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AND MORPHOLOGY]
141
HYDROMEDUSAE


pelagic organisms, floating on the surface of the open sea, propelling themselves feebly by the pumping movements of the umbrella produced by contraction of the sub-umbral musculature, and capturing their prey with their tentacles. The genera Cladonema (fig. 20) and Clavatella (fig. 21), however, are ambulatory, creeping forms, living in rock-pools and walking, as it were, on the tips of the proximal branches of each of the tentacles, while the remaining branches serve for capture of food. Cladonema still has the typical medusan structure, and is able to swim about, but in Clavatella the umbrella is so much reduced, that swimming is no longer possible. The remarkable medusa Mnestra parasites is ecto-parasitic throughout life on the pelagic mollusc Phyllirrhoe, attached to it by the sub-umbral surface, and its tentacles have become rudimentary or absent. It is interesting to note that Mnestra has been shown by J. W. Fewkes [15] and R. T. Günther [19] to belong to the same family (Cladonemidae) as Cladonema and Clavatella, and it is reasonable to suppose that the non-parasitic ancestor of Mnestra was, like the other two genera, an ambulatory medusa which acquired louse-like habits. In some species of the genus Cunina (Narcomedusae) the youngest individuals (actinulae) are parasitic on other medusae (see below), but in later life the parasitic habit is abandoned. No other instances are known of sessile habit in Hydromedusae.

EB1911 Hydromedusae - Cladonema radiatum - the medusa walking.jpg

From Allman’s Gymnoblastic Hydroids, by permission of the Council of the Ray Society.

Fig. 20.Cladonema radiatum, the medusa walking on the basal branches of its tentacles (t), which are turned up over the body.


EB1911 Hydromedusae - Clavatella prolifera - ambulatory medusa.jpg

From Allman’s Gymnoblastic Hydroids, by permission of the Council of the Ray Society.

Fig. 21.Clavatella prolifera, ambulatory medusa. t, tentacles; oc, ocelli.


EB1911 Hydromedusae - Corymorpha nutans - adult female medusa.jpg

After E. T. Browne, from Proc. Zool. Soc. of London.

Fig. 22.Corymorpha nutans, adult female Medusa. Magnified 10 diameters.

Hydromedusae23.jpg

After O. Maas, Die craspedoten Medusen der Plankton Expedition, by permission of Lipsius and Tischer.

Fig. 23.Aeginopsis hensenii, slightly magnified, showing the manner in which the tentacles are carried in life.

The external form of the Hydromedusae varies from that of a deep bell or thimble, characteristic of the Anthomedusae, to the shallow saucer-like form characteristic of the Leptomedusae. It is usual for the umbrella to have an even, circular, uninterrupted margin; but in the order Narcomedusae secondary down-growths between the tentacles produce a lobed, indented margin to the umbrella. The marginal tentacles are rarely absent in non-parasitic forms, and are typically four in number, corresponding to the four perradii marked by the radial canals. Interradial tentacles may be also developed, so that the total number present may be increased to eight or to an indefinitely large number. In Willia, Geryonia, &c., however, the tentacles and radial canals are on the plan of six instead of four (figs. 11 and 26). On the other hand, in some cases the tentacles are less in number than the perradii; in Corymorpha (figs. 3 and 22) there is but a single tentacle, while two are found in Amphinema and Gemmaria (Anthomedusae), and in Solmundella bitentaculata (fig. 67) and Aeginopsis hensenii (fig. 23) (Narcomedusae). The tentacles also vary considerably in other ways than in number: first, in form, being usually simple, with a basal bulb, but in Cladonemidae they are branched, often in complicated fashion; secondly, in grouping, being usually given off singly, and at regular intervals from the margin of the umbrella, but in Margelidae and in some Trachomedusae they are given off in tufts or bunches (fig. 24); thirdly, in position and origin, being usually implanted on the extreme edge of the umbrella, but in Narcomedusae they become secondarily shifted and are given off high up on the ex-umbrella (figs. 23 and 25); and, fourthly, in structure, being hollow or solid, as in the polyp. In some medusae, for instance, the remarkable deep-sea family Pectyllidae, the tentacles may bear suckers, by which the animal may attach itself temporarily. It should be mentioned finally that the tentacles are very contractile and extensible, and may therefore present themselves, in one and the same individual, as long, drawn-out threads, or in the form of short corkscrew-like ringlets; they may stream downwards from the sub-umbrella, or be held out horizontally, or be directed upwards over the ex-umbrella (fig. 23). Each species of medusa usually has a characteristic method of carrying its tentacles.

EB1911 Hydromedusae - Rathkea octonemalis.jpg

After O. Maas, Craspedoten Medusen der Siboga-Expedition, by permission of E. S. Brill & Co.

Fig. 24.Rathkea octonemalis.
EB1911 Hydromedusae - Aeginura grimaldii.jpg

After O. Maas, Medusae, in Prince of Monaco’s series.

Fig. 25.Aeginura grimaldii.

The sub-umbrella invariably shows a velum as an inwardly projecting ridge or rim at its margin, within the circle of tentacles; hence the medusae of this sub-class are termed craspedote. The manubrium is absent altogether in the fresh-water medusa Limnocnida, in which the diameter of the mouth exceeds half that of the umbrella; on the other hand, the manubrium may attain a great length, owing to the centre of the sub-umbrella with the stomach being drawn into it, as it were, to form a long proboscis, as in Geryonia. The mouth may be a simple, circular pore at the extremity of the manubrium, or by folding of the edges it may become square or shaped like a Maltese cross, with four corners and four lips. The corners of the mouth may then be drawn out into lobes or lappets, which may have a branched or fringed outline (fig. 27), and in Margelidae the subdivisions of the fringe simulate tentacles (fig. 24).

The internal anatomy of the Hydromedusae shows numerous variations. The stomach may be altogether lodged in the manubrium, from which the radial canals then take origin directly as in Geryonia (Trachomedusae); it may be with or without gastric pouches. The radial canals may be simple or branched, primarily four, rarely six in number. The ring-canal is drawn out in Narcomedusae into festoons corresponding with the lobes of the margin, and may be obliterated altogether (Solmaris). In this order the radial canals are represented only by wide gastric pouches, and in the family Solmaridae are suppressed altogether, so that the tentacles and the festoons of the ring-canal arise directly from the stomach. In Geryonia, centripetal canals, ending blindly, arise from the ring-canal and run in a radial direction towards the centre of the umbrella (fig. 26).

Histology of the Hydromedusa.—The histology described above for the polyp may be taken as the primitive type, from which that