in all species of the
family, is essentially a secondary complication of the hectocotylus of the
Octopodidae. The hectocotylus is detached by autotomy during copulation
and remains alive in the mantle cavity of the female, where it fertilizes the
eggs in the oviduct. The penis probably enters the oviduct and ejects the
sperm from the spermatophore. Parts of the penis have been found in the
proximal part of the oviduct of Argonauta. Because of this internal
fertilization, the eggs can develop further in the long, winding oviduct
which functions as an uterus (see Argonaut a and Ocythoe).

729 The other arms are normal, but absolutely and relatively shorter than
those of the female and bear markedly fewer suckers. This and their
smaller size gives them the character of inhibited growth.

The coloration of the Argonautidae is bright and iridescent. Blue and
violet metallic tones give them a striking and colorful appearance. The
skin is smooth (but see Ocythoe). These characters can be derived from
those of the Octopodidae and have to be considered as adaptations to nektonic
life. The presence of several hectocotyli in the mantle cavity of females
proves that the dwarf males are far more numerous than the females,
which is very useful in view of the difficulty of finding a specimen of the
other sex in not numerous nektonic -pelagic animals and also because of
the higher mortality of the females during their longer development (p. 730).

d. POSTEMBRYONIC DEVELOPMENT OF THE
ARGONAUTIDAE

The sexes do not differ at hatching also in the extreme forms
(Argonauta), and the sex of a freshly hatched animal has therefore to
be determined by microscopic examination. Figures 437 and 455 show
such young stages. They are typical larvae of Octopodidae, with short

705

arms which usually bear 3 small suckers (rarely 2) and have spines on
mantle, head, funnel and the bases of the arms. The dorsal arms are
slightly longer but otherwise resemble the others.

The interbrachial membrane is characteristic, if the membrane which
connects the bases of the arms can already be considered as such. If the
arms are extended, this membrane shows no special characters except the
contrast between the skin of the free and connected parts of the arms; the
first part is thin and bare, while the second is tough and covered with
spines. If the arms are contracted, the tougher skin forms a kind of cuff
for the terminal parts as it is not contracted but remains erect (Figures 437
and 455; cf. Vol. II: Plates XXXI, XXXVI). This gradually disappears
later (but see Figures 456 and 457). Other characters of the larvae are
the short, almost ovoid body, the small size (1 — 3 mm) and the very few
chromatophores. There are differences between the various genera and
they can nearly always be identified.

The hectocotylus at first resembles the other arms, but it soon becomes
detached from the interbrachial membrane, which forms a direct connection
between the 2nd and 4th arms. It is coiled inside the pocket and grows
730 rapidly longer, becoming differentiated in the different genera by various
specializations which are not yet completely explained (Figures 444 and
454). The male reaches maturity rapidly (within a year), while the female
probably has a longer development.-' Growth of the arms of the male
slows down, and its whole appearance is neotenic.

e. VARIATION OF THE TYPICAL STRUCTURE

The typical relationships between the 4 genera which form the family
are shown in the following graph: i. e. the genera developed by secondary

specialization from forms I — IV in a direct
series.

The ideal or hypothetical form I corresponds
to the type of the family; form II corresponds
to the type of the 3 genera, form III to the type
of the 2 highest genera. Form IV is the ancestral
form of Argonauta. The forms derived from
II and III are clearly natural groups which can
be defined, despite the specialization of their
recent representatives. The important
morphological relationships, however, call for
a division into 2 subfamilies by drawing a line
between II and III, i. e. a distinction between
a lower (I) and a higher (II) stage of variation.
This is, of course, arbitrary (see p. 16), and
is only intended to introduce order for practical purposes by stressing the
essential and omit characters of less importance. This division stresses
the distance between forms II and III and creates the two natural
subfamilies: Tremoctopodinae and Argonautinae.

Argonauta

Argonauta probably lives several years.

5110/2

706

1. SUBFAMILY TREMOCTOPODINAE NOV.

Diagnosis. A large interbrachial membrane present at least between
2 dorsal pairs of arms. Hectocotylus (as far as is known) on the right
side, situated below the eye, with fringelike processes on both sides and
with a cutaneous, sac -shaped or lobelike spermatophore reservoir at the
end of the part with suckers. Basal stump of hectocotylus with only one
sucker (if Verrill’s figure is correct). Paramedian teeth of radula
markedly reduced.'-'

The subfamily contains the genera Tremoctopus and Alloposus.
They differ markedly from the type of the family, but show distinct
731 similarities, particularly in the structure of the hectocotylus. This
separates them from the other two genera, in which the hectocotylus
shows marked complication and modification.

2. GENUS ALLOPOSUS VERRILL, 1880

Diagnosis. Body gelatinous, swollen. Arms of different length (formula:
1, 2, 3, 4), otherwise uniform, normal and connected by a large inter-
brachial membrane. Funnel corners fused with the mantle. Water pores
absent (p. 739). Spermatophore with an open spermatophore groove and with a
sac -shaped spermatophore reservoir (which can be closed?) and with a short,
free terminal ’’penis"; fringes present on entire length of hectocotylus on
both sides (Figure 444).

Early stages of Alloposus are unknown and we do not know how much
they differ from those of Tremoctopus. The fusion between mantle
and funnel (Figure 435 on p. 727) is certainly secondary. This genus is
thus apparently the least specialized in the family in many characters.

Alloposus occurs in the Atlantic, but not in the Mediterranean.

The following brief description is only intended to clarify our concept of
the general type. Its habitus resembles that of Octopus, but its body is
very plump, short, gelatinous and swollen. The funnel gland is W- shaped.
Sexual dimorphism is less marked than in the other forms; the male is
relatively large (about 30 cm), but the female reaches an enormous size.
The hectocotylus is situated in a pocket below the right eye, as in
Tremoctopus and differs from that of other Argonautidae in the
presence of an open spermatophore groove and a simpler reservoir
(Figure 444) which is situated terminally, as in Tremoctopus. The
penis forms the free end of the hectocotylus. I assume that the penis is
connected with the reservoir of the spermatophore as in other Argonautidae
(p. 728), although this is not stated in the original description of Verrill.

Compare Plate 16, Figure 6, Joubin, 1893 for Tremoctopus; also Guerne, 1895, p. 114, Figure 3 for
Alloposus. To the latter form also belongs "Bolitaena”microcotyla Hoyle, 1904, Plate 4,
Figure 1.

707

3. SUBFAMILY ARGONAUTINAE

Diagnosis. Interbrachial membrane reduced or weakly developed between
bases of all arms, or absent. Hectocotylus situated on the right or left
side; fringes of Tremoctopodinae replaced by skin folds which are
contracted like a sac over the inward -curved juvenile arm, but they are
modified and form a pocket on the detached hectocotylus, which is curved
outward. Spermatophore groove, except the proximal entrance, closed
into a tube situated on upper side of hectocotylus and differentiated into a wide,
proximal, muscular spermatophore reservoir and into a narrow, thin-walled
distal spermatophore duct continuing in the penis canal. Basal stump of
hectocotylus with 2 normal suckers.

732 The genera of this subfamily, Ocythoe and Argonauta, closely
resemble each other in all these characters but they differ markedly in
external appearance because the female Ocythoe lacks homologues of
the shell of Argonauta. We will have yet to examine whether this lack
is primary or secondary. At any rate, the highly specialized resemblances
described in the diagnosis prove a close relationship between the two
genera.

The funnel gland is characteristic for the subfamily. It shows in
principle the same conditions as in the Decapoda. The outer arms of the
W- shaped figure of the Polypodoidea have again become separated, leaving
a A -shaped median figure and two lateral stripes (Jatta, Plate 18,

Figure 21; Plate 19, Figure 7).

FIGURE 436. Diagram of form of hectocotylus in the Argonautinae. a) Arm before use, coiled inside the
pocket, b) Hectocotylus beginning to detach itself, before copulation. Its terminal part projects from the
opening of the sac (5), which is widened. The arrow is directed toward the inside, c) The arm is ready to
function. The pocket is turned inside out and forms a secondary sac. (4, "accessory spermatophore reservoir")
on the dorsal side of the hectocotylus:

1 — basal stump; 2 — position of (primary) spermatophore reservoir on dorsal side of arm; 3 — inner space of
pocket of hectocotylus; 4 — cavity of secondary reservoir of spermatophore; 5 — margin of folds of sac;

6 — penis.

708

The Argonautinae are good swimmers and have a long, strong funnel.

The hectocotylus is of particular interest. Despite the marked external
differences (Figures 454 and 469) there are striking resemblances. We
begin the description with the ancestral form of the Argonautidae (p. 72 8).

The hectocotylus has been detached from the connection with the interbrachial
membrane and consists of a short, more or less normal basal stump with
2 suckers behind each other, and of a very long part with 2 rows of suckers
which is later detached. However, the spermatophore groove does not pass
laterally on the arm (its "morphological position" is the ventral outer edge
of the arm) but is displaced to the dorsal side of the hectocotylus. At least,
we must assume that the "anlage" of the groove develops there. Only the
tube has been established, which is certainly formed by constriction of the
groove from the skin, i. e. as we have assumed for the penis (p. 728) from
which the process probably continues in a proximal direction. The
spermatophore groove is thus transformed into a tube which still opens
733 proximally in a pore near the base of the arm and continues distally in the
canal of the penis. This is the basic form of the hectocotylus of the
Argonautinae which shows further specializations. As in the Tremoctopo-
dinae, part of the spermatophore groove, i. e. a canal in this case, develops
into a reservoir, because the hectocotylus of the Argonautidae has to carry
the large spermatophores. However, the reservoir is formed in the
proximal part in the form of an oblong, muscular sac which continues
distally in the thin-walled part of the canal and ends blind proximally. The
entrance is here situated not at the beginning but almost at the end of the
reservoir; we do not know whether this is primary or secondary.

The hectocotyli of A r gonaut a and Ocythoe show a number of
further resemblances. Instead of the fringes of Tremoctopus and
Alloposus, there is on each side a skin fold which is homologous to the
fringes. The undeveloped hectocotylus is curved toward the mouth; the
folds enclose it and are contracted above it like a sac. These folds are
named sac folds, and the structure the sac of the hectocotylus. If the
hectocotylus frees itself from the sac, straightens and curves outward (this
takes place perhaps later, during detachment), the folds are turned back and
become contracted again into a sac on the dorsal side of the arm, so that a
structure is formed which probably serves as an accessory reservoir of the
spermatophore; at any rate, sperm -containing parts of spermatophores are
usually found there in Ocythoe and sometimes in Argonauta.

Other skin folds at the base of the penis form a penis sac which may be
complete (Ocythoe) or less complete (Argonauta). It is not certain
whether this formation should be considered as a remnant of the spoon of
the Octopodidae (cf. p. 728). There are thus a number of highly specialized
resemblances between these two genera, which differ apparently so
markedly, proving their close relationship and justifying the assumption
of a common ancestral form. This cannot be defined before a detailed
description of the different forms. Compare particularly the part
describing the derivation of the shell of Argonauta (Chapter 56).

709

734 Chapter 54

GENUS TREMOCTOPUS
Delle Chiaje, 1829

Contents; a. Diagnosis. — b. Trent octopus violaceus (p. 735). — 1. Diagnosis (p, 735). —

2. Literature (p. 735). — 3. Indifferent young stages (p. 736). — 4. Development of the female (p. 737). —
5. Development of the male (p. 744).

a. DIAGNOSIS

Dorsal arms markedly predominant, connected by a large interbrachial
membrane and with very small suckers. Lateroventral arms markedly
retarded in growth. Funnel corners folded back and fitting into pocketlike
pits in the mantle. Water pores opening on base of dorsal and ventral arms,
leading into large subcutaneous cephalic cavities. Mature hectocotylus
without spermatophore groove, with a vesicular reservoir of the spermato-
phore at the end; penis whiplike, originating proximal to the reservoir and
at least partly retractible into a basal pocket, from which it emerges
through a short passage which is directed toward the mouth and opens on
the inner side of the arm.

This genus probably contains several closely related local forms, which
cannot be exactly determined owing to the insufficient data in the older
literature. None of the numerous names proposed, except T. violaceus,
are valid because the authors did not know the differences from T. viola-
ceus (if there are such differences). I therefore placed all descriptions
and drawings of species of this genus in T. violaceus. I later examined
specimens from the Atlantic which differ slightly but distinctly from the
Mediterranean forms. Berry (1914) described specimens from Hawaii,
which he considered as T. violaceus, with some reservations, since the
specimens were not identical with the Mediterranean material. The
differences observed, however, are not distinct enough to justify the
establishment of a species. They are rather gradual differences which
extend the range of variation and heterochronisms in the differentiation
of the hectocotylus, which will not be discussed here. These variations
may be connected with the wide distribution of the species which includes
735 the Mediterranean, the Atlantic and the Pacific. However, they are not
important for understanding the typical morphogenesis of the genus as a
whole which will be described according to the Mediterranean forms.

My interpretation of the definition of this type confirms in principle the
view of Jatta (1896, p. 207), who considered most specimens of
Tremoctopus known to him as stages of the same species. Jatta based
his view on substantial material which I later obtained, although he had

710

no deeper understanding of their morphology. Above all, he apparently
overlooked the o c e 1 la t u s stage (T r e m o c t o pu s ocellatus Brock,
1882) in the development of the female and he had no strictly genetic concept
of the morphological variation. Numerous authors made unjustified
distinctions within the genus, but some species of "T r e m o c t o p u s"
described do not belong to this genus (Naef, 1912, pp. 199 — 200). The
following forms have been wrongly placed in this genus: 1. "Tremocto-
pus scalenus" (Hoyle, 1904, Plate 4, Figures 6 — 9) resembles
Octopus macropus, and does not belong to the Argonautidae.

2, "Tremoctopus hyalinus" (Joubin, 1900, Plate II, Figures 1—4) is
probably a young stage of Bolitaena or a related form. 3. "Trem-
octopus hirondellei" (Joubin, 1895, Plate I, Figures 1 and 2) is a
young male of an Argonauta, in which the hectocotylus is situated on the
right side, which is unusual but possible; this form may also be an unknown
species of Argonautidae oranAlloposus. 4. "Tremoctopus
doderleini" (Ortmann, 1888, Plate 20) is a typical Ocythoe, possibly
O. tuberculata, or another species. The openings described as water
pores on the dorsal side of the head are probably artifacts; at any rate,
their position is not that of the water pores in Tremoctopus.

b. TREMOCTOPUS VIOLACEUS
Delle Chiaje, 1829

1. DIAGNOSIS

Dorsal arms long, with whiplike ends, with broad margins; their distal
parts are always lost in the maturing female. Hectocotylus differentiated
into a proximal part and distal part without fringes and with modified
suckers. Position of funnel gland occupied by numerous glandular
longitudinal folds which develop on the area of the W- shaped formation.

2. LITERATURE

1830 (29) Delle Chiaje (Vol. I, Plates 70, 71), Tremoctopus violaceus.

1830 Ferussac, Octopus velifer (illustrated after a specimen obtained from Verany. Detailed description
in the following work):

1835 Ferussac and d’Orbigny (pp. 91, 96, 98, 100, 104, Plates 16, 23, 5, 10, 18, 20, 33), Octopus
(Philonexis) velifer, quoyanus, atlanticus, brevipes.

1835 d’Orbigny (2) (p. 17), Octopus (Philonexis) quoyanus, atlanticus. brevipes.

736 1836 Owen (Plate 21), Octopus (Philonexis) se mipalmatus.

1837 Rang (p. 60), Octopus velatus.

1840 V6rany (p. 513), Octopus kollikeri,

1841 Delle Chiaje (Vol. I, p. 5; Vol. VI, p. 66), Tremoctopus.

1844 Kolliker (p. 161), Tremoctopus violaceus.

1844 Philippi (p. 201), Octopus velifer.

1849 Gray (p. 27), Tremoctopus violaceus, quoyanus.

1851 Verany (pp. 33, 41, Plates 11, 14, 15, 16), Octopus (Tremoctopus) violaceus, kollikeri.

1852 Eydoux and Souleyet (pp. 13, 15, Plate I), O ctopus gracilis anddubius.

1855 (1845) d’Orbigny (pp. 202, 204, 205), Philonexis velifer, quoyanus, microstomus,
atlanticus, hyalinus.

1857 Troschel (pp. 44, 50), Philonexis m i cr os t om u s (= k 51 li ker i ) and v i ol a c e u s .

711

1860 Steenstrup (p. 332), Trem octopus v i ol a c e u s and q u oy a n u s .

1866 Keferstein (p. 1449), Tremoctopus.

1869 Targioni-Tozzetti (p. 16). Tremoctopus violaceus.

1871 Woodward (p. 165), Tremoctopus violaceus, velifer, quoyanus.

1879 Tryon (pp. 130, 131, Plate 43), Tremoctopus violaceus, quoyanus, m i c r os t o m u s (?),
gracilis, dubius.

1880 Tiberi (p. 14), Tremoctopus violaceus.

1880 Brock (p. 98), Tremoctopus violaceus.

1882 Brock (p. 583), Tremoctopus violaceus, ocellatus.

1885 Ninni (p. 1), Tremoctopus violaceus.

1886 Hoyle, Tremoctopus violaceus, quoyanus, m i c r os t o m u s (?).

1887 Fischer (p. 335), Tremoctopus violaceus.

1889 Jatta (p. 64), Tremoctopus qu o y a n u s (= ko 1 1 i ke r i ).

1890 Colombatovic, Tremoctopus violaceus.

1890 Carus, Tremoctopus violaceus (m i c r os t o m u s, kdllikeri?).

1894 Pelseneer (p. 207), Tremoctopus violaceus.

1896 Jatta (p. 204, Plates 6, 20), Tremoctopus violaceus,

1904 Hoyle (p. 418), Tremoctopus quoyanus.

1912 Naef (p. 199), Tremoctopus violaceus.

1914 Berry (p. 281, Figs. 8, 9, 10), Tremoctopus violaceus.

1915 Chun (p. 478, Plate 77, Figs. 1, 4, 5, 6), Tremoctopus hyalinus.

1916 Naef (p. 17), Tremoctopus violaceus.

1920 Wiilker (pp. 56, 51), Tremoctopus violaceus.

1921 Naef (p. 538), Tremoctopus violaceus.

3. INDIFFERENT YOUNG STAGES

The description of these variable animals can only be embryological.

We begin with the freshly hatched form (Figure 437). There are no sexual
differences and it closely resembles this stage of Argonauta (Figure 455).

FIGURE 437. Freshly hatched stage of Tremoctopus violaceus. 20x. Compare with the similar and
earlier stages shown on Plate X, Figure 4, also Vol. II: Plate XXXI. The eggs from which these forms hatched
were probably from Messina and belong to the material of A. Kolliker; this material is now in the Bavarian
State Collection in Munich. Compare also with Figure 443. Note the resemblance to the larvae of
Argonauta (Figure 455):

1 — cutaneous spines; 2 — primary lid; 3 — LV-arms, transformed into hectocotylus in the male.

The more or less ovoid form of this young specimen does not differ from
the youngest Argonauta (see p. 729). Even the relative length of the arms
is strikingly similar (formula: 1, 4, 2, 3 instead of 1, 2, 4, 3) because the
dorsal arms are by far the largest, and the lateroventral arms are retarded.

712

On the other hand, it resembles Ocythoe in that the 2nd arms are
slightly shorter than the 4th, although the difference is very slight. The
specimens are about 2.5 mm long, twice as long as the youngest
Argonauta; it should be remembered that they were kept in alcohol for
many years and were not preserved by modern methods. The arms are
737 much thicker than those of a well preserved Argonauta, which may also
be due to preservation. They were probably longer and thinner in the live
animal. The suckers differ slightly from the typical condition: The dorsal
arms already show 2 distinct "anlagen" of secondary suckers, in addition
to the 3 well developed primary larval suckers, of which the third is
particularly large. The 2nd and 4th arms bear the 3 normal suckers, the
3rd arms only a single sucker. The definitive proportions are thus already
prepared at this stage. The funnel ^Flate X, Figure 4) already shows the
special character of the funnel corners, which are folded sharply anteriorly
and fit inside the pockets of the inner mantle surface. This is characteristic
for Tremoctopus and perhaps for the whole family; at any rate, it is
typical for the 3 higher genera.

The mantle cavity already shows the typical conditions of young
Argonautidae: the mantle adductor is restricted to the anterior part, near
the anus, the gills have only about 7 or 8 lamellae, and the stellate ganglia
are situated even farther posteriorly, near the base of the gills. The
youngest stages obtained from the plankton are much larger and further
developed (Figures 438 — 440 on pp. 738 and 739). They already show
external sexual differences; the right arm of the 3rd pair of the male is
more or less completely transformed into a hectocotylus, while it
resembles the left arm in the female. The form of the animals is at first
unchanged.

4. POSTEMBRYONIC DEVELOPMENT OF THE FEMALE

Figure 438 shows a young female of T. violaceus from the plankton
in Naples. Jatta (1896, Plate 7, Figure 23) described this stage alive and
Verany named it "Oc t opu s kollikeri." Such stages will be named
kollikeri stages. Their habitus is very characteristic; except for the
relative length of the arms and the funnel-mantle connection, they show no
resemblance to the preceding stages. The mantle sac is very short, almost
cup-shaped, and the head is broadened because of the marked enlargement of
the eyes, which are still directed slightly anteriorly and not laterally as
usually in young Dibranchiata. The arms differ markedly in length, but the
primary proportions are unchanged; they are now much longer and bear
more numerous suckers. The contrast between the two long dorsal pairs
and the two short ventral pairs is more marked. The lateroventral arms
are mere stumps with only a few suckers which form 2 rows, as on the
other arms. The suckers have a characteristic form, like bulging pots.

738 Sausage -shaped bodies (Figure 438) adhere to the suckers which are

drawn inside the suction chambers. The epithelial wall of these bodies is
filled with nematocysts. They are probably parts of Cnidaria, probably

* On the allegedly completely atypical funnel gland (Jatta, 1896, p. 22, Figure 51) see p. 743.

713

tentacles of a medusa. It is not certain whether they are accidentally
attached after contact with a medusa or have been captured by a host to
which the nematocysts are useful, but the latter seems more probable.

All arms have a weakly developed interbrachial membrane at the base
which may become slightly more prominent by contraction of its muscles.
This is not more marked on the dorsal arms.

FIGURE 438. A female k 6 11 i ke r i stage ofTrem
octopus vulgaris. 4x. From the plankton in
Naples. Note the characteristic form of head and
mantle sac; proportions of arms; form of suckers;
rudimentary interbrachial membranes; attached
tentacles of a medusa.

FIGURE 439. A male k 5 1 1 i ke r i stage of
Tremoctopus violaceus. 4x. The only
difference from the specimen in Figure 438 is
the hectocotylus. Note position and form of
the pocket of the hectocotylus (3) and the form
of the long, thin hectocotylus with its slight
terminal swelling (2). 1 — interbrachial
membrane.

All my specimens are old, alcohol-preserved, with completely discolored
chromatophores. The tissue is delicate, almost gelatinous. The live
animal probably has reddish brown and dark brown chromatophores (Jatta,
Plate 7, Figure 23). The liver is visible through the skin, yellowish brown,
as in most small Cephalopoda. The eyes are dark brown and the body is
hyaline.

The arms become further differentiated: an interbrachial membrane
develops between the 2 dorsal pairs, but the suckers remain retarded. On
the other hand, the interbrachial membrane between the ventral arms
remains small while the suckers of these arms rapidly become as large
as or larger than those on the dorsal arms. The mantle sac becomes
longer, and the head expands laterally to its definitive form. Small
epithelial depressions develop into the characteristic water pores at the
base of the dorsal and ventral arms. These processes lead to the following
stage (Figure 440), which is about 4 times as large as the preceding stage.
Its general appearance has markedly changed and is very characteristic
for the genus. All later changes have to be related to this stage, which
739 certainly passed directly to the definitive stage in the predecessors, i. e.
by growth, and is certainly more typical for the genus than the adult
T. violaceus.

In addition to the above relative changes of mantle and head, the arms
and the "anlage" of the water pores are interesting. The water pores have
already attained their definitive form and position: they are rounded
openings situated at the base of the dorsal and ventral arms and lead at

714

first into shallow skin pockets which later become wider. The arms have
the usual proportions. The two dorsal pairs are longer and bear very
small suckers, the shorter ventral arms bear large and very prominent
suckers the form of which varies slightly according to the state of
contraction. The normal biserial arrangement becomes uniserial in the
distal part of the dorsal arms, while suckers are absent on the differentiated
terminal part, which develops into a flagellum in the later stages.

FIGURE 440. Young female of Trem octopus violaceus, natural size. Note form of mantle and head,
development of the interbrachial membrane, especially between the 4 dorsal arms, on which the membrane
extends far toward the apex (6). The suckers differ in size on the different arms. The arrangement is
uniserial (6) in the distal part of the dorsal arms, which end in a thin, slightly flattened end part (1). The
smaller figure shows the end part of an arm of a slightly younger specimen, enlarged l.Stimes. 4— water cavity.

The 4 dorsal arms are now connected by a much larger interbrachial
membrane which extends from the deep indentations as broad margins
toward the apex of the arms, only on the inner side of the laterodorsal
arms and still further on both sides of the dorsal arms, on which the margins
remain wide to near the end part and then suddenly end at its base. This
forms a sharply differentiated end part, in contrast to the earlier stages
(small drawing in Figure 440), in which the margins are narrow, like
ridges. The interbrachial membrane is weakly developed between the
ventral arms but also between the 2nd and 3rd arms. The lid apparatus
already shows its definitive form (Figure 387 on p. 660).

Jatta (Plate 20, Figures 4, 6) shows similar stages, but does not describe
the interesting details. The specimen in Figure 441 is slightly more
advanced. There are no new developments; the form of the body is the
same, slightly changed by contraction, but the arms are relatively longer,
especially the dorsal arms, and their uniserial distal part has become
relatively longer, while the end part forms a long, whiplike appendage.

740 The dorsal arms are very extensible, and their margins are very variable.
These arms can be flattened into thin veils and may form sacs in which
small prey is perhaps caught. Brock (1882) published a drawing of this
stage under the name T. ocellatus; see also p. 700.

The dorsal arms are usually damaged in slightly larger specimens
(Figures 442 and 443); a more or less large distal part is usually absent.
The loss is apparently due not to accidental injury but to a sharp tear along
the transversely striated musculature. The mutilation is often symmetrical

715

on both arms; and this is usually the rule in larger animals (Figure 443)
(see Jatta, 1896, Plate 20, Figures 7 and 8).

FIGURE 441. Young female of Trem octopus violaceus,
natural size, (ocellatus form). This stage differs from the
preceding mainly in the larger arms, especially the dorsal arms,
the end part of which (1) has developed into a whiplike appendage.
The small figure (c) shows a kollikeri stage, natural size.

2, 3 — water pores.

Figure 442 shows an animal which is already partly mutilated: the left
dorsal arm has lost the greater part of the terminal flagellum, but the rest
of the arm is normal and has reached the peak of its development. This
arm and the stump of the other arm bear some stripes of glandular
epithelium at the edge of the margins. The epithelium forms a glandular
stripe with small interruptions. I did not examine the finer structure of
this epithelium because the specimen had been kept in alcohol for a long
time; their function and that of the distal part of the dorsal arms remain
obscure. This glandular epithelium is of particular interest since its
position corresponds to that of the shell glands of Argonauta and is
perhaps homologous to them. The left dorsal arm lost its distal part; the
stump, however, is well scarred and has a characteristic margin: the
transverse tear of the margin passes exactly along their distinct transverse
striation; the tear of the arm itself is slightly contracted and forms an
indentation of the margin.

741 The progress of the mutilation is illustrated in Figure 443. The dorsal
arms are always more or less symmetrical. The asymmetry in the
transitional stages suggests accidental loss of the ends of the dorsal arms,
although it is prepared and regulated by the structure of the arm. On the
other hand, it seems justified to assume that the animal later completes the
mutilation actively. This is certainly a very peculiar method of achieving
the definitive form during normal development of a highly organized animal

742 and the only analogue is the loss of larval organs. These characteristic
and atypical dorsal arms which the animal discards are part of the typical
organization of the Octopoda. This unique phenomenon is of great interest
and will be discussed again in connection with the phytogeny of the shell of

716

Argonauta (Chapter 56). A relation to the development of A r go n au t a
is not excluded a priori because the ends of the arms of T. violaceus are
homologous to those which produce the problematic shell of Argonauta
and also bear glands which certainly correspond to the shell glands of

Argonauta.
{741) 1

w

FIGURE 442. Young female of Tremoctopus
violaceus, natural size (transitional stage). The
dorsal arms have lost some parts; the dotted lines
indicate the assumed primary form of the arms. The
terminal flagellum was probably much longer in life
(FI). Note the points mentioned in Figure 441. Dr —
arm glands; Et — lost part of arm; Wp — water pores.

The form of the adult female is determined mainly by these developments.
(Figure 443). The mantle sac is relatively larger than in the young forms,
longer and wider; its width approximates that of the head also in contracted
specimens, so that the animals appear strikingly undifferentiated in dorsal
view. The mantle sac tapers slightly posteriorly and has a rounded end.

The olfactory tubercle is an oval, smaU, transverse papilla, which is
usually not covered by the mantle margin. It is prominent, not embedded
in the gelatinous skin as in the other Octopoda. The eyes are large, rounded
as in other Argonautidae, and project only slightly. The lens is usually in
direct contact with the water; the lid apparatus is half open.

717

(741)

FIGURE 443. a) Diagrammatic dorsal view of an adult female of Tremoctopus violaceus, after a
complete mutilation of the dorsal arms. 0.5 x. b) Younger animal. Compare with Jatta (Plate 20,
Figures 1 and 12, which are drawn after intermediate stages), c) Form of funnel gland of the ocellatus
stage. 1 — stump of arm; 2 — water pore.

FIGURE 444. Egg mass of T r e m oc t o pu s violaceus. 0.5 x. For details
of its origin see Figure 437. It resembles that of Argonauta argo, but the
eggs are much larger (on the embryos, see Plate XXXI in Vol. II). The eggs
are of different age, like the egg masses in the "boats" of Argonauta. The
stalks of the egg membranes are entwined and cemented together into a
grapelike mass and its large stems are attached to the sticklike base which
replaces the shell of Argonauta. The breeding animal probably has 2 such
sticks. According to Kolliker, they are held by the dorsal arms, probably
enveloped by the membrane.

The proportions of the arms are determined by their development. The
arm formula is now 2, 1, 4, 3; the stumps of the dorsal arms are only
slightly longer than the ventral arms. After the loss of the distal part of
the dorsal arms, all arms bear 2 rows of suckers. Except for the most
proximal suckers, those of the two dorsal pairs are much smaller than those
on the two ventral pairs; they are largest on the ventral arms. All suckers
have long, gelatinous stalks. The stalks of the small suckers of the dorsal
arms are thicker than the suckers; those of the large suckers of the ventral
arms are strong but less thick. Between the stalks are cutaneous connec-
tions resembling the protective margins of the Decapoda. These connections

718

are characteristic for the family but are probably absent in Alloposus.
They are gelatinous and swollen on the dorsal arms and connect the rows
of suckers into pallisade-like elevations.

743 Of particular interest is the development of the head cavities which

develop in connection with the water pores and gradually attain a consider-
able size under the skin. The cavities of each side remain separated in the
middle, but the dorsal and ventral part of each cavity become united with
a dorsal and ventral exit. This system consists of a main duct which forms
a half- ring around one side of the head and extends from the dorsal to the
ventral pore. It passes between the eye and the bases of the arms, and has
secondary diverticula or tunnels between the bases of the arms. The tunnels
between the 1st and 2nd arms are especially deep and reach the mouth area;
those between the 2nd and 3rd and between the 3rd and 4th arms are less
developed. A large diverticulum descends behind the eye, while a narrower
tunnel extends ventrally between the eye and the funnel posteriorly. The
eyeball is. thus surrounded by a double system of cavities, an inner, the
orbital cavity, and an outer, the head cavities, the wall of which separates
them from the orbital cavity.

The funnel is characterized by the connection with the mantle described
on p. 726 and the atypical form of the funnel gland. This begins to develop
as a typical "anlage** and is still distinct as a broad, raised W- shaped stripe
also in the ocellatus stages. On its surface, on each side, about 8 — 10
not quite regular longitudinal folds develop which later replace the primary
glandular surface when this becomes indistinct in form (Jatta, Plate 20,
Figure 13). This structure is unique among the recent Dibranchiata and is
the extreme modification of this organ.

The organs of the mantle cavity and the jaws are characteristic for the
family (p. 727). The radula (Plate XVI, Figure 6) is very characteristic:
the median teeth are broad, with 2 large secondary cusps, the adjacent
teeth unicuspid and almost rudimentary.

The skin is apparently smooth. Sections, however, show numerous
characteristic elevations of the cutis, covered by the epidermis, so that the
surface is slightly rugose (Tippmar, 1913; Muller, 1853, Z.w.Z.,

Vols. 107 and 4). The animal has a beautiful coloration and a strong silvery
sheen (also preserved specimens). The coloration is more marked on the
dorsal side, the sheen on the ventral side. The chromatophores of the
dorsal side are reddish brown or purple in formol; they are usually deep
violet-gray in alcohol. The ventral surface shows a much looser
arrangement of orange-yellow to carmine red chromatophores. The dorsal
side and eyes of the live animal are deep blue with pink and purple tones
caused by the combined effect of the dark chromatophores and the iridocytes
(Jatta, 1896, p. 207); the ventral side is more orange. The protective
membrane between the dorsal arms is especially dark and it is usually
coiled in the live animal (Jatta, Plate 6, Figure 2). This membrane is
dark brown-violet in well preserved specimens, and only the proximal
glandular pads (Figure 442) are still visible as yellowish brown spots.

The adult female is about 15 cm long without arms (I saw such a large
specimen in the Museum of Natural History in Stuttgart),

719

744 5. POSTEMBRYONIC DEVELOPMENT OF THE MALE

The male also has a kollikeri stage which closely resembles that of
the female except for the "anlage" of the hectocotylus (Figure 439 on p. 7 38),
which shows a markedly aberrant development. It is recognizable already
from the outside, as the live animal is almost transparent and at least
translucent if preserved. The right 3rd arm has grown into a very long,
thin band, most of it coiled inside the pocket, except for a very short
basal part which is connected with the other arms and bears a single
normal sucker. The pocket has already become contracted over the arm.

Figure 445b shows the inner side of a straightened hectocotylus; a normal
segment with about 30 "anlagen" of suckers has to be inserted between the
two ends. The inner side of the long, band-shaped arm bears a single row
of about 70 "anlagen" of suckers of which the distal suckers are quite
young, and the proximal suckers are further differentiated; most suckers,
however, are low, oval, transverse papillae which are the preceding stage
of typical suckers. There is the normal sucker at the base which remains
outside the pocket and at the apex a low swelling, the "anlage" of the
spermatophore reservoir, the penis and the penis pocket. The long
persisting uniserial arrangement of the suckers is characteristic as
I found it otherwise only in embryos of Sepia (Vol. II). The transition to
an alternating biserial pattern is apparently inhibited, which may be
connected with the rapid growth of the whole formation as a preparation
for its further differentiation. I could not observe the whole development
of the hectocotylus, but the two stages described give a fairly exact idea
of the process.

In Figure 445 e, the terminal part of the hectocotylus is already biserial
and bears differentiated suckers, but mainly the terminal part shows
important- changes: the apex has grown into a whiplike appendage of the
penis, and proximal to it is a lobelike process facing the ventral edge of
the arm. This process probably developed from the spoon and bears a
shallow depression or groove on its inner side which begins near the last
sucker and leads to the base of the penis. This groove probably continues
on the (averted) dorsal side of the bent appendage of the penis and becomes
closed into a canal. I did not observe this, as I had not made sections,
but the subsequent development cannot be explained otherwise.

We have to refer here to the structure of the corresponding parts in
Alloposus mollis. A similar assumption has to be made here, although
Verrill, who did not make a detailed comparison with Tremoctopus, did
not record anything similar. In any case, comparison of the hectocotyli
of these two forms (cf. Figure 445f) gives interesting results: the
745 spermatophore groove has evidently been lost in Tremoctopus, and it
is doubtful whether it ever existed. However, this would make it
impossible to understand the development of the higher Argonautidae which
are partly related to A 11 o p o s u s and partly to Tremoctopus. The
following should also be noted. At the point where the groove of Alloposus
ends and opens in the sac -shaped reservoir of the spermatophore, there are
traces of such a groove also in Tremoctopus, which lead to the
depression and then to the penis. Since this depression is the "anlage" of
the reservoir of the spermatophore, it must be homologous to the organ of
Alloposus, in which it probably has a concealed communication with
the penis.

720

In the more advanced stage shown in Figure 445d, the terminal part of
the hectocotylus is folded inward. The depression has become a laterally
open pocket which is connected openly with the penis in its depth (dotted
line). The penis thus appears as a continuation of a pear-shaped basal
swelling, which is the "anlage" of the penis sac and is already
differentiated from the penis by a narrow groove. If this becomes deeper
and the base of the penis inside the swelling is contracted, a sheath is
formed which at first surrounds the base of the penis (analogous to the
sheath of the tentacles of Nautilus); the base of the sheath is later
widened into the sac of the penis.

In the still further developed stage in Figure 445c, the reservoir of the
spermatophore occupies the end of the hectocotylus, where it reaches its
definitive structure. The opening of the reservoir is still lateral, but
strongly reduced, while the proximal part of the slit formed by the folding
(Figure 445c) has become fused. The small pocket to which the opening
leads certainly opens in the penis canal. The connection of the parts of
the completely developed organ is thus established (Figure 445g). The
reservoir of the spermatophores gradually becomes a terminal sac with a
terminal opening which is later glued together; this sac communicates by
a short duct with the canal of the penis. The reservoir of the spermatophore
contains parts of empty spermatophores in the mature animal, particularly
their long, coiled sperm canaliculi and an apparently chitinous formation
the shape and position of which are shown in Figure 445g. It is not clear
whether the spermatophore enters through the persisting terminal opening
or through the opening of the penis, but the latter is not excluded. The
penis sac is situated proximal to the reservoir of the spermatophore, and
a large part of the coiled and contracted penis can be contained in this sac.
The outlet of the penis sac forms a long tunnel on the inner side of the arm
between the two rows of suckers, toward the base of the arm; a longer or
shorter part of the penis usually projects from the tunnel.

The suckers form 2 widely separated rows and have short but probably
very extensible stalks. The suckers differ markedly in the proximal and
in the distal part, and are transitional in the middle. The proximal suckers
generally resemble the normal suckers, but their openings are narrowed to
short longitudinal slits. On the other hand, the distal suckers have markedly
746 widened and flattened openings; particularly their lateral margin is extended
in the form of a dish. They are connected by skin folds. The sides of the
arm bear fringes proximally to slightly beyond the middle which are usually
arranged in 2 more or less regular longitudinal rows. In Berry's specimen
from Hawaii, the fringed part is much longer, nearly of the arm, which
is also relatively longer; Berry mentions 46 pairs of suckers, whereas
my specimens have 33—40.

The penis is the direct continuation and the morphological end of the arm,
like the corresponding terminal part of the hectocotylus of Eledone
cirrosa (Figure 433 on p. 723). The displacement of the reservoir of
the spermatophore to the apex forms a special connection between it and
the base of the penis which is shown in Figure 445g, i. e. a duct passing
proximally in the wall of the penis sac. The end of the penis shows a
similar differentiation as in Ocythoe (Figure 454) — it is awl-shaped and
longitudinally striated and apparently facilitates the introduction of the organ
into the oviduct. I could not determine the position of the opening; it is

721

perhaps situated at the base of this end part. The other part of the penis
747 contains the sperm duct which may be more or less undulate according to
the contraction.

(746)

FIGURE 445. Structure, development and homologies of the hectocotylus of Tremoctopus violaceus.
a) Mature hectocotylus, with one spermatophore, 3x. b) "Anlage" of hectocotylus of a kollikeri stage,
40 X (middle half omitted), c), d), e) Three successive stages of development of terminal part,
f) Hectocotylus of A 11 opos us mollis, after Verrill, diagrammatic, lower magnification, g) Terminal
part of mature hectocotylus, diagrammatic median section with an empty spermatophore reservoir,
h) Apex of penis of a mature animal, highly magnified (cf. Figure 454). Note in a); the polymorphous
suckers, fringes, the point of constriction, penis sac, penis and reservoir of spermatophore. In b): uniserial
arrangement of the "anlagen" of the suckers, undifferentiated terminal part and normal basal stump.

