Order 2. Flexibilia.—Dicyclica in which proximal brachials are incorporated in the dorsal cup, either by their own sides, or by interbrachials, or by a finely plated skin, but never rigidly; plates may occur between RR. Tegmen flexible, with distinct ambulacrals and numerous small interambulacrals; mouth and food-grooves remain supra-tegminal and open. Top columnal a persistent proximale, often fusing with IBB, which are frequently atrophied in the adult.
All the Palaeozoic representatives have non-pinnulate arms, while the Mesozoic and later forms have them pinnulate. There are other points of difference, so that it is not certain whether the latter really descended from the former. But assuming such a relationship we arrange them in two grades.
|Fig. 14.—A living Pentacrinid, Isocrinus asteria; the first specimen found, after Guettard’s figure published in 1761.|
Order 3. Dicyclica Camerata.—Dicyclica in which the first, and usually the second, orders of brachials are incorporated in the dorsal cup by interbrachials, at first loosely, but afterwards by close suture. IBB always the primitive 5. An anal plate always rests on the posterior basal; mouth and tegminal food-grooves closed; arms pinnulate. Families: Reteocrinidae, Dimerocrinidae, Lampterocrinidae, Rhodocrinidae, Cleiocrinidae.
Class IV. Edrioasteroidea.—Pelmatozoa in which the theca is composed of an indefinite number of irregular plates, some of which are variously differentiated in different genera; with no subvective skeletal appendages, but with central mouth, from which there radiate through the theca five unbranched ambulacra, composed of a double series of alternating plates (covering-plates), sometimes supported by an outer series of larger alternating plates (side-plates or flooring-plates). In some forms at least, pores between (not through) the ambulacral elements, or between them and the thecal plates, seem to have permitted the passage of extensions from the perradial water-vessels. Anus in posterior interradius, on oral surface, closed by valvular pyramid. Hydropore (usually, if not always, present) between mouth and anus. Families: Agelacrinidae, Cyathocystidae, Edrioasteridae, Steganoblastidae. All Palaeozoic. The structure and importance of Edrioaster have been discussed above (figs. 11, 12).
Grade B. ELEUTHEROZOA—Echinoderma in which the theca, which may be but slightly or not at all calcified, is not attached by any portion of its surface, but is usually placed with the oral surface downwards or in the direction of forward locomotion. Food is not conveyed by a subvective system of ciliated grooves, but is taken in directly by the mouth. The anus when present is typically aboral, and approaches the mouth only in a few specialized forms. The aboral nervous system, if indeed it be present at all, is very slightly developed. The circumoesophageal water-ring may lose its connexion with the exterior medium; the podia (absent only in some exceptional forms) may be locomotor, respiratory or sensory in function, but usually are locomotor tube-feet.
The classes of the Eleutherozoa probably arose independently from different branches of the Pelmatozoan stem. The precise relation is not clear, but the order in which they are here placed is believed to be from the more primitive to the more specialized.
Class I. Holothurioidea.—Eleutherozoa normally elongate along the oro-anal axis, which axis and the dorsal hydropore lie in the sagittal plane of a secondary bilateral symmetry. The calcareous skeleton, which may be entirely absent, is usually in the form of minute spicules, sometimes of small irregular plates with no trace of a calycinal or apical system; to these is added a ring of pieces radiately arranged round the oesophagus. Ambulacral appendages take the form of: (1) circumoral tentacles, (2) sucking-feet, (3) papillae; of these (1) alone is always present. The gonads are not radiately disposed.
The comparative anatomy of living forms, combined with the evolutionary hypothesis sketched above, suggests that the early holothurians possessed the following characters: subvective grooves entirely closed; 5 radial canals, proceeding from the water-ring, gave off branches furnished with ampullae to the podia on each side of them, the 10 anterior podia being changed into cylindrical tentacles; the transverse muscles of the body-wall formed a circular layer, probably interrupted at the radii (though Ludwig believes the contrary); longitudinal muscles as paired radial bands, without those special retractors for withdrawing the anterior part of the body which occur in many recent forms; a hydropore connected with the water-ring by a canal in the dorsal mesentery; a gonopore behind the hydropore connected by a single duct with a bunch of genital pouches on each side of the mesentery; gut dextrally coiled, with a simple blood-vascular system, and with an enlargement at the anus for respiration, this eventually producing branched caeca called “respiratory trees”; skeleton reduced to a ring of 5 radial and 5 interradial plates round the gullet, and small plates, with a hexagonally meshed network, dispersed through the integument. Such a form gave rise to descendants differing inter se as regards the suppression of the radial canals and of the podia, the form of the tentacles, and the development of respiratory trees. These anatomical facts are represented in the following classification by H. Ludwig:—
Order 1. Actinopoda.—Radial canals supplying tentacles and podia.
|A. With respiratory trees.|
|(a) With podia||Fam. 1, Holothuriidae.|
|Fam. 4, Cucumariidae.|
|(b) Without podia||Fam. 5, Molpadiidae.|
|B. Without respiratory trees.|
|(a) With podia||Fam. 2, Elpidiidae.|
|(b) Without podia||Fam. 3, Pelagothuriidae.|
Order 2. Paractinopoda.—Neither radial canals nor podia. Tentacles supplied from circular canal. Fam. Synaptidae.
|Fig. 15.—An Aspidochirote Holothurian of the family Holothuriidae, showing the mouth surrounded by tentacles, the anus at the other end of the body, and three of the rows of podia.|
It is admitted, however, that this scheme does not represent the probable descent or relationship of the families. Consideration of the views of Ludwig himself, of H. Östergren, and especially of R. Perrier, suggests the following as a more natural if less obvious arrangement.
Order 1. Aspidochirota.—Tentacles more or less peltate; calcareous ring when present simple and radially symmetrical; no retractors; stone-canal often opens to exterior; genital tubes sometimes restricted to left side in consequence of altered position of gut (Fig. 15.) Families: Elpidiidae (deep-sea forms, with sub-famm. Synallactinae, Deimatinae, Elpidiinae, Psychropotinae), Holothuriidae (shallow water), Pelagothuriidae (pelagic).
Order 2. Dendrochirota.—Tentacles simple or branched, never peltate; calcareous ring well developed, often bilaterally symmetrical; retractor muscles usually present; stone-canal opens internally; genital tubes in right and left tufts.
Class II. Stelliformia (＝Asteroidea sensu lato).—Eleutherozoa with a depressed stellate body composed of a central disk, whence radiate five or more rays; this radiate symmetry affects all the systems of organs, including the genital. The radial water-vessels lie in grooves on the ventral side of flooring-plates (usually called “ambulacrals”); they and their podia are limited to the oral surface of the body and their extremities are separated from the