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686
CEPHALOPODA

mantle-skirt. The shell consists of a series of chambers, the last-formed of which is occupied by the body of the animal, the hinder ones (successively deserted) containing gas (fig. 1). The pair of cephalic eyes are hollow chambers (fig. 14. A), opening to the exterior by minute orifices (pin-hole camera), and devoid of refractive structures. A pair of osphradia are present at the base of the gills (fig. 4, olf). Salivary glands are wanting. An ink-sac is not present. Branchial hearts are not developed on the branchial afferent vessels.

EB1911 Cephalopoda Fig. 1.— Lateral view of the female Pearly Nautilus.jpg
Fig. 1.—Lateral view of the female Pearly Nautilus, contracted by spirit and lying in its shell, the right half of which is cut away (from Gegenbaur, after Owen).

a, Visceral hump.

b, Portion of the free edge of the mantle-skirt reflected on to the shell,—the edge of the mantle-skirt can be traced downwards and forwards around the base of the mid-foot or siphon i.

l, l, Superficial origin of the retractor muscle of the mid-foot (siphon), more or less firmly attached to the shell, of which a small piece (s) is seen between the letters l, l.

s, (farther back) points to the siphuncular pedicle, which is broken off short and not continued, as in the perfect state, through the whole length of the siphuncle of the shell, also marked s and s′.

 o, points to the right eye.

 t, is placed near the extremities of the contracted tentacles of the outer or annular lobe of the fore-foot—the jointed tentacles are seen protruding a little from their long cylindrical sheaths.

 v, The dorsal “hood” formed by an enlargement in this region of the annular lobe of the fore-foot (m in figs. 2, 3).

 V, A swelling of the mantle-skirt, indicating the position on its inner face of the nidamental gland (see fig. 4, g.n.).

Visceral Hump and Shell.—The visceral hump of Nautilus (if we exclude from consideration the fine siphuncular pedicle which it trails, as it were, behind it) is very little, if at all, affected by the coiled form of the shell which it carries, since the animal always slips forward in the shell as it grows, and inhabits a chamber which is practically cylindrical (fig. 1). Were the deserted chambers thrown off instead of being accumulated behind the inhabited chamber as a coiled series of air-chambers, we should have a more correct indication in the shell of the extent and form of the animal’s body. Amongst Gastropods it is not very unusual to find the animal slipping forward in its shell as growth advances and leaving an unoccupied chamber in the apex of the shell. This may indeed become shut off from the occupied cavity by a transverse septum, and a series of such septa may be formed, but in no Gastropod are these apical chambers known to contain a gas during the life of the animal in whose shell they occur. A further peculiarity of the nautilus shell and of that of the allied extinct Ammonites, Scaphites, Orthoceras, &c., and of the living Spirula, is that the series of deserted air-chambers is traversed by a cord-like pedicle extending from the centro-dorsal area of the visceral hump to the smallest and first-formed chamber of the series. No structure comparable to this siphuncular pedicle is known in any other Mollusca. The siphuncle does not communicate with the coelomic cavity; it is a simple vascular process of the mantle, whose cavity consists of a venous sinus, and whose wall contains a ramification of the pallial artery. There appears to be no doubt that the deserted chambers of the nautilus shell contain in the healthy living animal a gas which serves to lessen the specific gravity of the whole organism. This gas is said to be of the same composition as the atmosphere, with a larger proportion of nitrogen. With regard to its origin we have only conjectures. Each septum shutting off an air-containing chamber is formed during a period of quiescence, probably after the reproductive act, when the visceral mass of the nautilus may be slightly shrunk, and gas is secreted from the dorsal integument so as to fill up the space previously occupied by the animal. A certain stage is reached in the growth of the animal when no new chambers are formed. The whole process of the loosening of the animal in its chamber and of its slipping forward when a new septum is formed, as well as the mode in which the air-chambers may be used as a hydrostatic apparatus, and the relation to this use, if any, of the siphuncular pedicle, is involved in obscurity, and is the subject of much ingenious speculation. In connexion with the secretion of gas by the animal, besides the parallel cases ranging from the protozoon Arcella to the physoclistic fishes, from the hydroid Siphonophora to the insect-larva Corethra, we have the identical phenomenon observed in the closely allied Sepia when recently hatched. Here, in the pores of the internal rudimentary shell, gas is observable, which has necessarily been liberated by the tissues which secrete the shell, and not derived from any external source (Huxley).

The coiled shell of Nautilus, and of the majority of extinct Tetrabranchiata, is peculiar in its relation to the body of the animal, inasmuch as the curvature of the coil proceeding from the centro-dorsal area is towards the head or forwards, instead of away from the head and backwards as in other discoid coiled shells such as Planorbis; the coil is in fact absolutely reversed in the two cases. Such a shell is said to be exogastric. But in some extinct forms, e.g. Phragmoceras, Cyrtoceras, Ptenoceras, the shell is coiled towards the ventral side, when it is termed endogastric. Amongst the extinct allies of the nautilus (Tetrabranchiata) we find shells of a variety of shapes, open coils such as Scaphites, leading on to perfectly cylindrical shells with chamber succeeding chamber in a straight line (Orthoceras), whence again we may pass to the corkscrew spires formed by the shell of Turrilites. In some extinct genera, e.g. Gomphoceras, among the Nautiloidea the aperture of the shell is contracted and the edge of the aperture is lobed. In these cases the animal was probably able only to protrude its appendages and not its whole head. The ventral part of the aperture corresponding to the funnel is separated from the dorsal part by a constriction. Hence it is possible to distinguish the ventral and dorsal sides of the shell and to decide whether it was exogastric or endogastric. The direction of the coil of the shell cannot be determined by the position of the siphuncle, which traverses the septa centrally, ventrally or dorsally. Contracted shell apertures occur also in Ammonitoidea, the condition reaching an extreme in Morphoceras, where the original aperture is subdivided by the ingrowth of the sides, so that only five small separate apertures remain. Of these the central probably corresponded to the mouth, two lateral to the eyes, and the remaining two to the pedal appendages.

Head, Foot, Mantle-skirt and Sub-pallial Chamber.—In the pearly nautilus the ovoid visceral hump is completely encircled by the free flap of integument known as mantle-skirt (figs. 2, 3, d, e). In the antero-dorsal region this flap is enlarged so as to be reflected a little over the coil of the shell which rests on it. In the postero-ventral region the flap is deepest, forming an extensive sub-pallial chamber, at the entrance of which e is placed in fig. 3. A view of the interior