existence, by taking food in directly through the mouth, survived, and their podia gradually specialized as sucking feet. Such a form as this, when once its covering-plates had atrophied, would be a starfish without more ado (fig. 12, B); but the sea-urchins present a more difficult problem, on which Bothriocidaris sheds no light. An Upper Silurian echinoid, however, Palaeodiscus, is believed by W. J. Sollas and W. K. Spencer to have had in its ambulacra an inner as well as an outer series of plates. If this be correct, the only change from Edrioaster, as regards the ambulacra, was that in Palaeodiscus the covering-plates could no longer open, but closed permanently over the whole groove, while the podia issued through slits between them. In more typical echinoids the covering-plates alone remained to form the ordinary ambulacral plates, while the flooring-plates disappeared, the canals and other organs remaining as before. In any case we have to admit a closure of the integument over the ciliated groove (fig. 12, D, e) just as in holothurians, since this is necessitated by anatomical evidence. The genital organs in both Asteroidea and Echinoidea would retain the interradial position they first assumed in Edrioaster; and in Echinoidea their primitive temporary openings to the exterior were converted into definite pores, correlated with five interradially placed plates at the aboral pole. The anus also naturally moved to this superior and aboral position. In the Echinoidea the water-canals and associated structures, ending in the terminal plates, stretched right up to these genital plates; but in the Asteroidea they never reached the aboral surface, so that the terminals have always been separated from the aboral pole by a number of plates.
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| Fig. 11.—Edrioaster. A, upper or oral surface of E. Bigsbyi, with the covering-plates on the anterior and left posterior food-grooves, but removed from the others, which show only the flooring-plates, between which are pores; B, under surface of E. Buchianus, with covering-plates on right posterior and right anterior food-grooves (left hand in the drawing). The * denotes the position of the anal interradius. |
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| Fig. 12.—Diagrammatic sections across the ambulacra of A, C, Pelmatozoa, and B, D, Eleutherozoa, placed in the same position for comparison. S, Blood-spaces, of which the homology is still uncertain. |
Analysis of Echinoderm Characters.—Regarding the Echinoderms as a whole in the light of the foregoing account, we may give the following analytic summary of the characters that distinguish them from other coelomate animals:—
They live in salt or brackish water; a primitive bilateral symmetry is still manifest in the right and left divisions of the coelom; the middle coelomic cavities are primitively transformed into two hydrocoels communicating with the exterior indirectly through a duct or ducts of the anterior coelom; stereom, composed of crystalline carbonate of lime, is, with few exceptions, deposited by special amoebocytes in the meshes of a mesodermal stroma, chiefly in the integument; reproductive cells are derived from the endothelium, apparently of the anterior coelom; total segmentation of the ovum produces a coeloblastula and gastrula by invagination; mesenchyme is formed in the segmentation cavity by migration of cells, chiefly from the hypoblast. Known Echinoderms show the following features, imagined to be due to an ancestral pelmatozoic stage:—Increase in the coelomic cavities of the left side, and atrophy of those on the right; the dextral coil of the gut, recognizable in all classes, though often obscured; an incomplete secondary bilateralism about the plane including the main axis and the water-pore or its successor, the madreporite, often obscured by one or other of various tertiary bilateralisms; the change of the hydrocoel into a circumoral, arcuate or ring canal; development through a free-swimming, bilaterally symmetrical, ciliated larva, of which in many cases only a portion is transformed into the adult Echinoderm (where care of the brood has secondarily arisen, this larva is not developed). All living, and most extinct, Echinoderms show the following features, almost certainly due to an ancestral pelmatozoic stage:—An incomplete radial symmetry, of which five is usually the dominant number, is superimposed on the secondary bilateralism, owing to the outgrowth from the mouth region of one unpaired and two paired ciliated grooves; these have a floor of nervous epithelium, and are accompanied by subjacent radial canals from the water-ring, giving off lateral podia and thus forming ambulacra, and by a perihaemal system of canals apparently growing out from coelomic cavities. All living Echinoderms have a lacunar, haemal system of diverse origin; this, the ambulacral system, and the coelomic cavities, contain a fluid holding albumen in solution and carrying numerous amoebocytes, which are developed in special lymph-glands and are capable of wandering through all tissues. The Echinoderms may be divided into seven classes, whose probable relations are thus indicated:—
Brief systematic accounts of these classes follow:—
Grade A. PELMATOZOA.—Echinoderma with the viscera enclosed in a calcified and plated theca, of which the oral surface is uppermost, and which is usually attached, either temporarily or permanently, by the aboral surface. Food brought to the mouth by a subvective system of ciliated grooves, radiating from the mouth either between the plates of the theca (endothecal), or over the theca (epithecal), or along processes from the theca (exothecal: arms, pinnules, &c.), or, in part, and as a secondary development, below the theca (hypothecal). Anus usually in the upper or oral half of the theca, and never aboral. An aborally-placed motor nerve-centre gives off branches to the stroma connecting the various plates of the theca and of its brachial, anal and columnar extensions, and thus co-ordinates the movements of the whole skeleton. The circumoesophageal water-ring communicates indirectly with the exterior; the podia, when present, are respiratory, not locomotor, in function.
Class I. Cystidea.—Pelmatozoa in which radial polymeric symmetry of the theca is developed either not at all or not in complete correlation with the radial symmetry of the ambulacra (such as obtains in Blastoidea and Crinoidea); in which extensions of the food-grooves are exothecal or epithecal or both combined, but neither endothecal nor pierced by podia (as in some Edrioasteroidea) All Palaeozoic.
This class shows much greater diversity of organization than any other, and the classifications proposed by recent writers, such as E. Haeckel, O. Jaekel and F. A. Bather, start from such different points of view that no discussion of them can be attempted here. Following the narrative given above, we recognize a primitive group—Amphoridea—represented by Aristocystis (fig. 8). From this are derived the orders Diploporita (fig. 9) and Rhombifera (fig. 10) and the class Edrioasteroidea, all which have already been described as steps in the evolution of the phylum. But there were also side-branches leading nowhere, and therefore placed in separate orders—Aporita and Carpoidea.
Order 1. Amphoridea.—Radial symmetry has affected neither food-grooves nor thecal plates; nor, probably, nerves, ambulacral vessels, nor gonads. Canals or folds when present in the stereom are irregular. Families: Aristocystidae (fig. 8); Eocystidae.
