Limnocodium by S. Goto, who considers the genus to be allied to
Olindias. Allman, on the other hand, referred Limnocodium to the
Leptomedusae.
In this connexion must be mentioned, finally, the medusae budded from the fresh-water polyp Microhydra. The polyp-stages of Limnocodium and Microhydra are extremely similar in character. In both cases the hydranth is extremely reduced and has no tentacles, and the polyp forms a colony by budding from the base. In Limnocodium the body secretes a gelatinous mucus to which adhere particles of mud, &c., forming a protective covering. In Microhydra no such protecting case is formed. In view of the great resemblance between Microhydra and the polyp of Limnocodium, it might be expected that the medusae to which they give origin would also be similar. As yet, however, the medusa of Microhydra has only been seen in an immature condition, but it shows some well-marked differences from Limnocodium, especially in the structure of the tentacles, which furnish useful characters for distinguishing species amongst medusae. The possession of a polyp-stage by Limnocodium and Microhydra furnishes an argument against placing them in the Trachylinae. Their sense-organs require renewed investigations. (Browne [10] and [10a].)
Order VI. Siphonophora.—Pelagic floating Hydrozoa with great differentiation of parts, each performing a special function; generally regarded as colonies showing differentiation of individuals in correspondence with a physiological division of labour.
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Fig. 68.—Diagram showing possible modifications of medusiform and hydriform persons of a colony of Siphonophora. The thick black line represents endoderm, the thinner line ectoderm. (After Allman.) |
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n, Pneumatocyst. k, Nectocalyces (swimming bells). l, Hydrophyllium (covering-piece). i, Generative medusiform person. g, Palpon with attached palpacle, h. e, Siphon with branched grappling tentacle, f. m, Stem. |
A typical Siphonophore is a stock or cormus consisting of a number of appendages placed in organic connexion with one another by means of a coenosarc. The coenosarc does not differ in structure from that already described in colonial Hydrozoa. It consists of a hollow tube, or tubes, of which the wall is made up of the two body-layers, ectoderm and endoderm, and the cavity is a continuation of the digestive cavities of the nutritive and other appendages, i.e. of the coelenteron. The coenosarc may consist of a single elongated tube or stolon, forming the stem or axis of the cormus on which, usually, the appendages are arranged in groups termed cormidia; or it may take the form of a compact mass of ramifying, anastomosing tubes, in which case the cormus as a whole has a compact form and cormidia are not distinguishable. In the Disconectae the coenosarc forms a spongy mass, the “centradenia,” which is partly hepatic in function, forming the so-called liver, and partly excretory.
The appendages show various types of form and structure corresponding to different functions. The cormus is always differentiated into two parts; an upper portion termed the nectosome, in which the appendages are locomotor or hydrostatic in function, that is to say, serve for swimming or floating; and a lower portion termed the siphosome, bearing appendages which are nutritive, reproductive or simply protective in function.
Divergent views have been held by different authors both as regards the nature of the cormus as a whole, and as regards the homologies of the different types of appendages borne by it.
The general theories of Siphonophoran morphology are discussed below, but in enumerating the various types of appendages it is convenient to discuss their morphological interpretation at the same time.
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After A. Agassiz, from Lankester’s Treatise on Zoology. |
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Fig. 69.—Porpita, seen from above, showing the pneumatophore and expanded palpons. |
In the nectosome one or more of the following types of appendage occur:—
1. Swimming-bells, termed nectocalyces or nectophores (fig. 68, k), absent in Chondrophorida and Cystophorida; they are contractile and resemble, both in appearance, structure and function, the umbrella of a medusa, with radial canals, ring-canal and velum; but they are without manubrium, tentacles or sense-organs, and are always bilaterally symmetrical, a peculiarity of form related with the fact that they are attached on one side to the stem. A given cormus may bear one or several nectocalyces, and by their contractions they propel the colony slowly along, like so many medusae harnessed together. In cases where the cormus has no pneumatophore the topmost swimming bell may contain an oil-reservoir or oleocyst.
2. The pneumatophore or air-bladder (fig. 68, n), for passive locomotion, forming a float which keeps the cormus at or near the surface of the water. The pneumatophore arises from the ectoderm as a pit or invagination, part of which forms a gas-secreting gland, while the rest gives rise to an air-sack lined by a chitinous cuticle. The orifice of invagination forms a pore which may be closed up or may form a protruding duct or funnel. As in the analogous swim-bladder of fishes, the gas in the pneumatophore can be secreted or absorbed, whereby the specific gravity of the body can be diminished or increased, so as to cause it to float nearer the surface or at a deeper level. Never more than one pneumatophore is found in a cormus, and when present it is always situated at the highest point above the swimming bells, if these are present also. In Velella the pneumatophore becomes of complex structure and sends air-tubes, lined by a chitin and resembling tracheae, down into the compact coenosarc, thus evidently serving a respiratory as well as a hydrostatic function.
Divergent views have been held as to the morphological significance of the pneumatophore. E. Haeckel regarded the whole structure as a glandular ectodermal pit formed on the ex-umbral surface of a medusa-person. C. Chun and, more recently, R. Woltereck [59], on the other hand, have shown that the ectodermal pit which gives rise to the pneumatophore represents an entocodon. Hence the cavity of the air-sack is equivalent to a sub-umbral cavity in which no manubrium is formed, and the pore or orifice of invagination would represent the margin of the umbrella. In the wall of the sack is a double layer of endoderm, the space between which is a continuation of the coelenteron. By coalescence of the endoderm-layers, the coelenteron may be reduced to vessels, usually eight in number, opening into a ring-sinus surrounding the pore. Thus the disposition of the endoderm-cavities is roughly comparable to the gastrovascular system of a medusa.
The difference between the theories of Haeckel and Chun is connected with a further divergence in the interpretation of the stem or axis of the cormus. Haeckel regards it as the equivalent of the manubrium, and as it is implanted on the blind end of the pneumatophore, such a view leads necessarily to the air-sack and gland being a development on the ex-umbral surface of the medusa-person. Chun and Woltereck, on the other hand, regard the stem as a stolo prolifer arising from the aboral pole, that is to say, from the ex-umbrella, similar to that which grows out from the ex-umbral surface of the embryo of the Narcomedusae and produces buds, a view which is certainly supported by the embryological evidence to be adduced shortly.
In the siphosome the following types of appendages occur:—
1. Siphons or nutritive appendages, from which the order takes its name; never absent and usually present in great numbers (fig. 68, e). Each is a tube dilated at or towards the base and containing a mouth at its extremity, leading into a stomach placed in the dilatation already mentioned. The siphons have been compared to the manubrium of a medusa-individual, or to polyps, and hence are sometimes termed gastrozoids.
2. Palpons (fig. 68, g), present in some genera, especially in Physonectae; similar to the siphons but without a mouth, and purely tactile in function, hence sometimes termed dactylozoids. If a distal pore or aperture is present, it is excretory in function; such varieties have been termed “cystons” by Haeckel.