In c), d), e); differentiation of reservoir of spermatophore, its connection with the penis, "anlage" of
penis sac. In f): basal stump, fringes, spermatophore groove, reservoir of spermatophore, penis. In g);
connection between reservoir of spermatophore, canal and sac of penis, entrance into reservoir of spermato-
phore. In h); longitudinally striated terminal part of penis and undulate canal of penis;

1 — entrance to receptacle of spermatophore (2); 3 — penis sac; 4 — opening of penis sac; 5 — free part of
penis; 6 — fringes of hectocotylus, corresponding to folds of the sac of Argonautinae (Figure 436 on p. 732);

7 — basal part of hectocotylus; 8 — undifferentiated terminal part of young hectocotylus; 9 — "anlagen" of
suckers; 10 — omitted part of hectocotylus; 11 — spermatophore duct, connecting part; 12 — same in the
penis; 13 — spermatophore groove; 14 — spermatophore (empty); 15 — opening of penis; 16 — terminal part
of penis (longitudinally striated).

722

The whole organ resembles that of Alloposus in the following
details: 1. The complete inclusion of the hectocotylus in a pocket formed
by the modified interbrachial membrane, which is closed over the coiled
arm, leaving only a small pore (the hectocotylus has no membranes),

2. The formation of lateral fringes; these are replaced in the Argonautinae
by a continuous skin fold which may have developed from the fringe.

3, The terminal position of the reservoir of the spermatophore, the
development of which is apparently related to conditions in Alloposus.
(The closer relationship is based on the fact that the spermatophore
reservoir of the higher Argonautinae develops from the spermatophore
groove, particularly from its proximal part, which therefore cannot be
strictly homologous with that of Alloposus and Tremoctopus. The
reservoir of the spermatophore of the higher Argonautinae therefore
probably developed from a primary form transitional between Alloposus
and Tremoctopus in which the spermatophore groove is still
completely developed, while the reservoir of the spermatophore is not

yet strictly localized, but which must have had a bent, threadlike penis
as in Tremoctopus and an "anlage" of the penis sac because these
characters are also found in Ocythoe, )

FIGURE 446. Mature male of T r em o ct opu s violaceus.2x
Note the interbrachial membrane; water pores; the large hecto-
cotylus (3), visible inside the pocket (1). For the sexuaUy mature
animal, a hectocotylus like that in Figure 445a would have to be
inserted instead near the mouth. Note also the slight asymmetry
of the ventral arms and the different suckers on the dorsal and

ventral arms. 2, 4 ~ water pores.

Except for the development of the hectocotylus, the growing male
shows at first the same changes as the female. The interbrachial
membrane between the 2 dorsal arm pairs becomes larger but the suckers
remain small. On the other hand, the suckers of the ventral pairs grow
rapidly and have the same form as in the female, while the membrane
between them remains very small. The mantle sac becomes longer, and
its form varies according to the state of contraction, as in the female.

The head develops to its normal form because the eyes become larger
and directed laterally. The water pores develop as in the female.

However, the growth of all these parts ceases at an early stage. The body
748 is barely 2 cm long without arms, and the arms remain slightly shorter

723

than in females of the same size. The differentiation of the apical part of
the dorsal arms described above does not take place. On the other hand,
the hectocotylus grows rapidly in its pocket, so that it forms a large
swelling which markedly disturbs the symmetry of the body.

In the maturing animal (Figure 446 on p. 747), the pocket of the hecto-
cotylus pushes the right ventral arm aside and inhibits mechanically the
development of the water pore; the equilibrium is only incompletely
restored when the hectocotylus of the mature animal leaves the pocket.

This does not involve tearing, since the arm leaves through the opening at
the ba*se, which is not closed during contraction of the pocket. It carries
one spermatophore and projects free for some time (until copulation) as a
large appendage which is markedly larger than the other arms. The
hectocotylus is detached during copulation. It is apparently regenerated
from the small basal stump. I have not seen a regenerated hectocotylus,
but I have seen a specimen without hectocotylus and without injury to the
pocket.

The coloration of the male probably resembles that of a female of the
same age and size; i. e. it is markedly duller than the adult female.

I have not seen fresh or well preserved specimens and can therefore give
no information about their natural coloration. A freshly preserved specimen
from Messina had reddish brown chromatophores; some of them, mainly
on the dorsal side, were dark brown.

724

749 Chapter 55

GENUS OCYTHOE

Rafinesque, 1814

a. DIAGNOSIS

Interbrachial membrane completely reduced. Two lateral pairs of arms
much shorter than median pairs. Funnel corners cartilaginous, coiled like
a snail and grasping firmly into corresponding depressions of mantle. Water
pores present only at base of ventral arms, absent at base of dorsal arms
and leading into large cephalic cavities in the female. Hectocotylus
enclosed in pocket until maturity, and forming a large, stalked club. End
of hectocotylus with a penis sac from which the penis projects proximally
through a pore between the last suckers. Eggs developing in the oviduct
until hatching.

This genus contains only one species.

b. OCYTHOE TUBERCULATA Raf., 1814
1. DIAGNOSIS

Arm formula: 4, 1, 2, 3. Hectocotylus on the right side. Pocket of
detached hectocotylus turned inside out and forming a narrow pore at the
proximal end. Ventral half of mantle of female with pointed, cartilagelike
tubercles, which are sometimes connected into a network of ridgelike
elevations.

2. LITERATURE

1814 Rafinesque, Ocythoe tuberculata (p. 29).

1826 Risso, Octopus tuberculatus (p. 3, tom. 4).

1828 Delle Chiaje, Octopus tuberculatus (Vol. 1, p. 68; Vol. 4, pp. 41.56).

1829 Wagner, Octopus veranyi (p. 225).

1830 Blainville, Octopus pictus (p. 8).

1835 FAussac and d'Orbigny, Philonexis tuberculatus (p. 67, Plates 6, 63, 63, 23).

1841 Cantraine, Octopus tuberculatus (p. 19).

1844 Philippi, Octopus catenulatus (p. 201).

1849 Gray, Ocythoe tuberculatus (p. 49). (Believes that the animal lives in the shell of Argonauta;
see p. 761 ).

725

1851 Verany, Octopus catenulatus (female, Plate 13) and O. catenae (male, Plate 14) (pp. 34-37).
Verany, Hectocotylus octopodis, Plate 41.

1851 Vogt and Verany, Tremoctopus catenulatus (large female) and T. catenae (small female
and male) (p. 157).

1855 D’Orbigny, Philonexis tuberculatus (p. 206).

750 1858 Adams, Octopus catenae and tuberculatus (Vol. 1, pp. 19, 22).

1860 Steenstrup, Parasira catenulata (p. 333).

1866 Keferstein, Parasira (Vol. Ill, p. 1449).

1869 Targioni-Tozzetti, Parasira catenulata and tuberculata (pp. 11,13).

1879 Tryon, Parasira catenulata and catenae and Ocythoe tuberculata (p. 132).

1880 Steenstrup, Ocythoe tuberculata (p. 104).

1880 Tiberi, Philonexis reticularis Pet. and tuberculatus (p. 13).

1880 Brock,, Tremoctopus catenulatus and catenae (p. 98).

1882 Verrill, Parasira catenulata and tuberculata (p.. 179).

1885 Ninni, Philonexis (= Parasira).

1886 Hoyle, Ocythoe tuberculata (p. 5).

1887 Fischer, Parasira tuberculata (p. 335).

1887 Steenstrup, Ocythoe tuberculata (p. 61).

1888 Ortmann, Tremoctopus dbderleini (Plate 20).

1890 Colombatovic, Ocythoe tuberculata.

1890 Carus, Ocythoe tuberculata (p. 457).

1894 Pelseneer, Philonexis catenae (p. 207).

1896 Jatta, Ocythoe tuberculata (p. 198).

1916 Berry, Ocythoe tuberculata (p. 3).

1916 Naef, Ocythoe tuberculata (pp. 17; 1922: p. 291).

3. EARLY YOUNG STAGES

I have not seen mature embryos or freshly hatched specimens. Jatta' s
material has been lost. He states (p. 201) that he found stages ready to
hatch in the oviduct of large females. I did not find such stages in the only
gravid female which I examined, but only embryos of young and middle
stages (which will be described in the embryological part). However,

Jatta' s and my data prove that this species is viviparous.

FIGURE 447A. Youngest known postembryonic stage
of Ocythoe tuberculata. 15x. (Atlantic
Ocean). Note the general form in comparison with
Figure 437 on p. 736 and Figure 455 on p. 765; pro-
portions of arms; oval primary opening of eyelid;
small eyes, olfactory papilla and funnel pocket.

Note also the contracted, pointed ends of the longer
arms.

The freshly hatched larvae probably resemble those of Tremoctopus
(Figure 437 on p. 736) and Argon auta (Figure 455 on p. 765), and probably
differ from Tremoctopus only in the stronger development of the ventral
arms. (The relative length of the arms of Argonautidae is already definitive
in the youngest postembryonic stages and in the embryo.) According to the

726

size of the eggs, the youngest larvae are about twice as large as those of
Argonauta and also larger than the larvae of Tremoctopus
(Figure 437 on p. 736).

I examined, by the courtesy of Chun, a young stage (Figure 447A) from
the material of the "Michael Sars." It shows the characters of very young
Argonautidae. The proportions of the arms are distinct (formula: 4, 1,2, 3).
The 4 median arms bear 7—9 suckers, the lateral arms only 3 suckers.

(The youngest larvae probably bear 3 suckers on the median arms and one
751 on the lateral arms.) This specimen is probably a female, because the right
3rd arm shows no special characters. On the other hand, this may be a still
indifferent stage because the hectocotylus bears 2 normal suckers at the
base, while the third sucker is just beginning to develop. The arm can
therefore not yet differ markedly from a developing hectocotylus. However,
as The hectocotylus usually develops early, a male would show a more
distinct beginning differentiation at this stage (Figure 467a).

FIGURE 447B. Youngest
female of Ocythoe tu-
bercular a. 2x. Note
752 the form of mantle sac and
head; the large* slightly
anteriorly directed eyes;
large funnel; water pores;
different length of the
arms and the markedly
projecting suckers.

4. DEVELOPMENT OF THE FEMALE

A very young female from Naples is shown in
Figure 447B. This is old material and the chroma-
tophores are discolored, but its form leaves no
doubt about the determination, in addition to the
resemblance to later stages.

The mantle sac is slightly reconstructed. It is
strongly contracted and appears too narrow in
comparison with the head, on which the eyes protrude
markedly and are still directed slightly anteriorly.
The lid apparatus shows the typical differentiation.
The water pores are already developing at the base
of the ventral arms. The proportions of the arms
are very characteristic. The arms bear prominent
suckers, the stalks of which are certainly much
further extended in life. There are slightly more
than 70 suckers on the ventral arms.

The skin is completely smooth.

Young animals of the size shown in Figure 448
were rather common near Naples in the past; they
disappeared around 1900. They have an O c t o pu s -
like habitus, and early observers therefore took them
for species of Octopus (p. 749). The mantle sac
and head in fact resemble those of a species of
Octopodidae (Figure 42 6 on p. 717), but the eyes are
very large and the anterior mantle margin reaches
close to the eyes. The funnel is very long and the
water pores are distinct. The arms show the
characteristic proportions, but the differences are
less marked than in the preceding stage. The body

727

is covered with reddish brown and dark brown chromatophores, which are
much denser on the dorsal than on the ventral side, and the skin is iridescent,
with reddish, violet and silver}'- tones, as in the adult. The ventral side of
the mantle bears numerous warts which are, however, apparently still
transient and depend on the state of contraction of the skin. There is no
regular network between the warts in these stages.

FIGURE 448. Young female of Ocythoe tuberculata, natural size. Note the Octopus-like form,
distribution of chromatophores, lid apparatus, olfactory tubercle, large funnel, water pore, and the different
length of the arms.

The suckers of each longitudinal row are connected by transient skin
folds which are formed by contraction of certain cutaneous muscles. The
longitudinal folds (Figure 449) on the outer side of the 1st and 4th arms which
are analogous to the swimming margins, are also not of constant form at
this stage; they often form a continuation of the rudimentary and not always
distinct interbrachial membranes which belong to the inner attachment of the
dorsal arms and the outer attachment of the ventral arms. The mantle sac
later grows larger with respect to the head and eyes but it keeps its ovoid
form. The head, however, loses its normal form because of the development
of the cephalic cavities. These develop as in T r e m o c to pu s, but remain
closed dorsally because dorsal water pores are absent.

The ventral pores lead into large pockets which at first surround the
inner muscular base of the funnel and from there extend in different
directions. One diverticulum extends ventral to the eyeball posteriorly,
to near the mantle margin; another, before the eye, extends dorsally to the
middle of the vertex behind the base of the dorsal arms. Both are super-
ficial and very extensible. Secondary branches of these cavities extend
between the bases of the arms to the skin of the mouth area, between the
first and second sucker of the arms at 10 points, i. e. 2 between the bases
of the dorsal and ventral arms and 1 between the bases of the other arms.
Chromatophores are present only near the outer opening and the whole
pockets are separated in the middle from each other and from the eye
chambers, which they partly enclose.

753 The arms become relatively shorter the larger the animals are. They
are very short in the largest, at least in preserved specimens; if they are
strongly shrunken, they are about as long as the body or even shorter. The
arms are often folded back along the mantle and the marginlike folds (p. 767)
of the median arms become very prominent; if the arms are regularly
curved (Figure 449), they form wide, continuous margins which taper toward

728

729

FIGURE 449. Large, almost mature female of O c y th o e tuberculata. 0.5X. (Drawn 11 April 1916 after a dying specimen.) Note the ovoid
body; the head, which is slightly swollen because of the cephalic cavities, the relatively small eyes, the typical lid apparatus, olfactory tubercle,
water pore (Wp),and large funnel. The arms are of different length; the suckers have long stalks and are connected by folds. Dorsal and ventral
arms have swimrning margins, and there is a rudimentary interbrachial membrane. The ventral side of the mantle bears tubercles.

SI - secondary lid; Pu - primary lower lid; Po - primary upper lid; Pp - pupil.

(754)

FIGURE 450. Large female of Oc y th o e t u b e rc u 1 a t a, ventral part of mantle removed. 0.5x.
Note the water pores (Wp); funnel bonds (Th, exposed by splitting the mantle bonds); concentration
of mantle adductor anteriorly (Ad); wide genital openings; large, slender gills; concentration of
renal sacs anteriorly; enormous development of the winding oviduct (Od), which functions as the
uterus.

Kh — branchial heart; Va — venous appendages; Kr — branchial retractor; Np — renal papilla;

Nv - visceral nerve; Kv - branchial vein; Ap - posterior aorta; Vc - vena cava; Ed ~ hind
intestine; Tr — funnel adductor; Mm — muscular mantle; Ht — skin; Hk — tubercles.

730

the apex and the base. The margins continue partly on the head, the
dorsal margins on the vertex, where they may be contiguous, which corres-
ponds to a rudimentary interbrachial membrane; the ventral margins end in
the lateral border of the water pores (Wp in Figure 449). The suckers of
each row are always connected by strong longitudinal skin folds passing on

754 their outer side. According to the contraction, these longitudinal folds may
rise to the level of the marginal ring or collapse, but they never disappear
completely. Figure 449 shows the typical position of the arms and the form
of a large, live specimen.

The funnel shows the connection between funnel and mantle which is char-
acteristic for Ocyt hoe, also for the males and younger stages (Figure 450).
As in T r e m o c t o p u s, the funnel corners are curved and project like hooks
into depressions of the mantle which open posteriorly. However, both the

755 corner and the depression are of complicated form which makes the
connection more solid. Considerable force is therefore necessary to break
the connection also in fresh specimens. The modified funnel corner can be
compared to the scroll of a violin; it is widened inside the depression, and
can only be removed by extending the latter, which the animal is certainly
not able to do itself. The funnel corner is cartilaginous and rigid. The
figure shows its form better than a description. On the funnel glands see

p. 672.

The mantle cavity shows the typical characters of the Argonautidae:

1. The median mantle adductor is situated anteriorly. 2. The superficial
gill retractors radiate into the ventral skin, 3. There are numerous
chromatophores on the ventral side. 4. The genital openings are situated
close to the base of the gills. The genital openings of Tremoctopus form
pointed, projecting papillae, whereas those of Ocythoe are wide, rounded
openings situated before and above the renal papillae; the gills or renal
papillae often cover the openings. The gills are very long and pointed and
resemble those of the Decapoda. Under the entire posterior part of the
ventral skin passes the winding oviduct, filled with eggs in the mature
animal (about 100,000 in the specimen in Figure 450). The eggs in the
distal part of the oviduct are in about the middle of their development,
while further proximally the eggs are less developed, finally uncleaved.

The eggs in their membrane (stalked chorion) resemble those of Octopus
vulgar is, but their stalks are not intertwined into grapelike clusters,
x^s in Octopus (Figure 408 on p. 685), the mature ovary pushes the renal
sacs anteriorly.

The jaws are large and shallowly convex; they show the typical characters
of the family (Plate VIII, Figure 11). The radula closely resembles that of
the Teuthoidea: the teeth of the median row have regular secondary cusps
on each side (Plate XVI, Figure 8), those of the adjacent row only a single
secondary cusp on the outer side. The two following rows on each side have
long, awl-shaped teeth without secondary cusps, and the marginal plates
are small, isolated and square. This is a very characteristic type, which
is unique in the structure of the paramedian teeth. Wherever the para-
median rows have secondary cusps in the Octopodidae, they are situated on
the median side (Plate XVI, Figure 4). The secondary cusp probably
developed into a main cusp. These teeth are usually unicuspid in the Argo-
nautidae (Plate XVI, Figures 6 and 7); they are also reduced in Tremocto-
pus (but compare A 1 1 op o s u s in Guerne, 1895, p. 1 14, Figure 3; and exotic
species of Argonaut a).

731

Characteristic skin tubercles cover the whole ventral side of the mantle.
They are pointed conical and hard like cartilage in the adult, although they do not
consist of cartilage (Jatta, p, 200) but of very firm cutaneous tissue (the skin
and muscles of these animals are very tough and hard). Contraction of the
fibers connecting the tubercles forms a netlike sculpture on the surface.

The surface of other parts of the body is smooth.

756 Coloration of the living animal.* The flesh is translucent, milky white,
also in the largest females, as in Argonaut a; the lateral and ventral
sides of the mantle have a bright silvery sheen. There are also greenish
and reddish reflexes, especially on head and arms, but the prominent tones
are blue and violet, mainly on the dorsal side of the mantle, head and
dorsal arms. If the chromatophores are contracted, the whole dorsal
surface becomes sky blue. Extension of the dark chromatophores, however,
gives a deep blue-violet to blackish neutral tone to the dorsal side, vertex
and dorsal arms, swimming margins and inner side of the 1st arms; this
is probably the normal coloration of the dorsal side. The violet tone is
caused by the combined effect of the dark brown chromatophores of the
deeper skin layers and the iridocytes situated above them, but examination
with the magnifying lens gives the impression that the effect is due entirely
to the chromatophores. The superficial chromatophores cause the blackish
discoloration of the beautiful blue violet. Dorsally and especially on the
lateral and ventral sides, there are, in addition to the brown chromatophores,
orange-red ones, which are usually contracted on the ventral side, and the
milky white tone of the flesh and the silvery -iridescent sheen become
prominent. The orange-red chromatophores are best visible near the
cartilagelike tubercles as reddish brown to orange rings around a light spot.

The animal attains a considerable size, 30 cm and more without arms.

5. DEVELOPMENT OF THE MALE

Figure 451 shows a young male from C. Chun's collection. Except for
the hectocotylus, this specimen probably does not differ from a female of
the same size. The arms are recurved as in Argonaut a. The lid
apparatus is still undifferentiated, but the primary opening already shows the
almond shape which is the beginning of the differentiation. Both median
pairs of arms bear 23 suckers in a normal arrangement; the laterodorsal
arms bear 17 much smaller suckers, and the left lateroventral arm 11.

Arm formula; 4, 1,2,3. The hectocotylus bears 2 normal suckers at the
base; I could not examine the part of the arm coiled inside the pocket.

Also older males do not differ distinctly from a female of similar size,
except for the hectocotylus. However, the development ceases earlier and
appears inhibited in some minor characters: the ventral side of the mantle
bears no tubercles also in fully mature animals; the secondary leveling of
the length of the arms does not take place; the longitudinal skin folds between
the suckers remain indistinct. Coloration is the same.

* Compare the good drawing in Verany, 1851, Plate 13.

732

757

FIGURE 451. Young male of Ocythoe tubercu-
lar a, collection "Michael Sars," 4x. Note form of
body; position of arms; length of funnel; form of
primary lid opening; different length of arms;
developing, club-shaped hectocotylus inside the sac.

FIGURE 452. Mature male of Ocythoe tubercular a. lx. Note the similarity with the slightly
larger female in Figure 448 on p. 752. The large hectocotylus is coiled inside the sac; the crescent-
shaped slit (1) marks the entrance to the spermatophore reservoir. 2 - point of closure of sac.

FIGURE 453. Mature male of Ocythoe tubercular a, natural size, after a specimen from the
University Museum in Naples. Note particularly the large, spread hectocotylus, with the charac-
teristic suckers which are connected by longitudinal folds (3); the penis sac (1) is terminal, while
the sac of the hectocotylus (4) is now situated on the dorsal side of the arm. A small opening
between the last suckers (2) is the exit of the penis from its sac. 5 - secondary opening of sac.
The hectocotylus is already extended but still not loaded; cf.,by contrast, p. 785.

733

The adult male thus differs from a female of the same size (Figures 452
and 448) in the characters mentioned above and in the large, club-shaped
hectocotylus. The structure of the hectocotylus is difficult to recognize if
the arm is situated inside the pocket, especially in preserved specimens.
However, the following details are of morphological interest. The stalk-
shaped basal part bears 2 suckers and a markedly lengthened part without
outer appendages. The club is situated on a looped proximal part, which
is connected on each side with the sac folds. The distal part of the arm is
coiled inside the sac. The folds of the sac could remain widely separated
in the opening of the loop, as may be assumed for the ancestral form of the
Argonautinae (Figure 436 on p. 732), and the arm would enter the opening.
However, because of the elasticity and contractility of all skin folds in the
Octopoda, this opening should be completely closed and perhaps even fused
or at least glued together, so that the opening would not be recognizable in
preserved specimens. Otherwise, it would be visible in Figure 452 at the
point labeled 2, as in the diagrammatic Figure 454d. The hectocotylus later
breaks through at this point (Figure 436 on p. 732).

The mature hectocotylus is extended and turns the sac inside out, so that
the opening of the sac is now situated on the dorsal side (Figure 453). It

758 remains open until the hectocotylus is detached; it later becomes contracted
again and glued together (Figure 454c). There is a crescent-shaped slit (l)
at the distal end of the club (p.757) which leads to the spermatophore reservoir
through a short duct which probably serves for the passage of spermato-
phores (p. 759). It is at first distinctly visible on the free dorsal side of
the hectocotylus, but it is later covered by the inverted sac (near 10 in
Figure 454c).

The hectocotylus leaves the sac before copulation, prior to being loaded
with a spermatophore, as numerous specimens (cf. Figure 453) show. When
the hectocotylus leaves, the sac is turned inside out, so that it is now situated
on the dorsal side of the arm and becomes again an empty pocket. However,
the spermatophore reservoir is still empty.

Except for the stalk part, the hectocotylus bears 2 rows of characteristic
suckers with thin, flattened, dishlike margin. A continuous longitudinal skin
fold connects all the suckers in the row. Similar folds are present on all
arms of large females. However, the folds are interrupted there by the
suckers, while the fold in the male has a continuous margin which in well
preserved specimens projects slightly above the suckers. The two folds
are connected before the first and beyond the last sucker. The difference
between the quite proximal and distal suckers, already observed in Tr e -
moctopus, is distinct also here. The openings of the suckers become
increasingly eccentric distally, i. e. their outer margin becomes wider,
while the openings are displaced inward. However, this contrast is not very
marked, and most suckers are of the distal type.

759 The penis at the end of the hectocotylus is not free as in Tremoctopus:
it is coiled inside a sac of the form of a lemon and projects through

a small opening between the last suckers (Figure 454 on p. 760). This
resemblance is so mairked that the question arises whether this suggests a
closer relationship, or how the absence of a similar structure in Argo -
nauta has to be explained. To solve the problem, I studied the development
of the penis sac. I made preparations of the terminal parts of the hecto-
cotylus of two young animals 14 and 16 mm long without arms, and observed
the following conditions (gh and ik in Figure 454). The suckers of the

734

younger animal are still small, not functional, embryonic. The suckers of
the older animal are typical but do not show the later specific characters.

The axis of the arm of both specimens still passes directly into the penis,
and there is still no trace of an envelope. There is also no curvature
posteriorly as in the stages of Tremoctopus (Figure 445 on p. 746), or
of the "anlage" of the penis sac as in Tremoctopus. However, there are
2 skin proliferations on the dorsal side of the ends of the arms which are
separate in the younger stage but are fused in the older animal. This is
the "anlage" of the penis sac and resembles the folds at the corresponding
point on the hectocotylus of Argonauta. We conclude, therefore, that the
further development of these skin folds is a convergence to Tremocto-
pus, which together with other primary and secondary resemblances may
mislead our views on the relationships in the Argonautidae. The marked
resemblance between the hectocotyli of Ocythoe and Tremoctopus
(Figure 445 on p. 746 and Figure 454 on p. 760) is caused by the presence of a
thin-skinned, lemon-shaped terminal vesicle which contains a coiled thread.
This vesicle is the spermatophore reservoir of Tremoctopus, but it is the
penis sac in Ocythoe! I do not know the functional significance of the
posteriorly curved penis or the method by which it leaves the sac. These
characters are probably connected with the activity of the hectocotylus in the
female. The penis probably does not leave the sac before the detachment
of the hectocotylus (Figure 454 on p. 760); this has not been observed directly
but probably takes place during copulation and at a predetermined place, i, e.
at the end of the stalk part, which remains attached to the body. This is
probably a case of autotomy, as in other Argonautidae. The arm must be
loaded before detachment, i. e. a spermatophore must enter the reservoir.
This may take place through the opening of the inverted sac of the hecto-
cotylus and then through the entrance of the reservoir described above.

I do not know whether this occurs, because I have seen only loaded hecto-
cotyli after they had become detached, i. e. they were found in the mantle
cavity of the female (Figure 454c). The inverted sac of these hectocotyli
is contracted and its elastic skin covers the point of detachment, and the
opening which was wide open before, has become narrowed again to a narrow
pore. The contraction of the sac is probably the reason that the proximal
part of the hectocotylus is always curved posteriorly (Figure 454a). The
penis is usually completely contained in its sac also in the hectocotyli found

760 in the mantle cavity of the female. If the penis is extended, it forms a very
long (Figure 454b), threadlike appendage, accompanied by 2 thin folds, and
the thin penis canal passes along it. As in Tremoctopus (Figure 746h),
the terminal part is differentiated and finely longitudinally striated.

I could not study the spermatophore of a fresh hectocotylus. Such
spermatophores are, of course, empty; the description of a very long sper-
matophore (Marchand) is due to confusion of the sperm duct with the
spermatophore. This is a thin tubule, which is coiled tightly inside the
spermatophore reservoir, and often extends into the spermatophore duct
and penis canal, and at last projects from it, I found that the projecting end
is often swollen, with strongly sclerotized walls, and asked myself, as in

761 Tremoctopus, whether the spermatophore does not perhaps enter through
the opening of the penis. The frequent presence of a second spermatophore
(or part of the first?) inside the inverted sac of the hectocotylus is also

735

problematic. This spermatophore closely resembles that situated in the
spermatophore reservoir and its end sometimes projects from the opening
of the sac (Figure 454c2i). Verany and Vogt (l852) illustrated an intact
spermatophore from the genital duct of the male (see also Tryon, 1879,
Plate 18, Figures 27-35).

(760)

FIGURE 454. Morphology of the hectocotylus of Ocythoe tuberculata. a) Detached hectocotylus
from the mantle cavity of an immature female, b) End of such a detached hectocotylus, with ex-

tended penis, c) Diagrammatic median section through a. d) Diagrammatic median section through a
hectocotylus which is still coiled inside the sac (without the end part), e) End of penis, magnified 6x,
lateral, f) As e, ventral, with skin margin (1). g,h) and i,k) Two young stages: dorsal and ventral view
of end part, higher magnification.

1 — longimdinal canal (spermatophore duct?); 2 — longitudinal striation on terminal part of penis; 3 —
undifferentiated main part of penis; 4 - penis sac or its "anlage"; 5 - projecting part of penis; 6 -
margin of hectocotylus (false protective margin); 7 - accessory spermatophore reservoir; 8 - as 20;

9 — entrance to true spermatophore reservoir; 10 — canal inside the reservoir; 12 — spermatophore;

13 — spermatophore duct; 14 — removed distal part of hectocotylus; 15 — point of amputation; 16 —
muscular axis of arm; 17 - sac of hectocotylus; 18 - sac fold; 19 - primary entrance to sac; 10 -
secondary entrance (after inversion); 21 - second spermatophore.

The males of Ocythoe are always found in an empty test of Salpa,
which is used as a floating habitation, like inPhronima sedentaria
(Jatta, 1896, Plate 7, Figure 8). It rests inside (Figure 453, on p. 758) like
Argonauta in its shell, the arms curved back and holding the test from
within. (l have often heard fishermen report that the female carries a

736

similar test but discards it when caught. These reports should be treated
with reservation, although a zoologist may confuse this species with
Argonauta more easily than a fisherman in Naples. On the other hand.
Gray (1849) stated that the species lives inside a shell of A r g o n a u t a, and
my material contains several females in shells of Argonauta, at least
preserved. This may be a mistake or the specimens might have been
placed in the shells as a joke.) At any rate, the males of Ocythoe have the
instinct to use alien housings as swimming habitation. This must be borne
in mind if the morphological consideration of the shell of Argonauta is
compared with the behavior of closely related forms, since it could explain
the phylogenetic development of this shell (see p. 770).

737

762 Chapter 56

GENUS ARGONAUTA
Linne, 1758

Contents: a. Diagnosis of the genus, -b. Argonauta argo(p. 763). - 1. Diagnosis (p. 763). -
2. Literature (p. 764). - 3. Early young stages (p. 765). - 4. Development of the female (p. 767). -
5. The adult female (p. 770). - 6. Biology (p. 777). - 7. Origin of the shell of Argonauta (p. 778). -
8. Development of the male (p. 783).

a. DIAGNOSIS

Interbrachial membrane distinct but weakly developed. Water pores
absent. Funnel corners slightly curved, forming a "funnel bond" as in the
Decapoda around a tubercle on the inner side of the mantle. Male very
small (less than 2 cm without hectocotylus); hectocotylus usually developing
on the left side and temporarily enclosed in a pocket between the 2nd and
4th arms below the eye; penis not contained in a special sac. Dorsal arms
of female curved back on themselves, with a skin fold in the loop and with a
zone of dense shell glands situated on the inner side of the distal part of
the arm and producing a spirally, calcified, paper-thin, boat-shaped shell
serving as a floating habitation which always contains the posterior part of
the body and in which the eggs are laid.

The genus Argonauta contains species which are incompletely known,
except for details of the sculpture of the shell. Some of them are of an
older geological age.''' A.hians Solander is known from the Pliocene of
Upper Italy. Other species have become extinct. The oldest form known
is A.johanneus from the Miocene of Styria (see also Fossile Tinten-
fische, 1922, pp. 291, 294). The only species which will be described in
detail is A.argo. According to Berry (1914), A. b 6 ttg e r i Maltzan shows
characteristic differences from A.argo in the soft parts (see below), but
these differences are not of general interest. The following data apply in
all important points to the whole genus, which is described here for the
first time more or less comprehensively. A.bottgeri Maltzan, 1881 is
much smaller than the mature A. argo (but A. argo is already mature

763 and bears eggs in the shell at the size of A.bottgeri). The eggs of
A.bottgeri are slightly larger, and its arms are shorter and bear fewer
suckers: the laterodorsal arms of A.argo bear about 150 suckers, those
of A.bottgeri about 60; the respective figures for the hectocotylus are
about 90 and 44. The shell and the hectocotylus (Berry, p. 278, Figure 4)

■> Fossil shells of Argonauta are very rare and the recent "paper nautili" are also rarely found complete
without the animal. Fragments are occasionally dragged. This would therefore not contradict our view
that the genus cannot be traced back to the Cretaceous, as our concept of its relationship to the Ammonites
requires (see p. 781).

5110/2

738

differ markedly, to judge from the figure (p. 280), although Berry did not
examine the hectocotylus in detail: the shred of skin at the base of the
penis (homologous to the penis sac of Ocythoe) is absent or was over-
looked by Berry, also the inverted sac of the hectocotylus. Particularly
important for identification is the arm formula, which is 1,2, 3,4 in A.bott -
geri, because of the short ventral arms. It is not certain that "Tre -
moctopus" hirondellei (p. 735) Joubin, 1895 belongs to Argonauta
(see p. 736). If it does, A r g o n a u t a includes a form in which the hecto-
cotylus is formed on the right side instead of the left. This would be the
only case in the genus. Except for specific characters, the description of
A. argo applies to the whole genus; at any rate, further distinctions cannot
be made before the morphology and development of the other species have
been studied. Chun (l915) placed a number of young specimens without shell,
including males, in A. hi a ns. However, there are no morphological data
for considering these forms as distinct from A. argo.

Argonauta has been the object of considerable interest for a long time,
but it still remains problematic. The dwarf males, the "wandering" hecto-
cotylus, the characteristic dorsal arms, which were considered as sails in
the past, the genetically obscure shell, which resembles those of the Ammo-
nites, pose a number of difficult questions. We shall have to consider
critically the homology and history of the shell of Argonauta to clarify
the morphology of the genus.

b. ARGONAUTA ARGO L., 1758
1. DIAGNOSIS

Brood shell with a narrow (3—7 mm wide), smooth keel bordered by
dense (2 — 6 mm) tubercles and with smooth, undulate, lateral ridges without
tubercles. Ventral arms distinctly longer than the LK and LV arms
(formula; 1, 4, 2, 3). Hectocotylus with about 90 suckers.

This is the diagnosis of the Mediterranean form. Very similar shells
from the Cape of Good Hope (Museum of Natural History in Stuttgart) are
much larger: length 26 cm, width of keel 10 mm, tubercles 9 mm apart on
one side. Monterosato distinguishes several species and varieties of
Mediterranean Argonauta by the structure of the shell. All my spe-
cimens correspond to his A. argo L. var. m e d i t e r r a n e a, and I cannot
decide whether this variety is valid, especially as no differences between
the animals are known.

739

764 LITERATURE

- Aristoteles, Navrikog (lib. IV, Cap. I, lib. IX., Cap. XX).

- Plinius, Nautilus, Pompilus orNauplius (lib. IX, Cap. XXIX, XXX).

1758 Linne, Argonauta argo (T. 1, pars VI, p. 3367).

1553 Belon, Nautilus (lib. II, p. 378).

1554 Rondelet, N a u ti 1 u s, P o 1 y p u s pars andl Argonauta argo (lib, XVII, p. 517).
1558 Gesner, N a u ti 1 u s, P o ly p us (lib.IV, Vol.IV, pp.732, 734).

1642 Aldrovandl, Nautilus (lib.III, Cap.III, p.257).

1650 Johnston, Nautilus (lib.III, Cap. I, p, 39, Plate X).

1658 Bontius, Nautilus (lib.V, Cap. XXVII).

1666 Clear, Nauplius, Pompilius (p. 66, Plate 32).

1677 Legato, Nautilus (p.l05).

1684 Bonanni, Nautilus, Nauplius (pp. 142, 436, Plate 1).

1685 Lister, Nautilus maximus (t.VII, lib.IV, tab. DLVII).

1687 Rumph, Nautilus tenuis (tabs. XII, XVIII).

1689 Lister, Nautilus (lib.IV, sect, IV, cap. II).

1702 Petiver, Nautilus (tab. VI).

1704 Valentini, Nautilus (Vol.II, p.l83, tab.XXV),

1716 Lochner, Nautilus, Pompilius (p. 70, 1716, tab. 19).

1724 Valentyn, N a u t i 1 u s minor (p.58, tab.l).

1726 Kindmann, N a u ti lus (p.l24).

1742 D'Argenville, Nautilus papyraceus (p. 250, tab. 8).

1742 Gualtieri, Cymbium maximum and minus (tabs. II, XII).

• 1748 Hill, Nautilus (t.III, p.l22).

1748 Lesser, Cymbium (p.l49).

1753 Klein, Nautilus sulcatus (p, 3, tab.l).

1755 Geve, Nautilus papyraceus (p. 11, tab. 2).

1758 Seba, Nautilus nitidus (III, tab. 84).

1762 Ginanni, Nautilus (tom. II, tab. III).

1766 Knorr, Ammonia (tom. I, p. 40),

1769 Martini, Ammonia p a py r a c e us (I, p.230, tab.XVII),

1773 Selis-Marschlins, Nautilus (p.360).

1775 Favart-d’Herbigny, Argonauta (tom.II, p.419).

1778 Da Costa, Nautilus (?) (Tab.III).

1780 Favanne, Nautilus papyraceus (tom. I, pp. 707, 709, 710, tab. 7).

1780 Born, Argonauta argo (pp. 119, 140).

1781 Schroter, Argonauta (Cap. XX, tab. V).

1781 Gronovius, Argonauta (p.284).

1784 Favanne, Argonauta (p.57).

1784 Schneider, Nautilus (p.l20).

1789 Bruguiere, Argonauta argo (p.l22, tab.LXVII).

1791 Shaw, Argo na u t a, N a u til us papyraceus (t.III, p. 101).

1791 Wolfen, Argonauta argo (p.235).

1792 Olivi, Argonauta argo (p.l29).

1797 Humphrey, Argonauta corrugata (p.6).

1800 Cubieres, Nautile papyrace (p. 43, pi. 4).

1802 Montfort, Argonaute papyrace (t.III, p. 119, pi. 25, 26).

1802 Bose, Argonaute papyrace (t.III, pi. 27).

1808 Montfort, Argonauta argo (t. II, p.6).

1811 Wood, Argonauta haustrum (p.62, tab.V).

1814 Rafinesque, Ocythoe tuberculata (?) (p.l9).

1816 Duvernoy, Argonauta argo (Vol.III, p.l02).

1817 Schumacher, Argonauta argo (p.268).

1817 Cuvier, Argonauta argo (Vol.2, p.8) (3rd ed.).

1817 Leach, Ocythoe antiquorum (p.l39).

1818 Burrow, Argonauta argo (p. 75, pi. 12).

740

1820 Ranzani, Argonauta argo (I.Dec., p.85).

1822 Lamarck, Argonauta argo (Vol.ll, p.355, 12th ed.)*

1825 Delle Chiaje, Argonauta a rg o (Vol.II, p.219). Hectocotylus a "worm": "Tricocephalus
ace ta b ul us."

1825 Ferussac, Argonauta argo (p, 160, pi. 14).

1825 Blainville, Argonauta argo and Ocythoe argonautae (p.366, pl.l).

1826 Payraudeau, Argonauta (p.l72).

1826 Risso, Argonauta argo (Vol.4,p.4).

1826 Blainville, Argonauta argo, c o m p r e ssa, O c y tho e tuberculatus and antiquorum
(tom,XLlII,p.l96).

1826 Poli, Argonauta argo (tom. Ill, pars 1, p.4).

1826 Ferussac, Argonauta argo (Vol.l, p.552).

1826 D’Orbigny, Argonauta argo (p.l37).

1827 Blainville, Octopus antiquorum (tom.XLIII,p.l92).

1827 Mauriani, Argonauta argo (p.390).

1828 Broderip, Argonauta argo (pp. 57, 224, pi. 5).

1829 Costa, Argonauta argo (p.61).

1829 Sangiovanni, Argona uta argo (p.322).

1832 Deshayes, Ocythoe argos (tom. Ill, p. 643).

1833 Oken, Argonauta argo (Vol.V, Abt.I, p.532).

1835 Ferussac and d’Orbigny, Argonauta argo (p.l58).

1836 Scacchi, Argonauta argo (p.l9).

1837 Rang, Argonauta argo (p.286).

1838 Potiez and Michaud, Argonauta argo (I,p.2).

1841 Cantraine, Argonauta argo (p.20).

1841 Costa, Argonauta argo (p.l84: considers the Hectocotylus as a spermatophore).

1844 Philippi, Argonauta argo (p.201).

1845 Power, Argonauta argo (p.369).

1846 Reeve, Argonauta argo (II, p. 305).

1848 Requier, Argonauta argo (p.87).

1849 Gray, Argonauta argo (shell) and Ocythoe tuberculata (animal).

1851 Verany, Argonauta argo (p. 48, Plates 17, 18. Hectocotylus Argonautae, Plate 49, Figs. 1-11).

1852 Dunker, Argonauta gunneri (p.48).

1855 D'Orbigny, Argonauta argo (pp. 226, 234).

1857 Troschel, Argonauta argo (p.42).

1858 Adams, Argonauta argo and gunneri (Vol. 1, p.25).

765 1858 Reeve, Argonauta argo, gunneri and haustrum (p.305).

