Anthozoa

Anthozoa (i.e. “flower-animals”), the zoological name for a class of marine polyps forming “coral” (q.v.). Although corals have been familiar objects since the days of antiquity, and the variety known as the precious red coral has been for a long time an article of commerce in the Mediterranean, it was only in the 18th century that their true nature and structure came to be understood. By the ancients and the earlier naturalists of the Christian era they were regarded either as petrifactions or as plants, and many supposed that they occupied a position midway between minerals and plants. The discovery of the animal nature of red coral is due to J.A. de Peyssonel, a native of Marseilles, who obtained living specimens from the coral fishers on the coast of Barbary and kept them alive in aquaria. He was thus able to see that the so-called “flowers of coral” were in fact nothing else than minute polyps resembling sea-anemones. His discovery, made in 1727, was rejected by the Academy of Sciences of France, but eventually found acceptance at the hands of the Royal Society of London, and was published by that body in 1751. The structure and classification of polyps, however, were at that time very imperfectly understood, and it was fully a century before the true anatomical characters and systematic position of corals were placed on a secure basis.

The hard calcareous substance to which the name coral is applied is the supporting skeleton of certain members of the Anthozoa, one of the classes of the phylum Coelentera. The most familiar Anthozoan is the common sea-anemone, Actinia equina, L., and it will serve, although it does not form a skeleton or corallum, as a good example of the structure of a typical Anthozoan polyp or zooid. The individual animal or zooid of Actinia equina has the form of a column fixed by one extremity, called the base, to a rock or other object, and bearing at the opposite extremity a crown of tentacles. The tentacles surround an area known as the peristome, in the middle of which there is an elongated mouth-opening surrounded by tumid lips. The mouth does not open directly into the general cavity of the body, as is the case in a hydrozoan polyp, but into a short tube called the stomodaeum, which in its turn opens below into the general body-cavity or coelenteron. In Actinia and its allies, and most generally, though not invariably, in Anthozoa, the stomodaeum is not circular, but is compressed from side to side so as to be oval or slit-like in transverse section. At each end of the oval there is a groove lined by specially long vibratile cilia. These grooves are known as the sulcus and sulculus, and will be more particularly described hereafter. The elongation of the mouth and stomodaeum confer a bilateral symmetry on the body of the zooid, which is extended to other organs of the body. In Actinia, as in all Anthozoan zooids, the coelenteron is not a simple cavity, as in a Hydroid, but is divided by a number of radial folds or curtains of soft tissue into a corresponding number of radial chambers. These radial folds are known as mesenteries, and their position and relations may be understood by reference to figs. 1 and 2. Each mesentery is attached by its upper margin to the peristome, by its outer margin to the body-wall, and by its lower margin to the basal disk. A certain number of mesenteries, known as complete mesenteries, are attached by the upper parts of their internal margins to the stomodaeum, but below this level their edges hang in the coelenteron. Other mesenteries, called incomplete, are not attached to the stomodaeum, and their internal margins are free from the peristome to the basal disk. The lower part of the free edge of every mesentery, whether complete or incomplete, is thrown into numerous puckers or folds, and is furnished with a glandular thickening known as a mesenterial filament. The reproductive organs or gonads are borne on the mesenteries, the germinal cells being derived from the inner layer or endoderm.  
  
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Fig. 1\. Diagrammatic longitudinal section of an Anthozoan zooid.  
  
m, Mesentery.

t, Tentacles.

st, Stomodaeum.

sc, Sulcus.

r, Rotteken’s muscle.

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s, Stoma.

lm, Longitudinal muscle.

d, Diagonal Muscle.

go, Gonads.  
  
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Fig. 2.—1, Portion of epithelium from the tentacle of an Actinian, showing three supporting cells and one sense cell (sc); 2, a cnidoblast with enclosed nematocyst from the same specimen; 3 and 4 two forms of gland cell from the stomodaeum; 5a, 5b, epithelio-muscular cells from the tentacle in different states of contraction; 5c, an epithelio-muscular cell from the endoderm, containing a symbiotic zooxanthella; 6, a ganglion cell from the ectoderm of the peristome. (After O. and R. Hertwig.)  
  
In common with all Coelenterate animals, the walls of the columnar body and also the tentacles and peristome of Actinia are composed of three layers of tissue. The external layer, or ectoderm, is made up of cells, and contains also muscular and nervous elements. The preponderating elements of the ectodermic layer are elongated columnar cells, each containing a nucleus, and bearing cilia at their free extremities. Packed in among these are gland cells, sense cells, and cnidoblasts. The last-named are specially numerous on the tentacles and on some other regions of the body, and produce the well-known “thread cells,” or nematocysts, so characteristic of the Coelentera. The inner layer or endoderm is also a cellular layer, and is chiefly made up of columnar cells, each bearing a cilium at its free extremity and terminating internally in a long muscular fibre. Such cells, made up of epithelial and muscular components, are known as epithelio-muscular or myo-epithelial cells. In Actinians the epithelio-muscular cells of the endoderm are crowded with yellow spherical bodies, which are unicellular plants or Algae, living symbiotically in the tissues of the zooid. The endoderm contains in addition gland cells and nervous elements. The middle layer or mesogloea is not originally a cellular layer, but a gelatinoid structureless substance, secreted by the two cellular layers. In the course of development, however, cells from the ectoderm and endoderm may migrate into it. In Actinia equina the mesogloea consists of fine fibres imbedded in a homogeneous matrix, and between the fibres are minute branched or spindle-shaped cells. For further details of the structure of Actinians, the reader should consult the work of O. and R. Hertwig.

The Anthozoa are divisible into two sub-classes, sharply marked off from one another by definite anatomical characters. These are the Alcyonaria and the Zoantharia. To the first-named belong the precious red coral and its allies, the sea-fans or Gorgoniae, to the second belong the white or Madreporarian corals.  
  
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Fig. 3.—An expanded Alcyonarian zooid, showing the mouth surrounded by eight pinnate tentacles. st, Stomodaeum in the the centre of the transparent body; m, mesenteries; asm, asulcar mesenteries; B, spicules, enlarged.  
Fig. 4.—Transverse section of an Alcyonarian zooid mm, Mesenteries; mb, muscle banners; sc, sulcus; st, stomodaeum.  
  
Alcyonaria.—In this sub-class the zooid has very constant anatomical characters, differing in some important respects from the Actinian zooid, which has been taken as a type. There is only one ciliated groove, the sulcus, in the stomodaeum. There are always eight tentacles, which are hollow and fringed on their sides, with hollow projections or pinnae; and always eight mesenteries, all of which are complete, i.e. inserted on the stomodaeum. The mesenteries are provided with well-developed longitudinal retractor muscles, supported on longitudinal folds or plaits of the mesogloea, so that in cross-section they have a branched appearance. These muscle-banners, as they are called, have a highly characteristic arrangement; they are all situated on those faces of the mesenteries which look towards the sulcus. (fig. 4). Each mesentery has a filament; but two of them, namely, the pair farthest from the sulcus, are longer than the rest, and have a different form of filament. It has been shown that these asulcar filaments are derived from the ectoderm, the remainder from the endoderm. The only exceptions to this structure are found in the arrested or modified zooids, which occur in many of the colonial Alcyonaria. In these the tentacles are stunted or suppressed and the mesenteries are ill-developed, but the sulcus is unusually large and has long cilia. Such modified zooids are called siphonozooids, their function being to drive currents of fluid through the canal-systems of the colonies to which they belong. With very few exceptions a calcareous skeleton is present in all Alcyonaria; it usually consists of spicules of carbonate of lime, each spicule being formed within an ectodermic cell (fig. 3, B). Most commonly the spicule-forming cells pass out of the ectoderm and are imbedded in the mesogloea, where they may remain separate from one another or may be fused together to form a strong mass. In addition to the spicular skeleton an organic horny skeleton is frequently present, either in the form of a horny external investment (Cornularia), or an internal axis (Gorgonia), or it may form a matrix in which spicules are imbedded (Keroeides, Meistodes).

