The rudiment of the bud is in typical cases composed of two vesicles, an outer derived from the ectoderm of the parent and enclosing free blood-cells (mesodermal) between its wall and that of the inner vesicle—which is usually of endodermal origin, but in Botryllidae is derived from the peribranchial sac, an ectodermal structure. The inner vesicle, derived in the two cases from different germ-layers, forms the same organs of the bud, and these organs may be of widely different origin in the larva. Moreover, free cells of the blood may play in the bud a very important part, and give rise (Perophora) to such important systems as pericardium and heart, neural tube and ganglion, the gonads and their ducts, some of which are of ectodermal and others of endodermal origin in the larva.

In some cases of precocious budding (blastogenetic acceleration) the young buds begin to appear during the tailed larval stage. The larva may even contain a first blastozooid (bud) with a branchial sac as large as that of the oozooid (derived from the egg); and in the Diplosomatidae the larva (see Fig. 42, F), when it settles down, may be already a small colony of three young ascidiozooids.

The larvae in most Compound Ascidians, in place of adhering papillae, have several or even a considerable number of ectodermal tubes or prolongations from the body (see Fig. 42, E and F) into the surrounding test. These apparently aid in the formation of the common test of the young colony, which grows over and adheres to foreign objects.

There are many irregularities in the larval development of Compound Ascidians, due to the very different amount of food-yolk present in the ova in different genera. In some cases there is even dimorphism, two forms of larvae being found in the same colony.

Compound Ascidians are amongst the most varied and brilliant of sessile animals seen at low tide on our own and most other coasts. Some are stalked and form club-shaped or knob-like outgrowths. Others again form flat gelatinous expansions attached to sea-weeds or stones, and are symmetrically marked with bright spots of colour in the form of circles, meandering lines, or star-like patterns. In such colonies each spot of colour or ray of a star represents an ascidiozooid or member of the colony, equivalent to the whole animal in the case of the solitary Simple Ascidian.

Group A. MEROSOMATA.

Viscera posterior to branchial sac; budding stolonial.

Fam. 1. Distomatidae.—Ascidiozooids divided into two regions, a thorax, containing the branchial sac, and an abdomen, with the remaining viscera (Fig. 47, B); testes numerous; vas deferens not spirally coiled. The chief genera are—Distoma, Gaertner, with some British species; Chondrostachys, Macdonald, Cystodytes, v. Drasche, with calcareous plate-like spicules in the test (Fig. 50, A); Distaplia, Della Valle, and Colella, Herdman, forming a pedunculated colony (Fig. 47, A), in which the ascidiozooids (Fig. 47, B) are provided with large incubatory pouches, opening from the peribranchial cavity, but also connected, as Bancroft^[101] has recently shown, with the end of the oviduct (see Fig. 47, B). In these pouches the embryos undergo their development, and are set free by the decay of the top of the colony. The stolons pass from the ascidiozooids in the upper part of the colony down into the stalk, and there produce buds which gradually work up to the top of the stalk, where they take their places as young ascidiozooids. At the top of the colony the old ascidiozooids die and are removed (see Fig. 47, A). Caullery has shown that in this genus there may be dimorphism in the buds, some of them placed deeply in the stalk having a large amount of reserve food-matter in their ectoderm, and remaining dormant until required to regenerate the "head" or upper part of the colony when it is lost. This genus was made known by the "Challenger" expedition. The species are mostly tropical, or from southern seas.

Fam. 2. Coelocormidae.—Colony not fixed, having a large axial cavity with a terminal aperture. Branchial apertures five-lobed. This includes one species, Coelocormus huxleyi, Herdman, which is in some respects a transition-form between the ordinary Compound Ascidians (e.g. Distomatidae) and the Ascidiae Luciae (Pyrosoma, see p. 90).

Fam. 3. Didemnidae.—Colony usually thin and incrusting. Test containing stellate calcareous spicules (Figs. 49 and 50, B). Testis single, large; vas deferens spirally coiled (Fig. 49). The chief genera are—Didemnum, Savigny, in which the colony is thick and fleshy, and there are only three rows of stigmata on each side of the branchial sac; and Leptoclinum, Milne-Edwards, in which the colony is thin and incrusting (Fig. 49), and there are four rows of stigmata. Colonies of Leptoclinum, forming thin white, grey, or yellow crusts under stones at low water, are amongst the commonest of British Compound Ascidians.

