zooids arise at intervals. In Clavularia and its allies each outgrowth contains several solenia, and the outgrowths may take the form of flat expansions, composed of a number of solenial tubes felted together to form a lamellar surface of attachment. Such outgrowths are called stolons, and a stolon may be simple, i.e. contain only one solenium, as in Cornularia, or may be complex and built up of many solenia, as in Clavularia. Further complications arise when the lower walls of the mother zooid become thickened and interpenetrated with solenia, from which buds are developed, so that lobose, tufted, or branched colonies are formed. The chief orders of the Synalcyonacea are founded upon the different architectural features of colonies produced by different modes of budding. We recognize six orders the STOLONIFERA, ALCYONACEA, PSEUDAXONIA, AXIFERA, STELECHOTOKEA, and

FIG. 6.-Portion of a colony of Corallium rubrum, showing expanded and contracted zooids. In the lower part of the figure the cortex has been cut away In the order ALCYONto show the axis, ax, and the longi- ACEA the colony consists tudinal canals, le, surrounding it. of bunches of elongate cylindrical zooids, whose proximal portions are united by solenia and compacted, by fusion of their own walls and those of the solenia, into a fleshy mass called the coenenchyma. Thus the coenenchyma forms a stem, sometimes branched, from the surface of which the free portions of the zooids project. The skeleton of the Alcyonacea consists of separate calcareous spicules, which are often, especially in the Nephthyidae, so abundant and so closely interlocked as to form a tolerably firm and hard armour. The order comprises the families Xeniidae, Alcyonidae and Nephthyidae. Alcyonium digitatum, a pink digitate form popularly known as "dead men's fingers," is common in 10-20 fathoms of water off the English coasts.

In the order PSEUDAXONIA the colonies are upright and branched,

99

consisting of a number of short zooids whose proximal ends are imbedded in a coenenchyma containing numerous ramifying solenia and spicules. The coenenchyma is further differentiated into a medullary portion and a cortex. The latter contains the proximal moieties of the zooids and numerous but separate spicules. The medullary portion is densely crowded with spicules of different shape from those in the cortex, and in some forms the spicules are cemented together to form a hard supporting axis. There are four families of Pseudaxonia the Briareidae,s Sclerogorgidae, Melitodidae, and Corallidae. In the first-named the medulla is penetrated by solenia and forms an indistinct axis; in the remainder the medulla is devoid of solenia, and in the Melitodidae and Corallidae it forms a dense axis, which in the Melitodidae consists of alternate calcareous and horny joints. The precious red coral of commerce, Corallium rub-d rum (fig. 6), a member of the family Corallidae, is found at depths varying FIG. 7.-The sea-fan (Gorgoniant from 15 to 120 fathoms indepe cavolinii). Teal bon the Mediterranean Sea, chiefly on the African coast. It owes its commercial value to the beauty of its hard red calcareous axis which in life is covered by a cortex in which the proximal moieties of the zooids are imbedded. Corallium rubrum has been the subject of a beautifully-illustrated memoir by de Lacaze-Duthiers, which should be consulted for details of anatomy.

The AXIFERA comprise those corals that have a horny or calcified sponds to the axis of the axis, which in position correPseudaxonia, but, unlike it, is never formed of fused spicules; the most familiar example is the pink sea-fan, Gorgonia cavolinii, which is found in abundance in 10-25 fathoms of water off the English coasts (fig. 7). In this order the axis is formed as an ingrowth of the ectoderm of the base of the mother zooid of the colony, the cavity of the ingrowth being filled by a horny substance secreted by the ectoderm. In Gorgonia the axis remains horny throughout life, but in many forms it is further strengthened by a deposit of calcareous matter. In the family Isidinae the axis consists of alternate segments of horny and calcareous substance, the latter being amorphous. The order contains six familiesthe Dasygorgidae, Isidae, Primnoidae, Muriceidae, Plexauridae, and Gorgonidae.

A, Portion of the surface of a colony of Heliopora coerulea magnified, showing two calices and the surrounding coenenchymal tubes. B, Single zooid with the adjacent soft tissues as seen after removal of the skeleton by decalcification. Z', the distal, and Z3, the proximal or intracalicular portion of the zooid; ec, ectoderm; ct, coenenchymal tubes; sp, superficial network of solenia.