Nearly all the Alcyonaria are colonial. Four solitary species have been described, viz. Haimea funebris and H. hyalina, Hartea elegans, and Monoxenia Darwinii; but it is doubtful whether these are not the young forms of colonies. For the present the solitary forms may be placed in a grade, Protal-cyonacea, and the colonial forms may be grouped in another grade, Synalcyonacea. Every Alcyonarian colony is developed by budding from a single parent zooid. The buds are not direct outgrowths of the body-wall, but are formed on the courses of hollow out growths of the base or body-wall, called solenia. These form a more or less complicated canal system, lined by endoderm, and communicating with the cavities of the zooids. The most simple form of budding is found in the genus Cornularia, in which the mother zooid gives off from its base one or more simple radiciform outgrowths. Each outgrowth contains a single tube or solenium, and at a longer or shorter distance from the mother zooid a daughter zooid is formed as a bud. This gives off new outgrowths, and these, branching and anastomosing with one another, may form a network, adhering to stones, corals, or other objects, from which 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 Cornothecalia.  
  
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Fig. 5.  
  
A. Skeleton of a young colony of Tubipora purpurea. st, Stolon; p, platform.

B. Diagrammatic longitudinal section of a corallite, showing two platforms, p and cup-shaped tabulae, t. (After S.J. Hickson.)  
  
Fig. 6.—Portion of a colony of Coralinum rubrum, showing expanded and contracted zooids. In the lower part of the figure the cortex has been cut away to show the axis, ax, and the longitudinal canals, lc, surrounding it.  
  
In the order Stolonirera the zooids spring at intervals from branching or lamellar stolons, and are usually free from one another, except at their bases, but in some cases horizontal solenia arising at various heights from the body-wall may place the more distal portions of the zooids in communication with one another. In the genus Tubipora these horizontal solenia unite to form a series of horizontal platforms (fig. 5). The order comprises the families Cornulamdae, Syringopordae, Tubipondae, and Favositidae. In the first-named, the zooids are united only by their bases and the skeleton consists of loose spicules. In the Tubipondae the spicules of the proximal part of the body-wall are fused together to form a firm tube, the corallite, into which the distal part of the zooid can be retracted. The corallites are connected at intervals by horizontal platforms containing solenia, and at the level of each platform the cavity of the corallite is divided by a transverse calcareous partition, either flat or cup-shaped, called a tabula. Formerly all corals in which tabulae are present were classed together as Tabulata, but Tubipora is an undoubted Alcyonarian with a lamellar stolon, and the structure of the fossil genus Syringopora, which has vertical corallites united by horizontal solenia, clearly shows its affinity to Tubipora. The Favositidae, a fossil family from the Silurian and Devonian, have a massive corallum composed of numerous polygonal corallites closely packed together. The cavities of adjacent corallites communicate by means of numerous perforations, which appear to represent solenia, and numerous transverse tabulae are also present. In Favosites hemisphaerica a number of radial spines, projecting into the cavity of the corallite, give it the appearance of a madreporarian coral.

In the order Alcyonacea the colony consists 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.  
  
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Fig. 7.—The sea-fan (Gorgonia cavolinii).  
Fig. 8.  
  
A. Colony of Pennatula phosphorea from the metarachidial aspect. p, The peduncle.

B. Section of the rachis bearing a single pinna, a, Axis; b, metarachidial; c, prorachidial; d, pararachidial stem canals.  
  
In the order Pseudaxonia the colonies are upright and branched, 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, 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 rubrum (fig. 6), a member of the family Corallidae, is found at depths varying from 15 to 120 fathoms 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 axis, which in position corresponds to the axis of the Pscudaxonia, 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 families—the Dasygorgidae, Isidae, Primnoidae, Muriceidae, Plexauridae, and Gorgoniaae.