1867 Weinkauff, Argonauta argo (tom. II, p. 432).

1869 Targioni-Tozzetti, Argonauta argo (p.9).

1879 Tryon, Argonauta argo (p.l39).

1880 Tiberi, Argonauta argo (p.5).

1882 Verrill, Argonauta argo (p.l82).

1885 Hoyle, Argonauta argo (p.4).

1887 Fischer, Argonauta argo (p.335, Figs. 117-119).

1890 Colombatovic, Argonauta argo (p.5).

1890 Carus, Argonauta argo (p.457).

1894 Pelseneer, Argonauta argo (p.207).

1896 Jatta, Argonauta argo (p. 191, Plates 8, 18).

1907 Marchand, Argonauta argo (p. 73, Fig. male).

1912 Naef, Argonauta argo (p.l98).

1915 Pfefferkorn, Argonauta argo (p.444).

1916 Naef (Syst.), Argonauta argo (p.l7).

1920 Wiilker, Argonauta argo (p. 56). (Red Sea).

1921 Naef (Syst.), Argonauta argo (p.56).

741

3. TYPICAL STRUCTURE OF THE EARLY YOUNG STAGES

A strictly ontogenetic description is advisable, since the very charac-
teristic formations of both sexes develop only during postembryonic
development. We begin with the freshly hatched larvae (Figure 455), of which
a large number were obtained from a captured female. They are lively and
slightly longer than 1 mm (with arms), and they move in leaps. The habitus
is typical for the earliest stages of Argonautidae (p. 729), but they show
some distinct characters. The chromatophores are very few and of constant
position (Vol. II, Plate XXXVI, Figures 4, 5, 6), Those situated on the bottom
of the dorsal mantle sac appear to be external because of the transparent
body (Plate X). Superficial chromatophores are present on the posterior
end and at the anterior margin of the mantle. Arms and funnel bonds are
particularly characteristic. The arms are markedly different already at
this early stage, and show the formula: 1, 2, 4, 3; the dorsal arms are
markedly longer. However, the difference in length is caused almost
entirely by the development of the whiplike terminal part, while the basal
part of all arms usually shows 3 fully developed suckers, as in Octopus.

If the arms are spread (Vol. II, Plate XXXVH, Figure l), a uniformly
developed interbrachial membrane becomes visible which almost always
takes the form of a "cuff" if the arms are held close together (Figure 455).

FIGURE 455. Freshly hatched Argonauta argo. 40 x.
Compare with the youngest larvae of Tremoctopus
(Figure 437 on p. 736) and Ocythoe (Figure 447A on
p. 750), particularly the relative length of the arms and
the closure apparatus of the funnel, which is detached and
is visible when the mantle margin is folded back. Typical
characters: the cuff-shaped interbrachial membrane; the
small eyes, which are directed slightly ventrally and
anteriorly, and the oval opening of the primary lid; the
few chromatophores; the shining spines, which are visible
only on the dark parts of the body and are rarely split;
large, flat olfactory tubercle; dorsal suture of mantle;
funnel and funnel pockets and insertion of funnel retractor
on funnel bond.

766 The closure apparatus of the funnel already resembles the definitive

structure, as in Tremoctopus and as was assumed for Ocythoe. The
funnel corner fits into a funnel bond which is formed, as in the Decapoda,
like a dishlike disk with a margin. However, the funnel corner is curved
around a transverse, sharply projecting tubercle of the inner side of the
mantle as in all Argonautidae (A1 1 o p o s u s?). The same structure is
observed in the adult.

The freshly hatched animal usually bears the rest of the external yolk
sac which firmly closes the fully developed mouth (Plate X, Figures 5 and 6).
The sac disappears completely in a few hours; sections of larvae at this
stage show yolk and membrane inside the dilated fore intestine ("crop").

This is the youngest form of Argonauta known from the plankton. The
animals differ from other young Octopoda in their small size and they are
the smallest, free young stages of Cephalopoda; they are slightly longer than
1 mm with extended arms.

742

FIGURE 456. Young female (a) and
male (b) of A rg o n aut a argo
from the plankton in Naples, 10 x.

(c) shows the right 3rd arm of the
female, 50 X, with 3 larval suckers
(4) and with "anlagen" of 4 further
suckers (3, 2, 1). Compare with
Figure 455, and note: 1) absence of
spines; 2) strong arms; 3) detach-
ment of the 3rd left arm of the male
from the interbrachial membrane.
The structure has little changed
otherwise .

FIGURE 457. Young stages of Argonauta
argo from the plankton in Naples, a) Male
or female? 12 x. The arms are retracted deep
into the cuff (they are often not visible in
badly preserved specimens), and the body is
contracted, b) and c) Female and male of
Figure 456, 10 x. Note the invaginated hecto-
cotylus of the male; closure apparatus; funnel
retractors; deeply situated stellate ganglia;
vena cava, anal papilla and origin of median
mantle adductor. d,e) - intact male of simi-
lar size, 12 X. Note the fused and free mantle
margin, invaginated hectocotylus, and the
funnel, which is situated deep in the swollen
tissue of the head.

767 The later external sexual dimorphism is not yet recognizable, but the

inner genitalia are already beginning to develop. I found only a single such
larva in the plankton (p. 7 65).

The dimorphism is already distinct in a planktonic larva about 3 times
the size of the above specimen (Figures 456 and 457) in the differentiation
of the hectocotylus, but the male resembles the female in other characters.
The hectocotylus becomes detached early from the other arms and is not
visible externally, except that it can be recognized through the transparent
interbrachial membrane between the 2nd and 4th arms (Figure 470).

The greater part of the funnel is embedded in the soft subcutaneous
tissue, so that only its apex projects. The eye still lacks a secondary lid
fold. The primary lid fold is transparent and can be contracted to a small
pore, so that it functions as a cornea. It is moderately dilated in the
drawing. Upper and lower lid are not yet differentiated. These stages
show the beginning of a typical development of the arms: further "anlagen" of

743

suckers develop and are displaced alternately to the right and left. This
also takes place with the earliest suckers, which later form a zigzag row
from the first sucker onward. The two sexes differ later markedly and will
therefore be described separately.

4. DEVELOPMENT OF THE FEMALE

In a female about 5 times the size of the youngest larvae, the terminal
parts of the dorsal arms, which are still without suckers, are curved
together in loops, and there is a skin fold between the loops which is

contracted in preserved specimens
and causes the coiling of the arms
(Figure 458, l). The picture is
similar in slightly older stages, in
which the development of the suckers
has advanced to the apex, as on the
other arms. Shell glands are still not
recognizable in total preparations, and
there is no shell. The primary lid
is divided at this stage into an upper
and a lower lid, which are crossed at
the corners. The secondary lid is
distinct around the opening if the
circular muscles are contracted. The
arms are further developed, and the
cuff is still indicated (2) but it is about
to disappear. The ventral arms have
grown larger than the dorsolateral arms, so that the definitive formula is:

1, 4, 2, 3.

Figure 459 shows the smallest Argonauta which may have had a shell,
but which was lost. The shell arms are completely developed, with shell
glands and suckers which extend to the apex (Figure 465). The shell is
absent, but Jatta published drawings of similar animals situated in the shell.
The drawing of the juvenile shell (Figure 459c, dorsal view) is based on
Jatta's drawings. Figure 459b corresponds to the nucleus of a slightly
older shell (Figure 460), which can be recognized by the growth lines.

The shell illustrated corresponds to the size of the animal; however,
it is not the primordial shell but a shell the margins of which have grown
considerably according to the modus of the older shells, as shown by the
growth lines. The primordial shell is only the nearly hemispherical,
usually irregularly compressed initial part, which is always distinctly
delimited (5) from the later formed part. The reproduced drawing of Jatta
(Figure 459c) is apparently not quite exact (cf. Jatta, 1896, Plate 18,

Figure 27) and its scale is not given. At any rate, its form does not
correspond to that of the typical shell nucleus. Hoyle (1904, Plate 10,

Figure 12) published a better drawing. Surprisingly, there is hardly a trace
of the bipartite structure which could be expected from the activity of two
shell arms, except perhaps in the often slightly irregular form of the apex in

FIGURE 458. Young female of Argonauta argo,
still without a shell (?),but already with differen-
tiated shell- producing arms (1). 9x. It does not
differ much from the younger stages (Figures 456
and 457), but the arms are stronger, the ventral arms
have grown longer, the lid apparatus is differen-
tiated, and the reddish brown chromatophores are
more numerous. 2 - cuff.

768

744

the median line. Its formation is not known. It is undoubtedly produced
by the same shell glands which form the later developing parts, because the
substance is identical. There is also a fine striation, caused by the continued
growth of an at first small "anlage." The striation is concentric, but not
769 distinctly eccentric as in the later parts (Figure 459b).

FIGURE 459. Young female of Argonauta argo.

6x. At the bottom is its shell, reconstructed after the
growth lines of a slightly older shell. Figure c shows
a dorsal view of a similar young shell (after Jatta,

Plate 18, Figure 27). Note the growth lines of the shells;
the boundary (5) of the hemispherical "primordial
shell"; the always strongly shrunken dorsal arms (1)
with the "shell membrane" (2). Note in comparison with
Figure 458: the arms are longer; the cuff has disappeared
and is replaced by a normal interbrachial membrane;
the more numerous chromatophores; the increase in size.

3 - growing dorsal margin of shell; 4 - transition of
dorsal margin into the ventral margin.

FIGURE 460. Young female of Argonauta argo.
3x. Drawn after a preserved specimen. Note parti-
cularly the sculpture of the further developed shell,
the dorsal margin (2) of which is characteristically
curved and strengthened, and the lateral corners (1)
are sharply defined. Ridges (4) on the sides of the
shell end in tubercles (5) on the ventral side. These
ridges cross the growth lines and the anterior shell
margin (3).

The development of the primordial shell forms a special problem,
distinct from further growth. How does the primordial shell attain its form,
from which further growth can continue? It can be assumed that the
developed shell arms of a slightly younger stage are curved back and are
spread on the soft posterior part of the body in the same manner as they are

745

later spread on the shell. The arms could secrete shell substance inward
and form a shell which covers the posterior end of the body in the middle.
The main characters of the future form are already recognizable in the
juvenile shell shown in Figure 459. The ventral margin extends anteriorly
by eccentric growth, and the shell becomes boat-shaped, with the opening
directed anteriorly and upward. A small, flat process of the shell margin
develops on each side of the widest part. These processes, called "lateral
corners," divide the margin of the shell into an anterior and a dorsal
margin. The lateral corners later become more prominent. The shells
of these stages are not quite smooth but have an indistinct sculpture which
becomes more distinct later (Figure 460).

All arms have become longer. The suckers are now more numerous
and extend to the apex of the arms, also on the shell arms. The inter-
brachial membrane is still large, but quite normal. The funnel has become
much longer and projects distinctly, as in the older stages. An animal
about twice the size of this stage, shows further development in this direction
but no distinct external changes of the soft body (Figure 460). On the other
hand, the shell has developed markedly and bears regularly spaced lateral
ridges which end in tubercles on the ventral margin. Since the ridges cross
the anterior shell margin and the growth lines obliquely, their position
relative to the margin changes during the development: they begin at first
dorsally and reach to the tubercles at the ventral side. This is an interesting
problem. The ridges are irregular at the margin, especially in the later
770 stages (Plate XI, Figure 2 ), but become regularly spaced near the tubercles,
so that there is the impression that the formation of the tubercles follows
a deliberate pattern, the morphodynamic importance of which is of par-
ticular interest (p. 771). The tubercles at first appear in more or less
regular pairs, more rarely alternating, and leave between them a flat ventral
stripe which is at first relatively wide and appears more like the bottom of
a boat than as a keel, although it is called a "keel" in the later stages. If
the tubercles have become distinct, the two rows are separated by 3—4 mm;
this distance later increases slowly and only slightly. The lateral corners
are now very prominent and become even more prominent in the later
stages; they may form horn-shaped processes which are directed markedly
laterally and resemble the "auricles" of ammonites. I never found such
structures in older specimens or in very small mature animals. To judge
from the growth lines, however, all specimens had such auricles temporarily.

A new character appears in slightly larger shells: the blackish brown
coloration of the rows of tubercles and their immediately adjacent parts
(Figure 465) in the earliest, dorsally directed parts of the shell. This
coloration is irregular and asymmetrical, and can therefore not be used for
identification of the species; the same applies to the auricles. The colo-
ration closely resembles the deposition of black substance on the shell of
Nautilus (Figure 7 on p. 55 ). The dorsal margin of the shell also shows
a brownish pigmentation at an early stage.

5. THE ADULT FEMALE

Adult females differ markedly in size. I found mature animals with a
shell only 4.4 and 5,2 cm long and already with eggs in the shell.

746

Figures 461,462,463,465 and Plate XI show the largest specimen I studied
and observed alive; some of my embryological material was obtained from
this animal, which had a shell 13 cm long. I saw a specimen in the collection
of the Zoological Station that was about twice as large, but I did not measure
it. The description of the adult female is based on the above specimen in
comparison with numerous others, most of which were preserved (cf. p. 763).

'771)

FIGURE 461. Female of Argonauta argo in swimming position. 0.5X.
Note particularly that the dorsal arms are placed around the shell along the
keel and cover the shell with the shell membrane. Also note the curvature
of the other arms, the prominent eyes, the large funnel, and the loose position
in the shell.

(771)

FIGURE 462. Live Argonauta argo (photograph).
Above: normal swimming position. The shell arms are
retracted, like the other arms, and partly shine through.
The animal is strongly contracted. Below: lying on the
bottom of a small vessel, agitated, the arms groping on
the walls of the vessel instead of holding the shell. VgX.
Note particularly the natural position of the live Argo-
nauta in view of the numerous unfortunate drawings
which show the animal "sailing" or in an impossible
position, like swimming on the back (Lessona, 1867,
Figure on p.l5). Such illustrations have been repeatedly
published.

747

Plate XI and the following figures show the natural posture of the
swimming animal. All these drawings are based on photographs and
drawings of live animals. Exact illustrations are especially necessary in
view of the impossible figures in manuals and textbooks (e. g. in the Zool.
Handwbrterbuch of H. E. Ziegler, 2nd ed., p. 46). Some of the drawings show
the animal with the ventral side upward inside the shell and with the shell
arms extended upward, like sails.

771 The structure of the shell (Plate Xl), has to be derived from the
morphology of the juvenile shell (Figure 459). It resembles the shell of the
Ammonites in form and in its position toward the medium during the
swimming. However, in contrast to the shell of Ammonites, a closed shell
duct is not developed (Figure 460). There is no dorsal cover, and the shell
forms a boatlike cavity which is completely open until the earlier formed
parts are folded over in the posterior dorsal region as a result of their
spiral growth. Growth is by marginal apposition, as in the shells of
molluscs, which are secreted by the mantle. The anterior margin differs
distinctly from the dorsal margin. The anterior margin grows very uni-
formly; only the ridges and tubercles (p. 770) complicate the otherwise
simple process. A causal explanation would be as follows: All arms,
including the shell arms, grow for a long time at the apex, as in other
Octopoda. The arm follows the shell margin, and it must gradually lag
behind (Figure 4 63), perhaps in proportion to the displacement of the ridges
on the shell margin. Each ridge would then correspond to a certain part

772 of the arm, about the interval between 2 suckers (p. 770). However, this
cannot be correct, because the number of ridges is much larger than that
of the pairs of suckers which advance gradually from the ventral margin of
the shell (Figure 465). On the other hand, the number of suckers along the
shell margin and in the zone of the shell gland is very large. Such a
mechanical explanation is therefore not acceptable. As in the typical shells
of molluscs, the formation of a geometrically complex and definitely curved
shell has to be explained by the activity of glands for which a simple
mechanical determination cannot be assumed. The problem can only be
mentioned here; we will discuss it elsewhere. This obviously involves
fundamental problems of organic morphogenesis.

The course of the ridges shows many interesting details, particularly
their sequence. In addition to the main ridges, which begin at the dorsal
margin and end in tubercles, there are intercalated or secondary ridges
which also end in tubercles but do not reach the dorsal margin. The
secondary ridges can be divided more or less distinctly into ridges which
occupy the ventral quarter, half, or about three quarters of the main ridges.
The sequence of ridges seems at first rather irregular.

The tubercles, except for the first, which are indistinct, show an individu-
ally characteristic form in each animal which, however, varies considerably
in the different specimens. Most tubercles have a blunt, hollow main part
and a solid apex which is slightly transversely compressed. The distance
between the tubercles of one side varies from 2 to 5 mm, and it increases
gradually but decreases in relation to the size of the shell. The two rows
of tubercles are at first separated by an interval of 3—4 mm, which increases
gradually to 7 mm. This stripe forms a slightly undulate bottom of a boat
or keel the relative width of which decreases with the growth of the shell.

748

and the shell appears more markedly laterally compressed. Except for a
light median stripe, the dorsal, anterior and posterior parts of the keel are
covered with a black substance which at first covers the tubercles. The
dorsal margin is also slightly pigmented, not its extreme edge, but a stripe
slightly distant from it; this coloration becomes most distinct during
maximal development of the auricles (Figure 4 63). To judge from the
growth lines, this stripe corresponds to the auricles, i. e. to the differen-
tiated upper part of the anterior margin; it is not added from above. At
the lateral corners, the stripe reaches not only the anterior but also the
dorsal margin; the growth of the dorsal margin adds new shell substance
in the form of a large ridge which resembles the columella of the shells of
Gastropoda. The other part of the shell is paper-thin; its detailed
structure shows no resemblance to shells of the Tetrabranchiata, although
it consists of a conchinlike basic substance with calcareous deposits, like
the shells of molluscs. The calcareous substance can be dissolved from
773 fragments of shell without changing their form. The wall of the shell

consists of an outer and a thinner inner prismatic layer. The layers are
formed by a bilateral deposition of shell substance on a fibrous basal layer.
Both sides of the shell are smooth and shining, except the ridges and minor
irregularities of the growth lines.

FIGURE 463. Adult female of Argonauta a rg o in normal swimming position, 0.5 x.
The shell is shown transparent (diagrammatic). The habitus is like that of the Octo-
poda. The arms are curved back as in Ocythoe and bear suckers which are longi-
tudinally connected. Note the large funnel. The eyes are prominent, with typical lid
apparatus and rounded pupil. The mantle sac is fitted into the shell. An egg mass is
visible in the dorsal part of the shell (4). Note also the form of the shell. The left
shell arm is removed, the right arm is visible at its passage to the shell.

1 - mouth; 2 - false protective margins; 3 - shell membrane; 5 - columella.

Except for the shell arms, the body does not differ markedly from that
of the typical Octopoda and is situated in the shell in the normal swimming
position as shown in Figure 463. The larger the animal, the closer the
adaptation of the form of the mantle sac to the space of the shell. In large

749

animals, the mantle sac is swollen in the middle, while it continues pos-
teriorly in an upward curved diverticulum and is slightly laterally com-
pressed. The anterior margin is situated near the eye. The eye is very
prominent, especially in life, and is directed laterally. Its position relative
to the shell closely resembles that in Nautilus (Figure 10 on p. 59).
Water pores are absent, but skin folds in their position often form pockets
which are comparable to the water pores but are not constant in form. The
water pores probably disappeared secondarily. The funnel becomes very
large. Only the extreme end of the funnel projects in the male (Figure 468
on p. 785) and in the young female. It forms here a' tube which can be
markedly projected and is inflated like a sac during the expulsion of water.
Its ventral side usually bears 2 ridges which probably function as keels.

(774)

FIGURE 464. Mouth region of Argonauta argo,
2x. Note: margin of lower jaw, inner lip, outer lip;
8 circumoral suckers; bilateral arrangement of the
8 arms; size of suckers on the arms; interbrachial
membranes. Same specimen as in Figure 466.

1 - lower jaw; 2 - upper jaw; 3 - inner lip;

4 — outer lip; 5 — interbrachial membrane; 6 —
false protective margin.

The arms are typically arranged around the mouth and their proximal
parts (Figure 464) are still normal. The very contractile interbrachial
connections are characteristically different. Also the suckers show the
typical differences in size (Figure 464), according to the length and thickness
of the arms. Longitudinal folds (2) connect the suckers of each row and
disappear only at the base of the arms (Figure 466). Except for the specific
structure of the dorsal arms, the formula is still: 1, 4, 2, 3. As Figure 463
shows, the normal position of the arms resembles that also frequently
observed in Ocythoe: all arms are curved posteriorly and, except for the
774 dorsal arms, they are situated inside the shell, their suckers firmly attached
to the inner surface. (A healthy, undisturbed animal never takes the ends of
the arms out of the shell or moves them about; only dying specimens do this in

750

the aquarium, but they may also leave the shell completely.) For feeding
and crawling, the female uses only the basal and middle parts of the arms.

A disturbed animal will retract also the dorsal arms into the shell, like the
other arms (Plate XI, Figure 2; Figure 465a).

FIGURE 465. Shell arms of A rgonauta argo, J.5x. 1) Left shell arm, in the process of leaving the
shell with assistance of the suckers, to spread itself on the outer side of the shell. Note the markedly
lengthened stalks, b) Right shell arm, spread over the shell, with spread margins, seen from the inner side.
Compare with Figure 463 on p. 773. Identify the parts of the arm following the keel, the anterior and the
dorsal margins. In the living animal, the latter two parts of the arms are folded over.

1 — median false protective margin; 7, 8 “lateral false protective margin; 2 — shell membrane (free
margin); 3 - crossing; 4 - end of arm folded over the columella (9); 5 - marginal shell glands; 6 - sub-
marginal shell glands; 10 — growing anterior margin of shell; 11 - black substance of keel; 12 - stalk of
sucker; 13 ~ ridge; 14 — tubercles of keel.

The structure of the dorsal arms is best recognizable when the arms
are spread on the shell, as during undisturbed swimming (Plate XI, Figure 1;
Figures 463 and 465b). They are folded back over the dorsal side, they reach
the dorsal part of the keel and continue along it, medially to the tubercles,
with the suckers facing the shell. The dorsal arms then follow the anterior
margin, one of the rows of suckers attached to the inner side, the other to
the outer side, or both rows attached to the inside, to the lateral corners.

The terminal part of the arm follows the dorsal margin of the shell. The
whole lateral surface of the shell would then be exposed if not for the
presence of a thin skin fold in the loop, the "shell membrane." This is
homologous to the skin which fills the loop of the dorsal arms in Ocythoe,
and also begins from the median attachment of the interbrachial membrane
(Figure 449 on p. 753). However, the membrane does not simply fill the
loop in A r g o na u t a, but its free margin has been slightly displaced from
the apex of the arm and ends slightly further anteriorly and ventrally on the
outer side of the surface. The terminal lobe, which is folded over the dorsal
margin and toward the inner side of the shell, becomes then so independent
that it can reach the more distant median parts of the columella (p. 772).

751

The membrane adheres normally to the outer side of the shell (Plate XT)

775 and reproduces most of its rugosities. During its spreading, the suckers
hold and gradually stretch the membrane until it covers the sides of the
shell completely. They finally extend beyond the tubercles of the keel, and
come in contact with the arms of the other side, so that the margins slightly
overlap, and the keel is also more or less completely covered. The anterior
margin of the shell is covered by the folded over shell membrane, while the
suckers are situated on the inner side. (The fold is spread in Figure 465 to
show the shell glands.)

FIGURE 466. Situs of mantle cavity of a young female of Argonauta argo. 2x. The mantle is
opened and spread. The "closure bond" (12) of the funnel is shown artificially opened, not curved inward
as in the preceding figures, which show the natural condition. The dorsal and ventral margins surround the
mantle bonds (13) like a clasp in the live animal, but are usually detached after death. The funnel (16)
is maximally contracted by fixation in formol. Note following superficial structures: vena cava (9); anal
papilla; median mantle adductor (11), reduced to a narrow band and cut; funnel retractors; anterior mantle
adductors; stellate ganglia; gill ligaments; gills; branchial hearts; superficial gill retractors; renal and
genital papillae (5). The following organs are visible below the surface: venous appendages (6); vena
cava; hind intestine (7); visceral nerves (10); oviducts (3), still only slightly undulate, with a median
entrance and vesicular widenings (1). The ovary (4), which fills the entire posterior part of the body. Muscles
cause the formation of skin folds from the arms to the funnel (18) and eyes (17) and to the dorsal side of the
head. These folds provide the explanation of the water pores of Tremoctopus and Ocythoe (Figure 450
on p. 754).

8 — median mantle artery; 14 — false protective margin; 15 — interbrachial membrane.

752

The membrane is usually irregularly contracted in the preserved animal.
The shell arms bear 2 rows of rather normal suckers at the margin of the
membrane. They have different functions, as their structure and arrange-
ment show. The suckers of the outer (median) rows have long stalks and
their main function is to stretch the shell membrane (Figure 465a). The in-
ner (lateral) rows consist of very low, sessile suckers, except the normal basal
suckers. The suckers of this row apparently only attach the shell membrane
to the shell. The suckers of the outer row are connected by a continuous
fold which projects slightly above the suckers and thus forms a continuation
of the shell membrane. The proximal suckers are used for creeping,
feeding, etc., like those on the other arms. However, the majority of the
suckers of the dorsal arms in the region of the shell membrane function in
connection with the shell. The folded parts of the other arms also take part
in holding the shell.'’'

776 The shell glands are of particular interest but they have apparently
escaped the attention of scientists. They occupy a wide stripe on the part
of the shell arm which normally encloses the margin of the shell
(Figure 465b). Examination of a good preparation (fixation with chromic
acid) under the magnifying lens shows dense glandular tubules which open
toward the margin but may extend for a considerable distance inward. The
tubules are partly arranged in groups with convergent openings. Between
the longer glands are shorter, flask-shaped glands, especially in the outer
part (see Vol. Ill), situated between every 2 suckers on the margin. How-
ever, the structure of the two types of gland does not differ much, and all
transitional stages in form are found. In the live animal, the marginal
membrane is curved slightly inward over the shell margin, which explains
that the shell consists of two layers. This applies particularly to the end
part of the arm, which produces the columella and thickens it all around.

The funnel bonds described above are spread in Figure 466. The funnel
shows no special characters except its large size (but see p. 672). The
conditions in the mantle cavity are typical for the family and closely
resemble those in Ocythoe (cf. Figure 466 and Figure 450 on p. 754).

The mantle bonds are characteristic for the attachment of the funnel bonds
and also the extreme reduction of the median mantle adductor, which has

777 been reduced to a narrow band near the anus and is the last remnant of a
septum which once divided sagittally the whole (now undivided) mantle cavity
and is still present as "anlage" in the embryo. In contrast to Ocythoe,
the "anlage" of the genital openings is at a distance from the base

of the gills so that a large part of the distal oviduct projects from the
surface. The openings of the mature animal may be invaginated or everted,
i. e. they may form papillae. The proximal oviduct is visible under the
ventral skin, and shows the same arrangement as in Ocythoe. However,
the specimen illustrated is a young female which has only a few mature eggs.
The oviduct is therefore narrow and only slightly undulate. As in Ocythoe,
it covers later the whole ventral side of the posterior part of the body,
particularly the ovary. It may contain over 100,000 eggs in various stages
of development, the most advanced of them in the stage of gastrulation.

The dying animal often leaves the shell or lets it fall. The healthy animal can be taken out of its shell
without injury, but it always returns to the shell and probably never abandons it voluntarily.

753

(The eggs are deposited at widely different times.) The branchial hearts
are often completely separated from the visceral complex and are sus-
pended in the mantle cavity, while they only project into the mantle cavity
in all other Octopoda.

The size of mature animals varies considerably (see p. 763). They are
3.5— 15 cm long without arms. These extremes may be associated with
certain varieties or races, but my material does not permit systematic
conclusions. All specimens from Naples and Messina belong to the same
species, i. e. there are no constant, distinct and important differences.

The live animal has a very delicate and bright coloration because of
the constant interplay of chromatophores and iridocytes. The shell
membrane has a delicate silvery sheen with variegated reflexes. Areas of
metallic violet sheen with blue, reddish and greenish reflexes are present
on the eye, body and bases of the arms. The extended dark chromatophores
produce a warm reddish brown coloration, which may be modified by the
yellow chromatophores. On the characteristic groups of chromatophores
see Figure 455 on p. 765 and Plate XI.

6. BIOLOGY OF ARGONAUTA AND THE FUNCTION
OF THE SHELL

Fantastic ideas on this mysterious genus are found in the literature
which have often been revived and further enriched. Keller (1913, Volume II,
p. 517) reports ancient "absurd stories," and quotes the following from
Pliny:" 'One of the most remarl^able curiosities is the animal known as
Nautilus or Pompilus. This animal comes to the surface of the water
lying on its back, expelling the water through a tube, thus discharging its load,
which enables it to swim easily. It then curves its two anterior arms posteriorly
778 and spreads an extremely thin membrane between them. This membrane
enables it to sail ... 'In fact, the animal uses the lobed arms in calm
weather as oars, not as sails." Both views are, of course, wrong. Pliny
apparently described a stranded animal.

Argonauta leads a pelagic life in the open sea. The animals are
usually isolated drifting on the surface, although they are reported to occur
occasionally in large swarms. Most of the specimens caught are injured.

If air is trapped in the upper part of the shell during an accidental rise to
the surface, it prevents a return to the depth. If the animal holds on to
floating objects or creeps on similar bodies or on the bottom, it keeps the
mouth downward and the shell upward. This position, however, cannot be
considered normal (cf. Part III). Argonauta certainly moves normally
by swimming. The nektonic life makes it necessary to carry the eggs along,
if the typical care of the Octopoda for their progeny is not to be abandoned.
The shell can thus only be interpreted as a breeding device. Such a thin
shell can hardly serve as protection. The related genera and the males of
Argonauta show that the animals can live without a shell. The same
apparently applies to the species as soon as a different form of care for the
brood is found.

754

7. DERIVATION OF THE SHELL OF ARGONAUTA

From the systematic -morphological or phylogenetic point of view, the
problem of this mysterious structure is its apparently sudden appearance;
the typical relationships and gradual development of the genus through
systematic stages — Dibranchiata, Octopoda, Polypodoidea, Heteroglossa,
Argonautidae, Argonautinae, Argonauta — are fully established. We know
that this is a highly specialized form of Octopoda. The development of this
unique apparatus is most surprising. On the other hand, we have indicated
that the shell of Argonauta is not quite as isolated as it appears at first
and that it represents the culmination of tendencies of adaptation that occur
also in other groups and are typical for them. Tremoctopus (p. 740) has
even similarly modified, widened, glandular dorsal arms and structures
functioning as egg carriers which were probably formed by the arms.
Although these structures do not resemble the shell of A r g o n a u t a, they
must be considered as homologues (i. e. substitutes) of the shell, not only
from the morphological but also from the systematic -biological point of
view.

The detachment of the dorsal arms of Tremoctopus makes it
probable that this is a change of an ancient ecological habit and suggests
that its predecessors had a breeding device resembling that of Argonauta.

779 The male of Ocythoe has the peculiar instinct to use an alien habitation'-'
and swimming with it, and his body is situated in it like the female of
Argonauta in the shell. The female of Ocythoe has skin folds of a very
simple type, but which are homologous to the shell membrane of Argonau-
ta. The female of Ocythoe also has a tendency to fold these formations
backward over the body, like Argonauta (Figure 449 on p, 753). It is also
justified to assume for Ocythoe that a modus of care for the progeny
resembling that of Argonauta was abandoned with the transition to
complete vivipary, a condition that may easily have developed from the
typical ovovivipary of the Argonautidae. We assume, therefore, that the
ancestors of all Argonautidae, at least the females, made use of alien
dwellings and deposited eggs in them, and that they probably resembled those
used by the male of Ocythoe. A closely analogous mode of care for the
progeny exists in Phromina sedentaria (Crustacea) and it will be
remembered that all Heteroglossa have the tendency to retreat (with the
posterior end first) into various cavities.*''' Perrier and Rochebrune (1894)
report that Octopus digueti hides in shells of P e c t e n and Cytherea
to deposit her eggs and remains there until they hatch. The eggs are
attached above and below the female on the inner side of the brood cavity,
and she guards them like O. vulgaris, although the circumstances are
slightly different and O. vulgaris builds the cavity of stones. The
advantages of a movable habitation could lead to permanent adaptations,
perhaps as took place in hermit crabs.

This behavior is only the manifestation of general instincts and typical
habits of Polypodoidea. However, there are further similarities: Octopus
vulgaris also attaches the eggs with some sticky secretion which lines

The Octopodidae usually hide in cavities (shells Jugs, stone cavities) and assume a similar position if
in danger.

Italian fishermen exploit this habit; they tie jugs together and sink them in shallow water. If the jugs
are hauled up, Octopodidae are found in some of them.

755

parts of the brood cavity and bears the egg batches.''' It may be assumed
that the skin glands of the arms of Argonautidae also have a function in
the care of the progeny and have undergone a special modification. This
may have been indifferent at first (formation and attachment of the eggs,

780 cementing the brood cavity, etc. ), and such a development would be an

important precondition for the formation of the shell of Argonaut a. It
is no longer necessary to assume that a mysterious creative force suddenly
induced the animal to invent a complete brood shell from scratch, or that
Argonauta had preserved an exact reminiscence of its ancestral shell
through the whole sequence of ancestral Dibranchiata, Octopoda, Polypo-
doidea, Heteroglossa, Argonautidae and Argonautinae, or that it has kept the
memory as a secret or, at least without a trace. The enormous distance
between Tetrabranchiata and Dibranchiata should be realized and the great
difference between the normally chambered shell of Decapoda and the
rudiments which have persisted in the Octopoda most closely related to the
Decapoda.

The ancestral Argonautidae probably deposited the eggs in empty shells
of other molluscs, and this habit might have been established so firmly and
at such an early stage that further adaptations could have developed (as in
the hermit crabs, which behave similarly). Such an adaptation would have
been the enlargement of the alien habitation to avoid the frequent moves
which would be necessary because of the rapid growth of Octopoda (p. 699).
The Paguridae also provide an analogy in this respect, but (to my knowledge)
not a complete analogy: Eupagurus prideauxi Leach is often found in
much too small shells (e. g. Natica) in which it has place only after the
protecting anemone Adamsia palliata Boh. has enlarged the margin by
a chitinlike secretion. This may have been done by the activity of the glands
of the arms in the ancestors of Argonauta.

The above data make such an adaptation very probable, and the hypo-
thesis agrees well with the principles of systematic morphology. It is
necessary to prove the possibility of a continuous transition from normal
Octopoda to the shell-bearing A r g o n a u t a, and this has been done here.

It has only to be assumed that the acquisition of the definitive dwelling has
taken place at an ever earlier stage (as in Eupagurus) and that the activity
of the arm glands has increased correspondingly, until the conditions in
A. argo were reached. However, this apparently did not take place exactly
according to the above hypothesis. One would expect, according to the above,
that the early "anlage" of the shell of the recent Argonauta is also
connected with an alien nucleus. This is certainly not the case in A. argo.
The primordial shell of this species (p. 768) has the same origin as the other
part of the shell, although its formation has not been directly observed.
Embryological development provides no confirmation of this hypothesis,
because Argonauta builds its shell quite independently. However, this
does not prove that the ancestors of Argonauta were also able to do this.

In any case, the above derivation seems to me the only one possible, not only

* I could not determine the origin of this secretion. In Octopus vulgar is, the secretion forms a
rapidly hardening, granulate, grayish green slime (cf. Vol. II), which forms a crust on the egg batch and
cements the chorionic stalks into clusters. The oviduct gland (Figure 407 on p.684) is much too small
for the production of such quantities of secretion, if it is compared with the size of the nidamental glands
ill the Decapoda (p.l85). The participation of skin glands is also not very probable.

756

because of the analogies (Adamsia - Eupagurus) and the behavior of
other Argonautidae (female of Tremoctopus, Ocythoe of both sexes),
but also because of the basic postulate that the formation of this charac-
teristic structure should be derived in stages from the typical basic forms

781 in its general outline. This hypothesis probably has therefore its
scientific validity.

Some special problems remain. The very specialized character of the
shell of Argonauta requires a special explanation. This is not simply
a brood shell but a very peculiar nursery with characteristic "adornments"
which the swimming animal carries along. All these characters have to be
explained in their connection. The external resemblance to the shell of
Tetrabranchiata is not very surprising ecologically because this is the only
functional form adapted to a certain type of movement (p. 83) and at the
same time fulfills the geometrical requirement that it can grow indefinitely
while the mass is markedly concentrated. (This is the general significance
of the spiral shell of molluscs.)

An explanation of the special form of the shell of Argonauta has to
be given according to the general hypothesis. The selection of a suitable
alien shell is decisive. It must be very light, more or less symmetrical,
or resembling the definitive or juvenile shell of Argonauta. Such shells
are rare among the recent species, but Brachiopoda or mussels have
similar shells. Among the East Asiatic Cardiidae, for example, there are
some thin-shelled, oblong -conical, spiral forms resembling that of Argo-
nauta. Such a shell could have served as a model for Argonauta
(consider, for example, Cardissa c a r d i s s u m (L. ), the boat- shaped
halves of which form together a conventional "heart form"). The search
for such a shell need not be restricted to recent forms. The high efficiency
of the shell-producing apparatus of Argonauta and the variety of forms
in the whole family suggest an older geological age. We have to consider
also extinct types, at least from the Lower Tertiary (cf. p. 762). The shell
of the Cretaceous Ammonites would fulfill the requirements of a brood
shell for the ancient Argonautidae and serve as a model for the recent forms.
The shell of these fossil forms is indeed perfectly suitable for this purpose.

It is difficult to assume that the Octopoda would not have used the numerous
shells of Ammonites scattered on the coasts throughout the world. The only
question is to explain how the ancestors of the recent Argonauta became
adapted and later replaced the borrowed shells of Ammonites and also to
determine how the other Argonautidae behaved. (A specific adaptation to
nektonic brood shells need not be assumed for other Octopoda or Octo-
podidae.)

The Ammonites disappeared almost suddenly toward the end of the
Mesozoic, and the care for the progeny of Argonauta must have been
already established in its main character at that time. On the other hand,

782 the closest relatives of Argonauta did not develop such a behavior and
had to or were able to use other methods. Ocythoe became viviparous
(p. 750); Tremoctopus found a makeshift solution (p. 742 ). A large
group of related forms (which is probable on the basis of morphological
divergence) probably became extinct.

Shells of Nautilus are also suitable for this use. However, they were
never very numerous, and their form is not equally suitable because of the
short living chamber. The sculpture of the shell of Argonauta resembles

757

that of the Cretaceous Ammonites (e. g. Ho pi it es). Not only the general
form but also the lateral ridges, keel and keel tubercles are very similar
and even vary as in the Cosmoceratidae and in Argonaut a. I admit that
I think of a direct imitation by transfer of a kind of "morphological instinct."
However, I will not discuss this metaphysical concept or formulate it as a
scientific hypothesis. In any case, it can be assumed that the shell of
Argonauta developed from such a "model."

