The manubrium of the Scyphozoa is usually quadrangular in section, and in those forms in which the shape is modified in the adult Medusa the quadrangular shape can be recognised in the earlier stages of development. The four angles of the manubrium are of importance in descriptive anatomy, as the planes drawn through the angles to the centre of the manubrium are called "perradial," while those bisecting the perradial planes and passing therefore through the middle line of the flat sides of the manubrium are called "interradial."
The free extremity of the manubrium in many Scyphozoa is provided with four triangular perradial lips, which may be simple or may become bifurcated or branched, and have frequently very elaborate crenate edges beset with batteries of nematocysts. In Pelagia and Chrysaora and other genera these lips hang down from the manubrium as long, ribbon-like, folded bands, and according to the size of the specimen may be a foot or more in length, or twice the diameter of the disc.
In the Rhizostomata a peculiar modification of structure takes place in the fusion of the free edges of the lips to form a suture perforated by a row of small apertures, so that the lips have the appearance of long cylindrical rods or tubes attached to the manubrium, and then frequently called the "oral arms." The oral arms may be further provided with tentacles of varying size and importance. In many Rhizostomata branched or knobbed processes project from the outer side of the upper part of the oral arms. These are called the "epaulettes."
Fig. 143.—Ulmaris prototypus. g, Gonad; I, interradial canal; M, the fringed lip of the manubrium; P, perradial canal; S, marginal sense-organ; t, tentacle. × 1. (After Haeckel.)
The lumen of the manubrium leads into a large cavity in the disc, which is usually called the gastric cavity, and this is extended into four or more interradial or perradial gastric pouches. The number of these pouches is usually four, but in this, as in other features of their radial symmetry, the jelly-fish frequently exhibit duplication or irregular variation of the radii.[[350]]
The gastric pouches may extend to the margin of the disc, where they are united to form a large ring sinus, or they may be in communication at the periphery by only a very narrow passage (Cubomedusae). In the Discophora the gastric pouches, however, do not extend more than half-way to the margin, and they may be connected with the marginal ring-canal by a series of branched interradial canals. Between the gastric pouches in these forms branched perradial canals pass from the gastric cavity to the marginal ring canal, and the system of canals is completed by unbranched "adradial" canals passing between the perradials and interradials from the sides of the gastric pouches to the ring-canal (Fig. 143).
In the Discophora there are four shallow interradial pits or pouches lined by ectoderm on the under side of the umbrella-wall. As these pits correspond with the position of the gonads in the gastric pouches they are frequently called the "sub-genital pits." In the Stauromedusae and Cubomedusae they are continued through the interradial gastric septa to the aboral side of the disc, and they are generally known in these cases by the name "interradial funnels." The functions and homologies of these ectodermic pits and funnels are still uncertain.
The Scyphozoa are usually dioecious, but Chrysaora and Linerges are sometimes hermaphrodite. The female Medusae can usually be distinguished from the male by the darker or brighter colour of the gonads, which are band-shaped, horseshoe-shaped, or circular organs, situated on the endoderm of the interradial gastric pouches. They are, when nearly ripe, conspicuous and brightly coloured organs, and in nearly all species can be clearly seen through the transparent or semi-transparent tissues of the disc. The reproductive cells are discharged into the gastric cavity and escape by the mouth. The eggs are probably fertilised in the water, and may be retained in special pouches on the lips of the manubrium until the segmentation is completed.[[351]] Asexual reproduction does not occur in the free-swimming or adult stage of any Scyphozoa. In some cases (probably exceptional) the development is direct. In Pelagia, for example, it is known that the fertilised egg gives rise to a free-swimming Medusa similar in all essential features to the parent.
In many species, however, the planula larva sinks to the bottom of the sea, develops tentacles, and becomes attached by its aboral extremity to a rock or weed, forming a sedentary asexual stage of development with a superficial resemblance to a Hydra. This stage is the "Scyphistoma," and notwithstanding its simple external features it is already in all essential anatomical characters a Scyphozoon.