In the order Stelechotokea the colony consists of a stem formed by a greatly-elongated mother zooid, and the daughter zooids are borne as lateral buds on the stem. In the section Asiphonacea the colonies are upright and branched, springing from membranous or ramifying stolons. They resemble and are closely allied to certain families of the Cornulariidae, differing from them only in mode of budding and in the dispostion of the daughter zooids round a central, much-elongated mother zooid. The section contains two families, the Telestidae and the Coelogorgidae. The second section comprises the Pennatulacea or sea-pens, which are remarkable from the fact that the colony is not fixed by the base to a rock or other object, but is imbedded in sand or mud by the proximal portion of the stem known as the peduncle. In the typical genus, Pennatula (fig. 8), the colony looks like a feather having a stem divisible into an upper moiety or rachis, bearing lateral central leaflets (pinnae), and a lower peduncle, which is sterile and imbedded in sand or mud. The stem represents a greatly enlarged and elongated mother zooid. It is divided longitudinally by a partition separating a so-called “ventral” or prorachidial canal from a so-called “dorsal” or metarachidial canal. A rod-like supporting axis of peculiar texture is developed in the longitudinal partition, and a longitudinal canal is hollowed out on either side of the axis in the substance of the longitudinal partition, so that there are four stem-canals in all. The prorachidial and metarachidial aspects of the rachis are sterile, but the sides or pararachides bear numerous daughter zooids of two kinds—(1) fully-formed autozooids, (2) small stunted siphonozooids. The pinnae are formed by the elongated autozooids, whose proximal portions are fused together to form a leaf-like expansion, from the upper edge of which the distal extremities of the zooids project. The siphonozooids are very numerous and lie between the bases at the pinnae on the pararachides; they extend also on the prorachidial and metarachidial surfaces. The calcareous skeleton of the Pennatulacea consists of scattered spicules, but in one species, Protocaulon molle, spicules are absent. Although of great interest the Pennatulacea do not form an enduring skeleton or “coral,” and need not be considered in detail in this place.  
  
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Fig. 9.  
  
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. Z1, the distal, and Z2, the proximal or intracalicular portion of the zooid; ec, ectoderm; ct, coenenchymal tubes; sp, superficial network of solenia.  
  
The order Coenothecalia is represented by a single living species, Heliopora coerulea, which differs from all recent Alcyonaria in the fact that its skeleton is not composed of spicules, but is formed as a secretion from a layer of cells called calicoblasts, which originate from the ectoderm. The corallum of Heliopora is of a blue colour, and has the form of broad, upright, lobed, or digitate masses flattened from side to side. The surfaces are pitted all over with perforations of two kinds, viz. larger star-shaped cavities, called calices, in which the zooids are lodged, and very numerous smaller round or polygonal apertures, which in life contain as many short unbranched 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.  
  
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Fig. 10.  
  
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.  
  
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 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 of mesenteries which are attached to the sides of the elongated stomodaeum the muscle-banners of each couple are turned towards one another, but in the sulcar and sulcular couples, known as the directive 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 entocoele; 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 exocoele 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.  
  
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Fig. 11.—A, Diagram showing the sequence of mesenterial development in an Actinian. B, Diagrammatic transverse section of Gonactinia prolifera.  
  
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 muscle-banners 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 stomodaeum 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 sulco-lateral 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.  
  
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Fig. 12.  
  
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.  
  
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 arrangement 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 stomodaeum, 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 exocoeles. 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 muscle-banners. 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 sixth pair inside the fifth. New mesenteries are continually added in the sulcar chamber, the seventh pair within the sixth, the eighth pair within the seventh, and so on (fig. 13). In the Cerianthidea, as in the Zoanthidea, much as the adult arrangement of mesenteries differs from that of Actinia, the derivation from an Edwardsia stock is obvious.  
  
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Fig. 13.  
  
A, Cerianthus solitarius (after A. Andres).

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

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

D, Transverse section of an older larva. The numerals indicate the order of development of the mesenteries.  
  
The order Antipathidea is a well-defined group whose affinities 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 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.  
  
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Fig. 14.  
  
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.  
  
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.  
  
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Fig. 15.—Corallum of Caryophyllia; semi-diagrammatic. th, Theca; c, costae; sp, septa; p, palus; col, columella.  
  
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 septa, 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 cup-coral 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 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 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.  
  
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Fig. 16.—Tangential section of a larva of Astroides calicularis which has fixed itself on a piece of cork. ec, Ectoderm; en, endoderm; 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.)  
  
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 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 entocoele 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 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.  
  
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Fig. 17.—Transverse section through a zooid of Cladocora. The corallum shaded with dots, the mesogloea represented by a thick line. Thirty-two septa are present, six in the entocoeles 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.)  
  
It is evident from an inspection of figs. 16 and 17 that every septum is covered by a fold of endoderm, mesogloea, and ectoderm, and is in fact pushed into the cavity of the zooid from without. The zooid then is, as it were, moulded upon the corallum. When fully extended, the upper part of the zooid projects for some distance out of the calicle, and its wall is reflected for some distance over the lip of the latter, forming a fold of soft tissue extending to a greater or less distance over the theca, and containing in most cases a cavity continuous over the lip of the calicle with the coelenteron. This fold of tissue is known as the edge-zone In some corals the septa are solid imperforate plates of calcite, and their peripheral ends are either firmly welded together, or are united by interstitial pieces so as to form imperforate theca. In others the peripheral ends of the septa are united only by bars or trabeculae, so that the theca is perforate, and in many such perforate corals the septa themselves are pierced by numerous perforations. In the former, which have been called aporose corals, the only communication between the cavity of the edge-zone and the general cavity of the zooid is by way of the lip of the calicle; in the latter, or perforate corals, the theca is permeated by numerous branching and anastomosing canals lined by endoderm, which place the cavity of the edge-zone in communication with the general cavity of the zooid.  
  
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Fig. 18.  
  
A, Schematic longitudinal section through a zooid and bud of Stylophora digitata. In A, B, and C the thick black lines represent the soft tissues; the corallum is dotted. s, Stomodaeum; c, c, coenosarc; col, columella, T tabulae.

B, Similar section through a single zooid and bud of Astroides calicularis.

C, Similar section through three corallites of Lophohelia prolifera. ez, Edge-zone.

D, Diagram illustrating the process of budding by unequal division.

E, Section through a dividing calicle of Mussa, showing the union of two septa in the plane of division and the origin of new septa at right angles to them.

(C original; the rest after von Koch.)  
  
A large number of corals, both aporose and perforate, are colonial. The colonies are produced by either budding or division. In the former case the young daughter zooid, with its corallum, arises wholly outside the cavity of the parent zooid, and the component parts of the young corallum, septa, theca, columella, &c., are formed anew in every individual produced. In division a vertical constriction divides a zooid into two equal or unequal parts, and the several parts of the two corals thus produced are severally derived from the corresponding parts of the dividing corallum. In colonial corals a bud is always formed from the edge-zone, and this bud develops into a new zooid with its corallum. The cavity of the bud in an aporose coral (fig. 18, A, C) does not communicate directly with that of the parent form, but through the medium of the edge-zone. As growth proceeds, and parent and bud become separated farther from one another, the edge-zone forms a sheet of soft tissue, bridging over the space between the two, and resting upon projecting spines of the corallum. This sheet of tissue is called the coenosarc. Its lower surface is clothed with a layer of calicoblasts which continue to secrete carbonate of lime, giving rise to a secondary deposit which more or less fills up the spaces between the individual coralla, and is distinguished as coenenchyme. This coenenchyme may be scanty, or may be so abundant that the individual corallites produced by budding seem to be immersed in it. Budding takes place in an analogous manner in perforate corals (fig. 18, B), but the presence of the canal system in the perforate theca leads to a modification of the process. Buds arise from the edge-zone which already communicate with the cavity of the zooid by the canals. As the buds develop the canal system becomes much extended, and calcareous tissue is deposited between the network of canals, the confluent edge-zones of mother zooid and bud forming a coenosarc. As the process continues a number of calicles are formed, imbedded in a spongy tissue in which the canals ramify, and it is impossible to say where the theca of one corallite ends and that of another begins. In the formation of colonies by division a constriction at right angles to the long axis of the mouth involves first the mouth, then the peristome, and finally the calyx itself, so that the previously single corallite becomes divided into two (fig. 18, E). After division the corallites continue to grow upwards, and their zooids may remain united by a bridge of soft tissue or coenosarc. But in some cases, as they grow farther apart, this continuity is broken, each corallite has its own edge-zone, and internal continuity is also broken by the formation of dissepiments within each calicle, all organic connexion between the two zooids being eventually lost. Massive meandrine corals are produced by continual repetition of a process of incomplete division, involving the mouth and to some extent the peristome: the calyx, however, does not divide, but elongates to form a characteristic meandrine channel containing several zooid mouths.