The general (p. 780) and specific (p. 781) interpretation of this hypothesis
may appear fantastic and unnecessary. However, the principles and the
factual data should be considered. It cannot be assumed that the arm glands
produced at first an indifferent body for the attachment of the eggs, as in
Tremoctopus (p. 742 ), and that this structure developed in the same family
into a complete dwelling of the perfection observed in A. argo. This
assumption cannot be disproved because of the incomplete material, but it
does not explain the actual development as well as our explanation. '•=

According to G. Steinmann (l 890 ), Argonauta is a direct descendant of
the Ammonites, and has retained their shell in an externally unchanged
form. The above discussion and the facts show that this concept cannot be
accepted. A retention of the form and sculpture after the complete loss of
the typical parts (phragmocone) is in any case impossible for the morphol-
783 ogist to accept.* ** Steinmann bases his view in addition on the following
factual errors:

1. He considers that the "hood" of Nautilus produces the black
substance of the shell, a mistake which we corrected on p. 62.

2. He assumes, according to Appellof, that the "shell gland," i. e. the
"anlage" of the shell sac of Argonauta, develops first but then dis-
appears and becomes the epithelium of the mantle surface. This is not
correct (cf. p. 108), as we shall show in greater detail in the embryological
part. A shell sac develops and becomes constricted in A r g o n au t a, though
in rudimentary form. However, it soon disappears as in all Argonautidae,
while the remnants of the shell of the Octopodidae are not formed at all.

3. He states, apparently as an observation (??), that the mantle of the
young Argonauta produces the "anlage" of the shell (he considers the
mantle of Argonauta as homologous to the mantle of Nautilus because
he has no knowledge of the nature of the "muscular mantle" of the Dibran-
chiata — p. 92) and he considers this statement as the strongest argument

* The phylogenesis of Argonauta can be considered as a sequence of the following stages: 1. Appro-
priation of an alien dwelling, as in the male of Ocythoe and the female of P h r o ni m a sedentaria.
2. Adaptation of the dorsal arms to the form of the alien shell. 3. Use of existing glands for the en-
largement of the shell. 4. Gradual displacement of the alien nucleus by development of an apparatus
for the formation of the shell by the animal itself. On the other hand, the independent development of
this apparatus remains obscure because the animal must acquire at the same time the shell and the
mechanism which forms it. It is miraculous for a soft organ to build a structure of such precise form
and contrary to any gross mechanical concept. We do not believe that nature achieves such perfection
otherwise than in stages and through an ascending series of closely similar forms. Systematic morpho-
logy teaches that this cannot take place in such a narrow range. Therefore, we have to assume the use
of alien material as the explanation.

We would like to point out that Steinmann's surprising views are based on a confusion between typical and
atypical similarities, i.e. between similarities between whole units and similarities between often
negligible parts. All this despite this author’s wide knowledge of facts. It must be made clear in
principle: it does not help to have different opinions on the "relationships of descendence" (see also
Fossile Tintenfische). It is quite clear that this is not a case of typical resemblance.

758

in favor of his theory. But this is wrong. There is no doubt that the
substance of the shell of Argonauta is secreted by the shell glands of the
dorsal arnas and the mantle does not take part in the process. This would
be rather surprising. It has to be remembered that the mantle of the
youngest Argonauta is covered with spines, as in other larvae of Octo-
poda, that it is a muscular mantle of extreme development (pp. 657, 727)
and that it does not correspond at all to the muscular mantle of the Tetra-
branchiata.

8. DEVOLOPMENT OF THE MALE

We described above (Figure 456 on p. 766) juvenile forms of male
Argonauta which have the same habitus as the female stages of the same
size and differ from them only in that their left lateroventral arm begins
to develop into a hectocotylus (Figure 467 on p. 784 and Figure 468 on
p. 785). My material did not permit a detailed study of its development.
However, the known stages resemble so closely the type of the family that
there are no important questions about the homologies.

(784)

FIGURE 467. Hectocotyli of small males of A rg o n a u t a, highly magnified (over 60 x);
stages intermediate between those shown in Figure 456 on p.766 and Figure 468b on p.785).
a) Indifferent stage (cf. Figure 456 on p.766), slightly swollen and widened at the end like
a spoon, with 3 rudimentary suckers (see Figure 445b on p.746). b) Beginning coiling inside
the sac of the hectocotylus, which is already completely contracted, c) Growth of apex
inside the sac. d) Further advanced stage, from the outside.

In the youngest Argonauta that can be recognized as a male, the hecto-
cotylus (Figure 456 on p. 766) begins to be curved inward and to become
detached from the common interbrachial membrane. The arm is at first
very short, blunt and characteristically widened (Figure 467 on p. 784a),
with 2 (rarely 1 or 3) normal suckers, developed or in the "anlage" at the
784 base. Its apex then rolls in (Figure 456b on p. 766), forming a clublike
formation. The sac of the hectocotylus develops very early (l could not
determine the transitional stages in detail) and is closed over the curved
apex (Figure 436 on p. 732). The third of the primary suckers is enclosed
in the sac; the other two remain at first outside, as in Ocythoe, and later
(Figure 456c on p. 766) become strongly compressed and reduced. The apex
grows inside the sac; it rapidly grows longer than the normal arms and
becomes coiled. The "anlage" of the ducts probably develops as described
on p. 728, but I could not follow its development; particularly, I did not

759

observe a state like a groove. In a stage slightly older than that shown
in Figure 4 67d, the spermatophore duct is already closed almost to the apex
of the penis, while the spermatophore reservoir is still absent (it is
apparently a late development in the subfamily). It is distinctly delimited
in the later stages, and forms a shining muscular tube on the dorsal side of
the basal part of the arm; it strengthens, so to say, the backbone of the
curved arm. Its entrance is visible near the end of the reservoir
(Figure 469) as a narrow slit the margins of which are often contracted
into a small papilla. The suckers develop according to the general pattern;
the normal arms of the male remain short and bear only 12 — 13 suckers
(Figure 470), while the hectocotylus bears about 95. The hectocotylus
varies in position: it is sometimes coiled inside the pocket (Figure 439 on
p. 738) formed by the new membrane between the 2nd and 4th arms, or else
it projects more or less free (Figure 468). The former condition is normal
for older but not quite mature animals. There is then a large swelling
below the left eye, so that the whole animal becomes asymmetrical
(Figure 470). In live specimens, the coiled hectocotylus is visible through
the” capsule and interbrachial membrane, if the chromatophores are con-
tracted.

FIGURE 468. Development of the male of Argonauta argo, a, b, c -- 4 X; d, e, f 2 x .
a,b) Young stage. Note the short, compact form; the 7 short arms; the hectocotylus, visible
below the left eye. c) Young animal with projecting hectocotylus (2). d) Young animal
with the hectocotylus enclosed in the sac (1). e) Mature animal with loaded hectocotylus
which is about to break out. Penis (4) and entrance (5) to the spermatophore reservoir (3)
are visible, f) Mature animal with free hectocotylus on which the suckers, the cutaneous
appendages (7) and the penis are visible. 6 — sac of hectocotylus.

There is no longer room for the hectocotylus below the eye of the mature
male, and it therefore projects free between the arms. Such animals are
apparently no longer able to lead a free active life and probably remain

760

785 on or inside the shell of the female, where they have been found occasionally.
When the animal reaches its definitive size (about 15 mm with extended
arms), the hectocotylus becomes fully developed and finally "loaded." The
hectocotylus in the specimen in Figure 4 68e is still completely enclosed
in the sac, but the muscular reservoir (3) already contains the coiled tube
of the spermatophore and therefore projects distinctly beyond the arms;
the arm apparently begins to break through the envelope, the club is very
large and could no longer be carried in its former place below the eye.

(786)

b >

FIGURE 469« Hectocotylus of a mature male of Argon auta argo, 2X.
a) Inner side of the extended arm, with the folds of the sac spread to show
their insertion, b) Outer side, with the spermatophore reservoir (9), its
entrance (6), and the spermatophore duct (7). c) Lateral view of same pre-
paration. d) Arm in natural condition, after copulation and detachment, sac
(10) inverted as secondary spermatophore reservoir, e) The arm still attached
to the animal, before copulation and before detachment, but prepared for it.

12 “ sac before inversion, f) Arm, fully developed, attest, attached to the
animal. The sac is still completely contracted.

1 “ penis; 2 -■ cutaneous margin as continuation of the rudimentary penis
envelope (3); 4 - folds of sac; 5 — point of detachment; 6 - entrance to
spermatophore reservoir; 7 - spermatophore duct; 8 -- insertion of the fold
of the sac; 9 “ spermatophore reservoir; 10 - secondary spermatophore
reservoir; 11 — entrance to 10; 12 “ sac of hectocotylus.

The male shown in Figure 46 8f is ready for copulation and the
hectocotylus has broken out of the reservoir.* We have to assume
that the hectocotylus is detached after entering the mantle cavity

* This takes place at the predetermined line of tear, at about the anterior end of the spermatophore
reservoir. This point is closed by a thin membrane, and is already recognizable in young stages;
it probably corresponds to the point of fusion of the sac (p. 757).

761

of the female and the sac is inverted (Figure 469 on p.786). Hecto-
cotyli have been found in the mantle cavity of the female, and torn
parts of its end also in the oviduct and capsule of the ovary.* The
penis differs markedly from that of Ocythoe; it is actively motile and has
a narrow margin at the base on each side. The margins of both sides are
united ventrally and form a small lobe. This very delicate structure probably
corresponds to the penis sac of Ocythoe (Figure 454b on p. 760).

Figure 4 69 shows the structure of the hectocotylus of the grown but not
completely mature animal (same specimen as in Figure 470).

In addition to their inhibited growth expressed in their small size, the
growing males differ so markedly from the female that they seem to belong

786 to a different species. The body is short and compact. The funnel is small
and only its apex usually projects in the preserved animal, while the rest

is fused with the head (Figure 470 on p. 787). The arms are of nearly equal
length; at any rate, the differences are very small in preserved specimens.
However, closer examination of numerous specimens shows a resemblance
to the female, in that the ventral arms are longer than the laterodorsal
arms (formula without hectocotylus: 1,4, 2, 3); the right lateroventral arm
is markedly shorter than the left. The normal arms bear only 12 — 13 distinct
suckers and often 1—2 indistinct "anlagen" of suckers at the apex.

All arms, except the hectocotylus, are connected by a well developed, rather
uniform interbrachial membrane. It is more strongly developed only between
the 2nd and 4th arm of the left side, where the separation of the hectocotylus
formed a direct connection and the margins of the membrane can enclose
the club of the hectocotylus more or less completely. The membrane may
remain uniform all around so that it gives the impression that there are
only 7 arms in small specimens.

The eyes show the same structure as in the female, but they are less
prominent. The funnel resembles at first that of the female, but never
reaches its large size. Its projecting part remains small, suggesting a
restricted capacity for swimming (nothing definite is known about the
biology of the male). The form of the funnel bond is slightly different from

787 that of the female; it is drawn unchanged in Figure 470a to show the cur-
vature of the posterior margin. It is directed anteriorly and outward,
corresponding exactly to the knob of the mantle, which is situated obliquely
to the mantle margin, not transversely as in the female. (Variety?)

The median mantle adductor is relatively stronger than in the adult
female, about as large as in a female of the same size. The male should
be compared with the young female, not with the adult female, because the .
various parts show inhibited growth. The anal region, stellate ganglia,
gills, renal sacs and renal papillae are as in the female and the posterior
part of the body is also occupied by the gonad, which is restricted in space
only by the large gonoduct. There is no free projecting part ("penis"),
and the gonoduct ends in a simple, slit-shaped opening; on the other hand,
it forms an enormous "penis diverticulum" which covers the left kidney
as a wide sac.

None of the numerous males examined is longer than 15 mm (with arms
but without hectocotylus). This is the smallest cephalopod existing except
for I d i o s e p i u s, which is of similar size. As pointed out above, the small

On fertilization, structure of spermatophore, etc. see Vol.III.

762

size has to be considered as a phenomenon of inhibition. (The marked
788 development of the adaptations in the female may have influenced the

reduction in the male. The male apparently only possesses the bare mini-
mum necessary for reproduction and is not an independent biological factor.)

(787)

FIGURE 470. a) Almost mature male of A rg o n a u t a arg o, with opened mantle,4 x. Note the
resemblance to the female (Figure 466 on p.776) in the following characters: form of funnel bond (6)
which is here of normal form,i»e- curved from behind and slightly from the inside; corresponding
form of mantle bond (7); projecting septum (5) between funnel and funnel pocket; short funnel;
arms and interbrachial membranes, especially between the 2nd and 4th arm of the left side (8);
hectocotylus (9), not yet fully developed but already large; the bare (larval) ends of the arms
(Figure 455 on p.765). Note in the mantle cavity: the enormously developed gonoduct (3) with
sac-shaped end (2) and slitlike opening (1), The other conditions (gills, anterior mantle adductors,
median mantle adductor, kidneys (4)) are typical.

b) Arms of a mature male of Argonauta argo. 4x. Semidiagrammatic. The hectocotylus is
shown much too small to show the other parts and to stress the topographical relationships to the other
arms. Note the short arms, the interbrachial membranes (5), especially between the 2nd and 4th arm
of the left side (3). Note on the hectocotylus: the transverse axis of the arm, the entrance (1) to the
spermatophore reservoir (4), the fold of the sac laterally (2). The normal mouth with lower jaw, inner
and outer lip.

The coloration resembles that of a female of the same size: blue and
violet sheen, especially on the dorsal side and eyes; yellow and reddish
brown chromatophores, dark brown to blackish if contracted. The length
varies markedly in animals with equally developed hectocotylus, which
suggests that growth continues also after maturity and that the lost hecto-
cotylus can be regenerated once or repeatedly.

763

789 CONCLUSION

1. REVIEW OF THE EVOLUTION OF THE SUBCLASS
INCLUDING PALEONTOLOGICAL DATA-

(SYSTEMATIC-PHYLOGENETIC SUMMARY)

The origin of the Dibranchiata is obscure. It is certain on
systematic -morphological grounds that they developed from the Tetra-
branchiata, but paleontology gives no certain information on the group of
Tetrabranchiata from which they developed. It is remarkable that the
phragmocones of the oldest forms resemble distinctly those of moderately
slender (5 — 10°) Orthoceratidae of the genus A t r a c t i t e s, which have a
strongly developed periostracum and a slender rostrum, like the Belemni-
tidae. Atractites occurs already in the Permian and can be followed
through the whole Triassic and deep into the Jurassic (Foss. T. , p. 273).

These very slender, long animals (some of them several meters long)
probably resembled the Belemnitidae also in other aspects. They were
thus already true Decapoda, like the entire family Aulacoceratidae (p. Ill),
and not Protodibranchiata, as defined on p. 91. The differentiation of the
Protodibranchiata into Octopoda and Decapoda must therefore have taken
place not later than the Upper Paleozoic (Carboniferous), although we have
no information on the possible predecessors of the Octopoda, and the only
form known from the Mesozoic is Palaeoctopus newboldi (cf. p. 671).
The early reduction of the shell of Octopoda (p. 656) explains this sufficiently.

It is not certain that A t r a c t i t e s-like forms are the ancestors of all
Decapoda. Already in the Triassic a very aberrant type occurs in the
genus Phragmoteuthis. This belongs to the Decapoda and can be
reconstructed after the ideal form described on p. 110.-"!' However, a
correction was necessary. Since we have proved (Foss. T. , pp. 166, 122, 183;
cf. above, pp. 147, 158) that neither the older Teuthoidea nor the Sepiidae had
790 hooks, there is no resemblance between the Teuthoidea and the Belemnoidea
(p. 127). However, the recent suborders show not only a similar differen-
tiation of the stalked tentacles, but also a far-reaching resemblance in the
whole structure and ontogenesis (Loligo-Sepia). These factors must
be considered and attributed to their common ancestral form. On the other
hand, the presence of hooks (which are very characteristic) in the belemnoid
Decapoda is not connected with the development of special tentacles. On the
contrary, it has been established with certainty that the known types

* Consult on this my monograph: "Fossile Tintenfische," Jena, 1922, quoted in the following as "Foss. T."
** This reconstruction was made in Foss. T., figure on p.l86c. However, I did not extend the peculiar

proostracum to the anterior margin of the mantle sac, but I think now that this (normal) condition is not
impossible if a marked elasticity is assumed.

764

(Belemnoteuthis, Acanthoteuthis, Phragmoteuthis) had 10 uni-
form arms with hooks, which is characteristic for this suborder. Neither
can we assume a marked differentiation of the arms in the "praebelemnoid"
ancestral foms, i. e. (forms without hooks) of the Decapoda (see p. 791 ).

Figure 37 on p. 110 is already corrected in this sense (Foss. T., p. 167d):
the arms are of about equal length and form, and all arms are assumed to
be rather extensible, as is characteristic for the young Octopoda (p. 678).

The common ancestral form of the recent groups of Decapoda (Xeuthoidea
and Sepioidea) is shown in Figure 37 on p. 110, with the special characters
defined on p. 117, i. e. this type had 8 arms without stalks and 2 tentacles
with stalks, with 4 and 8 rows of suckers respectively, at least in the
secondary (postembryonic) condition. We cannot decide whether the quadri-
serial arrangement on the arms preceded the differentiation of the tentacles
and has to be assumed also for the Belemnoidea because distinct fossil
remains of suckers have not been preserved, also not in the Teuthoidea
in which suckers were certainly present. Nor do we know whether the rows
of hooks of the Belemnoidea were accompanied by rows of suckers, as in
Gonatus (p. 248), although we assume that this was the case, because of its
general usefulness (p. 128). The description of the formation of hooks
(pp. 127—133) probably also applies to the undifferentiated arms of the
Belemnoidea and also to the Oegopsida with hooks (pp. 236, 793). However, a
special common ancestral form of the Belemnoidea and Teuthoidea cannot
be assumed, in contrast to the statement on p. 127.

This explanation is based on detailed paleontological studies in recent
years. One of its systematic weaknesses concerns the constitution of the
Belemnoidea. They are apparently Decapoda because they have hooks (p.66l).
However, we do not know whether hooks were already present in the Aula-
coceratidae, but it may be assumed with certainty that the hypothetical
Protodecapoda had only suckers, which are the ontogenetic preceding stages
of hooks. If the Belemnoidea are considered as the ancestral (central)
group of all Decapoda, the diagnosis on p. 12 7 must be abbreviated and
restricted to the shell, as follows: The Belemnoidea are Decapoda with a
straight, well developed phragmocone which, except for the rostrum, occupies
the end of the mantle sac, and the muscular mantle was inserted on the free
margin of the shell. As the predecessors of Sepioidea are identical with
those of Teuthoidea and are therefore not unchanged Belemnoidea (see above)
791 and such shells have to be assumed for the predecessors of Sepioidea, the
special diagnosis of the Belemnoidea becomes invalid, unless we charac-
terize the "Presepioidea" by the differentiation of the tentacles and define
their predecessors by the absence of such arms, which is necessary if the
typical relationships are to be expressed in a natural classification. This
would have the disadvantage that a direct establishment of the position of
the Belemnoidea (sensu lato) would be impossible and that their position
will have to be determined on the basis of special relationships with well
known species.

The classification of the Decapoda would be as follows according to the
above:

A. Decapoda with uniform arms and without transformation of suckers
into hooks. Suborder I: Praebelemnoidea. The Aulacoceratidae perhaps
belong here.

765

B. Decapoda with uniform arms, on which at least part of the suckers
are transformed into hooks during postembryonic development. Suborder 2.
Eubelemnoidea. Belemnitidae, Belemnoteuthidae, Phragmoteuthidae.

C. Decapoda with differentiated tentacles, with or without hooks.

Cl. Shell as in the Belemnoidea, i. e. with well developed, straight phrag-
mocone which occupies the end of the mantle sac. Suborder 3. Praese-
pioidea. The Diploconidae perhaps belong here.

C2. Shell as in the Teuthoidea, i. e. phragmocone completely reduced,
occupying the end of the mantle sac. Suborder 4: Teuthoidea.

C3. Shell as in the Sepioidea, i. e. phragmocone well developed but
strongly curved ventrally in the posterior part and the anterior part inserted
deep into the mantle sac or further modified from such a condition, or
reduced. Suborder 5. Sepioidea.

Suborders 1 and 3 are only of theoretical interest at this stage. In
practice, it would be advisable to broaden the diagnosis of the Belemnoidea
and to add (p. 790): "A large part of the suckers are transformed into
hooks and the LV arms are not distinctly modified into tentacles.''

The Teuthoidea must have become separated from the Presepioidea in
the Lower Lias e. The Teuthoidea show a typical character and a large variety
of forms with a completely reduced phragmocone already in Lias e
(Figure 53 on p. 136). On the other hand, the phragmocone has been pre-
served to the present in the Sepioidea.

The oldest types of Teuthoidea are broad, plump, apparently like
Sepia in habitus, with a large, posteriorly displaced ink sac (p. 144).
Already in the Lias but especially in the Malm appear also more
slender forms, which resemble the littoral Oegopsida (Figure 60)
on p. 143), some of them with shells which markedly resemble Loligo
(p. 146). We do not know Belemnoidea (Presepioidea) from which the
Teuthoidea can be derived; at any rate, not the Aulacoceratidae, and Calli -
792 conites dienerP' cannot be considered as a transitional form with a
reduced phragmocone, as could be assumed from the incomplete and in-
accurate quotations (p. 135; cf. p. 8OI).

It would be more likely that Phragmoteuthis was a predecessor of
the Teuthoidea because of its compact form, the relatively smaller phrag-
mocone and the blunt angle of its opening. The peculiar projection of the
lateral plates, which gives the impression of a tripartite proostracum
(Foss. T., pp. 172 and 186), also has its analogue in the oldest Teuthoidea, i. e.
in the Belopeltidae of Lias e (loc. cit,,p. 129). However, a closer relation-
ship can be excluded also here because Phragmoteuthis belongs to the
Belemnoidea with uniform arms and hooks, while the Belopeltidae apparently
resemble Geoteuthis simplex (loc. cit., p. 123 ) which has the same
geological age and has more normal lateral plates.'"'*

* Critical consideration of the original figures and the description of the specimens lead to the following
conclusion. Two widely different characters have been confused in C.dieneri; a) rostra with very
slender alveoles which resemble Atractites but should be considered as a special generic type (Foss.,T.,
p. 273, 318); b) phragmocones of a very slender Orthoceras with a long living chamber, which should
not be expected in the Dibranchiata. The living chamber is preceded by numerous normal air chambers
which are very narrow (short), not wide as in the Aulacoceratidae.

Its ecological character also speaks against such a relationship. Hooks are present only in nektonic forms
of the open sea, while the Prototeuthoidea were apparently benthic animals, in contrast to their descendants.

766

There is also no established relationship between the fossil and recent
Teuthoidea. The fossil forms do not belong to recent families and there
are no transitional series in the strict sense. Only the general derivation
of the type of Palaeololigo (Malm) from the type of the Beloteuthis
(Lias) is more or less confirmed by paleontological data (p. 146). However,
the differentiation of the numerous and varied recent families probably
began already in the Upper Cretaceous.

The typical character of all Decapoda permits a complete reconstruction
aiso of the fossil forms. This reconstruction was later confirmed by a
number of well preserved fossils. This applies also to the Sepioidea.

There is often a close resemblance in form and adaptations between fossil
and recent types, and a similar variety of adaptation at different degrees of
typical relationship. For example, the true Teuthoidea of the Maim include
a markedly Sep i a -like form (T rachyteuthis, Foss. T. , p. 139) which
shows, among other characters, that the lengthening of the fins anteriorly
and the displacement of the articulation of the fins to the muscular mantle
(p. 163) was prepared already in the Mesoteuthoidea, while the Prototeuthoi-
dea resembled in this respect the early young stages of the recent Teuthoi-
dea; they have transversely produced, very short fins attached to the cone
flag and restricted to the posterior end (pp. 143, 307; Foss. T,, figures on
pp. 114, 120).

All fossil types examined differ from the recent Metateuthoidea in the
insertion of the muscular mantle on the lateral plates of the gladius. The
793 mantle sac of the recent Metateuthoidea attempts to envelop the elastic

shell from the outside, so that it becomes a passive organ of movement which
facilitates the rhythmic dilatation. The fossil Teuthoidea show the opposite
(this applies at least to the Trachyteuthidae, Geoteuthidae, Plesioteuthidae
and Belopeltidae): the insertion of the muscular mantle is displaced to the
inner (ventral) side of the much tougher and less elastic (i, e. calcified)
shell and thus achieves at least its own, moderately restricted motility.

This takes place in Trachyteuthis also in the region of the cone flag,
which closely resembles the wings of the shell of Sepia (p. 530).

' The geological age of all recent types of families cannot be determined.
The exact time of the separation into Oegopsida and Myopsida (cf. p. 151)
is also not known. On the other hand, an explosive radiation of the numerous
recent forms is also improbable. We know natural groups in the Oegopsida
(p.236), the group Cranchiidae-Chiroteuthidae—Joubiniteuthidae—Brachio-
teuthidae, or the group Gonatidae—Enoploteuthidae— Onychoteuthidae-Octo-
podoteuthidae. The presence of hooks makes the second group particularly
interesting. It apparently represents the basis or nucleus (p. 236) also of
the first group (cf. Galiteuthis, p. 398 ), which shows a number of special
resemblances to the Onychoteuthidae in particular (p. 241 ), The modifi-
cation of the suckers thus appears as a typical character of at least a great
part of the Oegopsida, and it could be considered as a primary character of
the whole group (but cf. p. 157). It should be considered, however, that such
a modification does not occur in the group Psychroteuthidae— Ommato-
strephidae— Thysanoteuthidae nor in the group Architeuthidae— Histioteu-
thidae—Bathyteuthidae (Neoteuthidae? ). It would be more correct to con-
sider the formation of hooks in the Oegopsida as a specific character of the
above groups and to unite these groups under the name "Onychii," as

767

Steenstrup (1861 ) and Jatta (1896) did in a wider sense. The establishment
of definite groups is, however, not advisable because of the fluctuating
boundaries (e.g. Histioteuthidae— Enoploteuthidae). It has been stated
repeatedly that the transitions within the Onychii are especially distinct and
include a number of forms which lost the hooks secondarily (Ly c o t e u t h i s,
p. 262; Cy c loteuthi s, p. 312). This induced me to assume a multiple
inhibition of the formation of hooks for all Teuthoidea as long as it was
erroneously assumed that also the fossil forms had hooks (pp.47, 127, 147,
157). It later became clear that hooks appeared independently twice in
the development of the order, and that it was prepared by the clawlike
character of the suckers of Decapoda (Plate XIl). It cannot be disproved
that hooks appeared "polyphyletically" even in the Oegopsida because of a
similar general disposition. However, in a scientific interpretation of data
one has to choose the simplest explanation which agrees with the real
relationships. In this case, it is certain that the number of suckers modified
into hooks varies within the species, which proves a gradation of the process
794 but not its isolated occurrence in the single species. (Galiteuthis

(pp. 241, 400) has been shown to be closely related to the Onychoteuthidae
by special characters.)

The Sepioidea appear very late, in the Lower Cenozoic (Paleocene), at
first as forms which closely resemble the Belemnoidea, in which they were
placed in the past. Closer examination, however, proves their separate
character. They are certainly not transitional types; such types could be
sought in the Upper Mesozoic, for example among forms with still belemnoid
shells, such as Diploconus (Foss. T., p. 172 ). We do not know when the
change of the correlation of the shell typical for this suborder (p. 485) took
place. It probably occurred only at the end of the Cretaceous and would
explain the following differentiation in the Paleocene and Eocene. This
differentiation must have been very rapid. Belopterina and Belo-
sepia have been found in the Paleocene and their predecessors (Belem-
nosella, Spirulirostra, Spirulirostrina) were probably also
present, although they are known with certainty only from later strata
(p.497). The Spirulirostrinidae, Spirulidae and true Sepia apparently
appear only in the Miocene.'-' The biological age of the other recent families
(Sepiolidae and Idiosepiidae), is not known, because these forms could not
leave fossil remains (Miocene, Pliocene?).

The same applies to the Octopoda, with the restriction made on p. 789.

The development of the Octopoda will remain obscure forever. Isolated
fossils like Paleoctopus can be expected only in especially fine slates
(p. 671). Only optimists can hope to find paleontological data for the
development of the shell of Argonaut a. If our hypothesis (pp. 778—782)
is correct, secondarily patched-up shells of Ammonites could be found.
However, the likelihood of finding such fossils is small, because of the
scarcity of Argonauta and the delicate structure of the shells. We must
admit that paleontology cannot explain the derivation of the recent Octopoda
and that we have to use the data of comparative morphology and general
assumptions.

True Sepiae are absent in the Paleocene and Eocene (Foss.T., p.92). Numerous erroneous data have been
published about older species (loc.cit., p.93). Especially misleading is the Paleocene "Sepia" vera
Desh., which closely resembles Belosepia.but it is so distinct that 1 propose the new genus Pseudo-
sepia for this species.

768

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769

We have tried to summarize the knowledge on the systematic -mor-
phological relationships and the paleontological data in a "genealogical tree"
as a symbol of a hypothetical history. The purpose of this scheme is to
show the development of the typical relationships during the geological
periods, not to express opinions on their special descendance.

2. GENERAL CONCLUSIONS
a. PRINCIPLES

We undertook this large work with the object of deciphering a part of
phytogeny. This appeared to us the suitable point of view for a detailed
and comprehensive study of a natural group of animals (p. l). We intended
to formulate natural laws and to achieve progress in the understanding of
biology on the basis of history.

We must admit in retrospect that even the first objective was not
achieved as we had hoped, although we did overcome difficulties which we did
not anticipate at first. We could not give a detailed, comprehensive report
on the evolution of the Cephalopoda, but on the other hand, positive results
were achieved. The reader should not put down this book with the im-
pression of having learned numerous isolated facts instead of having
witnessed the building of a whole in which each detail has its natural
connection. It was our purpose to present a whole, not a multitude. To
achieve this, we based our work on a connecting idea, a synthetic fundamental
concept. This was discussed in detail in the introduction and will not be
repeated. Much appears clearer in retrospect, also in this work. A few
further remarks are therefore necessary.

At the beginning of the study we realized that the "naive phylogenetics"
have no basis, i. e. the theory which considers descendance as the clue to a
full understanding of the variety of organic life, confuses resemblance with
true consanguinity and attempts to build the overestimated "genealogical
trees" on the basis of existing or constructed classifications or even on the
basis of mere resemblances. It was not our purpose to determine whether
the Octopoda developed from the Decapoda, or the Myopsida from the Oegop-
sida, or the Sepiolidae from the Sepiidae, or the Cephalopoda from Gastro-
poda or Chiton, and we made not only reconstructions of ancestral forms
on the basis of a more fundamental interpretation of the process of descent,
but we formulated our questions in a progressively more general form. In
the final analysis, we consider the "natural order of the variety of organic
forms," unrestricted by secondary interests or hypothetical fantasies, as a
problem of systematic morphology. Also the historical trend of systematic
morphology, which is determined by the sequence of generations and the
metamorphosis of the animal world throughout geological history, cannot be
satisfied with the mere composition of genealogical trees, but it has to follow
the transformation of the structural forms (types) as such during the
geological periods (Foss. T., p. 296).

Views on special consanguinity are not sufficient as connecting principles
of a synthetic concept and description. It is necessary to find an inherent,
natural connnection between functional formative principles as it manifests

5110/2

770

itself. The fantastic pseudogenealogy must be replaced with critical
morphology. To recognize the inherent meaning and ideal relationship
between living forms gives deep satisfaction and promises more true
knowledge than the discussion of parentage, which cannot be proven, even
though its existence is considered as self-evident.

797 Our interest is thus mainly directed to the natural forms themselves,
their live content and the biological necessity of the norms which determine
the multiple replication of individual organisms. How far the actual meta-
morphosis agrees with the genealogical relationships is not our concern at
this stage. At any rate, this is a different problem and of minor importance
to us.

This type of consideration is the more justified because the principles
which govern the order of natural forms are derived from their actual pro-
perties and combine them into a logical system which includes the variety
of organic forms, as we discussed in general in the introduction and have
shown in detail in the special part. This can be done entirely within the
framework of idealistic morphology, without an uncritical realistic phytogeny.

The natural system is placed in geological time by paleontological data,
and is thus historically orientated. The theory of descent is based on
stages and the resemblance between systematic and ontogenetic stages
(Naef, 1917), and is connected with geographical distribution. However, the
reverse does not apply: the theory of descent cannot form the methodical
basis of morphology, as phylogeny has attempted to do during various
stages (also by the author in some early works).

These considerations prove the validity of the general theory of descent,
because it answers the following questions. 1. How can the graded variety
of forms (as in the natural system) be explained? 2. How was it possible
for the faunae and florae to reach by stages their recent condition? 3. How
did the resemblance develop between the ontogenetic and systematic stages
of the same animal or organ? What is the cause of the resemblance
between geographical and systematic groups? All these questions, and
therewith the theory of descent, only followed from the fact of a natural
system (in relation to others), and this system must therefore be based
logically on reality. It had to be shown that this can take place and we hope
to obtain further insight from the development of this knowledge. If the
preliminary draft of a natural system in 1859 created the theory of descent,
a more fundamental understanding of the relationships will make this theory
more perfect. This expectation has stimulated, often unconsciously, the
systematic -morphological research in post -Darwinian time, although the
biased and uncritical genealogical formulation of the research made success
rather unlikely. Instead of a thorough understanding followed extensive
accumulation of facts, a deterioration of the basic principles and a crumbling
of the old traditions of classical morphology. A multitude of data was
accumulated because of the view that it concerned the discovery of real

798 facts and relationships, like the descent of man, "progress," the (mechanical)
creation and variation of life, etc. This increased the knowledge of facts but
only hindered fundamental research.

We intend to achieve a new and deeper concept of nature and the
liberation of biology from the sterile thoughts of orthodox mechanism. We
consider nature as a real entity in the frame of space and time, but we keep

771

conciously in us the key to its conceptual understanding. We use in
systematic morphology the idea of the type for this purpose, i. e. the norm
which can be determined for every natural group of living forms. The norm
has the same importance as the law in nature, it creates order and connects
the components into a whole. Whether an objective (natural) reality corres-
ponds to the norm will remain an open question. On Dollo's law see p. 639.

At any rate, the type is not a consciously or unconsciously veiled
substitute of the "ancestral from," although we consider them as more or
less similar. The "ideal form" (the Platonic idea of a natural being) is
rather a symbol of the "hereditary mass" which persists throughout the
general line if we restrict ourselves to the natural circle of the narrowest
reproductive community. The hereditary mass can also be considered only
hypothetically as "genotype" and it can be shown only symbolically to be the
the cause of certain effects (p. 28). The hereditary mass, which consists of
agents that determine a measure and purpose, controls together with
external and internal factors the way in which the branches separate from
the common stem, from which they can depart as particular, not general,
circumstances permit. The type and its evolution are placed in parallel
with the basic concepts of modern genetics, which do not understand the
realities as directly as is generally believed. Systematic morphology is
thus recognized an active branch of modern biological science instead of
the late result of a dogma of the past alien to nature (cf. also p. 639 and
the following volumes and theoretical works).

b. THE ADAPTIVE CHARACTER OF THE
SYSTEMATIC STAGES

Systematic ecology will be treated in a special volume in an attempt to
establish its methodical basis, but it is necessary to stress that it forms a
complete parallel and supplement to morphology.

Wherever we succeed in clarifying the typical relationsips between
organisms (in the sense of Figure 2 on p. 12) and in determining the systema-
tic preceding stages of certain species, we realize that each stage represents
a definite "step" of an "adaptation" in a certain direction and to certain
conditions. In this sense, the adapted stage has gained an uncontestable
799 advantage over its predecessor in its technical equipment. We will discuss
this in greater detail for our group. In general, the following question
arises: How is it possible that the formative norms change in stages and at
the same time undergo a progressive course of adaptation and change?

This question will be based, with reason, on the theory of descent, although
it could also be given a more general formulation within the framework of
idealistic morphology because the above arguments against a complete
conversion of systematics into phylogeny do not apply. This problem is also
more urgent and of more practical and real interest, because it is still un-
solved. It is the central controversy between neo-Darwinism and neo-
Lamarckism.

All these relationships will be discussed in detail in Part III of this work,
but we will show here on a few examples of systematic -morphological im-
portance how we consider them. Adaptation in stages can be proved also in
all other parts and can be expressed in simple, very instructive formulae.

772

1. METAMORPHOSIS IN STAGES IN THE FORMATION
AND POSITION OF THE SHELL OF DIBRANCHIATA

I. The ancestral form of the shell of molluscs is a habitation (Figure 5
on p. 51 and Figure 6 on p. 53), i. e. a hollow, cuticular exoskeleton wich
accommodates and protects the greater part of the soft body. Its primary
stratification can be easily demonstrated in the Tetrabranchiata, althoug the
hypostracum shows already (or particularly) a special differentiation into
air chambers, septa, siphonal necks and substance of the annulus, which
together form the phragmocone (Figure 19 on p. 79; cf. the detailed
description in Foss. T., p. 14).

II. The basic form of the external shell of Cephalopoda is as in Ortho -
ceras. The basic form of the internal shell of Dibranchiata is best known
in the Belemnitidae, which have developed at least a stage further. Typical
variants of the well developed internal shell are known only among the
Decapoda, the ancestral form of which is thus also valid for all Dibranchiata
(Figure 23 on p. 91 and Figure 37 on p. IIO). (On the significance of the
reduction of the shell of Octopoda see p. 657.)

III. The shells of the Belemnoidea and Orthoceratidae do not differ
markedly in the phragmocone, as the numerous confusions in the past have
shown. The main topographical difference is that it is covered by a shell
fold so that it is situated completely inside the shell sac. The shell is here
under the influence of the chromatophores, which partly compensate for the
reduced protection by an internal shell. The stratification is the same,

800 except for the periostracum, which is formed by the secondary shell

epithelium and covers the prismatic layer in all Dibranchiata. The adaptive
character of the periostracum is quite evident in internal shells: while the
Orthoceratidae later discarded the delicate initial parts, they must here be
protected and strengthened. This gives another advantage: the center of
buoyancy at the posterior end is displaced anteriorly (p. IIO) and thus
facilitates in a varying degree the maintenance of the horizontal equilibrium
of the longitudinal axis, until a completely passive condition is attained. As
a result, the animal can swim independently from the surface, which the
Orthoceratidae could probably not do, because the phragmocone does not
function like the swim bladder of fish, which can be adapted; its hydrostatic
function is rigid, like its form, and permits at most slow changes.

The transition from the shell of Orthoceratidae to that of the Belemnoidea
is also characterized by a reduction of the ventral wall of the living chamber
and its replacement by the muscular mantle (p. 94). This is evidently the
most important development, because it creates a new active organ of move-
ment (p. lOO) which is very effective. The muscular mantle also deter-
mines further changes because it develops further at the expense of the
passive, protective internal shell.

The surrounding of the shell and the development of the muscular mantle
convert the Tetrabranchiata into Dibranchiata. The two processes are
evidently not independent because only the development of the shell fold
prepares a solid insertion for the muscular mantle ( Figure 20 on p. 85).
Other specific characters of the organization of the Dibranchiata are also
influenced by this development. Examples are the displacement of the
abdominal complex (p. 84); the reduction of the dorsal pair of gills (p. 101 );

773

the perfection of the statocysts (Vol. Ill), of the camera eyes (p. 96) and of
the adhesive organs (p. 63); fusion of the funnel (p. lOO); development of the
fins; and the increase of the protective adaptation given by the chromato-
phores by the inc sac. All these properties already gave the oldest Di-
branchiata a radius of action which is far greater than that of all other
invertebrates and equipped them for a continued and open competition with
marine vertebrates since the remote geological past.

IV. The type of Teuthoidea can be understood ecologically as follows:

The Belemnoidea show a tendency to compensate the buoyancy of the phrag-
mocone. This is most marked in the clavate forms, and characterizes them
as free swimming animals of the open sea. A better solution, however, would
be to eliminate both the buoyancy and its compensation by a simultaneous
development of the muscular mantle. This is the most marked character of
the more recent suborder, together with which the older suborder persisted
still for a long time, similarly as the Belemnoidea existed together with the
Orthoceratidae. Strangely enough, we do not know any transitional forms
also here, although they would have to be expected in the Upper Triassic,

801 Constructions like Conoteuthis dupini d'Orb. (Foss. T., pp. 171, 278),
which were considered as a connection between the Ommatostrephidae and
the Belemnitidae, can no longer be accepted. There is thus no paleontolo-
gical evidence for a gradual reduction of the phragmocone. On the other
hand, the "gladii" of the early Teuthoidea (Prototeuthoidea) are still well
developed proostraca which resemble those of the Belemnitidae, and the
young forms of the recent Teuthoidea still show large remnants of the
previous cone in the form of scoop-shaped "anlagen" of an initial chamber,
although this never attains typical development (Figure 67 on p. 161; cf. also
p. 791).