tubes, known as the coenenchymal tubes (fig. 9, A). The walls of the calices and coenenchymal tubes are formed of flat plates of calcite, which are so disposed that the walls of one tube enter into the composition of the walls of adjacent tubes, and the walls of the calices are formed by the walls of adjacent coenenchymal tubes. Thus the architecture of the Helioporid colony differs entirely from such forms as Tubipora or Favosites, in which each corallite has its own distinct and proper wall. The cavities both of the calices and coenenchymal tubes of Heliopora are closed below by horizontal partitions or tabulae, hence the genus was formerly included in the group Tabulata, and was supposed to belong to the madreporarian corals, both because of its lamellar skeleton, which resembles that of a Madrepore, and because each calicle has from twelve to fifteen radial partitions or septa projecting into its cavity. The structure of the zooid of Heliopora, however, is that of a typical Alcyonarian, and the septa have only a resemblance to, but no real homology with, the similarly named structures in madreporarian corals. Heliopora coerulea is found between tide-marks on the shore platforms of coral islands. The order was more abundantly represented in Palaeozoic times by the Heliolitidae from the Upper and Lower Silurian and the Devonian, and by the Thecidae from the Wenlock limestone. In Heliolites porosus the colonies had the form of spheroidal masses; the calices were furnished with twelve pseudosepta, and the coenenchymal tubes were more or less regularly hexagonal.

Zoantharia. In this sub-class the arrangement of the mesenteries is subject to a great deal of variation, but all the types hitherto observed may be referred to a common plan, illustrated by the living genus Edwardsia (fig. 10, A, B). This is a small solitary Zoantharian which lives embedded in sand. Its body is divisible into three portions, an upper capitulum bearing the mouth and tentacles, a median scapus covered by a friable cuticle, and a terminal

physa which is rounded. Both capitulum and physa can be retracted within the scapus. There are from sixteen to thirty-two simple tentacles, but only eight mesenteries, all of which are complete. The stomodaeum is compressed laterally, and is furnished with two longitudinal grooves, a sulcus and a sulculus. The arrangement of the muscle-banners on the mesenteries is characteristic. On six of the mesenteries the muscle-banners have the same position as in the Alcyonaria, namely, on the sulcar faces; but in the two remaining mesenteries, namely, those which are attached on either side of the sulcus, the muscle-banners are on the opposite or sulcular faces. It is not known whether all the eight mesenteries of Edwardsia are developed simultaneously or not, but in the youngest

A, Edwardsia claparedii (after A. Andres). Cap, capitulum; sc, scapus; ph, physa.

B, Transverse section of the same, showing the arrangement of the mesenteries. s, Sulcus; sl, sulculus.

C, Transverse section of Halcampa. d, d, Directive mesenteries; st, stomodaeum.

form which has been studied all the eight mesenteries were present, but only two of them, namely the sulco-laterals, bore mesenterial filaments, and so it is presumed that they are the first pair to be developed. In the common sea-anemone, Actinia equina (which has already been quoted as a type of Anthozoan structure), the mesenteries are numerous and are arranged in cycles. The mesenteries of the first cycle are complete (i.e. are attached to the stomodaeum), are twelve in number, and arranged in couples, distinguishable by the position of the muscle-banners. In the four couples o mesenteries which are attached to the sides of the elongated stomo daeum the muscle-banners of each couple are turned towards one another, but in the sulcar and sulcular couples, known as the directive d

opment in an Actinian. B, Diagrammatic transverse section of FIG. 11.-A, Diagram showing the sequence of mesenterial develGonactinia prolifera.

mesenteries, the muscle-banners are on the outer faces of the mesenteries, and so are turned away from one another (see fig. 10, C). The space enclosed between two mesenteries of the same couple is called an entococle; the space enclosed between two mesenteries of adjacent couples is called an exocoele. The second cycle of mesenteries consists of six couples, each formed in an exococle of the primary cycle, and in each couple the muscle-banners are vis-à-vis The third cycle comprises twelve couples, each formed in an exocoele between the primary and secondary couples and so on, it being a general rule (subject, however, to exceptions) that new mesenterial couples are always formed in the exocoeles, and not in the entocoeles.

While the mesenterial couples belonging to the second and each successive cycle are formed simultaneously, those of the first cycle

are formed in successive pairs, each member of a pair being placed on opposite sides of the stomodaeum. Hence the arrangement in six couples is a secondary and not a primary feature. In most Actinians the mesenteries appear in the following order: At the time when the stomodaeum is formed, a single pair of mesenteries, marked I, I in the diagram (fig. 11, A), makes its appearance, dividing the coelenteric cavity into a smaller sulcar and a large sulcular chamber. The muscle-banners of this pair are placed on the sulcar faces of the mesenteries. Next, a pair of mesenteries, marked II,II in the diagram, is developed in the sulcular chamber, its musclebanners facing the same way as those of I, I. The third pair is formed in the sulcar chamber, in close connexion with the sulcus, and in this case the muscle-banners are on the sulcular faces. The fourth pair, having its muscle-banners on the sulcar faces, is developed at the opposite extremity of the stomodacum in close connexion with the sulculus. There are now eight mesenteries present, having exactly the same arrangement as in Edwardsia. A pause in the development follows, during which no new mesenteries are formed, and then the six-rayed symmetry characteristic of a normal Actinian zooid is completed by the formation of the mesenteries V, V in the lateral chambers, and VI, VI in the sulcolateral chambers, their muscle-banners being so disposed that they form couples respectively with II, II and I, I. In Actinia equina the Edwardsia stage is arrived at somewhat differently. The mesenteries second in order of formation form the sulcular directives, those fourth in order of formation form with the fifth the sulculo-lateral couples of the adult.