Corals have been divided into Aporosa and Perforata, according as the theca and septa are compact and solid, or are perforated by pores containing canals lined by endoderm. The division is in many respects convenient for descriptive purposes, but recent researches show that it does not accurately represent the relationships of the different families. Various attempts have been made to classify corals according to the arrangement of the septa, the characters of the theca, the microscopic structure of the corallum, and the anatomy of the soft parts. The last-named method has proved little more than that there is a remarkable similarity between the zooids of all recent corals, the differences which have been brought to light being for the most part secondary and valueless for classificatory purposes. On the other hand, the study of the anatomy and development of the zooids has thrown much light upon the manner in which the corallum is formed, and it is now possible to infer the structure of the soft parts from a microscopical examination of the septa, theca, &c., with the result that unexpected relationships have been shown to exist between corals previously supposed to stand far apart. This has been particularly the case with the group of Palaeozoic corals formerly classed together as Rugosa. In many of these so-called rugose forms the septa have a characteristic arrangement, differing from that of recent corals chiefly in the fact that they show a tetrameral instead of a hexameral symmetry. Thus in the family Stauridae there are four chief septa whose inner ends unite in the middle of the calicle to form a false columella, and in the Zaphrentidae there are many instances of an arrangement, such as that depicted in fig. 19, which represents the septal arrangement of Streptelasma corniculum from the lower Silurian. In this coral the calicle is divided into quadrants by four principal septa, the main septum, counter septum, and two alar septa. The remaining septa are so disposed that in the quadrants abutting on the chief septum they converge towards that septum, whilst in the other quadrants they converge towards the alar septa. The secondary septa show a regular gradation in size, and, assuming that the smallest were the most recently formed, it will be noticed that in the chief quadrants the youngest septa lie nearest to the main septum; in the other quadrants the youngest septa lie nearest to the alar septa. This arrangement, however, is by no means characteristic even of the Zaphrentidae, and in the family Cyathophyllidae most of the genera exhibit a radial symmetry in which no trace of the bilateral arrangement described above is recognizable, and indeed in the genus Cyathophyllum itself a radial arrangement is the rule. The connexion between the Cyathophyllidae and modern Astraeidae is shown by Moseleya latistellata, a living reef-building coral from Torres Strait. The general structure of this coral leaves no doubt that it is closely allied to the Astraeidae, but in the young calicles a tetrameral symmetry is indicated by the presence of four large septa placed at right angles to one another. Again, in the family Amphiastraeidae there is commonly a single septum much larger than the rest, and it has been shown that in the young calicles, e.g. of Thecidiosmilia, two septa, corresponding to the main- and counter-septa of Streptelasma, are first formed, then two alar septa, and afterwards the remaining septa, the latter taking on a generally radial arrangement, though the original bilaterality is marked by the preponderance of the main septum. As the microscopic character of the corallum of these extinct forms agrees with that of recent corals, it may be assumed that the anatomy of the soft parts also was similar, and the tetrameral arrangement, when present, may obviously be referred to a stage when only the first two pairs of Edwardsian mesenteries were present and septa were formed in the intervals between them.  
  
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Fig. 19.—Diagram of the arrangement of the septa in a Zaphrentid coral. m, Main septum; c, counter septum; t, t, alar septa.  
  
Space forbids a discussion of the proposals to classify corals after the minute structure of their coralla, but it will suffice to say that it has been shown that the septa of all corals are built up of a number of curved bars called trabeculae, each of which is composed of a number of nodes. In many secondary corals (Cyclolites, Thamnastraea) the trabeculae are so far separate that the individual bars are easily recognizable, and each looks something like a bamboo owing to the thickening of the two ends of each node. The trabeculae are united together by these thickened internodes, and the result is a fenestrated septum, which in older septa may become solid and aporose by continual deposit of calcite in the fenestrae. Each node of a trabecula may be simple, i.e. have only one centre of calcification, or may be compound. The septa of modern perforate corals are shown to have a structure nearly identical with that of the secondary forms, but the trabeculae and their nodes are only apparent on microscopical examination. The aporose corals, too, have a practically identical structure, their compactness being due to the union of the trabeculae throughout their entire lengths instead of at intervals, as in the Perforata. Further, the trabeculae may be evenly spaced throughout the septum, or may be grouped together, and this feature is probably of value in estimating the affinities of corals. (For an account of coral formations see Coral-Reefs.)

