Volume 2, Slice 2 Part 12 (2/2)
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 h.o.r.n.y 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-ill.u.s.trated memoir by de Lacaze-Duthiers, which should be consulted for details of anatomy.
The AXIFERA comprise those corals that have a h.o.r.n.y 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 h.o.r.n.y substance secreted by the ectoderm. In _Gorgonia_ the axis remains h.o.r.n.y 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 h.o.r.n.y and calcareous substance, the latter being amorphous. The order contains six families--the _Dasygorgidae, Isidae, Primnoidae, Muriceidae, Plexauridae_, and _Gorgoniaae_.
[Ill.u.s.tration: FIG. 8.
A. Colony of _Pennatula phosph.o.r.ea_ 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 ca.n.a.ls.]
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 part.i.tion separating a so-called ”ventral” or prorachidial ca.n.a.l from a so-called ”dorsal” or metarachidial ca.n.a.l. A rod-like supporting axis of peculiar texture is developed in the longitudinal part.i.tion, and a longitudinal ca.n.a.l is hollowed out on either side of the axis in the substance of the longitudinal part.i.tion, so that there are four stem-ca.n.a.ls 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.
[Ill.u.s.tration: 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. Z, the distal, and Z, 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 ma.s.ses 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 part.i.tions 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 part.i.tions 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 h.o.m.ology with, the similarly named structures in madreporarian corals. _Heliopora coerulea_ is found between tide-marks on the sh.o.r.e 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 ma.s.ses; the calices were furnished with twelve pseudosepta, and the coenenchymal tubes were more or less regularly hexagonal.
[Ill.u.s.tration with caption: 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-cla.s.s the arrangement of the mesenteries is subject to a great deal of variation, but all the types. .h.i.therto observed may be referred to a common plan, ill.u.s.trated 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 s.p.a.ce enclosed between two mesenteries of the same couple is called an _entocoele_; the s.p.a.ce 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-a-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.
[Ill.u.s.tration: 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.
[Ill.u.s.tration: 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 l.a.b.i.al 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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