The derivation of the type of the Teuthoidea from that of Belemnoidea is
quite clear, despite the lack of intermediate forms. The stage between them
is characterized by the liberation from the passive and protective apparatus
of the chambered shell, which is replaced by the developing muscular mantle.
The gladius serves only as backbone of the mantle sac, and gives the
necessary support to its movements. In the Metateuthoidea, these develop-
ments are accompanied by the narrowing of the median plate into a "rhachis,”
which further strengthens the active parts, while the passive proostracum
facilitates this development by its decreased calcification (p. 149). Its
resulting elasticity contributes to the rhythmic opening of the mantle sac
during respiratory and swimming movements.

Further stages of reduction follow. Because of its elasticity, the flag
can be enveloped by the muscular mantle so that it becomes situated inside
it. When the muscular mantle becomes stronger (p. 356), the flag can dis-
appear completely, while the rhachis becomes an unimportant, pliable
supporting element. This takes place in particularly strong swimmers; in
other, more floating or even planktonic types, the process is apparently
inhibited at various stages (p.237).

V. The type of the Sepioidea is another hydrostratic experiment. Instead
of eliminating the buoyancy of the phragmocone, it is here displaced upward
and anteriorly until a passive equilibrium of the swimming position is
obtained. This makes the rostrum more or less unnecessary and it is more
or less reduced or lost in the recent forms (Spirula, Sepia), also if the

774

rest of the calcified shell is preserved (pp. 501 and 521). In the older types,
the rostrum develops to a large size (Figures 494 and 500 ) as a new adap-
tation caused by the need for a heavy posterior end in connection with a
certain mode of life (p. 480), or partly because of the need for a regulation
of the longitudinal axis (p.484) the maintenance of which is made more
difficult by the ventral curvature of the apex of the phragmocone. The
envelopment and reduction of the shell by the muscular mantle results in
some cases in its complete disappearance in the Sepiolidae and Idiosepiidae
(pp. 500 and 503).

The Sepiidae perfected the possibilities which the organization of the
Sepioidea provided. Instead of reduction, the chambered shell took on a
new correlation so that it became almost unrecognizable. Through the
802 stages of Spirulirostrina and Belosepia, however, the shell of the
Sepiidae appears as a variant of the general character of the Sepioidea, so
that a separation of the family from the other members of the group is not
justified. The special form of the shell of Sepia is due to the oblique
position of the septa and the connected flattening of the phragmocone.
Ecologically, this is a preservation of the improved hydrostatics of the
Sepioidea, but without the disadvantages which result from the displacement
of the shell opening into the soft body (p. 476) and the restriction of growth
caused by the early determination of the form of the posterior end (Spiru-
1 i r o s t r a, p. 500). The open curvature of the shell of Sepia permits an
unlimited enlargement at the same proportions, which is actually the case
in different species. The reduction of the shell in the Sepiolidae (Rossia)
permits a similar development so that growth is possible beyond the usual
size of the forms of this suborder (p. 479).

Comparison of these structural principles (the variation of which is
evidently markedly inhibited) clearly shows their character as stages.

This is not always the case with other structures of less fundamental form
and function. Such secondary structures often occur sporadically; their
scientific consideration is more ecological. This will become increasingly
evident in the two following sections, which deal with a number of similar
but not morphologically related phenomena.

2. WATER PORES AND WATER SACS

The Dibranchiata, especially the nektonic forms, have the peculiar tendency
to develop deep crevices and cavities which communicate with the environment.
These formations or their outer openings have been often named "water
pores" without regard to their homologies. I will try to avoid this. In my
terminology, water pores (ventral and dorsal) are present only in the Argo-
nautidae, also the "cavities of the head" into which they lead; "tentacle
pockets" are present in the Decapoda; "buccal pockets" particularly in the
Teuthoidea; "orbital cavities" in the Dibranchiata as a whole; "orbital
pores" in the Myopsida and Sepioidea; "lateral pockets" and "crescent-
shaped pockets" in the funnel pit of the Ommatostrephidae and Thysano-
teuthidae. All these crevices and cavities appear relatively late, usually
during postembryonic development; only the orbital cavities are typical for
the Dibranchiata in general and often appear already during embryonic
development.

775

Except for the mantle cavity, which is an important formation of varying
significance and a primary element of the type of molluscs in general, the
other cavities have a double function: on the one hand, they reduce the
specific weight and create a more streamlined form of the body, on the other,
803 they make the individual muscular parts (e. g. cephalopodium) mechanically
independent like the coelomic cavities or blood sinuses inside the body. The
latter is undoubtedly true for the deep "buccal pockets" which assure free
movement of the buccal organs, extension of the pharynx, etc., and thus avoid
strains and other inconveniences. This is probably also the function of the
pockets of the tentacles which expose the basis of the tentacle stalks situated
inside the tissues of the head (pp. 183, 566). This is also the case with the
orbital cavities: they maintain the mobility which the eye primarily has
(as stalked eye in Nautilus). The eye is thus protected inside the mus-
cular mass of the cephalopodium but it is not affected by any mechanical
changes in the cephalopodium.

The secondary development of these structures can be followed in stages,
at least in some groups (compare, for example, the tentacle pockets in the
Loliginidae, p. 178, and in the Sepiidae, Figure 289 on p. 526 and Figure 297
on p. 537). On the other hand, the dorsal and ventral water pores of the
Argonautidae (pp. 740 and 75 1) or the cavities in the tentacle clubs of some
Loliginidae (p. 194) are quite different and are only functional analogues.

3. DEVELOPMENT OF THE SEXUAL RELATIONSHIPS

The continuity of the adaptation in stages which characterizes not only
many other parts but also characters of behavior connected with a certain
structure, is absent in the relationship of the sexes. This is the more
striking because of the morphological expression of this relationship and
has claimed great interest since the systematic -morphological work of
Steenstrup (1856). The peculiar sexual dimorphism of many Dibranchiata
attained almost fantastic manifestations (Argonautidae), although it always
develops from typical juvenile conditions.

The proportions of the adult female change with maturity because of the
enormous development of the ovary. The posterior part of the body becomes
thicker, broader and sometimes also much longer (pp. 436, 499 and 551 ).

The posterior end is usually blunter and the whole body plumper than in a
male of the same size. Females are also always slightly larger than males,
although this can only be determined if a large number of mature individuals
are available, i.e. in Octopus, Eledone, Illex, Sepiola and Sepia,
Loligo vulgaris is perhaps an exception. I found mature and fertilized
females of a dorsal mantle length of only 13 cm, while the largest specimens
(mantle length over 50 cm) were always males. The broadening of the
posterior part of the body by the ovary affects also the shell. In specimens
of the same length, the females frequently have a broader shell. This is
especially striking in Sepia officinalis (p. 552) and Loligo vul-
garis (p. 200).

The sexual dimorphism of the body is associated with certain differences
in the mode of life which is certain in many forms: In the Oegopsida

776

804 (Abraliopsis scintillans), only the fertilized females come in large
numbers to the shore to deposit eggs. I have also often found large
numbers of mature female Sepiolidae without or with only a few males.

In other species (Loligo vulgaris), the males are not only present at
oviposition but show a marked interest and accompany the female in a state
of high excitation. Males often predominate in catches and many false
conclusions on the numerical proportion of the sexes have been drawn from
this.

Some females of Decapoda (pp. 434 and 586) have more strongly developed
tentacles which apparently have to provide food for the production of the
large egg masses. Other special characters assure the fertilization of the
eggs: The adult male shows changes mainly of the arms, which often become
very strong, and some of the suckers become enlarged. These changes are
clearly connected with copulation (overpowering of the female and transfer of
spermatophores). I call this phenomenon "hectocotylization, " in a wider
sense than used by Steenstrup (185 6). Usually one arm (rarely both)
becomes transformed for the transfer of the spermatophores. This arm is
known as "hectocotylus" although the hectocotyli of the different species are
not necessarily homologous throughout the subclass. Any arm can become
specialized into a hectocotylus, and there is no general type. Hectocotyli-
zation has never been observed in some families; it almost certainly does
not occur in such well known genera as Onychoteuthis, Ancistro-
teuthis and Gonatus. Formation of a special hectocotylus is therefore
not to be assumed for the ancestral form, but only hectocotylization in a
general sense, and the inherited normal form of copulation, i. e. an "embrace"
mouth-to -mouth and transfer of the spermatophores to the buccal funnel, or
more exactly, to its median, ventral and inner parts. A similar behavior
was established for Nautilus (p. 63).

There are numerous other forms of copulation: In the Sepiolidae, the
female is thrown on her back or lies supine on the male and is fertilized
from below; both positions are the same as regards the relationship of the
hectocotylus to the bursa copulatrix. This explains the displacement of the
hectocotylus (sensu stricto) to the dorsal side. This is probably the case
in many Teuthoidea (p. 354) in which the dorsal arms are hectocotylized.
Direct observations have not been made, but this is borne out by the behavior
of Loligo vulgaris, in which the sexes copulate not only in the typical
position (mouth against mouth) but also from above and from below by
animals swimming above each other. Drew (l91l) observed similar
behavior in Loligo pealei. The attachment of the spermatophores in
the female varies accordingly. The spermatophores can be attached in the
mantle cavity instead of in their typical position on the buccal membrane,
also in the above species. This assures an early and certain fertilization,
especially if the spermatophore is attached near the female genital opening

805 and if the area of attachment is specially adapted for the reception and
preservation of the spermatophore, as in the Sepiolidae (Figure 3 62 on
p. 617 and Figure 374 on p. 636).

In the Octopoda, the spermatophores are introduced directly into the
genital opening of the female. This probably stimulated further differen-
tiation of the hectocotylus and opened the way for transition to vivipary
by internal fertilization (Ocythoe, p. 750). The relationships here become

777

so complicated that only a development in stages can lead to the final
structure and its derivation (evolution) can be recognized systematic -
morphologically (Polypodoidea, p. 677; Octopodidae, p. 684; Argonautidae,
p. 728; Argonauta, p. 784).

In general, the tendency toward preservation of the species manifests
itself more distinctly in labile instincts than in rigidly defined types of
organization and is therefore only accessible to a restricted extent to mor-
phological methods.

3. TECHNIQUE

We succeeded in examining various structures which earlier authors had
overlooked, especially in small, young stages and embryos. This was due
in part to the availability of substantial fresh material. Knowledge of the
live animal and of its changes during death and preservation would have
prevented many observers from describing and illustrating various monsters
and from describing common characters as specific. Knowledge of the live
animal is the alpha and omega of morphological research — everything else
is secondary.*

At any rate, preparation is important also in studies which are consciously
and distinctly superficial and deal only with the external structure.

The fresh animal does not show the structure of the surface distinctly
because coloration and texture are prominent. To examine the finer
structure, the surface has to be made opaque and the coloration has to be
destroyed or weakened. This is obtained with precipitating reagents which
give a uniform, mat stain. Mixtures of chromic acid (e.g. chromic acetate
solutions are especially suitable for large preparations, of which a histolo-
gical study is not intended. However, they destroy the keratochitin of the
horny rings of the suckers of Decapoda so that they should not be used for
rare specimens, for which only strong formol solutions are suitable. If they
are added slowly during a longer period, they stain the keratochitin of the
horny rings and other cuticular structures of the suckers dark reddish brown,
806 which facilitates examination (see Plate XII, Figures 2, 5, 6).

The following procedure gives good results for small objects. After
any type of preservation, the specimen is placed for 24 hours or longer in
absolute alcohol and then in xylol for a similar time, where it becomes very
hard. The material is then dried. If the hardening was not excessive, the
specimen becomes opaque, chalk-white and only slightly smaller but it
retains its form and surface. Such preparations appear under the magni-
fying lens like gypsum models; they appear very plastic and show the finer
details of the surface. Chromic acid preparations should be used as controls
(because of shrinkage).

Such preparations should be stained previously (with hemalum or carmine);
sections can be made afterwards for topographical examination;** the
material is histologically damaged, but less than may be assumed. The

* About keeping animals in captivity see Vol.III.

** To dissolve air in cavities, the specimen is placed in alcohol through xylol.

778

preparations can also be embedded in paraffin and cut to show the body
cavities; after the paraffin has been dissolved in xylol, the interior can be
examined with the magnifying lens. This is much easier than preparing
models, and often more useful (Vols. II and III). I mount these preparations
on entomological pins. The specimen can then be examined under the
binocular microscope. The specimens are stored in entomological boxes
or on a cork in a tube.

Careful cocainization is an important preparatory step for the preser-
vation of small, delicate animals (like larvae of Oegopsida). Cocaine must
be used in a large dose to have a rapid effect; the slightly intoxicated animal
is then preserved in Zenker's solution. I place seawater in a glass dish of
6 cm^, then the larva and then 3 drops of a 5% solution of cocaine. When
respiration ceases, I transfer the larva with a wide pipette into the preser-
vation fluid. This usually gives excellent preparations. I sometimes made
drawings of deformed specimens, partly because fresh material was not
available, partly because of failures, but I then also gave reconstructions
(pp. 275, 305, 354) based on drawings of live animals, so that the natural form
of the deformed preparation can be determined (Figures 208, 209 on p. 421).

Special experience is necessary for the preservation or interpretation of
badly preserved Sepiola and Octopoda, because their body without a
skeleton tends particularly to deformations which are then often reproduced.
The inexperienced should send his material to a specialist. I found it at
first very difficult to distinguish deformations from the natural form, i. e.
in Sepiola, of which I studied large numbers. The usual methods of fishing
807 often provide defective and deformed material; the arms are curved, and the
suckers are lost during preparation; the head is retracted completely into
the mantle, so that its form becomes unnatural. The mantle margin, which
is of so specific form, is curved, and the skin becomes so strongly contracted
that even the organs in the mantle cavity are deformed and cannot be
examined in their natural position.

Such deformation can be prevented as follows: The live animal is slowly
narcotized or becomes intoxicated by its own carbonic acid inside a small
glass vessel. When it is motionless, the animal is placed in a flat dish with
a 1—2% solution of acetic acid in fresh water. The specimen then becomes
slowly stiff. Developing or already present contractions can be corrected
mechanically by rearranging the parts, stretching of the tentacles, etc. The
fins are spread by pressing them with a soft, brush against the bottom of the
dish. To correct the folding over of the mantle margin, a few strokes with
the brush are made from behind, which extends the skin. To preserve the
space of the mantle cavity, it is filled with very dilute acetic acid. If the
anterior margin of the mantle is contracted, it is extended mechanically.

Such preparations (pp. 569, 635, etc.) preserve the proportions and specific
characters of the live animal. This fixation prevents the frequent loss of
the suckers in the Sepiolidae. When the animals have the desired form, they
are placed for 24 hours in strong (4 — 10%) formol, then rinsed and placed
in alcohol.

On the whole, the usual methods of technique and some practical sense
are the best assurance for good results.

779

4. SYSTEMATIC REVIEW AND KEYS FOR DETERMINATION

The book is so simply arranged that the specialist will not lose his
orientation, despite the numerous facts. However, some keys for the
determination of species are desirable. We return to the list on p.46 and
complete it on the basis of studies made during the last 7 years.

808 a. FAMILIES AND GENERA OF THE

CEPHALOPODA DIBRANCHIATA

Order I: Decapoda Leach, 1818

Suborder a) + Belemnoidea (Zitt., 1885) Naef, 1912.'!"

1. Aulacoceratidae (Bernard, 1895: Aulacoceras Hauer, 1860,

D i c ty o c o n i t e s Mojs., 1902, Calliconites Gemm., 1904,
Atractites Giimb . , 1861.

2. Xiphoteuthidae Naef, 1921 : Xiphoteuthis Huxley, 1864.

3. Phragmoteuthidae Naef, 1 921 : P h r a g m o t e u t h i s Mojs., 1882.

4. Belemnitidae d'Orb., 1845: Hastitinae Naef, 1921 : Hastites Mayer,
1883, Rhabdobelus Naef, 1922. Coeloteuthinae Naef, 1922 :
Coeloteuthis Liss.,1915. Passaloteuthinae Naef, 1922 : Nanno-
belus Pawlow, 1913, P a s s a 1 o t e u t h i s Liss.,1915, Pseudo-
hastites Naef, 1922, Brachybelus Naef, 1921, Homaloteu-
this Stoll ey, 1919, Megateuthis Bayle, 1878, Gastrobelus
Naef, 1922, Pleurobelus Naef, 1922, Odontobelus Naef, 1922,

S a 1 p i n g o t e u t h i s Liss.,1915, D a c t y 1 o t e u t h i s Bayle, 1878.
Cylindroteuthinae: Cylindroteuthis Bayle, 1878, Pachyteuthis
Bayle, 1878, Oxyteuthis Stolley, 1911, Aulacoteuthis Stolley,
1911, Raphibelus Naef, 1922. Belemnopsinae: Belemnopsis
Bayle, 1878, Hibolites Mayer, 1883, P a r a h i b o 1 i t e s Stolley,

1911, Mesohibolites Stolley, 1 91 1 , Neohibolites Stolley, 1911,
Belemnitella d'Orb,, 1845, Actinocamax Miller, 1823,

Belem no conus Naef, 1922, Dicoelites Bohm, 1906. Duvaliinae
(Pavlow): Duvalia Bayle, 1878, Pseudobelus Blainv., 1825,
Conobelus Stolley, 1919, Rh o p a 1 o t e u t h i s Liss.,1915. Bayano-
teuthinae: B a y a n o t e u t h i s Mun.-Ch., 1872, Sty r a c ot e ut h i s
Crick, 1905.

5. Belemnoteuthidae (Zitt., 1 885 ) Naef, 1 922 : B e 1 e m n o t e u t h i s Pearce,
1842. (A c a nt h ot eu t h i s Wagner, 1832? )

6. Diploconidae Naef, 1922: Diploconus Zitt., 1868, Conoteuthis
d'Orb., 1842. (?Amblybelus Naef, 1922.)

7. Vasseuriidae Naef, 192 1 : Vasseuria Mun.-Ch., 1880.

Suborder b) Teuthoidea Naef, 1916.

a. 1 Prototeuthoidea Naef, 1921 .

1. Plesioteuthidae Naef, 1 92 1 : Paraplesioteuthis Naef, 1921,

P 1 e s i o t e u t hi s Wagner, 1860, Styloteuthis Fritsch, 1910.

* On the fossil groups, see Naef, 1922, "Die Fossilen Tintenfische. "

780

2. Leptoteuthidae Naef, 192 1 : Leptoteuthis H. v. M.,1834.

3. Geoteuthidae Naef, 1921 : Geoteuthis Milnst., 1843.

4. Belopeltidae Naef, 1921 : Belopeltis Voltz, 1840, Parabelopel-
tis Naef, 1921, Loliginites (Quenst., 1849) Naef, 1921 .

5. Lioteuthidae Naef, 1922: Lioteuthis Naef, 1922.

809 b. t Me sot euthoidea Naef, 1921 .

1. Trachyteuthidae Naef, 1921 : T r a c h y t e u t h i s H. v. M., 1846,
Glyphit euthis Reuss, 1870.

2. Beloteuthidae Naef, 1921 : Bel ot euthis Munst., 1843 .

3. Palaeololiginidae Naef, 1921: Palaeololigo Naef, 1921 (=Teu-
thopsis Wagn., I860), Tusoteuthis Logan, 1898, Phyllo-
teuthis Meek and Hayden, 1860 (?), Ptiloteuthis Gabb., 1869(?).

4. Celaenidae Naef, 1921 : Celaeno Munst., 1892, C e 1 a e n o t e u t h i s
Naef, 1922.

c. Metateuthoidea myopsida (d'Orb., 1845) Naef, 1921.

1. Loliginidae Steenstr., 1861 (as "Loliginei"): Loligo Lam., 1798,
Sepioteuthis Blainv., 1924, Loliolus Steenstr., 1856, Dory-
teuthis Naef, 1912, Alloteuthis (Naef, MS) Willker, 1920.

2. Promachoteuthidae Naef, 1912: P r o m a c h ot euth i s Hoyle, 1885.

3. Lepidoteuthidae Naef, 1912: L e p i d o t e u t h i s Joubin, 1895.

d. Metateuthoidea oegopsida (d'Orb., 1845) Naef, 1921.

1. Bathyteuthidae Pfeffer, 1900: Bathyteuthis Hoyle, 1885, Ctenop-
t e r y X Appellof , 1889.

2. Gonatidae (Hoyle, 1886) Pfeffer, 1900: Berryteuthis Naef, 1921,
Gonatus Gray, 1849, Gonatopsis Sasaki, 1920.

3. Onychoteuthidae Gray, 1849: Onychoteuthis Lichtenstein, 1818,

A n c i s t r o t eu t h i s Gray, 1849, Teleoteuthis Verrill, 1885,
Onykia Lesueur, 1821, Chaunot euthi s Appellof, 1891, Te-

t r o ny c h ot eu t h i s Pfeffer, 1900, Moroteuthis (Dahl, MS)
Verrill, 1881, Cycloteuthis Joubin, 1919.

4. Neoteuthidae Naef, 1921 : Neoteuthis Naef, 1921.

5. Octopodoteuthidae Berry, 1912; O c t o p o d o t e u t h i s Ruppell, 1844,
Cucioteuthis Steenstrup, 1882.

6. Histioteuthidae Verrill, 1880: H i s t i o t e u t h i s d'Orbigny, 1839,
Calliteuthis Verrill, 1880 (St i g m a t o t e u t h i s Pfeffer, 1900),
Histiopsis Hoyle, 1885, M ele ag r oteuthi s Pfeffer, 1900.

7. Architeuthidae Pfeffer, 1900: Architeuthis (Steenstr., 1857)
Verrill, 1875.

8. Enoploteuthidae Chun, 1910: Pyroteuthinae Pfeffer, 1912: Pyro-
teuthis Hoyle, 1904, P t e r y g i o t eu t h i s Fischer, 1896. Lyco-
teuthinae Pfeffer, 1912: Lycoteuthis Pfeffer, 1900, Lampadio-
teuthis Berry, 1916, Nematolampas Berry, 1913. Enoplo-
teuthinae s. lat. : Enoploteuthis d'Orbigny, 1839, Abralia Gray,
1849, Abraliopsis Joubin, 1896, A n c i s t r o c h i r u s Gray, 1849,
Thelidiot euthis Pfeffer, 1900.

9. Psychroteuthidae Thiele, 1920: P s y c h r o t e u t hi s Thiele, 1920.

10. Ommatostrephidae Gill., 1871 : Illex Steenstrup, 1880, Todarop-

sis Girard, 1890, Nototodarus Pfeffer, 1912, Ommato-
strephes d'Orbigny, 1839, Dosidicus Steenstrup, 1857,

781

810

Hyaloteuthis Gray, 1849, St h e n o t e u t h i s Verrill, 1880,

Sy m p 1 e c t o t e u t h i s Pfeffer, 1900, E u c 1 e o t e u t h i s Berry, 1916.

11. Thysanoteuthidae Keferstein, 1866: T hy s a n o t eu t h i s Troschel,
1857.

12. Brachioteuthidae Pfeffer, 1908: Brachioteuthis Verrill, 1881 .

13. Chiroteuthidae Gray, 1849: Chiroteuthinae Chun, 1908: Chiro-
teuthis d ' Orb igny, 1839, Chirosoma Joubin, 1912.’!' Mastigo-
teuthinae Chun, 1908: M a s tig ot eu th i s Verrill, 1881 (Chiro-
teuthoides Berry, 1920, Idioteuthis Sasaki, 1916). Grimaldi-
teuthinae Chun, 1908: Grimalditeuthis Joubin, 1898, Enoptro-
teuthis Berry, 1920.

14. Cranchiidae Gray, 1849: Cranchiinae Pfeffer, 1912: Cranchia
Leach, 1817, Leachia Lesueur, 1821, Pyrgopsis Rochebrune,
1884, Liocranchia Pfeffer, 1884, Liguriella Issel,1908.
Taoniinae Pfeffer, 1912: Pha s m at op s i s Rochebrune, 1884,
Toxeuma Chun, 1906, Taonius Steenstrup, 1861, Megalo-
cranchia Pfeffer, 1884, Taonidium Pfeffer, 1900, Chrystallo
teuthis Chun, 1906, Pha sm at ot euthion Pfeffer, 1912, Gali-
teuthis Joubin, 1898, Corynomma Chun, 1906, Verrilli-
teuthis Berry,1916, Desmoteuthis Verrill, 1882, Leuco-
cranchia Joubin, 1912, H e n s e n i ot e u t h i s Pfeffer, 1900,
Sandalops Chun, 1 906 (H e 1 i o c r a n c h i a Massy, 1907), Teu-
thowenia Chun, 1910, Bathothauma Chun, 1906.

15. Joubiniteuthidae Naef, 1922: Joubiniteuthis Berry, 1920.’!'’!'

Suborder c) Sepioidea Naef, 1916

1. t Belemnosidae Naef, 1921 : Belemnosis Edwards, 1849,
Belemnosella Naef, 1922, Sp i r u 1 i r o s t r e 1 1 a Naef,1921.

2. t Belopteridae Naef, 1921 : Beloptera (Desh. ) Blainv., 1825,
Belopterella Naef, 1921, Belopterina Mun.-Ch., 1872,

B e 1 o p t e r i d i u m Naef, 1922.

3. t Belosepiellidae Naef, 1921: Belosepiella Alessandri, 1905.

4. + Spirulirostridae Naef, 1921: Spirulirostra d'Orbigny, 1841,

Sp i r u 1 i r o s t r i d iu m Naef, 1922.

5. t Spirulirostrinidae Naef, 1 92 1 : Spirulirostrina Canavari, 1892.

6. Spirulidae (d'Orb., 1826) Owen, 1836: Spirula Lam., 1801.

7. Sepiidae Keferstein, 1866; t Belosepiinae Naef, 1921: Belosepia
Voltz, 1830, Stenosepia Cossman, 1913, Pseudosepia n. gen.
Sepiinae Naef, 1921 : Sepia L.,1758, Sepiella Gray, 1849,
Hemisepius Steenstrup, 1875, Metasepia Hoyle, 1885.

8. Idiosepiidae Appellof, 1898: Idiosepius Steenstrup, 1881.

9. Sepiolidae (Leach, 1817) Keferstein, 1866: Sepiadariinae, Naef, 1912:
Sepiadarium Steenstrup, 1881 ; S e p i o 1 o i d e a d'Orbigny, 1845.
Rossiinae Naef* 1912: Rossia Owen, 1834, Semirossia Steen-
strup, 1887. Heteroteuthinae Naef, 1912: Heteroteuthis Gray,
1849, Nectoteuthis Verrill, 1883, Iridoteuthis Naef, 1912,
Stoloteuthis Verrill, 1881 . Sepiolinae (Appellof, 1898 ) Naef, 1912 :
Sepiolina Naef, 1912, Euprymma Steenstrup, 1887, Sepiola
(Leach, 1817) Naef, 1912, Sepietta Naef, 1912, Rondeletiola
Naef, 1921.

* Its position is still problematic.

** Here belongs "Chiroteuthis" portieri Joubin, 1916 (Bull. Monaco, No.317, cf.p.227).

782

811 Order II: Octopoda Leach, 1818

Suborder a) + Palaeoctopoda Naef, 1921.

1. Palaeoctopodidae Dollo, 1912 : Palae octopus Woodw., 1896.

Suborder b) Cirroteuthoidea Berry, 1920.

1. Vampyroteuthidae Thiele, 1915: Vampyroteuthis Chun, 1913,
Watasella Sasaki, 1920, M e 1 a n ot eu th i s Joubin, 1912, Laet-
moteuthis Berry, 1913, Hymenoteuthis Thiele, 1916.

2. Cirroteuthidae Keferstein, 1866: Cirroteuthis Eschricht, 1836,

St au r ot eut hi s Verrill, 1879, Froekenia Hoyle, 1904, Cirro-
thauma Chun, 1911, Chunioteuthis Grimpe, 1916.

3. Opisthoteuthidae Verrill, 1896: Op i s th o t e u t h i s Verrill, 1883 .

Suborder c) Polypodoidea Naef, 1921. ^

a, Ctenoglossa Naef, 192p!'

1. Amphitretidae Hoyle, 1886: Amphitretus Hoyle, 1885.

2. Bolitaenidae Chun, 191 1 : Bolitaena (Steenstrup, 1859, Hoyle, 1886)
Chun, 1915, Eledonella Verrill, 1884 (Japetella Hoyle, 1885?),
(V i t r e 1 e d o n e 11a Joubin, 1818?).

b. Heteroglossa Naef, 1921.

1. Octopodidae d'Orb., 1845: Octopus Lam., 1798, Eledone Leach,
1817, Velodona Chun, 1915, Cistopus Gray, 1849, Pinnocto-
pus d'Orb., 1845, Paroctopus n.gen., (?Bathy polypu s,
Benthoctopus, Atlantoctopus Grimpe, 1921 ).

2. Argonautidae Naef, 1912: Argonautinae Naef, 1921 : Argonauta L.,
1758, Ocythoe Rafinesque, 1814. Tremoctopodinae Naef, 1921 :
Tremoctopus D.-Ch., 1829; Alloposus Verrill, 1880.

b. LIST OF MEDITERRANEAN SPECIES

1. Loligo vulgaris Lam., 1798.

2. Loligo forbesi Steenstr., 1856.

3. Alloteuthis media (L., 1758) Naef, MS. Wiilker, 1920 = "Loligo"
marmorae Verany, 1840.

4. Alloteuthis subulata (Lam., 1 798 ) nov. = "Lo 1 ig o" m e d i a
Jatta, 1896.

5. Ctenopteryx siculus (Ver., 1851 ) Pfeff., 1900.

6. Pyroteuthis m a r g a r it i f e r a (Riippell, 1844) Hoyle, 1904.

7. Abralia veranyi (Rlipp., 1844) Pfeff., 1912.

8. Abraliopsis morrisi (Ver., 1839) Pfeff., 1900.

9. Thelidioteuthis alessandrinii ( Verany, 1851 ) Chun, 1910.

10. Gonatus fabricii (Licht., 1818) Steenstr., 1880.

* Bolitaena d i a ph a n a (Chun, 1915, p.493) is the type of the group. Its special character is the presence
of multicuspid teeth in the lateral rows of the radula; this is also the case in Amphitretus (loc.cit.,
p.533). However, the name is so general that it fits any form of radula, and it is therefore also valid if a
simpler form of the radula is found in related forms.

783

812

11. Ony chot euth is banksi (Leach, 1817) d‘Orb., 1839.

12. Chaunoteuthis mollis Appellof, 1891.

13. A n c i s t r ot eu th i s 1 i c h t e n s t e i n i (d*Orb., 1839) Gray, 1849.

14. Octopodoteuthis sicula Riipp., 1844.

15. Calliteuthis reversa Verr., 1880.

16. H i s t i o t e u t h i s bonelliana (Fer., 1826) d*Orb., 1839.

17. B r a c h i o t e ut h i s riisei (Steenstr., 1882) Chun, 1910.

18. Chiroteuthis veranyi (Fer., 1835) d'Obr., 1839,

19. Galiteuthis armata Joubin, 1898.

20. Leachia cyclura Lesueur, 1821 (uncertain),

21. Liocranchia reinhardti (Steenstrup, 1856) Pfeff., 1884 (uncertain).

22. Illex coindeti (Ver., 1837) Steenstr., 1880.

23. Todaropsis eblanae (Ball, 1841) Posselt, 1893.

24. O m m a t o s t r e p h e s sagittatus (Lam., 1 798) d'Orb., 1848.

25. Sth e n ot eu th i s bartrami (Lesueur, 1821 ) Verr,, 1880.

26. Thy s anot euthi s rhombus Troschel, 1857.

27. Sepi-a officinalis L.,1758.

28. Sepia orbignyana Fer., 1826.

29. Sepia elegans d'Orb., 1839.

30. Spirula spirula (L., 1758) Hoyle, 1909.

31. Rossia macrosoma (Delle Chiaje, 1829) d'Orb., 1839.

32. H e t e r o t e u th i s dispar (Rupp., 1844) Gray, 1849.

33. Sepiola s t e e n s t r u p i a n a Levy, 1912,

34. Sepiola aurantiaca Jatta, 1896.

35. Sepiola ligulata Naef, 1912.

36. Sepiola rondeleti Steenstr., 1856.

37. Sepiola affinis Naef, 1912.

38. Sepiola intermedia Naef, 1912.

39. Sepiola robusta Naef, 1912.

40. Rondeletiola minor Naef (1912), 1921.

41. Sepietta oweniana (Pfeff., 1908) Naef, 1912.

42. Sepietta obscura Naef, 1916.

43. Sepietta neglecta Naef, 1916.

44. Octopus vulgaris Lam., 1798.

45. Octopus saluzzii Verany, 1837.

46. Octopus macropus Risso, 1826.

47. Octopus defilippii Verany, 1851.

48. Octopus tetracirrus Delle Chiaje, 1829.

49. Octopus unicirrus (Delle Chiaje, MS) d'Orb., 1839,

50. Eledone moschata (Lam., 1 798) Leach, 1817.

51. Eledone cirrosa (Lam., 1798) d'Orb., 1839.

52. Tremoctopus violaceus Delle Chiaje, 1829.

53. Ocythoe tuberculata Rafinesque, 1814.

54. Argonauta argo L.,1758.

c. KEY TO THE MEDITERRANEAN CEPHALOPODA
(Postembryonic stages)

la. Total length without tentacles or hectocotyli 1 — 10 mm 2

lb. Total length without tentacles or hectocotyli 10— 60 mm 19

784

(813)

FIGURE 472. Problematic larvae of Oegopsida which were considered as very young stages of Spirula and
are of great morphological interest. (About 10 x, slightly diagrammatic):

A — lateral view; A| - dorsal view of posterior body of embryo of "Spirula" collected near the Balearic
Islands (after Joubin, 1920, Plate 11, Figure 8, depth 2,000m); B - ventral view; B^ - dorsal view of a "young
form of Spirula" from the Atlantic, found among the "unidentifiable larvae" of C.Chun in Leipzig
(cf. Naef,1913,p.455).

B is undoubtedly a larva of Oegopsida (cf. pp.491,515); this form was still considered as unusual in 1913
because of the very large cone (c). Further specimens of this type have been found since then (Figure 67 on
p.l61 and Figure 112 on p.235). Such stages are typical for the youngest Oegopsida (p.233): they usually
develop inside the egg (Vol. II, Plates IX— X). The oegopsid character of the larvae is recognizable by the
inhibition of the lateroventral and ventral arms (3rd and 5th arms) so that the animals often seem to have
6 arms (p.234).

The cone of A is still larger, of extreme and unique form which distinctly resembles the incomplete initial
chamber of the Belemnoidea (Figure 36 on p.l07). Joubin’s assumption that this is a young Spirula is still
more probable than in B but it is also incorrect. As the separate "anlagen" of the LV and V arms show, this is
also a larva of Oegopsida. Another argument against Spirula is that freshly hatched larvae of this genus
(p.515) are much larger (p.470) and have more than one chamber and that these chambers should be smaller and
already calcified (Figure 279 on p.512). Nevertheless, a morphological relationship to Spirula is possible,
and even certain, but this is explained by the fact that the general form of the mantle sac is typical for all the
Dibranchiata. This formation is apparently primary because of the close resemblance of the shell of Decapoda
to the primary conditions in the Belemnoidea and because of the position of the shell in the soft body. The
mantle sac of the older embryos of the ancient Cephalopoda (i.e. Belemnoidea) cannot be assumed to have been
different. Compare Figure 36 on p.l07 (a), which was drawn long before this larva was known.

This applies in particular to the cone, the form of which could develop into the initial chamber of the
Belemnoidea or Sepioidea just as readily as it forms what is here the typical rudiment of the Teuthoidea. The
proostracum is already of the type of Teuthoidea and has reached a further stage of modification which we
consider as characteristic for the Mesoteuthoidea (p. 146, Figure 62). The form of the anterior end resembles
strikingly that of Beloteuthis acuta (Fossile Tintenfische,pp.l47, 148); the arms resemble those of the
Metateuthoidea, particularly the Oegopsida.

Of the Mediterranean forms, only Gonatus, Pyroteuthis, Calliteuthis and Histioteuthis have
such a young stage. All others can be recognized by their special characters already at this early stage. This
is probably a freshly hatched Pyroteuthis. Older stages of this genus have a very large cone (Figure 131
on p.275),at a time when the proostracum is already of the type of the Metateuthoidea and the retarded arms
have already developed.

B is probably a larva of Cranchiidae. This is shown by the very small arms, also the relatively uninhibited
arms. The related Chiroteuthidae and Brachioteuthidae (p.791) could also be considered. This is suggested by
the early narrowing of the proostracum, i.e. the rudimentary lateral plates (flag) and the long, narrow, free
rhachis (which is not always as distinct as shown in the drawing).

1 — "anlage" of lateroventral arms; v — "anlage" of ventral arms; i — funnel; x - insertion of the muscular
mantle (dotted) on the margin of the shell; p - proostracum, later divided into parts which become rhachis (r)
and lateral plate (s); c - cone; f - fin, displaced toward the ventral side; h - shell fold (skin) over the cone.

785

Ic. Total length without tentacles or hectocotyli over 60 mm 50

2a. Fins absent Larvae of Polypodoidea 16

2b. Fins forming a margin occupying over half of sides of mantle. Only

Sepia elegans (Figure 31 7 on p. 562 ); other species of Sepia
are larger from the beginning.

2c. Fins attached to sides of dorsum, widely separated, large, rounded,

with a narrower, more solid base. Young stages of Sepiolidae ... 14
2d. Fins situated near posterior end, usually more dorsal, more rarely
lateral-ventral, always at first small and occupying less than half

of sides of mantle 3

3a. Posterior end bluntly rounded, occupied by the translucent, scooplike
cone. Fins small, attached laterally to cone (often slightly more

813 dorsally or ventrally). Typical form of the youngest Dibranchiata

(Figures 67, p. 161, 13 1 , p. 275 and 473, p. 81 5) 4

814 3b. Posterior end modified, usually more pointed 5

4a. Cone calcified, about 0.7 mm in diameter, more than hemispherical,
curved ventrally. Arms without stalks, uniformly developed.
Youngest stages of Sp i r u 1 a, probably still embryonic (Figure 262
on p. 491 ).

4b. Cone soft, often larger or smaller, not curved ventrally. Arms

different, LV and V arms markedly retarded, stumpy, or still absent
(Figure 472 on p. 813). Youngest larvae of Oegopsida, undetermined.
D and DL arms at first with a single sucker, tentacles with several
suckers.

5a. Posterior end laterally compressed, very blunt, occupied by the

distinctly chambered, ventrally curved calcified shell (phragmocone).
Fins situated obliquely on sides of posterior end. Postembryonic
stages of Spirula ("larvae”) (Figure 253 on p.477 and Figure 279
on p. 512).

5b, Posterior end less blunt, containing a relatively smaller cone which

is often not recognizable from the outside 6

6a. Arms typical for the Oegopsida, i. e. as described in 4b; larvae of

very varying form 7

6b. LV and V arms well developed from the beginning, at any rate not
smaller than the others during postembryonic development
(Figure 88 on p. 188). Juvenile stages of Loliginidae. Markedly

larger animals with the same morphology belong to Loligo for-
b e s i, smaller animals to Alloteuthis; further determination is
not possible.