Fam. 4. Diplosomatidae.—Test reduced in amount (Fig. 51), rarely containing spicules. Vas deferens not spirally coiled. In Diplosoma, Macdonald, and other allied genera (Fig. 51), the larva is gemmiparous (Fig. 42, F). Some species are common British forms, especially on Zostera-beds and amongst seaweeds.

Fam. 5. Polyclinidae.—Ascidiozooids divided into three regions—thorax, abdomen, and post-abdomen (Fig. 46, C). Testes numerous; vas deferens not spirally coiled. The chief genera are—Pharyngodictyon, Herdman, with stigmata absent or modified, containing one species, Ph. mirabile (Fig. 44, C), the only Compound Ascidian known from a depth of 1000 fathoms; Polyclinum, Savigny, with a smooth-walled stomach (Fig. 52, A); Aplidium, Savigny, with the stomach-wall longitudinally folded (Fig. 52, B); Morchellium, Giard, with an "areolated" stomach (Fig. 52, D), bearing knobs on the outside; and Amaroucium, Milne-Edwards, in which the ascidiozooid has a long post-abdomen and a large atrial languet, and where the stomach-wall shows longitudinal ridges breaking up into knobs (pseudo-areolated, Fig. 52, C). The last four genera contain many common British species.

Many of the Compound Ascidians die down in winter; but amongst Polyclinidae, as in Clavelina, a form of hibernation is found, the old ascidiozooids dying, but some of the buds in the basal part of the colony accumulating a large store of reserve-material in their ectoderm, and lying dormant until spring, when they regenerate the colony.

Group B. HOLOSOMATA.

Body short, compact, with viscera by the side of branchial sac; budding parietal

Fam. 6. Botryllidae.—Ascidiozooids grouped in systems round common cloacal apertures (Fig. 53). Ascidiozooids having the intestine and reproductive organs by the side of the branchial sac (Fig. 46, A, p. 82). Dorsal lamina and internal longitudinal bars present in the branchial sac. Neural gland, as in Cynthiidae, dorsal to the ganglion in place of ventral as in the majority of Tunicata. The chief genera are—Botryllus, Gaertn. and Pall, with simple stellate systems (Fig. 53), and Botrylloides, Milne-Edwards, with elongated or ramified systems. There are many species of both these genera, which form brilliantly coloured fleshy crusts under stones and on sea-weeds at low tide. They are amongst the commonest and the most beautiful of British Ascidians. Both genera contain species remarkable for the rich profusion of ectodermal "vessels" which ramify and anastomose in the colonial test. On the margins of the colony these vessels end in knob-like dilatations, the ampullae (Fig. 46, A, t.k), which are said by Bancroft to pulsate rhythmically, and so aid in keeping up the colonial circulation. They are also storage reservoirs for the blood, doubtless help in respiration, and are organs for the secretion of the test-matrix.

Fam. 7. Polystyelidae.—Ascidiozooids not grouped in systems; branchial and atrial apertures four-lobed; branchial sac may be folded; internal longitudinal bars present. The chief genera are—Thylacium, Carus, with the ascidiozooids projecting above the general surface of the colony; Goodsiria, Cunningham, with the ascidiozooids completely imbedded in the investing mass (Fig. 54); and Chorizocormus, Herdman, with the ascidiozooids united in little groups which are connected by stolons. The last genus contains one species, Ch. reticulatus, in some respects a transition-form between the other Polystyelidae and the Styelinae amongst Simple Ascidians.

Budding in Holosomata—In the Polystyelidae, according to Ritter,^[102] the budding is of the same type as in Botryllidae, the bud arising in each case from the lateral body-wall of the parent.

In Botryllus^[103] the oozooid formed from the larva gives rise at a very early period to the first blastozooid of the future colony. This then forms the two buds of the second generation on its sides (see Fig. 55), and these in their turn form the third, and these the fourth generation, in which there are thus eight blastozooids; and so the process goes on, the buds of each generation arranging themselves in a circle to form a system. As each new generation makes its appearance, the preceding one undergoes degeneration, and is eventually absorbed. Consequently, in a system there can usually be seen, in addition to the adult members, certain older ones in various stages of degeneration and removal, and certain younger ones arising as buds on the sides of their predecessors, or just separated from them, and ready to take their places as young ascidiozooids in the system. Three distinct generations are thus commonly seen in a system. Now and again one or two young ascidiozooids become squeezed by the pressure of their neighbours out of a system into the surrounding test, and so give rise to new systems which add to the extent of the colony.

