Page:EB1911 - Volume 11.djvu/536

Fig. 36.

A, Veliger-larva of an Opisthobranch (Polycera). f, Foot; op, operculum; mn, anal papilla; ry, dry, two portions of unabsorbed nutritive yolk on either side of the intestine. The right otocyst is seen at the root of the foot.

B, Trochosphere of an Opisthobranch (Pleurobranchidium) showing—shgr, the shell-gland or primitive shell-sac; v, the cilia of the velum; ph, the commencing stomodaeum or oral invagination; ot, the left otocyst; pg, red-coloured pigment spot.

C, Diblastula of an Opisthobranch (Polycera) with elongated blastopore oi.

(All from Lankester.)

Fig. 37.—Phyllirhoë bucephala, twice the natural size, a transparent pisciform pelagic Opisthobranch. The internal organs are shown as seen by transmitted light. (After W. Keferstein.)
a,	Mouth.
b,	Radular sac.
c,	Oesophagus.
d,	Stomach.
c’,	Intestine.
f’,	Anus.
g, g′, g′′, g′′′,
	The four lobes of the liver.
h,	The heart (auricle and ventricle).
l,	The renal sac (nephridium).
l′,	The ciliated communication of the renal sac with the pericardium.
m,	The external opening of the renal sac.
n,	The cerebral ganglion.
o,	The cephalic tentacles.
f,	The genital pore.
y,	The ovo-testes.
w,	The parasitic hydromedusa Mnestra, usually found attached in this position by the aboral pole of its umbrella.

Order 1.—Opisthobranchia. Marine Euthyneura, the more archaic forms of which have a relatively large foot and a small visceral hump, from the base of which projects on the right side a short mantle-skirt. The anus is placed in such forms far back beyond the mantle-skirt. In front of the anus, and only partially covered by the mantle-skirt, is the ctenidium with its free end turned backwards. The heart lies in front of, instead of to the side of, the attachment of the ctenidium—hence Opisthobranchia as opposed to “Prosobranchia,” which correspond to the Streptoneura. A shell is possessed in the adult state by but few Opisthobranchia, but all pass through a veliger larval stage with a nautiloid shell (fig. 36). Many Opisthobranchia have by a process of atrophy lost the typical ctenidium and the mantle-skirt, and have developed other organs in their place. As in some Pectinibranchia, the free margin of the mantle-skirt is frequently reflected over the shell when a shell exists; and, as in some Pectinibranchia, broad lateral outgrowths of the foot (parapodia) are often developed which may be thrown over the shell or naked dorsal surface of the body.

The variety of special developments of structure accompanying the atrophy of typical organs in the Opisthobranchia and general degeneration of organization is very great. The members of the order present the same wide range of superficial appearance as do the Pectinibranchiate Streptoneura, forms carrying well-developed spiral shells and large mantle-skirts being included in the group, together with flattened or cylindrical slug-like forms. But in respect of the substitution of other parts for the mantle-skirt and for the gill which the more degenerate Opisthobranchia exhibit, this order stands alone. Some Opisthobranchia are striking examples of degeneration (some Nudibranchia), having none of those regions or processes of the body developed which distinguish the archaic Mollusca from such flat-worms as the Dendrocoel Planarians. Indeed, were it not for their retention of the characteristic odontophore we should have little or no indication that such forms as Phyllirhoë and Limapontia really belong to the Mollusca at all. The interesting little Rhodope veranyii, which has no odontophore, has been associated by systematists both with these simplified Opisthobranchs and with Rhabdocoel Planarians.

Fig. 38.—Three views of Aplysia sp., in various conditions of expansion and retraction. (After Cuvier.)
t,	Anterior cephalic tentacles.
t²,	Posterior cephalic tentacles.
e,	Eyes.
f,	Metapodium.
ep,	Epipodium.
g,	Gill-plume (ctenidium).
m,	Mantle-flap reflected over the thin oval shell.
os, s,	Orifice formed by the unclosed border of the reflected mantle-skirt, allowing the shell to show.
pe,	The spermatic groove.

In many respects the sea-hare (Aplysia), of which several species are known (some occurring on the English coast), serves as a convenient example of the fullest development of the organization characteristic of Opisthobranchia. The woodcut (fig. 38) gives a faithful representation of the great mobility of the various parts of the body. The head is well marked and joined to the body by a somewhat constricted neck. It carries two pairs of cephalic tentacles and a pair of sessile eyes. The visceral hump is low and not drawn out into a spire. The foot is long, carrying the oblong visceral mass upon it, and projecting (as metapodium) a little beyond it (f). Laterally the foot gives rise to a pair of mobile fleshy lobes, the parapodia (ep), which can be thrown up so as to cover in the dorsal surface of the animal. Such parapodia are common, though by no means universal, among Opisthobranchia. The torsion of the visceral hump is not carried out very fully, the consequence being that the anus has a posterior position a little to the right of the median line above the metapodium, whilst the branchial chamber formed by the overhanging mantle-skirt faces the right side of the body instead of lying well to the front as in Streptoneura and as in Pulmonate Euthyneura. The gill-plume, which in Aplysia is the typical Molluscan ctenidium, is seen in fig. 39 projecting from the branchial sub-pallial space. The relation of the delicate shell to the mantle is peculiar, since it occupies an oval area upon the visceral hump, the extent of which is indicated in fig. 38, C, but may be better understood by a glance at the figures of the allied genus Umbrella (fig. 40), in which the margin of the mantle-skirt coincides, just as it does in the limpet, with the margin of the shell. But in Aplysia the mantle is reflected over the edge of the shell, and grows over its upper surface so as to completely enclose it, excepting at the small central area s where the naked shell is exposed. This enclosure of the shell is a permanent development of the arrangement seen in many Streptoneura (e.g. Pyrula, Ovula, see figs. 18 and 32), where the border of the mantle can be, and usually is, drawn over the shell, though it is withdrawn (as it cannot be in Aplysia) when they are irritated. From the fact that Aplysia commences its life as a free-swimming veliger with a nautiloid shell not enclosed in any way by the border of the mantle, it is clear that the enclosure of the shell in the adult is a secondary process. Accordingly, the shell of Aplysia must not be confounded with a primitive shell in its shell-sac, such as we find realized in the shells of Chiton and in the plugs which form in the remarkable transitory “shell-sac” or “shell-gland” of Molluscan embryos (see figs. 26, 60). Aplysia, like other Mollusca, develops a primitive shell-sac in its trochosphere stage of development, which disappears and is succeeded by a nautiloid shell (fig. 36). This forms the nucleus of the adult shell, and, as the animal grows, becomes enclosed by a reflection of the mantle-skirt. When the shell of an Aplysia enclosed in its mantle is pushed well to the left, the sub-pallial space is fully exposed as in fig. 39, and the various apertures of the body are seen.