7a. Second typical stage of larvae of Oegopsida, resembling 4b. Cone

increasingly retarded, indistinct. Posterior end increasingly pointed
(Figure 199 on p.404, ci— -ca). Fins therefore situated close together
(Figure 146 on p. 305). G on at us, Onychoteuthidae, Chiroteuthis,

Cranchiidae, Ommatostrephidae 8

7b. Same, but cone flatter at an early stage, scooplike only at the

beginning, later spoon-shaped, often completely indistinct. Ct enop -

teryx, O c t o p o d o t e u t h i s, T hy s a no t e u t h i s 8

8a. Cone slender, pointed by apposition of a solid rostrum which projects
beyond the fins (Figure 147 on p. 306). Further specific stages of
Onychoteuthidae (and Gonatidae?) 9

786

815

8b. Cone short. Rostrum absent. Body with fleshy posterior apex
(Figures 141 on p. 291, 142 on p. 294, 212 on p. 427). Enoploteu-

thidae, Ommatostrephidae 9

8c. Cone lengthened and more pointed. Posterior end of body long,
slender, supported by hollow part of shell (Figures 127 on p. 270,

191 on p. 387, 193 on p. 393). Chiroteuthis, Brachioteuthis,

Galiteuthis, Pyroteuthis 9

9a. Tentacle stalks longitudinally fused. Clubs rudimentary, with only

4 small suckers. "Rhy n c hot eu th i s" larvae of Ommatostrephidae
without further identification. (Figure 209 on p. 421 and Figure 212
on p. 427.)

9b. Tentacles free, closely resembling the other arms so that the animal
apparently has 6 arms while the LV and V arms are still retarded.
Youngest Onychoteuthidae (and Gonatidae?). (Figure 145 on p. 305,
and Figure 148 on p. 307.)

9c. Tentacles early much longer than the other arms, with numerous

suckers 10

10a. Eyes markedly stalked (Figure 164 on p. 337 and Figure 199 on p. 404).

Cranchiidae (Galit euth is), O c t o p o d ot e u t h i s . 11

10b. Eyes more or less projecting but not stalked 11

11a. Arms situated together on long, pillarlike base (Figure 197 on

p. 400) Cranchiidae (Galiteuthis)

lib. Pillarlike base of arms indicated at most by a constriction before the

eyes 12

12a. Tentacle clubs with 7 rows of suckers distally. Fins rounded, pro-
jecting beyond posterior end (Figure 173 on p. 354 and Figure 177 on

p.357) Histioteuthidae

12b. Tentacle clubs with 4—8 distal rows of suckers. Fins supported at
first by a single muscular ray, later by several rays, and gradually
growing anteriorly on sides of mantle. Ctenopteryx (Figures 173 —
177 on pp. 353—357), cf. also No. 13.

12 c. Tentacle clubs with not more than 4 distal rows of suckers 13

13a. Fins relatively large, rounded, narrower at the base and never

reaching the pointed posterior end (Figure 131 on p. 275 and Figures

132, 133 on p. 277) Pyroteuthis

13b. Fins relatively large, growing early anteriorly on sides of mantle.
Mantle sac plump. Ventral protective margins of lateral arms

widened early T hy s a no t e ut hi s

13c. Fins small, rounded, completely terminal at an early stage

(Figure 112 on p.235). Arms very long at an early stage (Figure 140

onp.290) Abralia, Abraliopsis

13d. Fins the same, but arms not particularly long, however, neck region
very long between head and funnel (Figure 186c on p. 372; cf. also

Figures 190, 191 on p. 387) Brachioteuthis

14. (Sepiolidae). a. Fins of youngest stages (Plate VIIl) widely separated,

later with a distinct corner directed inward and posteriorly

Hete roteuthis (Plate VIII)

14b. Fins almost circular (Figure 327 on p. 570) 15

15a. Mantle cavity with ear -shaped luminous glands projecting laterally

from the ink sac (Figure 334 on p. 581). Sepiola (for determination
of the species, see pp. 606—609).

787

FIGURE 473. Grenacher’s larva of Oegopsida. 12 X. a) Freshly hatched animal, after Grenacher
(1874, Plate 40, Figure 12). b) Reconstruction and interpretation of the specimen on the basis of
my experience with the deformation of the mantle sac and arms of young Oegopsida during pre-
servation. The cone and insertion of the fins are exaggerated (diagrammatic). Note the natural
extension of funnel and arms (Figure 67 on p.l61 and Figure 131 on p.275).

CO - ventral margin of cone; fl - fin; vm - ventral corner of mantle margin; ro - olfactory
organ; tr - funnel; do - rudiment of yolk sac; ve - ventral arm; Id - laterodorsal arm; te -
tentacle; po - margin of proostracum; s - lateral plate; r - rhachis; c - cone flag; st - stellate
ganglion; If - later position of primary lid fold; x - posterior end of lateral margin of pillar of
tentacles, connected with the primary lid fold.

Like Figure 67 on p.l61 and the preceding figures, this drawing shows the typical conditions of very
young Decapoda, especially Oegopsida. The oegopsid character is shown by the relative develop-
ment of the arms, the LV and V arms are inhibited.

15b. Ink sac with "anlagen" of glands situated close together in the middle
between renal pores and replacing the accessory nidamental glands.

In preserved specimens, these "anlagen" appear at first as mat spots
in the young stages and in the form of a median lens (Plate XIX) in
larger animals. They are present in both sexes (Figure 371 on p. 631).

Length of young animals only about 2— 3 mm Eondeletiola

15c. Such "anlagen" of glands always absent in the male and lens-shaped

in the female. Length not less than 5 mm Sepietta

16. (Octopoda). a. Apparently 7-armed, i.e. with one LV arm markedly
abnormal, usually visible in a pocket below the eye or projecting
free like a club (hectocotylus). Male of Argonautidae (Figures 438 —
468 on pp. 738, 757, 766, 785); otherwise as 17.

16b. Eight arms, LV arms symmetrical 17

17a. Arms of almost uniform size (Figure 409 on p. 687 and Figure 410

on p. 688) Octopus

17b. Median arms longer than the lateral arms, LV arms markedly
retarded (Figure 447A on p. 750).

17c. D arms markedly longer than the others, LV arms markedly retarded

(Figures 437-439 on pp. 736, 738, 765)

Tr emoctopus, Argonauta 18

18a. Funnel bond only in form of a hooklike funnel corner (Plate X,

Figure 435 on p. 727) Tremoctopus

18b. Funnel bond dishlike, with distinct margin (Plate X, Figure 455 on

p.765) Argonauta

19a. Larger animals, 1 — 6 cm long without tentacles. 10 arms, 4th pair
more or less lengthened and separate from the others. Suckers on
thin stalks more or less bilateral (eccentric) Decapoda 20

19b.

20a.

20b.

20c.

20d.

21.

21a.

817

22.

22a.

22b.

23a.

23b.

24.

24a.

25.

25a,

Eight arms (rarely apparently only 7 arms), lateroventral arms
never detached on both sides from the others. Suckers more or
less radiate, situated on thick stalks or on a wide base . . . Octopoda 41
Fins rounded, large, attached at sides of dorsal surface. Posterior
part of body short, saclike, rounded posteriorly. Shell rudimentary
....................................... Sepiolidae 27

Fins rounded, small, subterminal on the saclike body (Figures 253,

277, 281 on pp.477, 509, 516). Spiral chambered shell visible through
dorsal and ventral skin.

Fins in the form of wide margins along greater part of sides of Spirula
mantle. Body saclike, supported by oval, calcified, air -containing
dorsal shell .............................. Sepia 33

Fins of small young stages subterminal or terminal, rounded, rela-
tively small, later growing more or less along sides of mantle.
Posterior part of body increasingly slender, supported by a thin
gladius, often with an already rudimentary cone at the end .......

...................................... Teuthoidea 21

Lens of eye covered by transparent skin (cornea)j convex like a watch-
glass in life and with a very small pore at the anterior margin.

Dorsal arms very weakly developed already in the youngest stages;
lateroventral arms very well developed and cone almost unrecog-
nizable (Plate I) .......................... Loliginidae 35

Lens of eye usually projecting anteriorly (always projecting in live
specimens), or covered partly or temporarily (Figures 118, 119 on
p. 254) by a skin fold (primary lid) forming a wide opening at rest
(p. 257). Youngest stages: V and LV arms still shorter than the
others (Figure, 148 on p. 307 and Figure 150 on p. 309). Cone usually
delicate, but in small specimens always well developed, spoon-shaped
to slender -conical .......................... Oegopsida 22

Buccal funnel consisting of 8 parts (Plate III, Figure 1 ) .........

.................................. Enoploteuthidae 36

Buccal funnel consisting of 7 parts ..................... 24

Buccal funnel consisting of 6 parts (Plate III, Figure 2) ........ 23

Tentacles large, with 7 rows of suckers. Body with luminous organs
on the ventral side, at least in the larger stages (Figure 183 on
p.365) .............................. Histioteuthis

Tentacles delicate, later absent (p. 339), with few (to 8) suckers in
2 rows. End of arms thickened. Body transparent, later translucent
and gelatinous, without luminous organs in the skin (Figure 164 on
p. 337 and Figure 167 on p. 342) . . . . . . . . . . . Octopodoteuthis
Funnel bonds and mantle bonds fused, also neck bonds and collar
bonds ................................ Cranchiidae 40

Above bonds free ................................. 25

Fins like in fish, supported by muscular rays. Mantle sac as in
Sepia, blunt posteriorly (Figures 120, 121 on pp. 256 and 257) ....

..................................... Ctenopteryx

Fins forming almost two circles, projecting behind posterior end of
mantle sac and connected by a frenulumlike, thin posterior apex of
body (Figure 174 on p. 355) .................. Calliteuthis

789

25b.

818

Fins of smaller stages still rounded, transverse, in the form of
subterminal or terminal lobes which gradually assume a transverse
oval and finally transverse rhomboidal, terminal form (Plate V), . . 26
25c. Fins also transverse-oval (Figure 190, 191 on p. 387). Cone lengthened
like a tube, supporting a tail-like appendage of the mantle sac and
projecting beyond the fins. Neck region strikingly lengthened ....

Chiroteuthis (**Do r a tops i s*’ stage)

26. Arms with 4 rows of suckers, tentacles with 8 or more rows ......

Gonatus

26a. Arms with 2 rows of suckers 27

27. Hand part of tentacle with 4 rows proximally. Body very narrow,
membranous; neck region markedly lengthened in the smaller animals.
Funnel bond long, narrow (Figure 186 on p. 372). . .Brachioteuthis

27a. Hand part of tentacle with 4 rows of suckers. Neck not lengthened . .

28

28. Funnel bond long, narrow (Figure 157 on p. 319). . . Onychoteuthidae 38
28a. Funnel bond in the form of a wide triangle with inward projecting

tubercles (Figure 215 on p. 427) . . Ommatostrephidae 39

29a. (Sepiolidae). Suckers on arms in 4 rows; mantle margin free

dorsally (Figure 338 on p. 592) Rossia

29b. Suckers on arms biserial; mantle margin free dorsally (Plate VIIl)

Heteroteuthis

29c. Suckers biserial, mantle margin connected with head by neck bond

dorsally Sepiolinae 30

30. Cf. at No. 15 about determination of genera, also at No. 31 for
Sepiola and No. 32 for Sepietta.

31. (Sepiola). Determination of male after Figure 344 on p. 604, female
after Figures 355—370 on pp. 611 — 628.

32. (Sepietta). Determination of the male after Figure 378 on p. 644,
the female after Figure 380 on p. 647 and Figure 384 on p. 653.

33. (Sepia), a. Coloration gray, grayish brown, except on dorsal side.

Suckers in 4 rows. Posterior end blunt S. officinalis

33b. Coloration yellowish brown to reddish brown or reddish white . ... 34
34a. Suckers biserial at least at base of arms, then in 2—4 rows.

Posterior end blunt S. elegans

34b. Suckers distinctly quadriserial in greater part of arms. Posterior
end with central apex caused by the (usually perforated) rostrum
S.orbignyana

35. (Loliginidae). a. Animals less than 2 cm long are difficult to
determine. Larger forms are recognized by the following characters*.

35b. Posterior end pointed. Dorsal margin of mantle with angular
process (Figure 89 on p, 190): Alloteuthis.

35c. Posterior end blunt. Dorsal margin of mantle with rounded process:
Loligo. For determination of species see No, 67 below.

36. (Enoploteuthidae). a. Fins almost circular, large (Figure 127 on

p. 270 and Figures 132, 133 on p. 277), leaving the gradually tapering
posterior end of the body free. Head large. Mantle sac short,

almost conical Pyroteuthis

36b. Same, but fins at first transverse- oval, then transverse-rhomboidal

(Figures 136, p. 284; 141, p. 291; 142, p. 294) ............... 37

790

36c.

Fins of oldest specimens very large, transverse-oval, occupying to
four-fifths of sides of mantle, still terminal in the youngest speci-
mens (Figure 143 on p. 299), indented posteriorly. Distribution of
luminous tubercles and structure of club later characteristic (p. 300)
Thelidiot euthi s

37. Ventral arms with distinct apical swellings (Figure 141 on p. 291)

. . . . Abraliopsis

37a. Such structures (luminous organs) absent. (See Figure 138 on

p. 288 about the protective margin of the tentacle club .... Abralia

38. (Onychoteuthidae). a. Rhachis of gladius not visible as a dark line

in the dorsal median line (Figure 159 on p. 328). Gladius with very
short, rudimentary cone Ancistroteuthis

38b. Rhachis visible as a dark line only in anterior part of dorsal median
line. Consistency of flesh gelatinous (Figure 158 on p. 324) ......

Chaunoteuthis

38c. Rhachis visible externally as a distinct, dark, dorsal median line

(p. 318). Cone well developed (p. 319). Consistency of body fleshy
to membranous Ony choteuthi s

39. (Ommatostrephidae). a. Stages 1 cm long are still larvae (p. 427)
and difficult to identify with certainty. The following characters
apply to older stages:

39b. Illicinae: Crescent-shaped fold absent (p. 427). Todaropsis:
body compact; 111 ex: figure graceful, end of club with 8 rows of
suckers.

39c. Ommatostrephinae: crescent-shaped fold present (Plate V). Omma-
tostrephes (pp. 450, 453): chromatophores sparse; Sthenoteu-
this (p.462): chromatophores dense and development retarded

(p.46l).

40. Cranchiidae: most, perhaps all forms belong to Galiteuthis

(pp. 400, 404). However, compare also Liocranchia and Leachia
(pp. 404 and 407-408).

41. (Octopoda 1—6 cm long), a. Solid mantle bond absent (Figure 435a on

p. 727). Benthic forms. Male identical to female .... Octopodidae 42
41b. Solid mantle-funnel bonds present. Pelagic forms. Males much

smaller than mature females; one arm of 3rd pair strongly modified

819 in male (hectocotylus) Argonautidae 47

42a. Suckers uniserial Eledone 43

42b. Suckers biserial Octopus 44

43 a. Coloration gray to grayish brown dor sally in life. Mantle sac without
lateral ridges. Musky odor (Figure 426 on p. 717)

Eledone moschata

43b. Coloration dorsally yellowish brown to reddish brown in life. Sides

of mantle with distinct ridges or thin skin margins at a length of about

4cm Eledone cirrosa

44. (Octopus), a. Body slender. Arms long, markedly different in size

45

44b. Body plump. Arms shorter, rather uniform 46

45a. Coloration yellowish brown to reddish brown dorsally. Arm formula:

1, 2, 3, 4 (Figure 419 on p. 706) . . . Octopus macropus

791

45b.

46a.

46b.

46c.

46d.

47.

47b.

48a.

48b.

48c.

49.

49a.

49b.

820

50a.

50b.

51a.

51b.

51c.

Yellowish brown to grayish brown.

on p. 708) ....... .............

Reddish brown to grayish brown. Arm formula:
on p. 688)

Arm formula: 3, 2, 4, 1 (Figure 420
............ O. defilippii

2, 3, 4, 1 (Figure 410
..... O. vulgaris

Yellowish brown to orange reddish. Arm formula: 3, 2,1=4; inter-
brachial membrane extending anteriorly along outer edges of arms.

Skin swollen ..O. saluzzii

Orange-yellow to orange reddish, very plump, gelatinous. Arms
short. Dorsal side with soft, constant warts . . . . O. tetracirrus
Yellowish brown, fleshy and tough. Dorsal side with hard, constant

warts O. unicirrus

(Argonautidae). a. One arm of 3rd pair absent or visible and
markedly modified (cf. No. 16). Males .................. 49

Arm apparatus complete, 3rd pair rather normal and symmetrical.

Females 48

Animal situated in boatlike, open, calcified shell which later grows
into a spiral (Figures 460—462 on pp. 769, 771 ). If the shell is lost,
the dorsal arms are curved toward each other, and a membrane with

shell -producing glands extended in the loop of the arm

Argonaut a, female

Form as in a normal Octopus (Figure 447B on p. 751), but arms
markedly different, formula: 4, 1; 2, 3. Water pores at base of V arms

Ocythoe, female

Younger stages with very short body; ventral arms retarded, dorsal
arms rapidly developing (Figure 440 on p. 739). A large membrane
between the dorsal arms. Water pores present also at base of Darms

Tremoctopus, female

Arms not of markedly different length (formula: 1, 2, 4, 3), except the
hectocotylus. Hectocotylus coiled inside bag below left eye, free in the
mature animal (p. 785), lost during copulation ... Argonaut a, male
Dorsal arms and water pores as in 48c; Hectocotylus always on the
right side, and with lateral fringes in the proximal part (Figure 445

on p. 746) Tremoctopus, male

Hectocotylus free between 2nd and 4th arm of right side, club-shaped
(Figure 452 on p. 757). Always found swimming in empty tests of

Salpa Ocythoe, male

Decapoda (p. 109) over 6cm ......................... 51

Octopoda (p. 655) over 6 cm 73

Primary lid open above lens, often contracted to an j.— shaped slit

Oegopsida 52

Primary lid continuously contracted above eye into a tough cornea
formed like a watchglass and closed except for a pore situated near
the anterior margin. Secondary lid in periphery absent; hooks
absent; suckers of arms always biserial, clubs with 4 rows ......

Loliginidae 67

Primary lid similarly modified. A distinct secondary lid usually
present around the cornea, especially along its ventral margin. If
a secondary lid is absent (Hete roteuthis), the general form of
Sepiolidae (Plate VIII) excludes the above groups. Fins not fused in
median line Sepioidea 70

792

821

51d.

52a.

52b.

52c.

53.

54a.

54b.

55a.

55b.

56a.

56b.

57a.

57b.

58a.

58b.

59a.

59b.

60a.

60b.

60c.

61a.

61b.

62a.

62b.

63a.

63b.

64a.

64b.

64c.

Primary lid often contracted above the lens into a more or less
narrow hole with a tough margin; otherwise open, as in the Oegopsida.
Habitus and translucent shell (Figure 253 on p,477) characterize

the genus Sp'irula

Buccal funnel consisting of 8 parts . Enoploteuthidae 36

Buccal funnel consisting of 7 parts 54

Buccal funnel consisting of 6 parts . 53

Cf. Histioteuthis (Figure 185 on p. 368) and Octopodoteuthis
(Figure 164 on p. 337).

Buccal funnel with median attachment to ventral arms 55

Buccal funnel with lateral attachment to ventral arms 60

Neck bond and funnel bond replaced by fusion Cranchiidae 65

Funnel bond and neck bond normal 56

Funnel bond rounded, more or less ear-shaped. Smaller specimens
as in 25c, later without tail, with short neck, thick head and especially
strong ventral arms (Figures 190—192 on pp. 387, 388)

Chiroteuthis

Funnel bond slender, long, typical 57

Fins terminal, rhomboidal-heart-shaped 58

Fins rounded, subterminal, occupying whole sides of mantle and
supported by muscular rays (Figure 121 on p. 257). . . Ctenopteryx

Suckers on arms biserial 59

Suckers on arms quadriserial Gonatus

Club with hooks or later absent, primarily with 4 rows

Onychoteuthidae 64

Club without hooks, primarily with numerous rows (Figure 186 on

p. 372) Brachioteuthis

Fins rounded, projecting beyond posterior end. Ventral side with

pearl-like luminous organs (p. 359) Calliteuthis

Fins transverse-rhomboidal, terminal. Funnel bond triangular, with

inward projecting tubercles (p. 457) Ommatostrephidae 61

Fins longitudinal -rhomboidal, occupying whole sides of mantle

(Figure 249 on p. 466) Thy sanoteuthis

Funnel pit with crescent- shaped fold and longitudinal folds

(Figure 240 on p.457) 62

Funnel pit without crescent -shaped fold and longitudinal folds ... 63
Tentacles with suckers on greater part of their length (Figure 232 on

p.448) Ommatostr ephes

Tentacles with suckers only in distal half (Figure 242 on p.458) . . .

Sthenot euthi s

End of club with 8 rows. Fins slightly longer than third of length of

mantle (Figure 216 on p. 431) Illex

End of club with 4 rows. Fins nearly half of length of mantle

(Figure 224 on p. 440) Todaropsis

(Cf. also 38). Clubs absent at an early stage. Rostrum rudimentary

Chau not euthi s

Clubs with 6—10 adhesive suckers on carpal pad; dorsal row with
one displaced and reduced median hook in dorsal row. Rostrum

short (Figure 154 on p. 316) Ony choteuthis

Clubs with 8 — 12 adhesive suckers, without such a displaced hook.
Rostrum long Ancistroteuthis

793

65a.

65b.

66a.

66b.

67.

67b.

68.

69a.

69b.

70.

70b.

71a.

71b.

72a.

72b.

73.

73b.

74a.

74b.

75a.

75b.

76a.

76b.

Club with hooks. Fins in form of a narrow leaf, tapering to a slender

apex (Figure 198 on p. 401) . . . . . . . . . . Galiteuthis

Club regularly lost, otherwise without hooks. Fins short, rounded . .

66

Ventral side of mantle sac with simple cartilaginous ridges on each

side (Figure 200 on p. 407) Leachia

Cartilaginous ridges forked posteriorly on each side (Figure 199 on

p.404) Liocranchia

(Loliginidae). a. Posterior end pointed (Figure 68 on p. 169) or pro-
duced into a tail (p. 222). Mantle with angular dorsal process. Buccal

corners without suckers Alloteuthis68

Posterior end usually blunt. Buccal corners with suckers .......

Loligo 69

(Cf. p. 215.)

Suckers of club 3—4 times larger in median rows than in marginal

rows Loligo vulgaris

Suckers of club of median rows only slightly larger than those of

marginal rows Loligo forbesi

(Sepioidea). a. Form Sepiola-like (cf. atNo. 20) .... Sepiolidae 71

Form Sepia-like (cf, No.20) Sepiidae 13

Suckers of arms normally quadriserial Rossia

Suckers of arms normally biserial Sepiolinae 72

Ink sac with ear-shaped luminous glands .... Sepiola rondeleti
Ink sac without ear- shaped luminous glands . . Sepietta oweniana
Octopoda over 6 cm long. a. One arm of 3rd pair (hectocotylus)

more or less abnormal, at least at the apex. Males 74

Such a modification absent. Females 41

Hectocotylus in the form of a large club or, if free of the envelope,
much longer and thicker than the opposite arm and with a vesicle at the

end (Figure 453 on p. 758) . Ocythoe, male

Hectocotylus with longitudinal groove which is lighter colored when
spread; hectocotylus otherwise modified only at the apex and always

shorter than the opposite arm Octopodidae 75

Hectocotylus on the left side, with a short spoon at the end

(Figure 421 on p. 710) 46c, d

Hectocotylus on the right side 76

Suckers uniserial. End of arms different, as shown in Figure 430 on
p. 720 and Figure 433 on p. 723. Cf. also 43.

Suckers biserial. Apex of hectocotylus different, as shown in
Figure 405 on p. 683. Cf. also under Nos. 45 and 46.

794

5. Bibliography

This list contains all publications used directly or indirectly. It
comprises works on recent Cephalopoda and fossil forms of special interest for
the relationships discussed. I have not excluded systematic publications
because a comprehensive list of references is more useful.

Abel, O. Palaeobiologie der Cephalopoden aus der Gruppe der Dibran-
chiaten (Paleobiology of the Cephalopoda Dibranchiata). Jena.

1916.

Abel, O. Die Methoden der palaeobiologischen Forschung (Methods of

Paleobiological Research). — In: Handb. d. biolog. Arbeitsmethoden
(Edited by Abderhalden), No. 35. 1921 .

Adams, C.B. Notice on a Fractured and Repaired Argonauta argo.—
Sillimans Journ., Vol. 6 (cf. also Ann. Mag. Nat. Hist. (2), Vol. 2.

1848.

Adams, H. and A. The Genera of Recent Mollusca. Vol. 1 . London. 1858.

A d a m s, H. and L. Reeve. Voyage of the "Samarang" in the Eastern Seas.
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Agassiz, A. On the Dredging Operations Carried out from December
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Mus. Comp. Z. Vol. 5. Cambridge, Mass. 1879.

Agassiz, A. Three Cruises of the U. S. Coast and Geodetic Survey Steamer
"Blake." - Ibid., Vol. 15. 1888.

Albini, G. Sui movimenti dei cromatofori nei cefalopodi. — Rendic. Acc.
Napoli, Vol. 24. 1885.

Alcock, A.Wm. Naturalist in Indian Seas. Mollusca, London. 1904.

Alder, F. Notes on Sepia biserialis and Sepia elegans.—

Ann. Mag. Nat. Hist. (2), Vol. 19. 1857.

Aldrovandi. Opera omnia. — Bononiae, Vol. 4. 1606.

Alessandri, G. de. Avanzi di un nuovo genere di Cefalopodi del Eocene
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Angas, G. On the Marine Molluscan Fauna of the Province of South
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Angerman, E. Uber das Genus Acanthoteuthis Miinst. (The Genus
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1902.

Apostolides, N. C. andY. Delage. Les Mollusques d'apres Aristote. —
Arch. Z. Exper., Vol. 9. 1881.

Appellof, A. Japanska Cephalopoder. ~ Kgl. Svenska Vet. Akad. Handl.,
Vol. 21. 1886.

795

Appellof, A. Om skalets bildning hos Sepia officinalis L. — Ofvers,
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Appellof, A. Teuthologische Beitrage (Xeuthological Contributions).

I. Ctenopteryx n. g., Veranya sicula Krohn, C a 1 1 i t e u t h i s
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Appelldf^A, Idem. 2. C h a u n o t e u t h i s n. g. Oegopsidarum. — Ibid,
f. 1890/1891.

AppellofjA. Idem, 3. Bemerkungen iiber die auf der norwegischen
Nordmeer-Expedition (1876—78) gesammelten Cephalopoden;

4. liber einen Fall von doppelseitiger Hectocotylisation bei Eledone
cirrhosa (Lam.) d'Orb. (3. Notes on the Cephalopoda Collected by
the Norwegian North Sea Expedition of 1876—1878; 4. A Case of
Bilateral Hectocotylization in Eledone cirrhosa (Lam. ) d'Orb. ). —
Ibid. f. 1892/93.

A p p e 1 1 o f. A, Die Schalen von Sepia, Spirula und Nautilus.

Studien uber den Bau und das Wachstum (The Shells of Sepia, Spi-
rula and Nautilus. Studies on their Structure and Growth).—

Kgl. Svenska Vet. Akad. Handl., Vol. 25 . 1894.

Appelldf,A. Cephalopoden von Ternate. 1. Verzeichnis der von Prof.

Kiikenthal gesammelten Arten. 2. Untersuchungen tiber Idiosepius,
Sepiadarium und verwandte Formen, ein Beitrag zur Beleuchtung
der Hectocotylisation und ihrer systematischen Bedeutung (Cephalo-
poda from Ternate. 1. List of the Species Collected by Professor
Kiikenthal. 2. Studies on Idiosepius, Sepiadarium and Related
Forms, with Particular Reference to Hectocotylization and its Syste-
matic Importance). — Abhandl. Senckenberg. naturf. Ges.,VoL24.

1898.

Appellof, A. tiber das Vorkommen innerer Schalen bei den achtarmigen
Cephalopoden (Octopoden) (The Occurrence of Internal Shells in
Octopoda). — Bergens Mus. Aarsber. f. 1899.

Aristoteles. De animalibus historiae libri X. With a commentary by

J. F. Schneider. Leipzig. — De Generatione Animalium, lib. V. 1811.

A s h w o r t h, J. H. and W . E. H o y 1 e. The Species of Ct e nopt e r yx, a

Genus of Dibranchiate Cephalopoda. — Mem, Lit, Phil. Soc. Man-
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Aubert, H. Die Cephalopoden des Aristoteles in zoologischer, anato-

mischer und naturgeschichtlicher Beziehung (Zoology, Anatomy and
Natural History of the Cephalopoda of Aristotle). — Engelmann,

Leipzig. 1862. (See also Zeitschr. wiss. Z., Vol. 12. 1863.)

Aubert, H. and F.Wimmer. Aristoteles Tierkunde (The Zoology of

Aristotle). — In Greek and German. Two Volumes. Leipzig. 1868,
Aucapitaine, M.H. Mollusques cephalopodes observes sur le littoral
de I'Algerie. — Revue Z. (Guerin-Meneville). 1863.

Audouin, J. Viet. See Savigny (explanation of plates). 1825.

Babor, J. F. Zur Histogenese der Bindegewebssubstanzen bei Weichtieren
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Baglioni, S. Sur Paction physiologique du poison des Cephalopodes.—

Arch. Ital. Biol., Vol. 51 . 1908.

796

Baglioni, S. Einige physiologische Beobachtungen an einem lebenden

Weibchen der Argonauta argo (Physiological Observations on a
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Baglioni, S. Zur Physiologic des Geruchssinnes und des Tastsinnes der
Seetiere. Versuche an Octopus und einigen Fischen (Physiology
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1909; also in: Arch. TtaL Biol. Vol. 52.

Baglioni, S. Zur Kenntnis der Leistungen einiger Sinnesorgane (Gesichts-
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Cephalopoden und Fische (On the Function of Some Sense Organs
(Optic, Tactile and Olfactory) and the Central Nervous System in
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Baker, H. An Account of the Sea Polypus (O c t o p u s). — Phil. Trans.
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Barrande. liber die innere Struktur der Nautilidenschalen (On the Inner
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Barts ch, P. Pirates of the Deep. Stories of the Squid and Octopus.—

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besonderer Beriicksichtigung der im Mittelmeer haufigen Formen
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Stat. NeapeL, Vol. 19. 1909.

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797

■B e 1 1 a r d i, L. Notice sur 1' A r g o n a u t a nit id a Lk. — Bull. Soc. Geol.

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Belon, P. De Aquatilibus. Paris. 1551.

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798

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Grimpe, G. Zur Systematik der achtarmigen Cephalopoden (Systematic s
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814

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815

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816

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Hoffmann, C.K. tiber die Stabchen in der Retina des Nautilus (On the
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817

Home, E. The Distinguishing Characters between the Ova of the
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Hoyle, W.E. Note on Loligo forbesii Steenstrup, the so-called
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Hoyle, W.E. 1. Supplement 1887-96. - Ibid., Vol. 12. 1897.

Hoyle, W. E. 2. Supplement 1 897-1 906. - Ibid., Vol. 1 7. 1909.

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Hoyle, W.E. Note on d'Orbigny's Figure of Onychoteuthis dussu-
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Hoyle, W. E. On the Generic Names Octopus, Eledone, and Histiop-
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H o y 1 e, W. E. On a New Species of Sepia and Other Shells Collected by
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Hoy le, W. E. On an Intrapallial Luminous Organ in the Cephalopoda. — -
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Hoyle, W.E. The Luminous Organs of Pt e ryg i o t e u t h i s marga-
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Vol. 46. 1902.

818

Hoyle, W. E. British Cephalopoda, their Nomenclature and Classification.—
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Hoyle, W.E. Cephalopoda. National Antarctic Expedition. 1907.

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819

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Ishikawa, C. Einige Bemerkungen iiber den leuchtenden Tintenfisch
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Ibid., Vol. 43. 1913.

Ishikawa, C. Uber eine Art von Enoploteuthis, Enoploteuthis
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Ishikawa, C. and Y. W a k i y a. Note on a Gigantic Squid, Obtained from
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820

Ishikawa, C. andY.Wakiya. On a New Species of M o r o t e u t h i s from
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Issel, R. Primo contribute alia conoscenza dello sviluppo dei cefalopodi

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Jaeckel, O. Uber neue Beobachtungen an Orthoceren (New Observations
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J a kub s k i, A. W. Studien iiber das Gliagewebe der Mollusken (Studies on
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834 Jatta, G. Sopra il cosi detto ganglio olfattivo dei Cefalopodi. — Boll. Soc.
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Jatta, G. La vera origine del nervo olfattivo nei Cefalopodi. — Ibid. 1887.

Jatta, G. La innervazione delle braccia dei Cefalopodi. — Ibid., Vol. 3.

1889.

Jatta, G. Elenco dei Cefalopodi della "Vettor Pisani." — Ibid. 1889.

Jatta, G. Sopra I'organo dell' imbuto nei Cefalopodi. — Ibid., Vol. 7. 1893.

J atta, G. Cefalopodi. — Fauna Flora Golf Neapel Mon. 23. 1896.

Jatta, G. Sopra alcuni Cefalopodi della "Vettor Pisani." — Boll. Soc.
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Jatta, G. A proposito di alcuni Cefalopodi del Mediterraneo. — Ibid., Vol. 17.
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821

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Joubin, L. Sur la ponte de I'Eledone et de la Seiche. - Arch. Z. Exper.
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Joubin, L. Recherches sur I'appareil respiratoire des Nautiles. — Revue
Biol. Nord France, Vol. 2. 1890.

Joubin, L. Recherches sur la coloration du tegument chez les Cephalo-
podes. — Arch. Z. Exper. (2), Vol. 10. 1892.

Joubin, L. Recherches sur I'appareil lumineux d'un Cephalopode:

H i s t i o t e u t h i s riippellii Verany. — Bull. Soc. Sc. Med. Guest.,
Vol. 2. 1893.

Joubin, L. Note complementaire. — Ibid. 1893.

Joubin, L. Voyage de la Goelette "Melita" sur les cotes orientates de
I'Ocean Atlantique et dans la Mediterranee. Cephalopodes. — Mem.

Soc. Z. France, Vol. 6. 1893.

Joubin, L. Quelques organes colores de la peau chez deux Cephalopodes
du genre C h i r o t e u t h i s. — Ibid. 1893.

Joubin, L. Note sur une adaptation particuliere de certains chroma-

tophores chez un Cephalopode. — Bull. Soc. Z. France, Vol. 18. 1893.

Joubin, L. Reponses a quelques critiques au sujet des chromatophores
des Cephalopodes. — Arch. Z. Exper. (3), Vol. 1. 1893.

J oubin, L. Note sur les Cephalopodes recueillis dans I'estomac d'un

Dauphin de la Mediterranee. — Bull. Soc. Z. France, Vol. 19. 1894.

Joubin, L. Cephalopodes d'i\mboine. — Revue Suisse Z., Vol. 1 . 1894.

J oubin, L. Note preliminaire sur les Cephalopodes provenant des

Campagnes du Yacht Hirondelle.— Mem. Soc. Z. France, Vol. 7.

1894.

Joubin, L. Nouvelles recherches sur I'appareil lumineux des Cephalopodes
du genre H i s t i o t e u t h i s. — Bull. Soc. Sc. Med. Guest. 1894.

Joubin, L. Note complementaire sur un Cephalopode d'Amboine. — Revue
Suisse Z., Vol. 3 . 1895.

Joubin, L. Cephalopodes recueillis dans I'estomac d'un Cachelot capture
aux lies Azores. — C. R. Acad. Paris, Vol. 121. 1895.

822

J oubin, L. Contribution a I'etude des Cephalopodes de I'Atlantique Nord. —
Res. Camp, Sc. Monaco, Vol. 9, 1895.

Joubin, L. Note sur les appareils photogenes cutanes de deux Cephalo-
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Joubin, L. Note sur divers fragments d'un Cephalopode: i\lloposus
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Joubin, L. Observations sur divers Cephalopodes. Ire note: Abraliop-
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1896.

Joubin, L. Idem. 2 e note: Octopus punctatus Gabb. — Mem. Soc.

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Joubin, L. Idem. 3e note: Cephalopodes du Musee polytechnique de
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Joubin, L. Sur quelques Cephalopodes du Musee royal de Leyde et

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Joubin, L. Observations sur divers Cephalopodes. Grimalditeuthis
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Joubin, L. Idem. Sur le genre C u c i o t e u t h i s. — Ibid. 1898.

Joubin, L. Note sur une nouvelle famille de Cephalopodes. — Ann, Sc. N.
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Joubin, L. Liste des Cephalopodes recueillis pendant les dernieres

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J oubin, L. Cephalopodes provenant des campagnes de la Princesse Alice
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1900.

Joubin, L. Revision des Sepiolidae. — Mem. Soc. Z. France, Vol. 15 ,

1902. See also Bull. Soc. Z. France.

J oubin, L. Sur quelques Cephalopodes recueillis pendant les dernieres
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Joubin, L. Observations sur divers Cephalopodes. 7e note: Hetero-
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J oubin, L. Note sur les organes lumineux de deux Cephalopodes. — Bull.
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J oubin, L. Note sur les organes photogenes de Poeil de Leachia
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J oubin, L. Description de deux Eledones provenant de 1' expedition du

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Joubin, L. Observations sur une jeune S p i r u 1 a. — Bull. Inst. Monaco,

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Joubin, L. Sur une jeune Spirule. - C. R. Acad. Paris, Vol. 150. 1910.

Joubin, L. Sur les Cephalopodes captures en 1911 par S. A. S. le Prince
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J oubin, L. Etudes preliminaires sur les Cephalopodes recueillis au cours
des croisieres de S. A. S. le Prince de Monaco. Ire note: Melano-
teuthis lucens n. g. n. sp. - Bull. Inst. Monaco, No. 220. 1912.

823

J oub i n, L. Idem.

No.226. 1912.
J oub i n, L. Idem.

Ibid., No. 275.
J o u b i n, L. Idem.

No.317. 1916.
J ou b i n, L. Idem.

No. 339. 1916.
J ou b i n, L. Idem.

Ibid., No. 340.
J ou b i n, L. Idem.

Ibid., No. 351.
J ou b i n, L. Idem.

2me note: Cirroteuthis grimaldii n. sp. ~ Ibid.,

3me note:
1913.

4me note:

Mastigoteuthis magna n, sp. ~

Chiroteuthis portieri n. sp
5me note: Moschites verrucosa (Verrill)

Ibid.

Ibid.,

6me note: Vitreledonella richardi Joubin. —

1918.

7me note: Cycloteuthis sirventi n. g. et sp. —

1919.

8e note: Quelques points de I'anatomie d'un Cephalopode
abyssal: Eledonella diaphana Hoyle. — Ibid., No. 382 . 1920.

Joubin, L. Cephalopodes provenant des Campagnes de la Princesse Alice
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Kent, W. S. A Further Communication upon Certain Gigantic Cephalopoda
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Kirk, T.W. Description of New Cephalopoda. — Ibid., Vol. 14 . 1881.

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Kirk, T.W. Brief Description of a New Species of Large Decapoda
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824

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826

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833

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5110/2

834

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835

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847

S t e e n s t r u p, Jap. Idem. 6. Species generis Sepiolae maris me-
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852

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Verrill, A.E. Idem. - Ibid., Vol. 22. 1881.

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853

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1835,

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854

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855

« « «

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See also the following early literature: Bibliotheca zoologica II (Taschenberg),
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For more recent publications, see the later issues (1879—1913) of the Zool.
Jahresbericht.

856

6. SUBJECT INDEX WITH REMARKS AND LIST OF FIGURES

The following list contains the systematic -morphological terms in this
volume. Some of the terms carry explanatory remarks. The numerals
indicate the [German] page on which the term is discussed, an asterisk
indicates a figure or its explanation. An exclamation mark indicates a place
of particular importance. The letter F refers to my monograph on the fossil
Cephalopoda (l922).