Sub-Order 3. Ascidiae Luciae.

Free-swimming pelagic colonies having the form of a hollow cylinder closed at one end (Fig. 56). The ascidiozooids forming the colony are imbedded in the common test in such a manner that the branchial apertures open on the outer surface and the atrial apertures on the inner surface next to the central cavity of the colony. They are placed with their ventral surfaces towards the closed end (Fig. 56, C). The first ascidiozooids of a colony are produced by gemmation from a stolonic prolongation of an imperfect oozooid or rudimentary larva (the "cyathozooid"), developed sexually. The subsequent ascidiozooids are formed from these as buds on a ventral stolon.

This sub-order includes a single family, the Pyrosomatidae, containing one well-marked genus Pyrosoma, Péron, with about six species. They are found swimming near the surface of the sea, chiefly in tropical latitudes, and are brilliantly phosphorescent. A fully developed Pyrosoma colony may be from an inch or two to upwards of twelve feet in length.

The Colony.—The shape of the colony is seen in Fig. 56, A. It tapers slightly towards the closed end, which is rounded. The opening at the opposite end may be reduced in size (see B and C), by the presence of a membranous prolongation of the common test, which can be contracted or expanded by means of the muscle-bands it receives from the atrial siphons of neighbouring zooids. The branchial apertures of the ascidiozooids are mostly placed upon short (in some cases longer) papillae projecting from the general surface, and many of the ascidiozooids have long conical processes of the test extending outwards beyond their branchial apertures (Fig. 57, t′). There is only a single layer of adult ascidiozooids in the wall of the Pyrosoma colony, as all the fully developed ascidiozooids are placed with their antero-posterior axes at right angles to the surface and communicate by their atrial apertures with the central cavity (Fig. 56, C). Their dorsal surfaces are turned towards the open end of the colony, and the buds are given off from their ventral edges (Fig. 57).

Anatomy.—The more important points in the structure of the ascidiozooid of Pyrosoma are shown in Fig. 57. A circle of tentacles, of which one, placed ventrally (tn), is larger than the rest, is found just inside the circular branchial aperture. From this point a wide cavity, with a few circularly placed muscle-bands running round its walls, leads back to the large branchial sac (br.s.), which occupies the greater part of the body. The large stigmata are elongated transversely (dorso-ventrally), and are crossed by internal longitudinal bars running antero-posteriorly. The dorsal lamina is represented by a series of eight or ten languets. The nerve-ganglion (on which is placed a small pigmented sense-organ, the unpaired "eye"), the neural gland, the dorsal tubercle, the peripharyngeal bands and the endostyle are placed in the usual positions. On each side of the anterior end of the branchial sac, close to the peripharyngeal bands is a mass of rounded mesodermal gland-cells (l.o), which are the source of the phosphorescence. They are apparently modified leucocytes lying in blood-sinuses. The alimentary canal is placed posteriorly to the branchial sac, and the anus opens into a large peribranchial or atrial cavity, of which only the median posterior part (cl), is shown in Fig. 57. The heart (Ht) lies between the posterior end of the branchial sac and the intestine, close to where the endostyle is prolonged outwards to form the inner tube of the ventral stolon. The reproductive organs are developed from a cord of germinal tissue which forms a part of every budding stolon, and so establishes a continuity of origin between the ova of successive generations of Pyrosoma. On the ventral edge of the body, immediately behind the stolon, with part of which it is continuous, a portion of this germinal tissue gives rise to a lobed testis (tes), and to a single ovum surrounded by indifferent or follicle-cells.

