Regarded from the standpoint of the evolutionist, we have here an interesting series of gradations. At one end of the series we have the Echinus with its rays all united in a box-like rigid shell. At the other end of the series we have the Brittle-stars and Comatulæ with their highly muscular and mobile rays. Midway in the series we have Astropecten and the common Star-fish, where the rays are flexible and mobile, though not nearly so much so as in the Brittle-stars. Now, the point to observe is, that in correlation with this graduated difference in the mobility of the rays, there is a correspondingly graduated difference in the development of the ambulacral system of suckers. For in Echinus this system is seen in its most elaborate and efficient form; in the common Star-fish the suckers are still the most important organs of locomotion, though the muscularity of the rays has begun to tell upon the development of the specially ambulacral system, the suckers not being so long or so powerful as they are in Echinus. Lastly, the Brittle-stars and Comatulæ have altogether discarded the use of their sucking feet in favour of the much more efficient organs of locomotion supplied by their muscular rays; and, as a consequence, their feet have dwindled into useless rudiments, while the rays have become limb-like in their activity.

Fig. 47.—Natural righting movements of common Star-fish.

There is only one other point in connection with the natural movements of the Echinodermata which it is necessary for me to touch upon. All the species when turned upon their backs are able again to right themselves; but seeing, as I have just observed, that the organs of locomotion in the different species are not the same, the methods to which these species have to resort in executing the righting manœuvre are correspondingly diverse. Thus, the Brittle-stars can easily perform the needful manœuvre by wriggling some of their snakelike arms under the inverted disc, and heaving the whole body over by the mere muscularity of these organs. The common Star-fish, however, experiences more difficulty, and executes the manœuvre mainly by means of its suckers. That is to say, it twists round the tip of one or more of its rays (Fig. 47) until the ambulacral feet there situated are able to get a firm hold of the floor of the tank (a); then, by a successive and similar action of the ambulacral feet further back in the series, the whole ray is twisted round (b), so that the ambulacral surface of the end is applied flat against the floor of the tank (c). The manœuvre continuing, the semi-turn or spiral travels progressing all the way down the ray. Usually two or three adjacent rays perform this manœuvre simultaneously; but if, as sometimes happens, two opposite rays should begin to do so, one of them soon ceases to continue the manœuvre, and one or both of the rays adjacent to the other takes it up instead, so assisting and not thwarting the action. The spirals of the co-operating rays being invariably turned in the same direction (Fig. 47, a, b, and c), the result is, when they have proceeded sufficiently far down the rays, to drag over the remaining rays, which then abandon their hold of the bottom of the tank, so as not to offer any resistance to the lifting action of the active rays. The whole movement does not occupy more than half a minute. As a general rule, the rays are from the first co-ordinated to effect the righting movement in the direction in which it is finally to take place—the rays which are to be the active ones alone twisting over, and so twisting that all their spirals turn in the same direction.

A Star-fish (Astropecten) which is intermediate between the Brittle-star and the common Star-fish, in that its ambulacral feet are partly aborted (having lost their suckers, as shown in Fig. 44) and its rays more mobile than those of the common Star-fish, rights itself after the manner shown in Fig. 48, where the animal is represented as standing on the tips of four of its rays, while the fifth one is just about to be thrown upwards and over the others, in order to carry with it the two adjacent rays, and so eventually to overbalance the system round the fulcrum supplied by the tips of the other two rays, and thus bring the animal down upon its ventral surface.

Fig. 48.—Righting movements of Astropecten.

Fig. 49.

But it is in the case of Echinus that these righting movements become most interesting, from the fact that they are so much more difficult to accomplish than they are in the case of the Star-fishes. For while a Star-fish is provided with flat, flexible, and muscular rays, comprising a small and light mass in relation to the motive power, an Echinus is a rigid, non-muscular, and globular mass, whose only motive power available for conducting the manœuvre is that which is supplied by its relatively feeble ambulacral feet. It is, therefore, scarcely surprising that unless the specimens chosen for these observations are perfectly fresh and vigorous, they are unable to right themselves at all; they remain permanently inverted till they die. But if the specimens are fresh and vigorous, they are sooner or later sure to succeed in righting themselves, and their method of doing so is always the same. Two, or perhaps three, adjacent rows of suckers are chosen out of the five, as the rows which are to accomplish the task (Fig. 49). As many feet upon the rows as can reach the floor of the tank are protruded downwards and fastened firmly to the floor; their combined action then serves to tilt the globe slightly over in their own direction, the anchoring feet on the other or opposite rows meanwhile releasing their hold of the tank to admit of this tilting (Fig. 50). The effect of this tilting is to enable the next feet in the active ambulacral rows to touch the floor of the tank, and, when they have established their hold, they assist in increasing the tilt; then the next feet in the series lay hold, and so on, till the globe slowly but steadily rises upon its equator (Fig. 51). The difficulty of raising such a heavy mass into this position by means of the slender motive power available can be at once appreciated on witnessing the performance, so that one is surprised, notwithstanding the co-ordination displayed by all the suckers, that they are able to accomplish the work assigned to them. That the process is in truth a very laborious one is manifest, not only from the extreme slowness with which it takes place, but also because, as already observed, in the case of not perfectly strong specimens complete failure may attend the efforts to reach the position of resting on the equator—the Echinus, after rearing up a certain height, becoming exhausted and again falling back upon its ab-oral pole. Moreover, in some cases it is interesting to observe that when the equator position has been reached with difficulty, the Echinus, as it were, gives itself a breathing space before beginning the movement of descent—drawing in all its pedicels save those which hold it securely in the position to which it has attained, and remaining in a state of absolute quiescence for a prolonged time. It then suddenly begins to protrude all its feet again, and to continue its manœuvre. At any time during such a period of rest, a stimulus of any kind will immediately determine a recommencement of the manœuvre.