In the present state of our knowledge the Zoantharia in which a primary cycle of six couples of mesenteries is (or may be inferred to be) completed by the addition of two pairs to the eight Edwardsian mesenteries, and succeeding cycles are formed in the exocoeles of the pre-existing mesenterial cycles, may be classed in an order Actiniidea, and this may be divided into the suborders Malacactiniae, comprising the soft-bodied Actinians, such as Actinia, Sagartia, Bunodes, &c., and the Scleractiniae, comprising the corals. The Scleractiniae may best be divided into groups of families which appear to be most closely related to one another, but it should not be forgotten that there is great reason to believe that many if not most of the extinct corals must have differed from modern Actiniidea in mesenterial characters, and may have only possessed Edwardsian mesenteries, or even have possessed only four mesenteries, in this respect showing close affinities to the Stauromedusae. Moreover, there are some modern corals in which the secondary cycle of mesenteries departs from the Actinian plan. For example, J.E. Duerden has shown that in Porites the ordinary zooids possess only six couples of mesenteries arranged on the Actinian plan. But some zooids grow to a larger size and develop a number of additional mesenteries, which arise either in the sulcar or the sulcular entocoele, much in the same manner as in Cerianthus. Bearing this in mind, the following arrangement may be taken to represent the most recent knowledge of coral structure:—

Group A.

Family I. Zaphrentidae.—Solitary Palaeozoic corals with an epithecal wall. Septa numerous, arranged pinnately with regard to four principal septa. Tabulae present. One or more pits or fossulae present in the calicle. Typical genera—Zaphrentis, Raf. Amplexus, M. Edw. and H. Streptelasma, Hall. Omphyma, Raf.

Family 2. Turbinolidae.—Solitary, rarely colonial corals, with radially arranged septa and without tabulae. Typical genera— Flabellum, Lesson. Turbinolia, M. Edw. and H. Caryophyllia, Lamarck. Sphenotrochus, Moseley, &c.

Family 3. Amphiastraeidae.—Mainly colonial, rarely solitary corals, with radial septa, but bilateral arrangement indicated by persistence of a main septum. Typical genera—Amphiastraea, Étallon. Thecidiosmilia.

Family 4. Stylinidae.—Colonial corals allied to the Amphiastraeidae, but with radially symmetrical septa arranged in cycles. Typical genera—Stylina, Lamarck (Jurassic). Convexastraea, D’Orb. (Jurassic). Isastraea, M. Edw. and H.(Jurassic). Ogilvie refers the modern genus Galaxea to this family.

Group B.

Family 5. Oculinidae.—Branching or massive aporose corals, the calices projecting above the level of a compact coenenchyme formed from the coenosarc which covers the exterior of the corallum. Typical genera—Lophohelia, M. Edw. and H. Oculina, M. Edw. and H.

Family 6. Pocilloporidae.—Colonial branching aporose corals, with small calices sunk in the coenenchyme. Tabulae present, and two larger septa, an axial and abaxial, are always present, with traces of ten smaller septa. Typical genera—Pocillopora, Lamarck. Seriatopora, Lamarck.

Family 7. Madreporidae.—Colonial branching or palmate perforate corals, with abundant trabecular coenenchyme. Theca porous; septa compact and reduced in number. Typical genera— Madrepora, Linn. Turbinaria, Oken. Montipora, Quoy and G.

Family 8. Poritidae.—Incrusting or massive colonial perforate corals; calices usually in contact by their edges, sometimes disjunct and immersed in coenenchyme. Theca and septa perforate. Typical genera—Porites, M. Edw. and H. Goniopora, Quoy and G. Rhodaraea, M. Edw. and H.

Group C.

Family 9. Cyathophyllidae.—Solitary and colonial aporose corals. Tabulae and vesicular endotheca present. Septa numerous, generally radial, seldom pinnate. Typical genera—Cyathophyllum, Goldfuss (Devonian and Carboniferous). Moseleya, Quelch (recent).

Family 10. Astraeidae.—Aporpse, mainly colonial corals, massive, branching, or maeandroid. Septa radial; dissepiments present; an epitheca surrounds the base of massive or maeandroid forms, but only surrounds individual corallites in simple or branching forms. Typical genera—Goniastraea, M. Edw. and H. Heliastraea, M. Edw. and H. Maeandrina, Lam. Coeloria, M. Edw. and H. Favia, Oken.

Family 11. Fungidae.—Solitary and colonial corals, with numerous radial septa united by synapticulae. Typical genera— Lophoseris, M. Edw. and H. Thamnastraea, Le Sauvage. Leptophyllia, Reuss (Jurassic and Cretaceous). Fungia, Dana. Siderastraea, Blainv.

Group D.

Family 12. Eupsammidae.—Solitary or colonial perforate corals, branching, massive, or encrusting. Septa radial; the primary septa usually compact, the remainder perforate. Theca perforate. Synapticula present in some genera. Typical genera—Stephanophyllia, Michelin. Eupsammia, M. Edw. and H. Astroides, Blainv. Rhodopsammia, M. Edw. and H. Dendrophyllia, M. Edw. and H.

Group E.

Family 13. Cystiphyllidae.—Solitary corals with rudimentary septa, and the calicle filled with vesicular endotheca. Genera—Cystiphyllum, Lonsdale (Silurian and Devonian). Goniophyllum, M. Edw. and H. (In this Silurian genus the calyx is provided with a movable operculum, consisting of four paired triangular pieces, the bases of each being attached to the sides of the calyx, and their apices meeting in the middle when the operculum is closed). Calcecla, Lam. (In this Devonian genus there is a single semicircular operculum furnished with a stout median septum and numerous feebly developed secondary septa. The calyx is triangular in section, pointed below, and the operculum is attached to it by hinge-like teeth.)

Authorities.—The following list contains only the names of the more important and more general works on the structure and classification of corals and on coral reefs. For a fuller bibliography the works marked with an asterisk should be consulted: * A. Andres, Fauna und Flora des Golfes von Neapel, ix. (1884); H.M. Bernard, “Catalogue of Madreporarian Corals” in Brit. Museum, ii. (1896), iii. (1897); * G.C. Bourne, “Anthozoa,” in E. Ray Lankester’s Treatise on Zoology, vol. ii. (London, 1900); G. Brook, “Challenger Reports,” Zoology, xxxii. (1899) (Antipatharia); “Cat. Madrep. Corals,” Brit. Museum, i. (1893); D.C. Danielssen, “Report Norwegian North Atlantic Exploring Expedition,” Zoology, xix. (1890); J.E. Duerden, “Some Results on the Morphology and Development of Recent and Fossil Corals,” Rep. Brit. Association, 1903, pp. 684-685; “The Morphology of the Madreporaria,” Biol. Bullet, vii. pp. 79-104; P.M. Duncan, Journ. Linnean Soc. xviii. (1885); P.H. Gosse, Actinologia britannica (London, 1860); O. and R. Hertwig, Die Actinien (Jena, 1879); R. Hertwig, “Challenger Reports,” Zoology, vi. (1882) and xxvi. (1888); * C.B. Klunzinger, Die Korallthiere des Rothen Meeres (Berlin, 1877); * G. von Koch, Fauna und Flora des Golfes van Neapel, xv. (1887); Mitth. Zool. Stat. Neapel, ii. (1882) and xii. (1897); Palaeontographica, xxix. (1883); (also many papers in the Morphol. Jahrbuch from 1878 to 1898); F. Koby, “Polypiers jurassiques de la Suisse,” Mem. Soc. Palaeont. Suisse, vii.-xvi. (1880-1889); A. von Kölliker, “Die Pennatuliden,” Abh. d. Senck. Naturf. Gesell. vii.; * “Challenger Reports,” Zoology, i. Pennatulidae (1880); Koren and Danielssen, Norske Nordhaus Exped., Alcyonida (1887); H. de Lacaze-Duthiers, Hist. nat. du corail (Paris, 1864); H. Milne-Edwards and J. Haime, Hist. nat. des coralliaires (Paris, 1857); H.N. Moseley, “Challenger Reports,” Zoology, ii. (1881); H.A. Nicholson, Palaeozoic Tabulate Corals (Edinburgh, 1879); M.M. Ogilvie, Phil. Transactions, clxxxvii. (1896); E. Pratz, Palaeontographica, xxix. (1882); J.J. Quelch, “Challenger Reports,” Zoology, xvi. (1886); * P.S. Wright and Th. Studer, “Challenger Reports,” Zoology, xxxi. (1889).

(G. C. B.)

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