The Index is divided into a morphological and a systematic part. The
first covers the structural elements and their combination into a typical
whole; the second is a review of the species (communities of reproduction).
Systematic morphology is based on both concepts, because the "homologous"
parts are followed throughout the system and the system is based on
typical similarities, i. e. homologies. About the basic methodical concepts
see the introduction.

a. MORPHOLOGICAL INDEX

Abdominal complex 69.', 70*, 85*, 103.'. Also Anal complex 68*. See also
Roof of mantle cavity
A c a n t h o t e u t h i s cf. p. 858

Accessory nidamental glands (on those of Nautilus cf. Naef, 1913, also
p. 106. As in the Dibranchiata, these glands are situated between the
nidamental glands and the anus, at the border between the abdominal
complex and the mantle; they form a stripe of skin with numerous pores
the homology of which is not certain). 105*, 125*, 183*, 205*!, 259,

Plates II, VII
Adductor — see Muscle

Adhesive apparatus of tentacle club 229*, 273*, 316*, 346*, 458*
ring 98*, 121, 662*

Air chambers 79*, 80, 112*, etc., also F. p. 157, 218*

Alveoles, i. e. cavities in the periostracum or the anterior part of the
rostrum of Belemnoidea, if this is well preserved — 111*

Alveolite — the phragmocone (q. v. ) in the alveole
Anal complex — see Abdominal complex
— papilla 102. Typical form: 140*

Annulus 59*. This formation is assumed to be present also in the Belemnoi-
dea, in which it also passes along the base of the last septum. In the
Teuthoidea and Sepioidea, the areas of connection between shell and soft
body are usually markedly modified. 185*, 509*! 530*

857

Aorta anterior 91*, 136* ff.

— posterior ibid.

Apex 51*, 53*. Apex of the shell, also the primordial shell (q. v.)

Apical growth of arms 107, 235, 462*, 667*, 687*, Plate X

— line — see Axial thread.

Arm apparatus — a collective term for the "anlagen" of the arms and the
structures which develop from them
— pillar 82 (the bases of the arms of Nautilus, which project elbow-
like posteriorly. Figure 10 on p. 59)

Arms, differentiation of, in Nautilus 59*, 61. In the Dibranchiata, this
term applies usually to the arms without stalks (sessile), in contrast
to the stalked arms or tentacles (or T arms) (q. v.). There are dorsal
(D), dorsolateral (DL), ventrolateral (VL) and ventral (V) arms. Only
the ventral arms are homologous in the Octopoda and Decapoda

Artery, branchial 73*, 665*

— genital 79*, 91*

— lateral pallial (mantle) 664*

— median pallial (mantle) 105*, 124*

— posterior pallial (mantle) 103, 124*

Asymptote lines, median and lateral (cf. Hyperbolic lines) 112*, 137*, 146*,
160. They are ideal in some cases, distinct in others; they begin from
the apex and pass along the conotheca. The growth lines turn back
toward these lines. Their anterior parts belong to the lateral plates
of the proostracum and formed its lateral margin in the past. Cf.
growth lines (2)!

Auxiliary fins 389*

Axial thread or axial part of rostrum, also called apical line (l54) — the
continuation of the primordial rostrum to the apex of the developed
rostrum — often visible as a distinct line in split specimens (always
visible if the growing apex was pointed). However, there is also often
a ridgelike "axial thread" instead of an "apical line." This thread may
be transversely laminated or may differ from the other parts of the
rostrum in being less solid

Axis of arm ~ the firm, muscular central part of each arm, without the skin
and suckers. 115*, 662*, 726

Basal pad of suckers of Decapoda 662*. Also Pillar 384, 567

— platelets of teeth of radula 67*, 100*

Beak — see Jaws

Biting edges of jaws 83

— process — Apex of jaw 99*

Black substance 59*, 62, 770

Body, apex of, fleshy 22 6*, 234*, 286*, 295*

Branchial ganglia (also Osphradial ganglia or Pallial ganglia) 51*
gland 73, 526*, 664

Bristles, tufts of, in young Octopoda 668*. (Cf. also cutaneous spines)

Brush teeth of radula 67*

Buccal funnel of Decapoda — rudiments of buccal arms ("buccal corners,"
"buccal points," "buccal funnel supports") connected by a membrane
("buccal membrane," "buccal skin") ~ 58*, 99, 119*, 122, 179*, 537

858

Buccal ganglia 91*, 110*, 656*

- pockets 119*, 159, 179*

Bursa copulatrix of the Eusepiolinae — the female organ which corresponds
to the copulatory apparatus (q. v.) of the male -- 580, 617*, 636*, 647

Calcified necks 58,510*

Capitulum “■ thickening of the periostracum near the initial chamber in the
Sepiolidae ~ 487*, 492*, 495*, 496*, 498*, 501*, 520*

Carpal pad 3 1 6*

— part of tentacle club of Oegopsida 229*

Cartilaginous rods 658, 664*, 675
Cephalopodium, 53 (cf. Mantle sac)

~ retractors of, 51*, 59*, 60, 69

Cerebral ganglia 51*, 63*

Chamber, formation of, 88

Chambered snails, i, e. Teuthoidea, belonging to the Tetrabranchiata;

cf. Dibranchiata
Cheek, tubercle of,140, 181*

Chromatophores 91
Cirri 58*, 61

Clasping hooks 303, 331* — short hooks with thin stalks, articulated on a
revolving base

Claw character of suckers of Decapoda 130, 793, Plate XII
Closure, apparatus of, between mantle and funnel, completely developed in
the Decapoda, permitting gliding movements 123, cf. also 741
Club-shaped rostrum 111*, 112 (cf. Clavirostridae Abel)

Coelomic ducts 53, 72
Collar bond 123

Conchin cap of initial caecum at the siphuncle (better: "initial calotte") 512*

— (Conchiolin) 52

— necks 58

Cone 135, 137*, 160, 307*, 319*, 373*, 413*. The cone (still? ) bears septa-
like fillings in the Gonatidae and Chiroteuthidae, Figure 66 on p. 157*

- flag 137*, 146*, 225*, 247*

Conotheca 104,112*; cf. 510* (os + hy), 512* (6)

Constrictor — see Muscle

Copulation, mode of, 63, 76, 106, 126, 580, 677

Copulatory apparatus ~ a specially differentiated part of the hectocotylus
of the Eusepiolinae, which comes in contact with the female genitalia
580, 604*, 636*, 644*

Cornea 153, 167, 183*, 185*, 201, 227, 533, 567, Plate XIX, Figure 10

Corneal fold — cf. Primary lid

Crista of gladius 301

Ctenidia (typical gills of Mollusca) 52

Cutaneous mantle 52 (cf. Primary mantle, i. e. the typical mantle of the
Mollusca) 58* (a) 79*, 89*, 92

— spines of larvae of Octopoda 668

Descent, theory of (general concept) 1
Dibranchiata. See also Tetrabranchiata
— , type of, 5 8*

859

Distal part of tentacle club 120, 22 9-'

"Doratopsis" stage of Chiroteuthidae 38 7''!"

Dorsal shield 521 (layers: 531-0

"Earlobes" 114, 199-% 257-% 270-, 286-% 315, 433, 466-% 503-, 548-% 570-% 635-
Eccentricity of suckers of Decapoda 120, 121-% 662-

— of rostrum 1 1 1-% 484, 485, 500-

— of shell, growth of 4 74, 477-

Embryonic chamber (correctly: initial chamber) 5 6, 57*, 88*, 89*, 112*

— rostrum (Stolley) 111; (correctly: juvenile rostrum; cf. F)

— shell — cf. Primordial shell. Scar!

"Embryonic thread" — cf. — Primordial rostrum!

End part of tentacle club 120, 229*

"Entomopsis" stage 375

Envelope of head 62

Equilibrium, maintenance of (see 75*, 483*; Hydrostatics)

Esophageal ring 51* — basic type of the central nervous system of Mollusca
Esophagus 51*, 79*, 91*, 110*, 656*

Eye, chamber of (better: orbital cavity — q. v.), 159

— development of 96*

— pit 61, 59*, 6n01 (cf. Orbit)

Eyeball — the projecting main part of the eye of Nautilus and embryos
of Dibranchiata; the eyeball is later more or less deeply situated in
the eye pit (q. v. )

Fin, support of, 658

Fins, membranous part of, 270 (cf. 388* Ht)

— of Dibranchiata (insertion, articulation) 95*, 114*, 139*, 163*

Flag — the broadened lateral plate together with the cone flag (q. v.) in the
Metateuthoidea 14 6*, 225*

Fork, forked part of the shell of Sepiidae 519, 521*, 523*

— margin of 520 (free margin of conotheca of Sepiidae)

Funnel 83

— adductors (inner, outer, lateral, cf. Muscle)

— apparatus 83, 99

— bonds of Decapoda; adhesion and gliding device indifferent (l24*)

or specialized 205*, 317*, 341*, 384*, 425*, 460*, 469*, 558*, 593*,
754*, 765*

— cartilage — (cf. Funnel bond) 122

— corner, at posterior margin, 102, 306*, 663*, 727*

— , function of, 69, 84, 93, 100, 123

— glands 102 (type of Decapoda 181*, type of Octopoda 672)

— , halves of (instead of funnel lobes), 100. They may be fused or separate

— , lobes of, 100 (cf. Funnel halves of)

— pit, directing the water jet in the Decapoda! 414. Fold of funnel pit 428

— pit, edge of, 140 (cf. 431*)

— , pockets of, 68, 83

— retractors 68*, 100*

— septum 101

— valve 58*, 110* (absent in Octopoda and Cranchiidae)

860

Ganglion

— buccal 91''s no*, 656*

— cerebral 71*, 63*

— gastric 91*, 110*, 656*

— pedal 51* . Their homologue in the Cephalopoda is apparently at
least the proximal part of the so-called visceral ganglion (q.v. )

— stellate 103, 104* — the nerve center of the muscular mantle in

the Dibranchiata

— visceral 51*. These ganglia should be named pleurovisceral

(cf. pleural ganglia) in the Cephalopoda (79*, 91*, 110*)

Genital organ, external — cf. Genital processes

— pocket 105. It is wide open in the Oegopsida. Cf. 186

— processes 102, 105*, 211*, 593*, 63 7*

Gill, axis of 73

” , lamellae of, 52, 52 6*, 665*

— , ligaments of, 70*, 173*, 185*, 526*

— , retractors of, 181*, 186*, 593*, 684*, 722*

— root, pockets of, 102

Gills (ctenidia) 73, 185*, 526*, 664*, Plate IV
Gladius 136, 137*, 141*, 146*, 164!, 166, 169*, 225*

Glandular lines, particularly distinct in young Decapoda (Plate I, Figure 6,
Plate VIII, Figure 4) 563*, 592*

Growth lines 52, 1 1 1*, 136 (cf. F. p. 322)

The formation of growth lines is of great morphological interest
because it gives information on the ontogeny of the adult and also
on fossil shells, and completes our knowledge on the development
of the shell.

There are several types of growth lines.

1. Marginal lines, caused by the steplike enlargement of the surface
of the shell. They are usually, but not always, more or less
parallel to the shell margin, and advance concentrically (l44*,
146*). The primordial shell is the nucleus.

2. Longitudinal lines, caused by the displacement of certain parts
of the shell-secreting epithelium. These lines radiate centri-
fugally from the primordial shell. The "asymptote lines"
belong to the same type (93*, 144*, 146*). The two types of line
may appear as differences in coloration, or plastic, i. e. as thin
ridges, edges or grooves.

3. Laminated lines, at breaks, cuts and points of polishing; they are
(as l) orientated around the primordial shell as a nucleus and
result from an irregular secretion of shell substance — more
calcified or more organic, or of varying density or strength (53*).

4. Radial structure of Belemnites (ill*). This is a phenomenon
of crystallization, the aragonite prisms are situated perpendi-
cularly on the growth (thickening) layers.

5. Transverse lines (striation) of gradually thickening shells. Like
the longitudinal lines (2) these lines can radiate like a fan from
the primordial shell (171*); because of the growth of the shell
epithelium, the parts of the matrix are gradually pushed apart

by their own secretion (see also F. p. 268, Figure d, p. 363).

Many, but not all, sculptures of the shell are connected with the growth
lines of Mollusca. Cf. Argonauta 768*, 769*, etc.

861

Growth, of shell, see Growth lines;

Growth rate of Octopoda 699

Hand part of tentacle club 120, 229*

Hectocotylization 86, 274*, 434*, 599*, 604*

Hectocotylus 106

- , sac of, 732*, 757*, 785*

Hind intestine 51*, 79*, 91*, 110*, 656*

Hood 58, 62

Hooks, development of, 127, 131, 331*, 790, 793
Horny rings of suckers 98*, 121*, 662*, Plates XII— XIII
Hydrostatics of the phragmocone. The air chambers lower the specific
weight of the shell from about 2.7 to nearly 1. The buoyancy of the
phragmocone can be localized in different places. It is primarily
active at the posterior end (Orthoceratoidea 87, Belemnoidea llO),
and is displaced anteriorly as a result of the coiling (75*) over the
soft body and farther anteriorly (in the Sepioidea, 487*, especially in
the Sepiidae 522). The Belemnitidae attained an equilibrium partly
because of the compensatory effect of the rostrum (llO, 111*, 112),
while the Teuthoidea and Octopoda (136*, 137, 65 6*) dispensed with the
heavy hydrostatic and protective shell in favor of active organs. Also
the Sepiolidae and Idiosepiidae (502*). Spirula (see p. 517 and 862)
reverted secondarily to a buoyant posterior end, like the Orthoceratidae
and Belemnoidea

Hyperbole, zone of — lateral plate of proostracum 112* (zone with hyper-
bolic lines)

Hyperbolic lines (or stripes) 93*. These lines are more or less hyper-
bolic only in the Belemnites (112*); also called "asymptotes"
Hypostracum 52, 53*, 510

Initial caecum 107* (cf. 512*)

"initial calotte" (cf. Conchin cap)

Initial chamber 75*, 80
Ink sac 103

Inner layer of periostracum — see Periostracum
Inner lip 58*, 65*

Inner plate of dorsal shield 531*

Inner plate of jaw 66*, 99*

Intercalated teeth of radula 67
Iridocytes 91
Iris flap 115;

Iris fold 96

Jaws 66*, 99*; consisting of keratin + chitin (according to Neri, 1896)
Juvenile rostrum ("embryonic rostrum" of Stolley) 111*

Kalmar, belonging to the Teuthoidea 135

Lanceola 157, 233

Larval forms — cf. young (juvenile) forms

862

Lateral edges 493^^ 495^^ 497*, 498*, 500*, 501*, 520*, 521*

— lines (ridges) on mantle sac of Sepiidae 538*, 553, 557*, 562* and

Polypodoidea 675*, 714*, 721*

— margins 534, 549*

— plates of proostracum 112*, 137;

— — of radula 66 (parts without teeth of the basal plate)

— pockets of funnel pit 457*

-- — of mantle cavity 418, 419* (x), 469* (y), 470, (719*)

— (side) wings 493 (cf. Lateral edges)

~ swellings or incrassations 492*

— teeth of jaws 419
Lens 96

Lid apparatus of Octopoda 659*, 660*

Ligament of attachment 119*, 177*, 179*, 537*

Living chamber 59*, cf. 75*, 94*, cf. 107*, this includes also the cavity of
the dorsal plate in the Dibranchiata
Lobed lines (better: suture lines) 59*, 60
Lower jaw 58

lid 96*, 659*, 660*

Luminous bacteria 231

- glands 517*, 574*, 581*, 632*, 636*, 639!

~ organs 230, 368*, 513; cf. 862!, Plate III

Main plate of radula 66 (cf. Lateral plates)

Mantle bond (incorrectly: mantle cartilage) 122, 205*, 317*, 418*, 530*, 593*,
727*, 754*

— cartilage — see Mantle bond
cavity 51*, 52

- ,roof of,51*, 52, cf. also 79*, 85*, 101*!

— ( ~ ) septum, median mantle (pallial) septum 91*, 102*, 183*, 419*

-- — , situs of. Plates I—V, VII, X; many figures in the text

— groove 51*, 52, 69, 103

— (primary, cutaneous mantle; cf. also Muscular mantle) 51*, 58*, 79*,

89*, 94*!

— sac 53, 59. One main region of the body of Mollusca divided from

the cephalopodium by connecting points 28 and 10 in Figure 5 on
p. 51 (also in Figures 9 on p. 58, 19 on p. 79, 25 on p. 91 ). The
vegetative organs can thus be separated from the animal organs by
severing the anterior aorta, vena cava, visceral nerves, fore intestine
and shell muscles (or retractors of cephalopodium)

— “ , dorsal (correctly: dorsal mantle cavity) 475* (ud), 658, 664*,

Plate X, Figure 6

Margin of muscular mantle 181*, 469*, 754**. Distinctly defined in strong
swimmers

Marginal edges of funnel pit 331 (cf. funnel, edges of)

— growth 52. Cf. Growth lines

— plates (or platelets) of radula 67*, Plates XIV— XVI

— ring of suckers 98*, 662*

Median mantle (pallial) septum — cf. Mantle (cavity) septum
- plate 112*, 137*

863

Membrane (or interbrachial membrane) 97, 116, 548'’% 576*, 699*, 700
(inner: 3 68*)

Metadecapus 128
Metamerism 72
Metateuthis 153
M eto rthoc e r a s 72
Midintestine 51*

Mouth (buccal) area 119*, 179*, 1 80*, 240*, 439*, 45 1*, 462*, 661*, 774*,

Plates III and VI

— - arms 58* (int. ) 62, 79*, 91, 99, 119*

— — cone 122, 159

Muscle (MuscuIus)

— adductor infundibuli internus 1 externally visible in the Oegop-

~ — — externus j sida 317*, 418*, 465*

— - - lateralis 537, 558*, 569*, 574*, 593*, 664*

— — pallii anterior (anterior mantle adductor) 664*

— — — medianus (median mantle adductor) 504, 570*, 574*,

581*, 582, 647*, 656*, 663*, 664*

— — — posterior (posterior mantle adductor) 664*

— constrictor pallii (mantle constrictor) 623

— rectus abdominis 103, 105*, 181*

— retractor branchialis superficialis — see Gill retractor

— — infundibuli — see Funnel retractors
Muscular mantle 93, 94*, 107* (primary insertion)

Nacreous layer (cf. Hypostracum) 52, 53*, 510*

Neck bond (incorrectly: neck cartilage), also "collar" 123!

— cartilage — see Neck bond

— , edges or folds of, 136, 140, 315*, 417*, 465*

— folds (better: edges of neck, q. v. )

— , ligament of, 578 (cf. 635*)

Nerve

— pallial (mantle) 104*

— stellate cf. pallial

— visceral (homologous to the pleurovisceral cord (51*) of the

Cephalopoda)

Nidamental glands 70*, 73, 105*, 106*, 124*, 125

Oegopsida, larvae of (typical, youngest) 234, 813*, 815* (cf. young (juvenile)
forms)

Olfactory organ 61, 107* (olfactory papilla, tubercle) 110*, 112*, 136*, 137*,
150*, 153*, 171*, 307*, 319*, 487*, 495*, 498*

Orbit (see Eye pit)

Orbital chamber, formed by the closure of the corneal fold (q. v. )

— pore 167, 185*, 502*, 527*, 567, 570*, (partly also "eye pore")
Osphradia 72
Ostracum 52, 53, 510*

Outer lip 58*, 65*

— margins — continuations of the folds of the membrane on the lateral

outer edges of the ventral arms of the Decapoda. Cf. Figure 3 7a
on p. 110

— plates (of jaws) 66*, 99
Oviduct, gland of, 71, 104, 123

864

Palps 9p:% 97, 116-

Parabolic lines 93- (Pv), 112- (7)

— zone (cf. Median plate) 137-
Pear-shaped vesicle 71
Penis 102, 728, 785-
Pericardial funnel 71,4 60-
Periostracum 53-, 510-

— of sheath 112-

Phragmocone. Chambered part of the shell of Cephalopoda (F., p. 14),

consisting of septa and septal necks and enveloped by the substance of
the annulus. A specially differentiated part of the hypostracum
Pillar, substance of 510-

Pleurovisceral cord 51- (cf. Visceral nerve)

Poison gland 91- (usually called salivary gland)

Praedecapus 117 — an ideal or hypothetical ancestral form of the
Belemnoidea (p. 790)

Praesepioides — an ideal or hypothetical form of the Praesepioidea
(791 ) and at the same time of the Praeteuthoidea (790)

Prehensile arms 62, 99. These arms are divided into stalked arms
(tentacles) and arms without stalks

— — , movement of, 1 19, 678

— hooks (cf. Clasping hooks) 303, 33 1-. Long, with thick stalks,

solidly attached

Primary lid 96''% 107-, (44 6-)» 509-

mantle 92, 107- (cf. Cutaneous mantle)
mouth — cf. Mouth
pupil 61, 96-

Primordial rostrum — the first pointed-conical deposition on the outer side
of the initial chamber in the Belemnites

— shell 52, 57, 107'^% The first shell rudiment of the Conchifera

which grows later concentrically
Prismatic layer (cf. Ostracum) 52)

Proostracum 91-, 93-, 94% 107% 110% 112% (l36-), 160, 161- (also: dorsal plate)
Prosiphuncle 80. 107-

Protective margin, supports of, 115'% 116''' (accessory 118)

— margins of arms of Decapoda 116'’' (false margins in Argonau-

tidae 726)

Protoctopus 656
Protodecapus 109'''

Protodibranchus 91'''

Protorthoceras 79'^'

P r o t o s e p i o i d e s 487
Prototeuthis 13 6'''

Pseudocornea 660'!'

Pupil 96'!' (opening of iris), 174'!', 533'!% 567'!% 682'!'

"Pyrgopsis" stage 407'!% 408'!'

Radula 67'!% 100'!% Plates XIV, XVI. Systematically important. Cf. Sepioidea
(473), Enoploteuthidae (2 61 ), Ctenoglossa (679 ), O c y t h o e (755), etc.

The radula consists of median, paramedian, lateral, outer (external)
and marginal teeth. There is also an intercalated and another para-
median row in Nautilus

865

Rasping teeth of radula 67*

Regulation of axis of shell 484, 499

Renal papillae 68^^ 70=^ 71, 85, 101*, 105*, 124*, 125*, 140*, 183*, 259*, 514*,
562*, 593*, 664*

Respiratory naovements — they take place like the swimming movements,
but are weaker; cf. funnel, function of
Retroflexion of abdominal complex 69

Rhachis — modified median plate of the Meso- and Metateuthoidea 145,
146*, 225*

R hy n c h o t e u t h i o n 420
"Rhy nchoteuthi s" stage 427*

Rostrum 110*, 112*, 136*, 137*, 150*, 153*, 171*, 307*, 319*, 487*, 495*, 498*

"Scar" of the shell of Na u t i lu s 57
Secondary lid 96*

Sensory arm 59*, 61

Sensory arms (cf. arms of Nautilus) 62

Septa — cf. Shell, septa of

Sexual dimorphism 65*, 72, 86, 106, 200, 552*

Shell epithelium 52, 107* (primary and secondary)

Shell folds 51*, 107*, 112*

Shell, growth of,88, 93*, 124*, 144* (cf. Growth lines)

Shell muscle (adductor) 51*, see Cephalopodium, retractor of
Shell sac — envelope of primary and secondary epithelium which surrounds
the shell 107*, 478*

Shell, septa of 57, 107

Shell, situs of, 104*, 124*, 141*, 169*, 173*, 530*

Shell, thickening of, 52

Sinus of eyelid 115, 130*, 136*, 449*

Siphonal or septal necks 57, 107* (cf. Calcified, Conchin necks)

Siphuncle 107

Siphuncle, structure of, 475*, 510*, 512

Skin, warts on, of Sepiolidae and Sepiidae 549, of Octopoda 668, 705
Snout 51*— typical formation in the Mollusca, often wrongly considered as
"head"

Spadix 62

Spermatophore gland — the ectodermal distal part of the male gonoduct,
corresponding grosso modo to the oviduct gland (cf. Naef, 1913)
Spermatophore pocket — differentiated, nonglandular, terminal part of
spermatophore gland

Spermatophore reservoir in Argonautidae 746*, 760*, 785*, 786* (accessory
reservoir 746* = secondary reservoir 732*)

Spirulisepia 520*, 522*, 524
Stalk margin of tentacle 158, 415*

Stalk part of tentacle club 229*

Statocysts 51*, 79*, 91*

Stretching (secondary) of mantle sac 436

Stretching (secondary) of phragmocone 494, 496!, 498*, 499, 501*, 510
Sucker, stalk of 98, differentiated into a stalk and a basal pad in the
Decapoda 662

5110/2

866

Suckers 98*, 121*, 662*!, Plates XII-XIII

Supporting bars 510*, a general character of the chambered shell
Surrounding of the gladius by the muscular mantle 156*, 157*, 161, 164, 171
Surrounding of the shell 51*, 85*, In the Dibranchiata, the shell fold covers
the primary shell epithelium early during embryonic development
Suture lines ~ lines of connection of the septa with the conotheca at the

anterior margin of the supporting bars. The supporting bars make the
suture appear as a broad double line if the conotheca is delicate and
translucent (as in S p i r u 1 a) or if it is detached. Strictly speaking,
suture and supporting bars are situated on the substance of the annulus,
not directly on the conotheca. This substance forms a fine lamella
between the phragmocone and the other parts of the shell
Swelling, swollen part 519, 521, 523 (cf. 521* )

Swimming margins 115*, 118 — probably a primary character of all

Dibranchiata. The glandular lines (q. v.) always are the "anlagen"
of the swimming margins
Swimming movement — see Funnel, function of

Tentacle pockets 120, 164, 177*, 537*, 566*

Tentacle (T arm) 115, 790 (differentiation 120, in the Oegopsida 229*,
cf. 204*, 487*)

Tentacles, function of,119 (cf. Prehensile arms, movement of)

Terminal organ (cf. S p i r u 1 a, p. 863 ) 507*, 509*, 516*, 565*, Sp,
Tetrabranchiata, character of, 58* (in contrast to the type of Dibranchiata)
Torsion of gills 73
Tubercle of cheek 140, 181*

Upper jaw 58
Upper lid 96

Vein

abdominal (correctly: posterior mantle vein q. v. )

— branchial 70*, (85*, 101*)

— cava 79*, 91* (not cephalic)

cephalic — vessels which reach the vena cava near the statocyst

— lateral mantle (pallii lateralis) 103*, 104*, 105*

~ posterior mantle (pallii posterior) 103, 105*

Venous appendages — clustered excretory evaginations of the main veins in
the kidneys 68*, 70*, 103, 105*

Ventral curvature of phragmocone 476, 477*, 478*, 480*, 483*, 485*, 487*,
491*, etc.

Ventral margin of conotheca 91*, 93, 136*, 137*, 485*, 491*

Ventral process 500*, 501*, 520*, 521*, 522*

Ventral ridge of the shell of Sepioidea 482*, 493*, 495*, 497*, 498*

Ventral wall of shell of Sepioidea 495*, 500*, 501*, 520* (not only of the
phragmocone)

Water pores of Argonautidae 739*, 741*, 752*, 754*. This term is wrongly
used also for the buccal pockets (q. v. )

867

Window 159, 210*, 227, 319-, 415=^

Wings of shell of Sepiidae 530, 556*, 559*, 561*, 563*

Young (juvenile) forms (also larvae) 107, 163, 188*, 223*, 253*, 275*, 290*,
305*, 336*, 354*, 365*, 372*, 387*, 393*, 404*, 427*, 477*, 608*, 668*,
736*, 750*, 765*, 766*, 813*, 815*

b. SYSTEMATIC INDEX

The following list contains the systematic names used in this book. The
valid fossil groups are indicated by a t , the bold type indicates units des-
cribed in detail or new entities. Invalid or fictitious names are given in
quotation marks.

A b r a 1 i a 48, 279, 280*, 282*, 284*, 809

Abraliopsi s 48, 235*, 239*, 285, 286, 288*, 289*, 291*, 294*, 809
"A canthoteuthi s" Wagner, 1832 (cf. pp. 1 12*, 147) is a still indispensable
fictitious generic name for fossils of Belemnoidea (bodies, proostraca and
phragmocones) the systematic position of which is still uncertain because
the decisive rostrum has not been found. The presence of a tongue-shaped
proostracum suggests the Belemnitidae, but the alleged absence of a pro-
ostracum in the Belemnoteuthidae is not likely, and this family must
therefore also be considered. Such uncertain forms of "Acantho -
teuthis" have provoked the discussion** on the number of arms in the
"Belemnites, " which was not in doubt at all. All Belemnoidea belong to
the Decapoda according to the shell and structure of the hooks (cf. pp. 12 7
and 662).

The position in the Belemnitidae is uncertain for the Lias forms of
Acanthoteuthis and also for those from the Malm. I proved in 1921
that these specimens had 10 uniform arms with hooks. The Belemnites
of Crick and his Acanthoteuthis ("B e 1 e m n ot e u th i s") m o nt e -
fiorei are identical. These forms had 6 (6—7) arms with hooks, while
I observed 8 (8— 9 ) arms with hooks in Acanthoteuthis speciosa.
Suppression of the formation of hooks is most probable on the V arms.

It is not justified to assume the absence of arms (cf. also p. 790), The
absence of a rostrum in Acanthoteuthis cannot be explained, as
Prell (l. c. p. 305) thinks, by an intact detachment from the initial shell,
but the initial shell disappears at the same time. The best preserved
phragm.ocones of Acanthoteuthis have always lost the initial parts;
the rest of the periostracum is also lost.

Cf. Crick, 1902 and 1907; Stromer, 1912; Abel, 1916; Prell, 1922. To my surprise, Prell, who had no
observations of his own, used my verbal communications to explain his views on the fossil Cephalopoda,
without mentioning this and without reference to my recent publications.

868

Acanthosepia 546
tAcroteuthis HP'S 808

"A c r o t e u th i s/* "A c r u r o t e u t h i s, " Acrololigo for Alloteuthis
209

tActinocamax 808
Alloposus 50,727^^731,811

Alloteuthis 47, 169"% 171"% 176"% 190"% 207*, 210*, 211*, 213*, 216*, 217*,
218*, 220*, 222*, 223*, 809
tAmblybelus 808; F. p. 279
Amphineura 52

Amphitretidae, Amphitretus 50, 673, 677, 795*, 811
A n c i s t r o c h i r u s 48,267,809
Ancistroteuthis 48,326, 316*, 331*, 333*, 809
Apteri 50, 669

Architeuthidae, A r c h i t e u this 48, 237, 795*, 809

Argonauta 50, 727*, 762, 765*, 767*, 769*, 771*, 773*, 775*, 776*, 785*,

787*, 811, Plates X, XI
Argonautidae 50, 673, 725, 727*, 795*, 811
Argonautinae 732*, 811
t Ascoceratidae cf. F. p. 95
tAtractites 46, 789, 795*, 808

tAulacoceras, Aulacoceratidae 46, 789, 795*, 808
t A u 1 a c o t e u t h i s 808

Bathothauma 49,397,810

Bathyteuthidae, Bathyteuthis 48, 251, 795*, 809

t Bayanoteuthinae, B a y a n o t e u t h i s 47,818; incorrectly as Neobelemnitidae
in the genealogical tree (795*)
t B e 1 e m n i t e 1 1 a 795*, 808
tBelemnitidae 47, 795, 808*; cf. F. p. 224;
tB e 1 e m n o c o nu s 808

tBelemnoidea 46, 117, 127, 151, 790, 795*, 808
t Belemnopsinae, Belemnopsis 795*, 808
t B e 1 e m n o s e 1 1 a 481,482,810

tBelemnosepia 47,144*, 148 (cf. F. pp. 109, 169)

1 Belemnosepiidae 47, 147, 143
t Belenanosidae 493, 795*, 810
iBelemnosis 49,492*, 810

tBelemnoteuthidae, B e 1 e m n ot eu th i s 47, 795*, 808 (cf. F. p. 276)

t Belopeltidae, Belopeltis 4 7, 147, 148, 795*, 808

tBeloptera 49,494*, 810

tB e 1 o p t e r e 1 1 a 147,810

tBelopteridae 49, 147, 493, 795*, 810

tBelopteridiuna 14 7,810

tBelopterina 49, 493*, 810

tBelos epia 49, 477*, 521*, 523*, 524*, 525, 810

tBelosepiella, Belosepiellidae 147, 795*, 810

tBelosepiinae 526,810

tBeloteuthidae 47, 145, 147, 792, 795, 809

Benthoteuthidae, Benthoteuthis cf. Bathyteuthis

869

Berryteuthis 244,809

Bolitaenidae, Bolitaena 50, 673, 677, 680, 79 5'-', 811
Brachioteuthidae 48, 370, 795'!', 810
Brachioteuthis 48, 370, 373'!', 810
tBrachybelus 808

"Calais" 50,147,i. e. Palaeoctopus
t Calli conite s 4 6, 134, 791-792, 795'!', 808

Calliteuthis 48, 344'!', 347=:c^ 349-c^ 35o>:^^ 352, 355'!', 357=:^, 359=:<^ 309.

Deegener (1922, Z. Anz. p. 220) and Pfeffer consider Loligo mene-
ghini as a Calliteuthis. This form cannot be identified with
certainty. Homologous stages of Calliteuthis are quite different.

My view (p. 339 and 352) is based on a thorough knowledge of both series,
t Celaenidae 47, 795'!', 809
t C e 1 a e n o t e u t h i s 809
Cephalopoda (General) 46, 77, 79'!'

C h a u n o t e u th i s 48, 322, 324'!', 809
Chirosoma 810
Chiroteuthidae 48, 376, 795'!', 810
Chiroteuthinae 378, 810

Chiroteuthis 48,381, 383'!', 334'!', 337^:=^ 333^ 310
Chiroteuthoides 393'!', 810
Chirothauma 48, 378, 810

Chondrophora Keferstein, 1 866 = Teuthoidea and Sepiolidae
C h u n i o t e u t h i s 50,672,811
"Cirrata" Grimpe 50. 148, 669 = Cirroteuthoidea
Cirroctopus n. g. 675,811

Cirroteuthidae, Cirroteuthis 50, 672 , 795'!', 811
Cirroteuthoidea 46, 50, 148, 670, 674, 795'% 811
Cirrothauma 674,811
Cistopus 50,692,811
"Clavirostridae" 111'!', cf. Conirostridae
^ Coeloteuthinae, Coeloteuthis 795'!', 808

"Conirostridae" 111'!'. Belemnites with a conical juvenile rostrum, in contrast
to the Clavirostridae, in which the rostrum is club-shaped. This dis-
dinction is more or less artificial; cf. Abel, 1916 and Naef, 1922
i Conobelus 808

iConoteuthis 795'!', 808; belongs perhaps to Diploconus (cf. F. p. 279)
Corynomma 49,397,810
Cranchia 49, 397, 810

Cranchiidae 49, 392, 395'% 396'% 404'% 795'% 810
Cranchiinae 397, 810
t "Crassicarinati" Quenstedt 145
C r y s t a 1 1 o t e u th i s 49,397,810
Ctenoglossa 50, 673, 679, 795, 811

Ctenopteryx 48, 233,252, 253'% 254, 256'% 257'% 259'% 809
Cucioteuthis 48,234,809
Cycloteuthinae, Cycloteuthis 312, 809
t Cylindroteuthinae, Cylindroteuthis 795'!', 808
t Cyrtoceras 75'!'

5110/2

870

t Dacty loteuthi s 808

Decapoda 46, 109, 111=:% 112'% 115=:% 116=:% 119=:% 121=:% 125=:% 128=:% 130=:% 662=:%
795=:% 808

D e s m o t e u t h i s 49,397,810

Dibranchiata 46, 58=:% 90, 91*, 93=:% 94*% 95^% 95^% 99=:% loi*, 105=:% io7=:% 789, 808
t Dicoelite s 808
t D i c ty o c on it e s 46, 795=:^, 808
t Diploconidae, Diploconus 795=:^, 808
“D o r a t op s i s" 378,387=:*

Doratosepia 546
Doryteuthis 47,191,193,809
Dosidicus 49,428,809
tDuvalia, Duvaliinae 795=:*, 808

Eledone 50, 659=% 667=-% 676=:*, 716, 717=:*, 719, 721=% 723=:*, 811

Eledonella 50,673,811

"Eledonidae” 716

Eledoninae 692

Endoceratidae 79

Enoploteuthidae 48, 261, 795=:*, 809

E noplot euthinae 265, 267, 809

E noploteuthi s 48,267,809

Enoptroteuthis 393=:*, 810

Entomopsis 375

t Eubelemnoidea 791

Eucleoteuthi s 49,428,809

"Eumalakia" 51; correctly; Conchifera Gegenbaur

Euprymna 49,573,810

E u s ep ia 545

Eusepiola 603

Eusepiolinae 579

"Euzygaena" 408=:*

Froeckenia 50, 672, 811

Gal iteuthi s 49, 393=«% 397, 398, 399=% 401=% 404, 810
tGastrobelus 808

t Geoteuthidae, Geoteuthis 143, 147, 795=-'*, 808

t Glyphiteuthis 47,809

Gonatidae 48, 242, 795=% 809

Gonatopsis 250,809

Gonatus 48,244, 246=% 247=:*, 249=% 809

"Gonoteuthis" = Actinocamax, q. v.

Graneledone 148,716,811
"Grimalditeuthidae" 48. Subfamily
Grimalditeuthinae, Grimalditeuthis 378, 393=:*, 810
tGyroceras 75=:*

Hastatidae Stolley = Belemnopsinae
Hastites 808
t Hastitinae 795=% 808
Heliocranchia 810
Hemisepius 49,543,810
H e n s e n i o t eu t hi s 810
Heteroglossa 50, 673, 680, 795=:*, 808

871

Heteroteuthinae 572, 574, 576, 810

Heteroteuthis 49, 595, 599-S 573, 810, Plates VIII, XIX

t Hib elites 111- (Fig. 38c), 112"% 808

Histiopsis 360-,351,809

Histioteuthidae 48, 343, 795-, 809

Histioteuthis 48, 363, 365"% 366"% 368"% 809

"Histiochromius" 250

t H o m a 1 o t e u t h i s 808

Hyaloteuthis 49,428,809

H y m e n o t e u t h i s 50,672,811

Idiosepiidae 49, 147, 503, 795"% 810

Idiosepius 49, 502"% 503"% 81 0. Here probably also belongs: Tilesius,
1814, Plate 88, Figs. 32— 34; cf. Krusenstern
"l d i o t e u t h i s" 48 to M a s t i g o t e u t h i si
Inioteuthis 573,603

II lex 49, 226"% 240"% 413"% 429, 431"% 433"% 435"% 437"% 809
Illicinae 428, 809

"incirrata" 50, 148, 669 = Polypodoidea
Iridoteuthis 49, 576"% 573, 810

Japetella 811

Joubiniteuthidae, J oubiniteuthis 147,237,810

L a e t m o t e u t h i s 50,672,811

L a m p a d i o t e u t h i s 48,266,809

Leachia 49, 397, 406, 407"% 810

Lepidoteuthidae, Lepidoteuthis 48, 795"% 809

t Leptoteuthidae, Leptoteuthis 47, 111, 143"% 147, 148, 795"% 808

L e u c o c r a n c h i a 810

Liguriella 49,397,810

Liocranchia 49, 402, 404 -, 810

"Lioglossa" = Cirroteuthoidea 50, 148, 669, 670

^ Lioteuthidae, L i o t e u t h i s 795"% 808

t L i s t r o t e u t h i s cf. F. p. 193

Loliginei Steenstr. = Loliginidae 168

Loliginidae 47, 149, 168, 795"% 809

"Loliginites” Queenstedt 47, 142,808. F. p. 102, 130
Loligo 47, 169^% 170"% 173"% 174''% 175*, 179*, 180*, 181*, 183*, 184*, 185*,
186*, 187*, 188*, 193, 195*, 199*, 200*, 201*, 204*, 205*, 809, Plates I-TI
"Loligopsis" 410*

Lolliguncula 191,809

Loliolus, Loliolinae 47, 191, 809

Lophosepia 546

Ly c o t e u t h i n a e 266,809

Lycoteuthis 48, 232*, 261,2 62*, 264*, 809

Mastigoteuthinae, Mastigoteuthis 48, 378, 393*, 810
M e g a 1 o c r a n c h i a 49,397,810
t M e g a t e u t h i s 808
Melanoteuthis 50,672,811
M e 1 e a g r o t e u t h i s 48,351,809
t M e s o h i b i 1 i t e s 808
+ Mesoteuthoidea 47, 145, 795*, 809

872

Metasepia 49,546,810
"Metasepioidea" 147
Metateuthoidea 4 7, 149, 795''% 809

— myopsida 1 65, 795''% 809; cf. Myopsida

— oegopsida 224, 795, 809; cf. Oegopsida
Moroteuthis 48,311,809

"Moschites" 50, 148, 669 (Note)

"M u c r o t e u t h i s" 111", cf. F.