Development and Life-History.—The development takes place within the body of the parent, in a part of the peribranchial cavity. It is a "direct" development, the tailed larval stage being omitted. The segmentation is incomplete or "meroblastic," and an elongated embryo is formed on the surface of a mass of food-yolk. Follicle-cells, or kalymmocytes, migrate into the embryo, where they aid in its nutrition. The embryo (or young oozooid),^[104] after the formation of an alimentary cavity, a tubular nervous system, and a pair of laterally placed atrial tubes, divides into an anterior and a posterior part (see Fig. 58). The anterior and ventral part, or stolon, then segments into four pieces (the tetrazooids or first blastozooids),^[104] which afterwards develop into the first ascidiozooids of the colony, while the posterior part remains in a rudimentary condition, and is what was called by Huxley the "cyathozooid" (Fig. 58, cy). This is really the degenerate oozooid, and eventually atrophies without having completed its development, but having precociously given rise to the budding stolon.

As the four ascidiozooids increase in size, they grow round the cyathozooid and soon encircle it (Fig. 58, B). In this condition the young colony leaves the body of the parent and becomes free. The cyathozooid absorbs the nourishing yolk upon which it lies, and distributes it to the ascidiozooids by means of a heart and system of vessels which have been meanwhile formed. When the cyathozooid atrophies and is absorbed, its original atrial aperture remains and deepens to become the central cavity^[105] of the young colony, which now consists of four ascidiozooids placed in a ring, around where the cyathozooid was, and enveloped in a common test. The test is at first formed by the ectoderm cells of the cyathozooid. Later it becomes invaded by mesoblast cells from the ascidiozooids in the usual manner. The colony gradually increases by the formation of buds from these four original ascidiozooids. The young colony is, in some species, at first male, and only becomes hermaphrodite when it has attained to some size.

Occurrence.—The half-dozen known species of Pyrosoma are widely distributed over the great oceans, although they are probably most abundant in tropical waters. Pyrosoma atlanticum, Péron, and P. giganteum, Lesueur, are the commonest forms. Although sometimes abundant in the Mediterranean and the North Atlantic they have apparently not been found in British seas. P. elegans, Lesueur, is a Mediterranean form allied to the last two; and P. minatum and P. aherniosum, Seeliger, were discovered during the German "Plankton" expedition in the tropical Atlantic. Finally, the enormous P. spinosum, Herdman, was found by the "Challenger" in both North and South Atlantic in 1873; and some years later (Perrier's P. excelsior) by the French "Talisman" expedition in the tropical Atlantic. The late Professor Moseley said of this ("Challenger") species, "I wrote my name with my finger on the surface of the giant Pyrosoma as it lay on deck in a tub at night, and my name came out in a few seconds in letters of fire." Bonnier and Pérez have recently recorded that they saw an enormous profusion of a large Pyrosoma (up to four metres in length) in the Arabian part of the Indian Ocean.

Order III. Thaliacea (Salpians).

Free-swimming pelagic forms of moderate size, which may be either simple or compound, and in which the adult is never provided with a tail or notochord. Consequently the whole body here corresponds to the trunk only of the Appendicularian without the tail. The test is permanent, and may be either well developed or very slight. In all cases it is clear and transparent. The musculature of the body-wall is in the form of more or less complete circular bands, by the contraction of which water is ejected from the body, and so locomotion is effected. The branchial sac has either two large, or many small, stigmata, leading to a single peribranchial cavity, into which the anus also opens. Blastogenesis takes place from a ventral, endostylar stolon. Alternation of generations occurs in the life-history, and may be complicated by polymorphism. The Order Thaliacea comprises two groups, Cyclomyaria (such as Doliolum) and Hemimyaria (such as Salpa).

Sub-Order 1. Cyclomyaria.

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.

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.

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.

(2) Some of the median buds become foster forms ("phorozooids"), which, like the preceding trophozooids, do not become sexually mature, but, unlike them, are eventually set free as cask-shaped bodies having the Doliolum appearance, with eight encircling muscle-bands, and having, moreover, a ventral outgrowth (not a stolon), which is formed of the stalk by which the body was formerly attached to the dorsal process of the oozooid. On this ventral outgrowth the "gonozooids" (3) are attached while still very young buds, and after the phorozooids are set free these reproductive forms gradually attain their complete development, become sexually mature, and are eventually separated off, finally losing all trace of their temporary connexion with the foster-forms. They resemble the foster-forms in having a cask-shaped body with eight muscle-bands, but differ in the absence of a ventral process, and in having the sexual reproductive organs fully developed.

