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Fig. 66.—Cunina rhododactyla, one of the Narcomedusae. (After Haeckel.) | |
| c, | Circular canal. |
| h, | “Otoporpae” or centripetal process of the marginal cartilaginous ring connected with tentaculocyst. |
| k, | Stomach. |
| l, | Jelly of the disk. |
| r, | Radiating canal (pouch of stomach). |
| tt, | Tentacles. |
| tw, | Tentacle root. |
The Narcomedusae exhibit peculiarities of form and structure which distinguish them at once from all other Hydromedusae. The umbrella is shallow and has the margin supported by a rim of thickened ectoderm, as in the Trachomedusae, but not so strongly developed. The tentacles are not inserted on the margin of the umbrella, but arise high up on the ex-umbral surface, and the umbrella is prolonged into lobes corresponding to the interspaces between the tentacles. The condition of things can be imagined by supposing that in a medusa primitively of normal build, with tentacles at the margin, the umbrella has grown down past the insertion of the tentacles. As a result of this extension of the umbrellar margin, all structures belonging to this region, namely, the ring-canal, the nerve-rings, and the rim of thickened ectoderm, do not run an even course, but are thrown into festoons, caught up under the insertion of each tentacle in such a way that the ring-canal and its accompaniments form in each notch of the umbrellar margin an inverted V, the apex of which corresponds to the insertion of the tentacle; in some cases the limbs of the V may run for some distance parallel to one another, and may be fused into one, giving a figure better compared to an inverted Y. Thus the ectodermal rim runs round the edge of each lobe of the umbrella and then passes upwards towards the base of the tentacle from the re-entering angle between two adjacent lobes, to form with its fellow of the next lobe a tentacle-clasp or peronium, i.e. a streak of thickened ectoderm supporting the tentacle. Similarly the ring-canal runs round the edge of the lobe as the so-called festoon-canal, and then runs upwards under the peronium to the base of the tentacle as one of a pair of peronial canals, the limbs of the V-like figure already mentioned. The nerve-rings have a similar course. The tentaculocysts are implanted round the margins of the lobes of the umbrella and may be supported by prolongations of the ectodermal rim termed otoporpae (Gehörspangen). The radial canals are represented by wide gastric pouches, and may be absent, so that the tentacles arise directly from the stomach (Solmaridae). The tentacles are always solid, as in Trachomedusae.
The development of the Narcomedusae is in the main similar to that of the Trachomedusae, but shows some remarkable features. In Aeginopsis a planula is formed by multipolar immigration. The two ends of the planula become greatly lengthened and give rise to the two primary tentacles of the actinula, of which the mouth arises from one side of the planula. Hence the principal axis of the future medusa corresponds, not to the longitudinal axis of the planula, but to a transverse axis. This is in some degree parallel to the cases described above, in which a planula gives rise to the hydrorhiza, and buds a polyp laterally.
In Cunina and allied genera the actinula, formed in the manner described, has a hypostome of great length, quite disproportionate to the size of the body, and is further endowed with the power of producing buds from a stolon arising from the aboral side of the body. In these species the actinula is parasitic upon another medusa; for instance, Cunoctantha octonaria upon Turritopsis, C. proboscidea upon Liriope or Geryonia. The parasite effects a lodgment in the host either by invading it as a free-swimming planula, or, apparently, in other cases, as a spore-embryo which is captured and swallowed as food by the host. The parasitic actinula is found attached to the proboscis of the medusa; it thrusts its greatly elongated hypostome into the mouth of the medusa and nourishes itself upon the food in the digestive cavity of its host. At the same time it produces buds from an aboral stolon. The buds become medusae by the direct method of budding described above. In some cases the buds do not become detached at once, but the stolon continues to grow and to produce more buds, forming a “bud-spike” (Knospenähre), which consists of the axial stolon bearing medusa-buds in all stages of development. In such cases the original parent-actinula does not itself become a medusa, but remains arrested in development and ultimately dies off, so that a true alternation of generations is brought about. It is in these parasitic forms that we meet with the method of reproduction by sporogony described above.
In other Narcomedusae, e.g. Cunoctantha fowleri Browne, buds are formed from the sub-umbrella on the under side of the stomach pouches, where later the gonads are developed.
Classification.—Three families of Narcomedusae are recognized (see O. Maas [40]):
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After O. Maas, Craspedoten Medusen der Siboga Expedition, by permission of E. S. Brill & Co. |
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Fig. 67.—Solmundella bitentaculata (Quoy and Gaimard). |
1. Cunanthidae.—With broad gastric pouches which are simple, i.e. undivided, and “pernemal,” i.e. correspond in position with the tentacles. Cunina (fig. 66) with more than eight tentacles; Cunoctantha with eight tentacles, four perradial, four interradial.
2. Aeginidae.—Radii a multiple of four, with radial gastric pouches bifurcated or subdivided; the tentacles are implanted in the notch between the two subdivisions of each (primary) gastric pouch, hence the (secondary) gastric pouches appear to be “internemal” in position, i.e. to alternate in position with the tentacles. Aegina, with four tentacles and eight pouches; Aeginura (fig. 25), with eight tentacles and sixteen pouches; Solmundella (fig. 67), with two tentacles and eight pouches; Aeginopsis (fig. 23), with two or four tentacles and sixteen pouches.
3. Solmaridae.—No gastric pouches; the numerous tentacles arise direct from the stomach, into which also the peronial canals open, so that the ring-canal is cut up into separate festoons. Solmaris, Pegantha, Polyxenia, &c. To this family should be referred, probably, the genus Hydroctena, described by C. Dawydov [11a] and regarded by him as intermediate between Hydromedusae and Ctenophora. See O. Maas [35].
Appendix to the Trachylinae.
Of doubtful position, but commonly referred to the Trachylinae, are the two genera of fresh-water medusae, Limnocodium and Limnocnida.
Limnocodium sowerbyi was first discovered in the Victoria regia tank in the Botanic Gardens, Regent’s Park, London. Since then it has been discovered in other botanic gardens in various parts of Europe, its two most recent appearances being at Lyons (1901) and Munich (1905), occurring always in tanks in which the Victoria regia is cultivated, a fact which indicates that tropical South America is its original habitat. In the same tanks a small hydroid, very similar to Microhydra, has been found, which bears medusa-buds and is probably the stock from which the medusa is budded. It is a remarkable fact that all specimens of Limnocodium hitherto seen have been males; it may be inferred from this either that only one polyp-stock has been introduced into Europe, from which all the medusae seen hitherto have been budded, or perhaps that the female medusa is a sessile gonophore, as in Pennaria. The male gonads are carried on the radial canals.
Limnocnida tanganyicae was discovered first in Lake Tanganyika, but has since been discovered also in Lake Victoria and in the river Niger. It differs from Limnocodium in having practically no manubrium but a wide mouth two-thirds the diameter of the umbrella across. It buds medusae from the margin of the mouth in May and June, and in August and September the gonads are formed in the place where the buds arose. The hydroid phase, if any, is not known.
Both these medusae have sense-organs of a peculiar type, which are said to contain an endodermal axis like the sense-organs of Trachylinae, but the fact has recently been called in question for