Myopsida (better: Metateuthoidea myopsida) 47, 150, 151, 165, 795''% 809

t Nannobelus 808
Nautiloidea 75'-'

Nautilus 55'% 57'% 59'% 63'% 65'% 67'% 68'% 70'% 73'%

Nectoteuthis 49, 573, 577'% 810
Nematolampas 48,266,809

"Neobelemnitidae" 795''' (incorrect for Bayanoteuthinae)

+ N e o h i b o 1 i t e s 808

Neoteuthis, Neoteuthidae 48, 237, 795''% 809
Nototodarus 49,428,809

Octopoda 50, 665, 656'% 661'% 663'% 665'% 794, 795'% 811, Plates X, XIX

Octopodidae 50, 681, 795'!", 811

Octopodinae 692, 811

Octopodoteuthidae 48, 334, 795''% 809

Octopodoteuthis 48, 334, 33 7'^% 341'% 342'!'% 809

O c t o p o d o t e u t h o p s i s 334

Octopus 50, 663"% 665'% 666'% 668'% 679'% 683'% 685'% 687'% 689'% 693, 694'%
699'% 701'% 703'% 705'% 706'% 708'% 711'% 712'% 714'% 727'% 811
Ocythoe 50, 749, 726'% 727'"-751'% 753'% 754'% 757'% 758'% 760'% 811
tOdontobelus 808

Oegopsida (Metateuthoidea oegopsida) 48, 149, 150, 161, 224, 225'% 229'% 231"%
333, 793, 795'% 809, 813'% 815'"

Ommatrostrephes 49, 415"% 417'!% 419=% 445^ 447"% 449'", 451"% 452'", 809
Ommatostrephidae 49, 233"% 411, 421"% 423'!% 425"% 427"% 795'% 809, Plates V,
VI, XIX

Ommatostrephinae 428, 809

Onychoteuthidae 48, 301, 305'% 307'% 309'% 310'% 795'% 809
Ony choteuthis 48, 311'% 313, 315'% 317, 319'% 809
Onychia, Onychiinae, cf . O n y k i a, Onykiinae
"Ony chii" 334, 793
O n y k i a, Onykiinae 311, 809

Opisthoteuthidae, Opi s thoteuthi s 50, 147, 672, 795"% 811
t Orthoceras 79'% 85"% 89"% 789
tOxyteuthis 47,111,808

t Pachyteuthis 808
t Palaeoctopoda 50, 672, 795"% 811

t Palaeoctopodidae, Palaeoctopus 50. 148, 671"% 672, 811
Palaeololiginidae 145, 147, 795"% 809
t Palaeololigo 146'% 792, 809

873

t Parabelopelti s 808
t Pa r ahibolite s 808

t P a r a p 1 e s i o t eu th i s 147, 808 (cf. 137'i')

Parasepia 545
Paroctopus n. g. 692

t Passaloteuthinae, Pa s s aloteuthi s 795^,808
P h a s m a t op s i s 49,697,810
Phasmatoteuthion (-teuthis) 49, 697, 810
Phragmoteuthidae, Phragmoteuthis 46, 789, 792, 795'**, 808
*’P hy 1 1 o t e u t h i s" 47 (cf. F. p. 149)

Pinnoctopus 50,669,692,811
Platysepia 546

1 Plesioteuthidae (Plesioteuthis 47, 143 ),147, 795*, 808

tPleurobelus 808

Polypodoidea 50, 670, 672, 675, 795*, 811

"Polypodidae, " "Polypus" 50, Note 696

t Praebelemnoidea 791

t Praesepioidea 791

Promachoteuthidae, Promachoteuthis 48, 149, 795*, 810
"Protosepioidea" 147

t Prototeuthoidea 47, 142, 791, 792, 793, 795, 808
^ Pseudobelus 808
t P s e u d o d u V al i a 808
1 P s e u d oh a s t i te s 808
t Pseudosepia 794,810

Psychroteuthidae, Psychroteuthis 227, 795*, 809
Pteroti 50, 669

Pterygioteuthis 48, 157*, 266, 809
"Ptilo teuthis" 47, 809
Pyrgopsis 48, 393*, 407*, 408*, 410*, 810
Pyroteuthinae 265, 809

Pyroteuthis 48, 267,269, 270*, 271*, 273*, 274*, 275*, 277*, 280, 809

t Rhabdobelus 808
t Rhaphibelus 808
t Rhopaloteuthi s 808

t "Rhynchoteuthis" 420, 423*, 425*, 427* (-t e u t h io n)

"R o n d e 1 e t i a’’ 49; correctly:

Rondeletiola 147, 629, 631*, 632*, 635*, 637*, 573, 810, Plate XIX
Rossia 49, 566*, 573 (To 575* Sasaki (l920) found similar rudiments of
luminous glands also in Rossia b i p a pi 1 1 a t a, and stated: "... a
peculiar papilliform organ of unknown function developed on either side
of the rectum") 589, 591*, 593*, 810
Rossiinae 572, 573, 574, 810

t Salpingoteuthis 808
Sandalops 810

"Scaeurgus" 50, 682*, 710, 711*, 712*, 714*

Semirossia 49,573,810

Sepia 49, 338*, 477*, 521*, 522*, 524*, 544,526*, 529*, 531*, 533*, 535*, 537*,
539*, 541*, 549*, 551*, 553*, 554*, 557*, 559*, 561*, 563*, 810, Plates VI, VII

874

Sepiadariinae 572, 573, 810
Sepiadarium 49,573,810
Sepiella 49,543,810

Sepietta 49, 565*, 569*, 570*, 573*, 583*, 631*, 640, 643*, 644*, 647*, 648*,
650*, 653*, 810, Plate XIX
Sepiidae 49, 519, 520*, 521*, 795*, 810
Sepiinae 527, 810

Sepioidea 49, 117, 150, 473, 478*, 483*, 485*, 487*, 489*, 491*, 794, 795*, 810
Sepiola 49, 573, 581*, 582*, 601*, 604*, 607*, 609*, 611*, 613*, 614*, 617*,
619*, 620*, 623*, 625*, 627*, 628, 810, Plate XIX
Sepiola-like Sepiolinae 584

Sepiolidae 49, 502*, 564, 565*, 567*, 568*, 573*, 575*, 795*, 810
Sepiolina 49, 573*, 810
Sepiolinae 572, 573, 578, 810
Sepiloidea 49, 573, 810

Sepioteuthinae, Sepioteuthis 47, 191, 196, 809

Spirula 49, 475*, 477*, 505, 507*, 509*, 510*, 512*, 514*, 516*, 810, ff.
Spirulidae 49, 505, 795*, 810

tSpirulirostra 49, 477*, 496*, 497*, 498*, 499*, 810
1 Spi ru 1 i r o s t r e 1 1 a 147,810

t Spirulirostridae, S p i r u 1 a r o s t r i d iu m 49, 496, 795*, 810
ISpirulirostrina 49,501*, 810
t Spirulirostridae 147, 795*, 810
S t a u r o t e u t h i s 50,675,811
"St ephanot euthi s" 595

Sthenoteuthis 49,455, 557*, 558*, 661*, 662*, 809

"S t i g m a t o t e u t h i s" 354

Stoloteuthis 49, 573*, 576*, 810

t Sty loteuthis 808

t S t y r a c o t e u t h i s 47,808

S y m p 1 e c t o t e u th i s 49,428,809

Taonidium 49,393,810

Taoniinae, T a o n i u s 49,397,810

"Teleoteuthinae" (cf. Onykiinae) 311

"T e 1 e o t e u t h i s" 48, 150*, 31 1 (correctly: Onykia)

"Tenuicarinati" Quenstedt 145
Tetrabranchiata 46, 55, 58*, 74, 76
T e t r o ny ch o t e u t h i s 48,312,809
"Teudopsis" 147 (cf. F. pp. 130, 136, 190)

"Teuthis," "Teuthos" 207,208

Teuthoidea 47, 97, 135, 137*, 138, 141*, 142*, 795, 808

"Teuthopsis" 47, 146*, 147

"Teuthos" cf. Teuthis

Teuthowenia 49,397,810

Thaum atolampa s 48 (cf. Lycoteuthis)

Thelidioteuthis 48, 296, 299*, 300*, 809
Thysanoteuthidae 49,463, 795*, 809
Thysanoteuthis 49,463, 465*, 466*, 4 69*, 809
T o d a r o p s i s 49, 438, 439*, 441*, 443*, 809, Plate XIX

875

Toxeuma 49,397,810

"T r a c h e 1 o t eu t h i s" (correctly: B r a c h i o t e u t h i s)
Trachyglossa 50,148,669

t Trachyteuthidae, T rachyteuthis 47, 145, 792, 795*, 809
Tremoctopodinae 730, 811

Tr emoctopus 50, 727*, 730, 734, 736*, 739*, 741*, 742*, 811, Plate X

Vampyroteuthidae, Vampyroteuthis 50, 148, 669, 672, 795*, 811
tVasseuria, Vasseuriidae 49, 147, 795, 808
Velodona 50, 691,811

Veranya 48 (correctly: O c t o p o d o t e ut h i s)

V e r r i 1 1 i t e u t h i s 810

V i t r e 1 e d o n e 1 1 a 148,680,811

atas ella 811

atasenia 285 (to A b r a 1 i o p s i s)

^ Xiphoteuthidae, Xiphoteuthis 47, 795*, 808

A general orientation on "The System of the Cephalopoda Dibranchiata
and their Mediterranean Species" with corrections and addenda to the present
part of the monograph can be found under the above name in "Mitteilungen
der Zoologischen Station zu Neapel" Vol. 22, pp. 527—542. 1921.

876

ABBREVIATIONS AND DESIGNATIONS

ad

adductor of mantle

ak

orbita

al

outer lip

an

anus

ap

posterior pallial artery

ar

arm crown

as

siphonal artery

at

arm-funnel, connection of

au

eye

ax

axis of body

ba

cheek tubercles

bt

support of buccal funnel

ch

chromatophore

cl

central cavity of embryo

CO

cone of gladius

da

yolk artery

de

dorsal corner of mantle

dh

yolk membrane

dk

dorsal edge of eye

dl

gland line

dm

dorsal mantle slit

do

yolk sac

dv

yolk vein

dz

yolk cells

ea

body of embryo

ed

hind intestine

es

apical point of mantle sac

fa

flag of gladius
"window"

fe

fl

fin

fr

frontal area

gb

ligament of girdle

gd

poison gland

gl

gladius

hb

posterior connection of
primary lid (read hi in
Plate II )

hi

posterior cavity of body
of embryo

ho

Hoyle's organ; 1 — median
branch; 2 — lateral branch

hz heart

il inner lip

ir iris

ka "anlage" of head

kb gill ligament

kh branchial heart
kl funnel valve

km gill

kn funnel bond ("cartilage")
kp head

kr retractor of gill

ks secondary margin of germinal

disk

kv branchial vein

lb liver

Is lens

ma mantle

mb median band of germinal disk
mf median area of germinal disk
mh mantle cavity

mk buccal cone

mr musculus rectus abdominis
ms mantle septum

mu mouth, primary margin of mouth
na navel

nd nidamental gland

ni kidneys

nk neck bond

np renal pore, renal papilla
nt suture

pd pericardial gland

pf 1—5 armpillars (counted dorso-
ventrally)

pk primary skin of head

pi primary lid

pr proostracum

pu pupil

ra marginal rays of germinal disk
rh rhachis of gladius

ro olfactory organ

rs marginal zone of fin

877

rt funnel retractor

se shell epithelium

sf shell fold

sg stellate ganglion

sh interbrachial membrane
sk secondary lid

si lateral line

sm pharyngeal mass

sn pallial nerve

sp pore of shell

sr subradular organ

ss swimming margin

st statocyst

su protective margin
ta tentacle pocket

tb ink sac

td funnel gland

te funnel corner

tk tentacle club

tm median part of funnel
tn tentacle

to funnel opening

tr funnel

ts funnel septum

tt funnel pocket

uk lower jaw

va venous appendages
vb anterior connection of
primary lid
VC vena cava

vd fore intestine or esophagus

ve

ventral corner of
mantle

vk

ventral edge of eye

vl

lateral pallial vein

vm

ventral mantle slit

vp

posterior pallial vein

vr

thickening ring of germinal
disk (meso-entoderm)

vs

vein branches

wk

white body

wz

skin warts

z, y, X, w, u, etc. are various points

in the embryo, to be explained
in each case (cf. Plate l),
and of different meaning
d, exceptionally, b — posterior end of
dorsal edge of eye
V — posterior end of ventral edge

of eye (cf. Plate VI, Figure 4)
p position of primary pupil.

Pore of primary
vesicle of eye (Plate 11,
Figure 5)
n scar

t funnel corner

I— V 5 arms of one side, counted
dorsoventrally (Plate XIX,
Figure 5)

X Medioventral rudiment (?)

(Plate II, Figure 5)

The data on time and age of the embryos are only approximate. They
do not apply to any particular egg mass, since the eggs deposited in the
aquarium never complete their development normally, and the duration of
the phenomena of development depends on the temperature (i. e. season).
Development of L o 1 i g o applies to April— May, to August for Octopus, i.e.
the normal spawning period.

878

Naef • Cephalopoda

PLATES TO VOLUME I

(SYSTEMATICS)

The plates supplement the mainly diagrammatic
drawings in the text, in which many important details
are omitted. The plates intend to show only the typical
of the reality, and disregard the accidental. We have
attempted, however, to show the typical in its natural
completeness, as far as the available techniques permit.
The coloration of living objects is shown only in
Plate XIX [here reproduced in black and white].

[Note. The plates have been reduced here to
of the original size.]

879

PLATE L Young stages of Loli go vulgaris
from the plankton of the Bay of Naples

FIGURE 1. Ventral view of an animal about 3 weeks (?) old.9X: 1 — lateral outer edge of V arm;

2 - insertion of eyeball, which is visible through the thin wall of the orbital chamber; 3 - funnel pocket;

4 — "anlage" ofmembrane between 3rd and 4th arm, delimiting the still shallow pocket of tentacle;

5 - funnel adductor; 6 - olfactory organ; 7 - funnel bond.

FIGURE 2. Right side of same animal: 8 - iris lid; 9 — tentacle; 10 — orbital pore; 11 — attachment
of fin.

FIGURE 3. Upper view of same animal: 12 — swimming margin of LV arm; 13 — lanceola, i.e. translucent
median plate of gladius. (Cf. Figure 62 on p.l46.)

FIGURE 4. Older animal, probably 5-6 weeks old. 6x. 14 - third longitudinal fold of "neck," protecting
the olfactory organ; 15 — a glandular ridge.

FIGURE 5. Further developed specimen, after removal of ventral part of muscular mantle. 9/2 X: 16 - pro-
tective margin; 17 - ventral supports of buccal funnel; 18 - ventral part of funnel gland; 19 - cephalic
vein; 20 — anal papilla; 21 — branchial vein; 22 — venous appendages visible through renal sac; 23 — bran-
chial heart with attached pericardial gland in the superficial coelomic pocket for the branchial heart;

24 - posterior pallial vein; 25 - median pallial septum; 26 - swimming margin of tentacle club; 27 - funnel
retractor; 28-inksac; 29 - hind intestine; 30 - gill retractor; 31 - median pallial artery.

FIGURE 6. Dorsal view of specimen in Figure 4: 15b and 15c — as 15a; 32 — cornea. The chromatophores are
partly expanded; they are light yellow, carmine red and dark reddish brown. (Cf. Plate XIX, Figure 3, in which
a closely similar specimen is shown.)

All these juvenile forms are almost completely transparent in life, except for the points of the jaws and
some dark inner organs which are covered by iridescent membranes (eyes, ink sac, liver). Expansion of the
chromatophores suddenly covers these reflexes. The next plate shows the further development of the fins.

880

881

PLATE IL Young Loligo vulgaris, 2X.

FIGURE 1. Half- grown male with opened mantle. For explanation see the corresponding figure in Chapter 7.
The borders of the funnel pit and neck folds are particularly distinct. Note the position of the soft parts in the
shell, the margin of which is sharply defined in the posterior part.

FIGURE 2. Region of gill base in a female of the same size, with juvenile nidamental and accessory glands.
(Cf. figures and explanations in Chapter 7.)

FIGURE 3. Head of a slightly smaller specimen with open tentacle pockets. Removal of the thin wall shows
the roots of the tentacle. Note the slitlike penetration of the pockets between the eyes and roots of the arms;
compare the corresponding figures and explanations in Chapters 6-8.

FIGURES 4 and 5. Younger specimen, preserved in formol but with retracted head and shortened arms. Note
distribution of chromatophores; compare the corresponding figure and explanation in Chapter 7.

882

883

PLATE III. Mouth area and mantle situs in young Oegopsida

FIGURE 1, Mouth area ofAbraliopsis morrisi (same stage as in Plate IV); 40 x. (For general orientation,
compare Figure 45b on p,119): I-III - buccal pockets; 1-4 - buccal pillars; 5 - ligament of attachment
(vena tentaculi); 6 - indication of 7th buccal pocket; 7 - outer attachment of ventral arms.

FIGURE 2. Mouth area of Histioteuthis b o ne 1 1 i a na, 25x. Note the fusion of the dorsal and ventral
buccal pillars (10), the invagination of the base of the tentacles together with the ligament of attachment (9);
11 — double attachment of ventral buccal pillar; 12 — adductor of the DL arm. Compare the figures and
explanations in the corresponding chapter.

FIGURES. Calliteuthis r e v e rs a, 16.3 x. Ventral part of mantle removed. Note the luminous organs
of the lid margin which are dense on the right (z) and sparser on the left (w); x,y — aberrant organs of the
posterior cavity of the heart; 13 — funnel adductor; 14 — olfactory organ; 15 — funnel valve; 16 — genital
process (female); 17 — renal papilla; 18 — posterior aorta; 19 — coelomic pocket for the branchial heart;

20 — posterior pallial vein; 21 - rudiment of nidamental gland; 22 — posterior pallial artery; 23 — apical
point of mantle sac, reduced to a frenulum between the fins.

FIGURE 4. Ctenopteryx sic ulus, 16. 3X: 24 — ventral buccal pillars; 25 — iris margin; 26 — third
longitudinal fold of neck, to which the olfactory tubercle is attached; 27 — renal papilla, anterior to it —
the wide, padlike, single (fused) accessory gland; 28 — connecting stripes between nidamental and accessory
glands (cf. Plate II, Figure 2); 29 — nidamental gland; 30 — "anlage " of oviduct; 31 — appendage of
branchial heart (34); 32 - heart; 33 - posterior aorta; 35 - lateral pallial artery.

The completed fins and arms are shown in a figure in the text.

884

885

PLATE IV. Mantle situs of Oegopsida

FIGURE 1. Onychoteuthis banks! juv. 6x, after removal of ventral parts of mantle. The figures
are explained in the adjacent drawing: Note form and size of fins; form of posterior end; form and size of

funnel part, especially the funnel cartilage (1). Note
on the head: closed lid folds; "window" (Fn); translucent
luminous organs (Oj); boundary of funnel pit; neck folds
(Lg, L4); olfactory tubercle (Rt), attached to the third
fold. In the mantle cavity: gills; branchial hearts (10)
with appendage (11), situated in the coelomic pocket (20);
venous branches (8): vena cava (6); renal papillae (7); hind
intestine (4); anus (Af); heart (9); posterior aorta (13);
posterior pallial veins (14); "anlage" of gonoduct outside
the left branchial heart (12); 2 - funnel retractor;

3 — characteristic depression near the insertion of the retractor
on the mantle; 5 - branchial vein; 15 - stomach (translucent);
16 - gladius (translucent); 17 - posterior pallial artery; 18 -
"anlage" of gonads (thin longitudinal ridge); Co — cone;

Rs - rostrum; Og - anal luminous organ; O3 - posterior
luminous organ on ink sac; Qu - transverse fold of neck;

Li — lens of eye; Te — base of tentacle.

FIGURE 2. Abraliopsis m orrisi juv., 18x. Mantle
cavity opened, as in 1; eyelids completely retracted from
eye (3); 1,2, 4, 5 - luminous organs; 6 - ink sac; 7 - branchial
gland; 8 - branchial retractor; 9 - "anlage" of gonoduct;

10 - posterior aorta; 11 - posterior pallial vein; 12 - stomach;
13 — caecum; 14 — posterior pallial artery; 15 — median pallial
artery (septum): 16 - apex (fleshy).

FIGURE 3. Chiroteuthis v e r a ny i. 3/2 x. (Material of C. Chun, 1910.): 17 - translucent luminous organ
of eyeball; 18 - funnel adductors; 19 - olfactory organ; 20 - mantle bond; 21 - luminous organ; 22 - nida-
mental gland; 24 — posterior pallial vein; 25 — long, translucent cone (I). Note the proportions of eyes and
ventral arms.

5110/2

886

5110/2

887

PLATE V. Young stages of Ommat os trephes sagittatus

FIGURE 1. Easily recognizable animal, 3 x. Explained in detail in the text.

FIGURES 2 and 3. Younger specimen, 16.3 x. Compare with Plate I, particularly the similar development
of the fins; 1 — tentacles, just separated: 2 — interbrachial membrane; 3 — gland line (cf. Plate I); 4 — lens
of eye; 5 — crescent- shaped fold; 6 — outer funnel adductor; 7 — swimming margin; 8 — sinus of lid margin
(9); 10 — fourth longitudinal fold; 11 — third longitudinal fold of neck; 12 — olfactory organ attached to this
fold.

FIGURES 4 and 5. Not definitely determined, indifferent stages of "Rhy nchoteuthion" of Ommatostrephidae,
anterior view, 33x. The tentacle stalks are fused with the "snout." On the left (3), the LV arms are still in
the "anlage," they begin to grow on the right (14), 1,2,3 - "anlagen" of arms; 4 - tentacle club (end part
of snout always with 4 suckers on each side); 5 - funnel; 6 - terminal sucker; 7 - iris margin; 8 - lid
margin; 12 - "anlage" of sucker; 13 - closed lid margin; 14 - LV arm; 15 - tentacle stalk as part of the
snout. .Note also; outer and inner lip and end of jaws in the mouth (cf. Vol.II, Plate XII).

888

1

889

PLATE VI. Morphology of mouth area in the Ommatostrephidae

FIGURE 1. Rhy nc ho teu thion stage, further developed (resembling Figure 2 of Plate V), 25x. The
tentacles begin to become separated; the buccal points appear; the ventral arms grow.

FIGURE 2. Sepia o f fi c i n a li s, recently hatched (compare with Figure 3), 10 X; 1-4 — buccal pillars;
a-d - their attachments; I— III - buccal pockets (both in typical arrangement); 5 - inner lip; 6 — buccal
membrane covering outer lip; 7,8 — tentacle clubs, invaginated, typical for the Sepiidae; 9 — protective
margins, united at the base of the arms as typical for the Sepiidae; 10 — outer margin of ventral arm;

11 - interbrachial membrane.

FIGURES. Ommatostrephes sagittatus juv., 24 x (same stage as Figure 2, Plave V): 1-4, 1-III as in
Figure 2; 5, 6, 7 - adductors of arms, externally visible in the Teuthoidea; 8 - fusion of adductors of arms;

9 - "anlage" of interbrachial membrane; 10 - lens; 11 - swimming margin; 12 - club, already normally
developed; 13 - lid margin.

890

891

PLA TE VII. Mantle situs of Sepia officinalis, natural size

Left — male; right — female. 1 — funnel; 2 — funnel pockets; 3 — funnel bond; 4 — stellate ganglion;

5 - cephalic vein; 6 - funnel retractor; 7 - anal papilla; 8 - wings (lobes) of anal papilla; 9 - renal
papilla; 11 — genital process; 12 — outlet of inc sac, near hind intestine; 13 — venous appendages, visible
through wall of renal sac; 14 — pericardial gland on branchial heart; 15 — gill; 16 — gill retractor; 17 —
posterior aorta; 18 - posterior pallial vein; 19 - gonoduct; 20 - posterior border of mantle cavity (very
extensible); 21 — median pallial artery (septum); 22 — posterior pallial artery; 23 — ink sac; 24 — posterior
end, containing spine of shell; 25 — fin; 26 — white marginal line, more distinct in the male; 27 - lateral
pallial vein; 28 - mantle bond; 29 - gill ligament; 30 - chromatophores of mantle margin; 31 - olfactory
organ; 32 - buccal funnel; 33 - funnel valve: 34 - funnel gland; 35 - median part of funnel gland; 37 -
section through funnel; 38 — tentacle stalk; 40 — accessory nidamental gland; 41 — outlet of accessory
nidamental gland, invaginated; 42 - median part; 43 - outlet; 44 - body of nidamental gland; 46 - as 23;
48 — ovary; 49 — siphonal artery; 50 — vein of nidamental gland. Mantle opened in both figures; funnel
opened only in the figure on the right.

892

893

PLATE VIII. H e ter oteuthis dispar

FIGURE 1. Male, 16/5 X

FIGURE 2. Female, 3x

FIGURES 3 and 4. Young stage, 8X

FIGURES 5 and 6. Very young animal, 16 x

Note the change of form, especially of the fins. 1 - enlarged sucker of male; 2 - funnel valve; 3 - funnel
gland; 4 — lateral funnel adductor, typical for the Sepioidea; 5 - luminous gland; 6 - mantle adductor, cut;

7 - ink sac; 8 - renal papilla; 9 - genital process ("penis"); 10 - spermatophore gland; 11 - mantle bond;

12 - atypical swimming margin of tentacle in the Heteroteuthinae; 13 - groove of gland; 14 - club; 15 -
base of tentacle in the "bag" formed by the membrane between the 3rd and 4th arm (Figure 3 near 31);

16 - outlet of luminous gland (17); 18 - separated part of accessory gland (19); 20 - genital process of female;
21 - oviduct gland; 22 - nidamental gland; 23 - posterior pallial vein, displaced (!); 24 - median pallial
septum; 25 - ovary; 26 - as 12; 27 - interbrachial membrane; 28 - gland line; 29 - lens; 30 - corner of
fin (typical for the Heteroteuthinael); 31 - interbrachial membrane; 32 - same around tentacle; 33 - orbit,
slit; 34 — eyeball; 35 — shovel-like part; 36 — margin; 37 — stalk part of fin; 38 — cornea; 39 — dorsal
border of cornea; 40 — eyeball; 41 - translucent ventral part of funnel gland; 42 - olfactory organ; 43 -
funnel bond; 44 - hind intestine, translucent; 45 - gills, same; 46 - margin of basal part of tentacle pocket.

894

895

PLATE IX. Young stages of Octopodidae
(cf. Plate XXX in Vol. ID

FIGURE 1. Planktonic stage of Octopus macropus, 18x. Specimen from Messina. Preservation in
formol has resulted in a marked shortening of funnel and arms. Note the glistening tufts of bristles; charac-
teristic distribution of chromatophores and small head.

FIGURE 2. Planktonic stage of Octopus vulgaris from Naples, 22 x. Chromatophores contracted. Note
the terminal flagella (vegetative centers of armsi).

FIGURES. Octopus vulgaris juv ., after transition to benthic life, 11 x.

FIGURE 4. Octopus v u 1 g a r i s, planktonic stage, soon after hatching. The arm bears 3 embryonic suckers
and the first 4 postembryonic "anlagen." 72 x.

FIGURE 5. Apex of arm of a half-grown E led one mo schat a, explaining the constant uniserial differen-
tiation of the suckers. 33 x.

FIGURE 6. Apex of arm of a young Octopus vulgaris, showing the primarily uniserial arrangement of
the "anlagen" of the suckers and their secondary displacement. 30 x.

896

897

PLATE X, Topography of mantle cavity of Oct opoda

FIGURE 1. Eledone mosc hat a, freshly hatched, 18X; 1 — funnel; 2 — funnel pockets; 3 — anal papilla;

4 — thickened mantle margin; 5 — depression for the funnel corner (6); 7 — gill margin; 8 ~ funnel retractor;

9 “ mantle adductors, cut from the mantle at 10; 11 — median pallial artery; 12 ~ hind intestine, translucent;
13 — vena cava, with the median pallial artery situated on it; 14 — branchial vein; 15 — branchial heart;

16 — appendages of lateral pallial vein; 17 — appendages of venous branches; 18 — superficial branchial
retractor (cf. Figure 1 in Plate II); 19 - heart; 20 - renal sac; 21 - posterior aorta; 22 - auricle; 23 - renal
septum; 24 - as 18; 25 ~ radiation of mantle adductor (10 - remnant of median pallial septum).

FIGURE 2. Octopus v u 1 g a ri s, young stage, 9/2X: 26 - branchial gland; 27 - adductor stellaris of mantle,
containing the pallial nerve; 28 - genital opening; 29 - renal papilla: 30 - oviduct.

FIGURE 3. Octopus v u 1 g a r i s, fully developed embryo, 25x (cf. Figure 7 of Plate XXX in VoLII).

31 - vena cava; 32 - stellate ganglion; 33 - gill; 34 - posterior mantle cavity; 35 - renal sac; 36 - funnel
retractor; 37 — slit perforating into dorsal mantle cavity; 38 — attachment of the funnel pocket on mantle.

FIGURE 4. Tremoctopus viol ace us, fully developed embryo, 25 x (cf. Plate XXXI in VoLII).

39 - olfactory organ in the mantle corner; 40 - branchial lamellae; 41 - depression for the funnel corner;

42 - as 25; 43 - posterior end; 44 - as 10.

FIGURES 5 and 6. Embryos of Argonauta argo (cf. Plate XXXVII in VoLII), 40x. Mantle cavity opened
dorsally and ventrally: 45 - yolk sac; 46 - as 27,i.e. pallial nerve for the stellate ganglion (37); 47 - mantle
septum, still completely preserved (!); 48 - ink sac; 49 - funnel bond; 50 - DL arm; 51 - inner view of
funnel pocket, which is situated dorsally on the mantle along the line of the cut; 52 — dorsal wall of body,
exposed by expansion of mantle cavity; 53 - chromatophores on dorsal wall, translucent in life; 54 - as 51.

898

899

PLATE XL Argonauta argo, after live specimens, natural size

FIGURE 1. Female in swimming position with closed shell and extended shell arms.

FIGURE 2. Same specimen with retracted shell arms.

FIGURE 3. Mature male, with retracted hectocotylus (below left eye).

FIGURES 1 and 2 are based on numerous observations, drawings and photographs. They are as realistic as
possible and should finally replace the monstrosities which appear in textbooks and manuals.

900

901

PLATE XII. Suckers of Teuthoidea

FIGURE 1. Loligo v u 1 g a r i s, sucker of middle of 3rd arm, 14x.

FIGURE 2. Ommatostrephes sagittatus, large sucker of club of tentacle, 6x.

FIGURES. Ancistroteuthis lichtensteini, 3rd arm, base, 20x.

FIGURE 4. Illex coindeti, 3rd arm, proximal part, 11 x.

FIGURES. Ommatostrephes sagittatus, 3rd arm, 12x.

FIGURES. Stenoteuthis bart rami, small sucker of proximal part of club of tentacle, 20x (?).

FIGURE 7. Illex coindeti, 3rd arm, distal part, 28X.

FIGURES. Stenoteuthis b a rt ra m i, distal sucker of 3rd arm, 40x.

FIGURE 9. Hypothetical hooklike sucker of Gonatus f a b ri c i i, almost identical with that occurring in
nature; cf. Appellof, 1892, Plate 3, Figure 6.

1 (cf. Figure 7, in Plate XIII) — supporting edge behind the small teeth (2); 3 — horny ring; 4 — main tooth;

5 — thickening of horny ring at base of tooth; 6 — chitinous ornament of the adhesive ring; 7 — supporting
edge, around the whole sucker; 8 - smooth adhesive ring; 9 - margin of adhesive ring; 10 - rounded
secondary tooth; 11 - same, smaller (8 - reduced adhesive ring); 12 - depression of suction pad; 13 - as 10;
14 - angular secondary tooth; 15 - reinforced base of main tooth; 16 - raised marginal ring which passes
above base of main tooth (hood); 17 — as 3.

902

903

PLATE XIII . Suckers of Oct op oda and Sepioidea
for comparison ivith the Teuthoidea

FIGURE 1, Octopus vulgaris, one of the enlarged suckers of the male, 8/3x. Median section
(cf. Figure 29 on p.98). The drawing explains the figures.

1 - musculature in the gelatinous tissue of the stalk
of the sucker; 2 - suction pad (bottom) ;3 - lateral
wall; 4 - musculature of adhesive ring; 5 - marginal
ring; 6 - skin; 7 - boundary of true (muscular) sucker.

FIGURE 2. Same sucker, opening: 1 — marginal ring; 2 — margin of adhesive ring.

FIGURE 3. Octopus tetracirrus, sucker from apex of arm, 35X: 1 - papilla of inner margin; 2-
papilla of marginal ring.

FIGURE 4. Sepia officinalis, sucker from middle of 3rd arm, 29 X; 1 - dense, often connected "incisor"
teeth of distal margin; 2 — horny ring; 3 — adhesive ring with chitinous papillae,

FIGURES. Sepiola robust a, large sucker of club of tentacle, 200 X: 1 - chitinous stripes; 2,3 -papillae
of adhesive ring, passing into true teeth (4).

FIGURES. Sepia officinalis, sucker from apex of tentacle, 120X; 1—4 — transition of papillae into
teeth.

FIGURE 7. Median section through a sucker ofLoligo vulgaris (Figure 1 in Plate XII). This sucker
is typical for the Decapoda (Teuthoidea); 1 - marginal ring; 2 - chitinous covering of adhesive ring;

3 - horny ring supporting lateral wall, distal part; 4 — circular muscle; 5 - deepest part of suction chamber;

6 — pit in suction pad for insertion of the stalk; 7 — pit in suction pad opposite insertion of stalk; 8 — as 3
but proximal part; 9 - as 2; 10 - as 1.

FIGURE 8. Sepia o f fi c i na 1 i s, large sucker of club of tentacle, 14 X; 1 - margin of adhesive ring;

2 — teeth; 3 — adhesive ring.

FIGURE 9. Sepia of f i c in a 1 i s, juvenile, apex of arm, 112 X; 4 - large tooth; 5 - small tooth; 6 - reduced
tooth. Note sculpture of the outer side.

904

905

PLATE XIV, Radulae of Teuthoidea

FIGURE 1. Loligo vulgaris, 44x.

FIGURE 2. Alloteuthis media, 68x.

FIGURES. Illex coindeti, 25x.

FIGURE 4. Illex coindeti, abnormal radula, 40x, Inner brush teeth on left side (xx) absent.

FIGURES. Todaropsis eblanae, 35x.

FIGURES. Omm atostrephes sagittatus, 44x.

FIGURE 7. Stenoteuthis bartrami, 26x.

FIGURES. Thysanoteuthis rhombus, 22x.

Such preparations (cf. pp.67 and lOO.Vol.II) are obtained by spreading fresh radulae on a glass slide and drying
them slowly. Most of the illustrations published show deformed objects, e.g. in Jatta (1896). Good drawings are
found in Targioni-Tozzetti (1869, Plate 6).

906

907

PLATE XV. Radulae of Teuthoidea (1-4) and Sepioidea (5—8)

FIGURE 1. Histioteuthis bonelliana, 27x.

FIGURE 2. Ancistroteuthis li ch tenstei ni, 43x,

FIGURES. Ony choteuthis banksi, 40x.

FIGURE 4. Chiroteuthis veranyi, 20x.

FIGURES. Sepia officinalis, 27x.

FIGURES. Sepietta oweniana, 31x,

FIGURE 7. Rossia macrosoma, 54x.

FIGURES. Sepia orbignyana, 35x.

All teeth of the Sepioidea are unicuspid but in the Teuthoidea they are unicuspid only in the Enoploteuthidae.
The marginal platelets are completely reduced.

908

55/4

909

PLATE XVI. Radulae of Nautilus and Octopoda

FIGURE 1.
FIGURE 2.
FIGURE 3.
FIGURE 4.
FIGURE 5.
FIGURE 6.
FIGURE 7.
FIGURE 8.

Nautilus pompilius, worn, 11 x.

Nautilus pompilius, right half. Teeth removed to show their position.
Nautilus pompilius, fresh specimen.

Eledone moschata, 40x.

Octopus vulgaris, 43x.

Tremoctopus violaceus, 43x.

Argonauta argo, 18x.

Ocythoe tuberculata, I8x.

910

oooo

911

O D O O

PLATE XVII . Jaws of Teuthoidea

Lateral view. Above — upper jaw; below — lower jaw. General explanation: VoLI,p.99: legend of
Figure 6; a - boundary between the dark, chitinized and the soft part; b - boundary between inner and
outer plates; c and d — lateral teeth; e - lateral ridge; f - median ridge. The other figures, in which
the lettering varies, can be interpreted by this; the lettering there only indicates characteristic details.

FIGURE 1. Loligo vulgaris, 3X.

FIGURE 2. Alloteuthis media, 20x.

FIGURES. Loligo forbesi, 4/3X.

FIGURE 4. Ancistroteuthis lichtenstei ni, 3x.

FIGURE 5. Onychoteuthis banksi, 8/3x.

FIGURES. Histioteuthis bonelliana, 20/3X.

FIGURE 7. Chiroteuthis verany, 4/3x.

FIGURES. Illex coindeti, 8/3x.

FIGURE 9. Todaropsis eblanae, 7/3x.

FIGURE 10. Ommatostrephes sagittatus, 5/3x.

FIGURE 11. Stenoteuthis bartrami, 2x.

FIGURE 12. Thysanoteuthis rhombus, 8/3X.

912

913

PLATE XVIII. Jaws of Sepioidea and Octopoda

FIGURE 1. Sepia officinalis, 2x.

FIGURE 2. Rossia m acrosoma, 4 x.

FIGURES. Sepietta oweniana, 19/3X.

FIGURE 4. Octopus vulgaris, 3/2X.

FIGURES. Octopus vulgaris (juv.), 11/3X.

FIGURES. Octopus macropus, 8/3X.

FIGURE 7. Octopus salutii, 8/3X.

FIGURE 8. Eledone cirrosa, 3X.

FIGURE 9. Eledone moschata, 11/3X.

FIGURE 10. Octopus ( " S c a e u rg u s " ) unicirrus, 25/3X.
FIGURE 11. Ocythoe tuberculata, 4/3x.

FIGURE 12. Argonauta argo, 3x.

914

915

PLATE XIX. Natural coloration of small Cephalopoda

and of some parts or products of Cephalopoda. Figure 7 was drawn by Com. Merculiano; Figures 1-3 were
drawn by V. Serino of the Zoological Station. The others were drawn by the author and colored by V, Serino.

FIGURES 1 and 2 are of larvae of Ommatostrephidae bred from a floating batch of eggs (cf. Vol.Il, Plates IX-XI).
After live animals, about 25x.

FIGURES. Young Alloteuthis spec., natural size.

FIGURE 4. Luminous glands ofRondeletiola minor, male, natural size (cf. Plate VIII, Figure 1),

(pit of opening).

FIGURES. Todaropsis eblanae, juvenile, alive, swimming position, natural size,

FIGURES. Luminous glands of Sepiola 1 igu 1 a ta, male, 3x.

FIGURE 7, Mantle situs of Sepietta obscura, natural size.

FIGURES. Sepietta oweniana, alive, natural size.

FIGURE 9. Calliteuthis r e ve rs a, juvenile, alive, 3 x.

FIGURE 10. Heteroteuthis dispar, juvenile, alive, 2x.

FIGURE 11. Sepiola ligulata, adult, alive, natural size.

FIGURE 12. Sepietta oweniana, as Figure 8 but dying or dead, injured during capture.

FIGURE 13. Rondeletiola minor, alive, namral size.

FIGURE 14. Sepiola a f f i n i s, alive, natural size.

FIGURE 15. Eggs of Rondeletiola minor, natural size; above — freshly hatched; below — further
developed.

FIGURE 16. Luminous glands and adnexes of a female of Rondeletiola. (Cf. Plate VIII, Figure 2 and
Figure 4 of this Plate). The lens- shaped organ is embedded in the yellowish red accessory nidamental gland.
Behind it and to the right is the bursa copulatrix. 4x.

FIGURE 17. Same complex in Sepiola ligulata, 3x. The ear- shaped glands are situated before the
accessory glands.

FIGURE 18. Eggs of Sepietta oweniana, natural size as Figure 15.

(Material from Naples).

916

917

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