Occurrence.—The best-known member of the genus is Doliolum tritonis, Herdman, which was captured in the tow-nets in thousands by Sir John Murray during the cruise of H.M.S. "Triton" in the summer of 1882 in the North Atlantic. Since then that species, or the closely allied D. nationalis, Borgert, have been found on more than one occasion in the English Channel and other parts of our south-west coast, and so Doliolum may be regarded as an occasional member of the British surface fauna.

It is probable that the occasional phenomenal swarms of Doliolum which have been met with in summer in the North Atlantic are a result of the curious life-history which, under favourable circumstances, allows of a small number of oozooids producing from minute buds an enormous number of phorozooids and gonozooids.

As the result of the careful quantitative work of the German "Plankton" expedition, Borgert thinks that the temperature of the water has more to do with both the horizontal and the vertical distribution of these Thaliacea in the sea than any other factor.

Other Genera.—Anchinia, of which only one species is known, A. rubra, Vogt, from the Mediterranean, has the sexual forms permanently attached to portions of the dorsal outgrowth from the body of the unknown oozooid ("nurse"). The stolon is probably much longer than in Doliolum, and curves round so as to reach and lie along the dorsal outgrowth, upon which it places the buds.

The body of the adult is elongated dorso-ventrally. The test is well developed and contains branched cells. The musculature is not so well developed as in Doliolum. There are two circular bands at the anterior end, two at the posterior, and two muscles on the middle of the body, which unite to form the characteristic S-shaped lateral bands. The stigmata are confined to the obliquely-placed posterior end of the branchial sac. The alimentary canal forms a U-shaped curve. The reproductive organs are placed on the right side of the body. The life-history is still imperfectly known. As in the case of Doliolum the sexual generation is polymorphic, and has three forms, two of which remain in a rudimentary condition so far as the reproductive organs are concerned. They are known as the first and second sterile forms, or "trophozooids." In Anchinia, however, the three forms do not occur, so far as we know, together at the same time on the one outgrowth, but are produced successively, or in different regions, the reproductive forms of the sexual generation being independent of the "foster-forms."^[106]

The third genus, Dolchinia, contains also only a single species, D. mirabilis, found by Korotneff^[107] in the Gulf of Naples. It must have three different forms in its life-history—oozooid, phorozooid, and gonozooid, but the first of these is still unknown. On what must be body processes detached from the oozooid are found phorozooids somewhat like those of Doliolum, bearing sexual forms attached to ventral stalks. Dolchinia is intermediate on the whole between Anchinia, the most simple member of the family, and Doliolum the most complex; and may eventually come to be united with the latter genus.

Sub-Order 2. Hemimyaria.

Free-swimming pelagic forms which exhibit alternation of generations in their life-history, and in the sexual condition form colonies. The body is more or less fusiform, with the long axis antero-posterior, and the branchial and atrial apertures nearly terminal and opposite. The test is well developed but transparent. The musculature of the body-wall is in the form of a series of transversely-running bands which do not usually form complete independent rings as in the Cyclomyaria. These partially-encircling muscles in the Salpidae (see Fig. 61, m.b) are probably to be regarded as modified branchial and atrial sphincters which have spread over the intervening body. The branchial and peribranchial (cloacal) cavities form a continuous space in the interior of the body, opening externally at the ends by the branchial and atrial apertures, and traversed obliquely from the dorsal and anterior to the ventral and posterior end by a long narrow vascular ciliated band, which represents the dorsal lamina, the dorsal blood-sinus, and the neighbouring parts of the dorsal edge of the branchial sac of an ordinary Ascidian. The alimentary canal is placed ventrally. It may either be stretched out so as to extend for some distance anteriorly, or, as is more usual, be concentrated to form along with the testis a rounded opaque mass near the posterior end of the body, known as the visceral mass or "nucleus." The embryonic development is direct, no tailed larva being formed. The embryo is united to the parent for a time by a "placenta."

This sub-order contains, in addition to its typical members, the Salpidae, another still somewhat problematical family the Octacnemidae, including a single very remarkable deep-water genus (Octacnemus), which in some respects does not conform with the characters given above, and exhibits a certain amount of affinity with the primitive fixed forms from which Salpidae have been derived.

