The medusa-buds, as already stated, are always produced from blastostyles, reduced non-nutritive polyps without mouth or tentacles. An apparent, but not real, exception is Halecium halecinum, in which the blastostyle is produced from the side of a nutritive polyp, and both are enclosed in a common theca without a partition between them (Allman [1] p. 50, fig. 24). The gonotheca is formed in its early stage in the same way as the hydrotheca, but the remains of the hydranth persists as an operculum closing the capsule, to be withdrawn when the medusae or genital products are set free (fig. 56).
The blastostyles, gonophores and gonothecae furnish a series of variations which can best be considered as so many stages of evolution.
Stage 1, seen in Obelia. Numerous medusae are budded successively within the gonotheca and set free; they swim off and mature in the open sea (Allman [1], p. 48, figs. 18, 19).
Stage 2, seen in Gonothyraea. Medusae, so-called “meconidia,” are budded but not liberated; each in turn, when it reaches sexual maturity, is protruded from the gonotheca by elongation of the stalk, and sets free the embryos, after which it withers and is replaced by another (Allman [1], p. 57, fig. 28).
Stage 3, seen in Sertularia.—The gonophores are reduced in varying degree, it may be to sporosacs; they are budded successively from the blastostyle, and each in turn, when ripe, protrudes the spadix through the gonotheca (fig. 57, A, B). The spadix forms a gelatinous cyst, the so-called acrocyst (ac), external to the gonotheca (gth), enclosing and protecting the embryos. Then the spadix withers, leaving the embryos in the acrocyst, which may be further protected by a so-called marsupium, a structure formed by tentacle-like processes growing out from the blastostyle to enclose the acrocyst, each such process being covered by perisarc like a glove-finger secreted by it (fig. 57, C). (Allman [1], pp. 50, 51, figs. 21-24; Weismann [58], p. 170, pl. ix., figs. 7, 8.)
Stage 4, seen in Plumularidae.—The generative elements are produced in structures termed corbulae, formed by reduction and modification of branches of the colony. Each corbula contains a central row of blastostyles enclosed and protected by lateral rows of branches representing stunted buds (Allman [1], p. 66, fig. 30).
| After Allman, Gymnoblastic Hydroids, by permission of the council of the Ray Society. |
| Fig. 57.—Diagrams to show the mode of formation of an Acrocyst and a Marsupium. In A two medusa-buds are seen within the gonotheca (gth), the upper more advanced than the lower one. In B the spadix of the upper bud has protruded itself through the top of the gonotheca and the acrocyst (ac) is secreted round it. In C the marsupium (m) is formed as finger-like process from the summit of the blastostyle, enclosing the acrocyst; b, medusa-buds on the blastostyle. |
The Leptomedusa in form is generally shallow, more or less saucer-like, with velum less developed than in Anthomedusae (fig. 55). The characteristic sense-organs are ectodermal otocysts, absent, however, in some genera, in which case cordyli may replace them. When otocysts are present, they are at least eight in number, situated adradially, but are often very numerous. The cordyli are scattered on the ring-canal. Ocelli, if present, are borne on the tentacle-bulbs. The tentacles are usually hollow, rarely solid (Obelia). In number they are rarely less than four, but in Dissonema there are only two. Primitively there are four perradial tentacles, to which may be added four interradial, or they may become very numerous and are then scattered evenly round the margin, never arranged in tufts or clusters. In addition to tentacles, there may be marginal cirri (Laodice) with a solid endodermal axis, spirally coiled, very contractile, and bearing a terminal battery of nematocysts. The gonads are developed typically beneath the radial canals or below the stomach or its pouches, often stretching as long bands on to the base of the manubrium. In Octorchidae (fig. 58) each such band is interrupted, forming one mass at the base of the manubrium and another below the radial canal in each radius, in all eight separate gonad-masses, as the name implies. In some Leptomedusae excretory “marginal tubercles” are developed on the ring-canal.
Classification.—As in the Gymnoblastea, the difficulty of uniting the hydroid and medusan systems into one scheme of classification is very great in the present state of our knowledge. In a great many Leptomedusae the hydroid stage is as yet unknown, and it is by no means certain even that they possess one. It is quite possible that some of these medusae will be found to be truly hypogenetic, that is to say, with a life-cycle secondarily simplified by suppression of metagenesis. At present, ten recent and one extinct family of Calyptoblastea (Leptomedusae) may be recognized provisionally: