1911 Encyclopædia Britannica/Sipunculoidea

by Arthur Everett Shipley

SIPUNCULOIDEA, marine animals of uncertain affinities, formerly associated with the Echiuroidea (q.v.) in the group Gephyrea. Externally, the body of a Sipunculoid presents no projections: its surface is as a rule even, and often glistening, and the colour varies from whitish through yellow to dark brown. The anterior one-quarter or one-third of the body is capable of being retracted into the remainder, as the tip of a glove-finger may be pushed into the rest, and this retractile part is termed the introvert. At the tip of the introvert the mouth opens, and is surrounded in Sipunculus by a funnel-shaped, ciliated lophophore (figs. 1 and 2). In Phascolosoma and Phascolion this funnel-shaped structure has broken up into a more or less definite group of tentacles, which in Dendrostoma are arranged in four groups. In Aspidosiphon and Physcosoma the tentacles are usually arranged in a horse-shoe, which may be double, overhanging the mouth dorsally. On the surface of the funnel-shaped lophophore are numerous ciliated grooves, and each of the tentacles in the tentaculated forms has a similar groove directed towards the mouth. These grooves doubtless serve to direct currents of water, carrying with them small organisms towards the mouth.

The skin consists of a layer of cuticle, easily stripped off, secreted by an ectodermal layer one cell thick. Within this is usually a sheath of connective tissue, which surrounds a layer of circular muscles; the latter may be split up into separate bundles, but more usually form a uniform sheet. Within the circular muscles is a layer of longitudinal muscles, very often broken into bundles, the number of which is often of specific importance. Oblique muscles sometimes lie between the circular and longitudinal sheaths. On the inner surface is a layer of peritoneal epithelium, which is frequently ciliated, and at the bases of the retractor muscles is heaped up and modified into the reproductive organs. The ectoderm is in some genera modified to form certain excretory glands, which usually take the form of papillae with an apical opening. These papillae give the surface a roughened aspect; the use of their secretion is unknown. They are best developed in Physcosoma.

When the body of a Sipunculoid is opened, it is seen that the body-cavity is spacious and full of a corpusculated fluid, in which the various organs of the body float. The most conspicuous of these is the long, white alimentary canal, crowded with mud. The mouth is devoid of armature, and passes without break into the oesophagus; this is surrounded by the retractor muscles, which are inserted into the skin around the mouth, and have their origin in the body-wall, usually about one-third or one-half of the body-length from the anterior end (figs. 1 and 2). Their function is to retract the introvert, which is protruded again by the contraction of the circular muscles of the skin; these, compressing the fluid of the body-cavity, force forward the anterior edge of the introvert. The number of muscles varies from one (Onchnesoma and Tylosoma) to four, the latter being very common. The alimentary canal is U-shaped, the dorsal limb of the U terminating in the anus, situated not very far from the level of the origin of the retractor muscles. The limbs of the U are further twisted together in a looser or tighter coil, the axis of which may be traversed by a “spindle” muscle arising from the posterior end of the body. No glands open into the alimentary canal, but a diverticulum, which varies enormously in size, opens into the rectum. As is so often the case with animals which eat mud and sand, and extract what little nutriment is afforded by the organic débris therein, the walls of the alimentary canal are thin and apparently weak. All along one side is a microscopic ciliated groove, into which the mud does not seem to enter, and along which a continuous stream of water may be kept up. Possibly this is respiratory—there are no special respiratory organs. A so-called heart lies on the dorsal surface of the oesophagus; it is closed behind, but in front it opens into a circumoesophageal ring, which gives off vessels into the lophophore and tentacles. The contraction of this heart, which is not rhythmic, brings about the expansion of the tentacles and lophophore. This system is in no true sense a vascular system; there are no capillaries, and the fluid it contains, which is corpusculated, can hardly have a respiratory or nutritive function. It is simply a hydrostatic mechanism for expanding the tentacles. The excretory organs are typical nephridia, with an internal ciliated opening into the body-cavity, and an external pore. One surface of the tube is prolonged into a large sac lined with glandular excretory cells. The organs are typically two, though one is often absent, e.g. in Phascolion. They serve as channels by which the reproductive cells leave the body, and they are sometimes spoken of as “brown tubes.” There is a well-developed brain dorsal to the mouth; this gives off a pair of oesophageal commissures, which surround the oesophagus and unite in a median ventral nerve-cord which runs between the longitudinal muscles to the posterior end of the body. From time to time it gives off minute circular nerves, which run round the body in the skin and break up into a very fine nerve plexus. There are no distinct ganglia, but ganglion cells are uniformly distributed along the ventral side of the cord. The whole is anteriorly somewhat loosely slung to the skin, so as to allow free play when the animal is extending or retracting its introvert. A pit or depression, known as “the cerebral organ,” opens into the brain just above the mouth; this usually divides into two limbs, which are deeply pigmented and have been called eyes.

