Free-swimming pelagic forms which exhibit alternation of generations in their life-history, but never form permanent colonies. The body is cask-shaped, with the branchial and atrial apertures at the opposite ends. The test is moderately well developed, never much thickened. The musculature is mostly in the form of complete circular bands surrounding the body. The branchial sac is fairly large, occupying the anterior half or more of the body. Stigmata are usually present in its posterior part only. The peribranchial cavity is mainly posterior to the branchial sac. The alimentary canal is placed ventrally, close to the posterior end of the branchial sac. Hermaphrodite reproductive organs lie ventrally near the intestine.
This group is clearly distinguished from the second sub-order, the Hemimyaria, by the condition of the muscle-bands and of the branchial sac, and by the life-history. The muscle-bands are complete rings (except in Anchinia), while in the Hemimyaria they are always more or less incomplete. The branchial sac in the Cyclomyaria is a distinct cavity, and communicates with the peribranchial cavity only by small slits or stigmata. The life-history is also very characteristic, as the sexual generation in the Cyclomyaria is always polymorphic, while in the Hemimyaria it consists of one form only.
Fig. 59.—Sexual generation of Doliolum tritonis, Herdman, from left side, × 10. at, Atrial aperture; at.l, atrial lobes; at.m, wall of atrium; br, branchial aperture; br.l, branchial lobes; br.s, branchial sac; d.t, dorsal tubercle; end, endostyle; h, heart; i, intestine; m, mantle; m1-m8, circular muscle-bands; n, nerve; n.g, nerve-ganglion; ov, ovary; p.br, peribranchial cavity; p.p, peripharyngeal bands; sg, stigmata; s.gl, neural gland; s.o, sense-organ; st, stomach; t, test; tes, testis; z, prebranchial zone. (After Herdman.)
Structure of Doliolum.—The single family Doliolidae includes three genera, Doliolum, Quoy and Gaimard, Dolchinia, Korotneff, and Anchinia, Eschscholtz. Doliolum, of which about a dozen species are known, from various seas, has a cask-shaped body (Fig. 59), usually from 1 to 2 cm. in length. The terminal branchial and atrial apertures are lobed, and the lobes are provided with sense-organs. The test is a thin but tough transparent layer, and contains no "test" cells. It is merely a cuticle covering the surface of the squamous ectoderm. The body-wall has eight or nine circular muscle-bands surrounding the body. The most anterior and posterior of these form the branchial and atrial sphincters. The wide branchial and atrial apertures lead respectively into branchial and peribranchial cavities separated by the posterior and postero-lateral walls of the branchial sac which are pierced by a considerable number of small stigmata; consequently there is a free passage for the water through the body along its long axis, and the animal swims by contracting its ring-like muscle-bands so as to force out the contained water posteriorly. When stigmata are found on the lateral walls of the branchial sac (see Fig. 59) there are corresponding anteriorly directed diverticula of the peribranchial cavity. There is a distinct endostyle on the ventral edge of the branchial sac and a peripharyngeal band surrounding its anterior end, but there is no representative of the dorsal lamina along its dorsal edge; and there are neither branchial nor atrial tentacles. The oesophagus commences rather on the ventral edge of the posterior end of the branchial sac, and runs backwards to open into the stomach, which is followed by a curved intestine opening into the peribranchial cavity. The alimentary canal as a whole is to the right of the middle line. The hermaphrodite reproductive organs are to the left of the middle line alongside the alimentary canal. They open into the peribranchial cavity. The ovary is nearly spherical, while the testis is elongated, and may be continued anteriorly for a long distance. The heart is placed in the middle line ventrally, between the posterior end of the endostyle and the oesophageal aperture. The nerve-ganglion lies about the middle of the dorsal edge of the body, and gives off many nerves. Under it is placed the neural gland, the duct of which runs forward and opens into the anterior end of the branchial sac by a simple aperture surrounded by the spirally twisted dorsal ends of the peripharyngeal bands.
Life-History.—The ova produced by the Doliolum of the sexual generation, after a complete or "holoblastic" segmentation, and normal invagination, produce tailed larvae with a relatively small caudal appendage, and a large body in which the characteristic musculature begins to appear (Fig. 60, A). These larvae after metamorphosis lose their tails and develop into oozooids, known as "nurses," which are asexual, and are characterised (Fig. 60, B) by the possession of nine muscle-bands, by the stigmata being few in number and confined to the posterior end of the branchial sac, by an otocyst on the left side of the body, by a ventrally-placed complex stolon or "rosette organ" near the heart, from which primary buds are produced by constriction, and by a dorsal outgrowth ("the cadophore") near the posterior end of the body. The buds (blastozooids) give rise eventually, after further division, to the sexual generation, which is polymorphic—having three distinct forms, in two of which the reproductive organs remain undeveloped.
Fig. 60.—Life-history of Doliolum. A, tailed larval stage; B, "nurse" or oozooid, showing buds (blastozooids) migrating from the ventral stolon to the dorsal process; C, posterior part of much later oozooid to show buds arranged in three rows on dorsal process; D, stolon segmenting; E, young migrating bud; F, trophozooid developed from one of the buds of a lateral row. At, Atrial aperture; b, buds; Br, branchial aperture; cl, cloaca; d.p, dorsal process; end, endostyle; ht, heart; l.b, lateral buds; m.b, median buds; n.g, nerve-ganglion; ot, otocyst; p.c, pericardium; sk, stalk; sto, stolon. (After Uljanin and Barrois.)
The primary buds are constricted off while still very young and undeveloped (Fig. 60, D, B, and E); they migrate from their place of origin on the stolon, over the surface (aided by large amoeboid test-cells which become attached to the buds) (Fig. 60, B), multiply by fission, and become attached (again by the help of amoeboid test-cells and ectoderm cells which form a slight "placenta") in three rows—a median and two lateral—to the dorsal outgrowth (Fig. 60, C) of the body of the nurse. This parent-form by this time has become greatly modified, and its structure is largely sacrificed for the good of the buds or growing zooids, for which it really forms a locomotory organ. Its muscle-bands become greatly developed in width (Fig. 60, C), and the branchial meshwork, endostyle, and alimentary canal disappear.
The three forms produced in the second generation are as follows:—(1) Nutritive forms ("trophozooids") derived from the lateral rows of buds, which remain permanently attached to the oozooid, and are sacrificed for the benefit of the rest of the colony. They serve merely to aid in respiration, and to provide the food for the nurse and the median buds. Their development is arrested; they have the body elongated dorso-ventrally with a large funnel-like branchial aperture (Fig. 60, F), and the musculature is very slightly developed.