Page:EB1911 - Volume 14.djvu/151

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AND MORPHOLOGY]
139
HYDROMEDUSAE


function of circulating and distributing nutriment through the colony.

The hydroid colony shows many variations in form and architecture which depend simply upon differences in the methods in which polyps are budded.

EB1911 Hydromedusae - Lar sabellarum and two stages of its Medusa, Willia stellata.jpg

After Hincks, Forbes, and Browne. A and B modified from Hincks; C modified from Forbes’s Brit. Naked-eyed Medusae.

Fig. 11.Lar sabellarum and two stages of its Medusa, Willia stellata. A, colony of Lar; B and C, young and adult medusae.

In the first place, buds may be produced only from the hydrorhiza, which grows out and branches to form a basal stolon, typically net-like, spreading over the substratum to which the founder-polyp attached itself. From the stolon the daughter-polyps grow up vertically. The result is a spreading or creeping colony, with the coenosarc in the form of a root-like horizontal network (fig. 5, B; 11, A). Such a colony may undergo two principal modifications. The meshes of the basal network may become very small or virtually obliterated, so that the coenosarc becomes a crust of tubes tending to fuse together, and covered over by a common perisarc. Encrusting colonies of this kind are seen in Clava squamata (fig. 5, A) and Hydractinia (figs. 9, 10), the latter having the perisarc calcified. A further very important modification is seen when the tubes of the basal perisarc do not remain spread out in one plane, but grow in all planes forming a felt-work; the result is a massive colony, such as is seen in the so-called Hydrocorallines (fig. 60), where the interspaces between the coenosarcal tubes are filled up with calcareous matter, or coenosteum, replacing the chitinous perisarc. The result is a stony, solid mass, which contributes to the building up of coral reefs. In massive colonies of this kind no sharp distinction can be drawn between hydrorhiza and hydrocaulus in the coenosarc; it is practically all hydrorhiza. Massive colonies may assume various forms and are often branching or tree-like. A further peculiarity of this type of colony is that the entire coenosarcal complex is covered externally by a common layer of ectoderm; it is not clear how this covering layer is developed.

EB1911 Hydromedusae - Colony of Bougainvillea fruticosa.jpg

Fig. 12.—Colony of Bougainvillea fruticosa, natural size, attached to the underside of a piece of floating timber. (After Allman.)

In the second place, the buds may be produced from the hydrocaulus, growing out laterally from it; the result is an arborescent, tree-like colony (figs. 12, 13). Budding from the hydrocaulus may be combined with budding from the hydrorhiza, so that numerous branching colonies arise from a common basal stolon. In the formation of arborescent colonies, two sharply distinct types of budding are found, which are best described in botanical terminology as the monopodial or racemose, and the sympodial or cymose types respectively; each is characteristic of one of the two sub-orders of the Hydroidea, the Gymnoblastea and Calyptoblastea.

EB1911 Hydromedusae - Portion of colony of Bougainvillea fruticosa.jpg

Fig. 13.—Portion of colony of Bougainvillea fruticosa (Anthomedusae-Gymnoblastea) more magnified. (From Lubbock, after Allman.)

In the monopodial method (figs. 12, 14) the founder-polyp is, theoretically, of unlimited growth in a vertical direction, and as it grows up it throws out buds right and left alternately, so that the first bud produced by it is the lowest down, the second bud is above the first, the third above this again, and so on. Each bud produced by the founder proceeds to grow and to bud in the same way as the founder did, producing a side branch of the main stem. Hence, in a colony of gymnoblastic hydroids, the oldest polyp of each system, that is to say, of the main stem or of a branch, is the topmost polyp; the youngest polyp of the system is the one nearest to the topmost polyp; and the axis of the system is a true axis.

EB1911 Hydromedusae - Diagrams of the monopodial method of budding.jpg

Fig. 14.—Diagrams of the monopodial method of budding, shown in five stages (1-5). F, the founder-polyp; 1, 2, 3, 4, the succession of polyps budded from the founder-polyp; a′, b′, c′, the succession of polyps budded from 1; a², b², polyps budded from 2; a³, polyp budded from 3.

In the sympodial method of budding, on the other hand, the founder-polyp is of limited growth, and forms a bud from its side, which is also of limited growth, and forms a bud in its turn, and so on (figs. 15, 16). Hence, in a colony of calyptoblastic hydroids, the oldest polyp of a system is the lowest; the youngest polyp is the top-