Fig. 1.—Sipunculus nudus, L., with introvert and head fully extended, laid open by an incision along the right side to show the internal organs.

a,	Mouth.
b,	Ventral nerve-cord.
c,	“Heart.”
d,	Oesophagus.
e,	Intestine.
f,	Position of anus.
g,	Tuft-like organs.
h,	Right nephridium.
i,	Retractor muscles.
j,	Diverticulum on rectum. The spindle-muscle is seen overlying the rectum.

Sipunculoids are dioecious, and the ova and spermatozoa are formed from the modified cells lining the body-cavity, which are heaped up into a low ridge running along the line of origin of the retractor muscles. The ova and the mother-cells of the spermatozoa break off from this ridge, and increase in size considerably in the fluid of the body-cavity. Fertilization is external; and in about three days a small ciliated larva, not unlike that of the Echiuroids, but with no trace of segmentation, emerges from the egg-shell. This little creature, which has many of the features of a Trochosphere larva, swims about at the surface of the sea for about a month and grows rapidly. At the end of this time it undergoes a rapid metamorphosis: it loses many of its larval organs, cilia, takes in a quantity of water into its body-cavity, sinks to the bottom of the sea, and begins life in its final form.

Fig. 2.—Right half of the anterior end of Sipunculus nudus, L., seen from the inner side and magnified.

a,	Funnel-shaped grooved tentacular crown leading to the mouth.
b,	Oesophagus.
c,	Strands breaking up the cavity of the tentacular crown into vascular spaces.
c′,	“Heart.”
d,	Brain.
e,	Ventral, and e′, dorsal retractor muscles.
f,	Ventral nerve-cord.
g,	Vascular spaces in tentacular crown.

The following genera of Sipunculoids are recognized:—(i.) Sipunculus. This, with Physcosoma, has its longitudinal muscles divided up into some 17-41 bundles. It has no skin papillae. The members of this genus attain a larger size than any other species, and the genus contains some 16-17 species. (ii.) Physcosoma (fig. 3) has its body covered with papillae, and usually numerous rows of minute hooks encircling the introvert. It is the most numerous genus, and consists for the most part of shallow-water (less than 50 fathoms) tropical and subtropical forms. They often live in tubular burrowings in coral-rock. The following three genera have their longitudinal muscles in a continuous sheath:—(iii.) Phascolosoma, with some 25 species, mostly small, with numerous tentacles. (iv.) Phascolion, 10 species, small, living in mollusc-shells and usually adopting the coiled shape of their house; only one kidney, the right, persists. (v.) Dendrostoma, with 4-6 tentacles, a small genus found in tropical shallow water. (vi.) Aspidosiphon, with 19 species, is easily distinguished by a calcareous deposit and thickened shield at the posterior end and at the base of the introvert, which is eccentric. (vii.) Cloeosiphon has a calcareous ring, made up of lozenge-shaped plates, round the base of its centric introvert. (viii.) Petalostoma, a minute form with two leaf-like tentacles, is found in the English Channel, (ix.) Onchnesoma, with 2 species, and (x.) Tylosoma, with 1 species, have no tentacles, only one brown tube, and only one retractor muscle. Both genera are found off the Norwegian coast. The last named is said to have numerous papillae and no introvert.

Fig. 3.—A semi-diagrammatic figure of the anterior end of half a Physcosoma, seen from the inner side. The introvert is fully everted and the lophophore expanded. The collar which surrounds the head is not fully extended. Two rows only of hooks are shown.

1,	Lophophore.
2,	Pigmented pit leading to brain.
3,	Section of dorsal portion of mesoblastic “skeleton.”
4,	Pit ending in eye.
5,	The brain.
6,	Blood-sinus of dorsal side surrounding brain and giving off branches to the tentacles.
7,	Collar.
8,	Retractor muscle of head.
9,	Hook.
10,	Sense-organ.
11,	Nerve-ring.
12,	Coelom of upper lip; it is continuous with 21.
13,	Mouth.
14,	Lower lip.
15,	Blood-sinus of ventral side, continuous with 6.
16,	Ventral portion of mesoblastic “skeleton.”
17,	Ventral nerve-cord.
18,	Coelom, continuous with 12 and 21.
19,	Oesophagus.
20,	Dorsal vessel arising from the blood-sinus 6.
21,	Coelom.

Authorities.—Selenka, “Die Sipunculiden,” Semper's Reisen (1883), and Challenger Reports, xiii. (1885); Sluiter, Natuurk. Tijdschr. Nederl. Ind. xli. and following volumes; Andrews, Stud. Johns Hopkins Univ. iv. (1887-1890); Ward, Bull. Mus. Harvard, xxi. (1891); Hatschek, Arb. Inst. Wien, v. (1884); Shipley, Quart. J. Micr. Sci. xxxi. (1890), xxxii. (1891), and xxxiii. (1892); P. Zool. Soc. London (1898), and Willey's Zoological Results, pt. 2 (1899); Horst, Niederland. Arch. Zool., Supplementary, vol, i.

(A. E. S.)