As far as the anatomy of the zooid is concerned, the majority of the stony or madreporarian corals agree exactly with the soft-bodied Actinians, such as Actinia equina, both in the number and arrange

A, Zoanthid colony, showing the expanded zooids. B, Diagram showing the arrangement of mesenteries in a young Zoanthid. C, Diagram showing the arrangement of mesenteries in an adult Zoanthid. 1, 2, 3, 4, Edwardsian mesenteries.

ment of the adult mesenteries and in the order of development of the first cycle. The few exceptions will be dealt with later, but it may be stated here that even in these the first cycle of six couples of mesenteries is always formed, and in all the cases which have been examined the course of development described above is followed. There are, however, several groups of Zoantharia in which the mesenterial arrangement of the adult differs widely from that just described. But it is possible to refer all these cases with more or less certainty to the Edwardsian type.

The order ZOANTHIDEA comprises a number of soft-bodied Zoantharians generally encrusted with sand. Externally they resemble ordinary sea-anemones, but there is only one ciliated groove, the sulcus, in the stomodaeum, and the mesenteries are arranged on a peculiar pattern. The first twelve mesenteries are disposed in couples, and do not differ from those of Actinia except in size. The mesenterial pairs I, II and III are attached to the stomodacum, and are called macromesenteries (fig. 12, B), but IV. V and VI are much shorter, and are called micromesenteries. The subsequent development is peculiar to the group. New mesenteries are formed only in the sulco-lateral exococles. They are formed in couples, each couple consisting of a macromesentery and a micromesentery, disposed so that the former is nearest to the sulcar directives. The derivation of the Zoanthidea from an Edwardsia form is sufficiently obvious.

The order CERIANTHIDEA comprises a few soft-bodied Zoantharians with rounded aboral extremities pierced by pores. They have two circlets of tentacles, a labial and a marginal, and there is only one ciliated groove in the stomodaeum, which appears to be the sulculus. The mesenteries are numerous, and the longitudinal muscles, though distinguishable, are so feebly developed that there are no musclebanners. The larval forms of the type genus Cerianthus float freely in the sea, and were once considered to belong to a separate genus, Arachnactis. In this larva four pairs of mesenteries having the typical Edwardsian arrangement are developed, but the fifth and sixth pairs, instead of forming couples with the first and second, arise in the sulcar chamber, the fifth pair inside the fourth, and the

A, Cerianthus solitarius (after A. Andres).

B, Transverse section of the stomodacum, showing the sulculus, sl, and the arrangement of the mesenteries.

C, Oral aspect of Arachnactis brachiolala, the larva of Cerianthus, with seven tentacles.

D, Transverse section of an older larva. The numerals indicate the order of development of the mesenteries.

are more obscure. The type form, Antipathes dichotoma (fig. 14), forms arborescent colonies consisting of numerous zooids arranged in a single series along one surface of a branched horny axis. Each zooid has six tentacles; the stomodaeum is elongate, but the sulcus and sulculus are very feebly represented. There are ten mesenteries in which the musculature is so little developed as to be almost indistinguishable. The sulcar and sulcular pairs of mesenteries are

A, Portion of a colony of Antipathes dichotoma. B, Single zooid and axis of the same magnified. m, Mouth; mf. mesenterial filament; ax, axis.

C, Transverse section through the oral cone of Antipathella minor. st, Stomodaeum; ov, ovary.

short, the sulco-lateral and sulculo-lateral pairs are a little longer, but the two transverse are very large and are the only mesenteries which bear gonads. As the development of the Antipathidea is unknown, it is impossible to say what is the sequence of the mesenterial development, but in Leiopathes glaberrima, a genus with twelve mesenteries, there are distinct indications of an Edwardsia stage.

There are, in addition to these groups, several genera of Actinians whose mesenterial arrangement differs from the normal type. Of

these perhaps the most interesting is Gonactinia prolifera (fig. 11, B), with eight macromesenteries arranged on the Edwardsian plan. Two pairs of micromesenteries form couples with the first and second Edwardsian pairs, and in addition there is a couple of micromesenteries in each of the sulculo-lateral exocoeles. Only the first and second pairs of Edwardsian macromesenteries are fertile, i.e. bear gonads. The remaining forms, the ACTINIIDEA, are divisible into the Malacactiniae, or soft-bodied sea-anemones, which have already been described sufficiently in the course of this article, and the Scleractiniae (= Madreporaria) or true corals.

