confinement. The alternative is to fish all stages of the medusa in its growth in the open sea, a slow and laborious method in which the chance of error is very great, unless the series of stages is very complete.
At present, therefore, classifications of the Hydromedusae have a more or less tentative character, and are liable to revision with increased knowledge of the life-histories of these organisms. Many groups bear at present two names, the one representing the group as defined by polyp-characters, the other as defined by medusa-characters. It is not even possible in all cases to be certain that the polyp-group corresponds exactly to the medusa-group, especially in minor systematic categories, such as families.
The following is the main outline of the classification that is Adopted in the present article. Groups founded on polyp-characters are printed in ordinary type, those founded on medusa-characters in italics. For definitions of the groups see below.
Sub-class Hydromedusae (Hydrozoa Craspedota).
- Order I. Eleutheroblastea.
- Order II. Hydroidea (Leptolinae).
- Sub-order 1. Gymnoblastea (Anthomedusae).
- Sub-order 2. Calyptoblastea (Leptomedusae).
- Order III. Hydrocorallinae.
- Order IV. Graptolitoidea.
- Order V. Trachylinae.
- Sub-order 1. Trachomedusae.
- Sub-order 2. Narcomedusae.
- Order VI. Siphonophora.
- Sub-order 1. Chondrophorida.
- Sub-order 2. Calycophorida.
- Sub-order 3. Physophorida.
- Sub-order 4. Cystophorida.
Organization and Morphology of the Hydromedusae.
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Fig. 1.—Diagram of a typical Hydropolyp. |
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a, Hydranth; b, Hydrocaulus; c, Hydrorhiza; t, Tentacle; ps, Perisarc, forming in the region of the hydranth a cup or hydrotheca (h.t),—which, however, is only found in polyps of the order Calyptoblastea. |
As already stated, there occur in the Hydromedusae two distinct types of person, the polyp and the medusa; and either of them is capable of non-sexual reproduction by budding, a process which may lead to the formation of colonies, composed of more or fewer individuals combined and connected together. The morphology of the group thus falls naturally into four sections—(1) the hydropolyp, (2) the polyp-colony, (3) the hydromedusa, (4) the medusa-colonies. Since, however, medusa-colonies occur only in one group, the Siphonophora, and divergent views are held with regard to the morphological interpretation of the members of a siphonophore, only the first three of the above subdivisions of hydromedusa morphology will be dealt with here in a general way, and the morphology of the Siphonophora will be considered under the heading of the group itself.
1. The Hydropolyp (fig. 1)—The general characters of this organism are described above and in the articles Hydrozoa and Polyp. It is rarely free, but usually fixed and incapable of locomotion. The foot by which it is attached often sends out root-like processes—the hydrorhiza (c). The column (b) is generally long, slender and stalk-like (hydrocaulus). Just below the crown of tentacles, however, the body widens out to form a “head,” termed, the hydranth (a), containing a stomach-like dilatation of the digestive cavity. On the upper face of the hydranth the crown of tentacles (t) surrounds the peristome, from which rises the conical hypostome, bearing the mouth at its extremity. The general ectoderm covering the surface of the body has entirely lost the cilia present in the earlier larval stages (planula), and may be naked, or clothed in a cuticle or exoskeleton, the perisarc (ps), which in its simplest condition is a chitinous membrane secreted by the ectoderm. The perisarc when present invests the hydrorhiza and hydrocaulus; it may stop short below the hydranth, or it may extend farther. In general there are two types of exoskeleton, characteristic of the two principal divisions of the Hydroidea. In the Gymnoblastea the perisarc either stops below the hydranth, or, if continued on to it, forms a closely-fitting investment extending as a thin cuticle as far as the bases of the tentacles (e.g. Bimeria, see G. J. Allman [1],[1] pl. xii. figs, 1 and 3). In the Calyptoblastea the perisarc is always continued above the hydrocaulus, and forms a cup, the hydrangium or hydrotheca (h, t), standing off from the body, into which the hydranth can be retracted for shelter and protection.
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From Allman’s Gymnoblastic Hydroids, by permission of the Council of the Ray Society. |
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Fig. 2.—Stauridium productum, portion of the colony magnified; |
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Fig. 3.—Diagram of Corymorpha. A, A hydriform person giving rise to medusiform persons by budding from the margin of the disk; B, free swimming medusa (Steenstrupia of Forbes) detached from the same, with manubrial genitalia, (Anthomedusae) and only one tentacle. (After Allman). |
The architecture of the hydropolyp, simple though it be, furnishes a long series of variations affecting each part of the body. The greatest variation, however, is seen in the tentacles. As regards number, we find in the aberrant forms Protohydra and Microhydra tentacles entirely absent. In the curious hydroid Monobrachium a single tentacle is present, and the same is the case in Clathrozoon; in Amphibrachium and in Lar (fig. 11, A) the polyp bears two tentacles only. The reduction of the tentacles in all these forms may be correlated with their mode of life, and especially with living in a constant current of water, which brings food-particles always from one direction and renders a complete whorl or circle of tentacles unnecessary. Thus Microhydra lives amongst Bryozoa, and appears to utilize the currents produced by these animals. Protohydra occurs in oyster-banks and Monobrachium also grows on the shells of bivalves, and both these hydroids probably fish in the currents produced by the lamellibranchs. Amphibrachium grows in the tissues of a sponge, Euplectella, and protrudes its hydranth into the canal-system of the sponge; and Lar grows on the tubes of the worm Sabella. With the exception of these forms, reduced for the most part in correlation with a semi-parasitic mode of life, the tentacles are usually numerous. It is rare to find in the polyp a regular, symmetrical disposition of the tentacles as in the medusa. The primitive number of four in a whorl is seen, however, in Stauridium (fig. 2) and Cladonema (Allman [1], pl. xvii.), and in Clavatella each whorl consists regularly of eight (Allman, loc. cit. pl. xviii.). As a rule, however, the number in a whorl is irregular. The tentacles may form a single whorl, or more than one; thus in Corymorpha (fig. 3) and Tubularia (fig. 4) there are two circlets; in Stauridium (fig. 2) several; in Coryne and Cordylophora the tentacles are scattered irregularly over the elongated hydranth.
As regards form, the tentacles show a number of types, of which
the most important are (1) filiform, i.e. cylindrical or tapering from