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101
ANTHOZOA

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 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.


1911 Britannica-Anthozoa-Zoanthid.png

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