HYDROMEDUSAE, a group of marine animals, recognized as belonging to the Hydrozoa (q.v.) by the following characters. (1) The polyp (hydropolyp) is of simple structure, typically much longer than broad, without ectodermal oesophagus or mesenteries, such as are seen in the anthopolyp (see article [Anthozoa]); the mouth is usually raised above the peristome on a short conical elevation or hypostome; the ectoderm is without cilia. (2) With very few exceptions, the polyp is not the only type of individual that occurs, but alternates in the life-cycle of a given species, with a distinct type, the medusa (q.v.), while in other cases the polyp-stage may be absent altogether, so that only medusa-individuals occur in the life-cycle.
The Hydromedusae represent, therefore, a sub-class of the Hydrozoa. The only other sub-class is the Scyphomedusae (q.v.). The Hydromedusae contrast with the Scyphomedusae in the following points. (1) The polyp, when present, is without the strongly developed longitudinal retractor muscles, forming ridges (taeniolae) projecting into the digestive cavity, seen in the scyphistoma or scyphopolyp. (2) The medusa, when present, has a velum and is hence said to be craspedote; the nervous system forms two continuous rings running above and below the velum; the margin of the umbrella is not lobed (except in Narcomedusae) but entire; there are characteristic differences in the sense-organs (see below, and [Scyphomedusae]); and gastral filaments (phacellae), subgenital pits, &c., are absent. (3) The gonads, whether formed in the polyp or the medusa, are developed in the ectoderm.
The Hydromedusae form a widespread, dominant and highly differentiated group of animals, typically marine, and found in all seas and in all zones of marine life. Fresh-water forms, however, are also known, very few as regards species or genera, but often extremely abundant as individuals. In the British fresh-water fauna only two genera, Hydra and Cordylophora, are found; in America occurs an additional genus, Microhydra. The paucity of fresh-water forms contrasts sharply, with the great abundance of marine genera common in all seas and on every shore. The species of Hydra, however, are extremely common and familiar inhabitants of ponds and ditches.
In fresh-water Hydromedusae the life-cycle is usually secondarily simplified, but in marine forms the life-cycle may be extremely complicated, and a given species often passes in the course of its history through widely different forms adapted to different habitats and modes of life. Apart from larval or embryonic forms there are found typically two types of person, as already stated, the polyp and the medusa, each of which may vary independently of the other, since their environment and life-conditions are usually quite different. Hence both polyp and medusa present characters for classification, and a given species, genus or other taxonomic category may be defined by polyp-characters or medusa-characters or by both combined. If our knowledge of the life-histories of these organisms were perfect, their polymorphism would present no difficulties to classification; but unfortunately this is far from being the case. In the majority of cases we do not know the polyp corresponding to a given medusa, or the medusa that arises from a given polyp.[1] Even when a medusa is seen to be budded, from a polyp under observation in an aquarium, the difficulty is not always solved, since the freshly-liberated, immature medusa may differ greatly from the full-grown, sexually-mature medusa after several months of life on the high seas (see figs. 11, B, C, and 59, a, b, c). To establish the exact relationship it is necessary not only to breed but to rear the medusa, which cannot always be done in 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. ” II. Hydroidea (Leptolinae). Sub-order 1. Gymnoblastea (Anthomedusae). ” 2. Calyptoblastea (Leptomedusae). Order III. Hydrocorallinae. ” IV. Graptolitoidea. ” V. Trachylinae. Sub-order 1. Trachomedusae. ” 2. Narcomedusae. Order VI. Siphonophora. Sub-order 1. Chondrophorida. ” 2. Calycophorida. ” 3. Physophorida. ” 4. Cystophorida. |
Organization and Morphology of the Hydromedusae.
| Fig. 1.—Diagram of a typical Hydropolyp. |
| 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.