Occurrence and Reproduction.—The family Salpidae^[108] includes the single genus Salpa, Forskål, which, however, may be divided into two well-marked groups of species—(1) those such as S. (Cyclosalpa) pinnata, in which the alimentary canal is stretched out ("ortho-enteric" condition) along the ventral surface of the body, and (2) those such as S. runcinata-fusiformis, in which the alimentary canal forms a compact globular mass (Fig. 61, v), the "nucleus" ("caryo-enteric" condition), near the posterior end of the body. About fifteen species altogether are known; they are all pelagic in habit, and are found in nearly all seas. Each species occurs in two forms (Fig. 61, A and B), the solitary asexual (proles solitaria), and the aggregated sexual (proles gregaria), which are in most species quite unlike one another, the aggregated form being usually more rounded, ovoid, or fusiform (Fig. 61, A), and the solitary more quadrangular, and often provided with conical processes or projecting points.

The solitary form gives rise, by gemmation at the posterior end of the endostyle (Fig. 63), to a complex tubular stolon, containing processes from the more important organs of the parent-body, which give rise to an endodermal tube, two peribranchial tubes, a neural tube, two blood-sinuses and mesoblast cells, a genital cord, and over all the ectodermal covering (see Fig. 64). This stolon becomes segmented (Fig. 63) into a series of buds or young "chain" individuals, of which there may be several hundreds. As the stolon elongates (Fig. 61, B, st″), the buds undergo lateral shifting, and rotation round their longitudinal axis, so as to acquire the relations seen in the "chain," which then emerges from the tube in the test through which it has been growing, so as to project to the exterior near the atrial aperture. The buds at its free end which have now become far advanced in their development are set free in groups, which remain attached together by processes of the test, each enclosing a diverticulum from the body-wall (Fig. 62), so as to form "chains." Each member of the chain is a Salpa of the sexual or aggregated form, and when mature may—either still attached to its neighbours or separated from them—produce one or several embryos (Fig. 61, A, emb), which develop into the solitary form of Salpa. Thus the two forms, different in appearance and structure and different in mode of origin, alternate regularly in the life-history of Salpa.

Structure.—The more important points in the structure of a typical Salpa are shown in Fig. 65. The branchial and atrial apertures are at opposite ends of the body, and lead into large cavities, the branchial and peribranchial sac respectively, which are in free communication at the sides of the obliquely-running dorsal lamina or "gill" (d.l). The transparent test is usually thick, and varies from a gelatinous to a stiff cartilaginous condition; it adheres closely to the surface of the mantle (ectoderm and body-wall). The muscle-bands (from 4 to about 20—usually 8 or 10) of the mantle do not in most cases completely encircle the body. They are present dorsally (Fig. 65, mus.bds) and laterally, but the majority do not reach the ventral surface. In many cases neighbouring bands join in the median dorsal line (Fig. 61). The muscle fibres are striated, and have rows of large equidistant nuclei. The anterior end of the dorsal lamina is in some cases prolonged to form a prominent tentacular organ, the languet or dorsal tentacle, projecting into the branchial sac, while near this opens a ciliated funnel corresponding to the dorsal tubercle, but having no connexion in the adult with either ganglion or subneural gland. The conjoined ganglion and subneural gland, the dorsal lamina, the peripharyngeal bands and the endostyle are placed in the usual positions. Eyes in the form either of a continuous horse-shoe-shaped pigmented ridge on the dorsal surface of the ganglion immediately below the ectoderm, or of one larger median and several smaller lateral ocelli are found in the various species of Salpa. These eyes have in most cases a retina formed of elongated cells, and a pigment-layer placed upon the ganglion.

The so-called otocysts of Salpa have been shown by Metcalf to be really glandular organs. They have been called lateral neural glands; they do not open at the dorsal tubercle, but separately into the pharynx. These lateral neural tubular glands have also been regarded as nephridia.

The large spaces at the sides of the dorsal lamina (often called the gill or branchia of Salpa), by means of which the cavity of the branchial sac is placed in free communication with the peribranchial cavity, are to be regarded as gigantic gill-slits formed by the suppression of the lateral walls and small stigmata of the branchial sac. The alimentary canal at the posterior end of the "gill" consists of oesophagus, stomach, and intestine, with a pair of lateral gastric glands or caeca. These viscera along with the reproductive organs, when present, make up the "nucleus" (Fig. 66, v).