All recent corals, as has already been said, conform so closely to the anatomy of normal Actinians that they cannot be classified apart from them, except that they are distinguished by the possession of a calcareous skeleton. This skeleton is largely composed of a number of radiating plates or sepla, and it differs both in origin and structure from the calcareous skeleton of all Alcyonaria except Heliopora. It is formed, not from fused spicules, but as a secretion of a special layer of cells derived from the basal ectoderm, and known as calicoblasts. The skeleton or corallum of a typical solitary coral-the common Devonshire cupcoral Caryophyllia smithii (fig. 15) is a good example-exhibits the followings parts:-(1) The basal plate, between the zooid and the surface of attachment. (2) The septa, radial plates of

are present the septa are correspondingly numerous. In some cases-e.g. in some species of Madrepora-only two septa are fully developed, the remainder being very feebly represented. Though the corallum appears to live within the zooid, it is morphologically external to it, as is best shown by its developmental history. The larvae of corals are free swimming ciliated forms known as planulae, and they do not acquire a corallum until they fix themselves. A ring-shaped plate of calcite, secreted by the ectoderm, is then formed, lying between the embryo and the surface of attachment. As the mesenteries are

which has fixed itself on a piece of cork, ec, Ectoderm; en, endoFIG. 16.-Tangential section of a larva of Astroides calicularis derm; mg, mesogloea; m, m, mesenteries; s, septum; b, basal plate formed of ellipsoids of carbonate of lime secreted by the basal ectoderm; ep, epitheca. (After von Koch.)

formed, the endoderm of the basal disk lying above the basal plate is raised up in the form of radiating folds. There may be six of these folds, one in each entococle of the primary cycle of mesenteries; or there may be twelve, one in each exocoele and entocoele. The ectoderm beneath each fold becomes detached from the surface of the basal plate, and both it and the mesogloea are folded conformably with the endoderm. The cells forming the limbs of the ectodermic folds secrete nodules of calcite, and these, fusing together, give rise to six (or twelve) vertical radial plates or septa. As growth proceeds new septa are formed simultaneously with the new couples of secondary mesenteries. In some corals, in which all the septa are entocoelic, each new system is embraced by a mesenteric couple; in others, in which the septa are both entocoelic and exocoelic, three septa are formed in I

FIG. 15.-Corallum of Caryophyllia; semi-diagrammatic. th, Theca: c, costae; sp, septa; p, palus; col, columella.

calcite reaching from the periphery nearly or quite to the centre of the coral-cup or calicle. (3) The theca or wall, which in many corals is not an independent structure, but is formed by the conjoined thickened peripheral ends of the septa. (4) The columella, a structure which occupies the centre of the calicle, and may arise from the basal plate, when it is called essential, or may be formed by union of trabecular offsets of the septa, when it is called unessential. (5) The costae, longitudinal ribs or rows of spines on the outer surface of the theca. True costae always correspond to the septa, and are in fact the peripheral edges of the latter. (6) Epitheca, an offset of the basal plate which surrounds the base of the theca in a ring-like manner, and in some corals may take the place of a true theca. (7) Pali, spinous or blade-like upgrowths from the bottom of the calicle, which project between the inner edges of certain septa and the columella. In addition to these parts the following structures may exist in corals:Dissepiments are oblique calcareous partitions, stretching from septum to septum, and closing the interseptal chambers below. The whole system of dissepiments in any given calicle is often called endotheca. Synapticulae are calcareous bars uniting adjacent septa. Tabulae are stout horizontal partitions traversing the centre of the calicle and dividing it into as many superimposed chambers. The septa in recent corals always bear a definite relation to the mesenteries, being found either in every entocoele or in every entocoele and exocoele. Hence in corals in which there is only a single cycle of mesenteries the septa are correspondingly few in number; where several cycles of mesenteries

corallum shaded with dots, the mesogloca represented by a thick line. FIG. 17.-Transverse section through a zooid of Cladocora. The Thirty-two septa are present, six in the entococles of the primary cycle of mesenteries, I; six in the entocoeles of the secondary cycle of mesenteries, II; four in the entocoeles of the tertiary cycle of mesenteries, III, only four pairs of the latter being developed; and sixteen in the entocoeles between the mesenterial pairs. D, D. Directive mesenteries; st, stomodaeum. (After Duerden.) every chamber between two primary mesenterial couples, one in the entocoele of the newly formed mesenterial couple of the secondary cycle, and one in each exocoele between a primary and a secondary couple. These latter are in turn embraced by the couples of the tertiary cycle of mesenteries, and new septa are formed in the exocoeles on either side of them, and so forth.

It is evident from an inspection of figs. 16 and 17 that every