Stylifer, the operculum is lost, animal fixed by a large proboscis which forms a pseudopallium covering the whole shell except the extremity of the spire, parasitic on all groups of Echinoderms. Entosiphon, visceral mass still coiled; shell much reduced, proboscis very long forming a pseudopallium which covers the whole body and projects beyond in the form of a siphon, foot and nervous system present, eyes, branchia and anus absent, parasite in the Holothurian Deima blakei in the Indian Ocean.
The Tornaria larva offers a certain similarity to larvae of Echinoderms (sea-urchins, star-fishes, and sea-cucumbers), and when first discovered was so described.
It is within the bounds of possibility that Tornaria actually does indicate a remote affinity on the part of the Enteropneusta to the Echinoderms, not only on account of its external form, but also by reason of the possession of a dorsal water-pore communicating with the anterior body-cavity.
Echinoderms. Class III.
Mailer's studies on the larval forms of Echinoderms and the discoveries of Vaughan Thompson were appreciated.
For edible purposes the most valuable of the Japanese echinoderms is the sea-slug or bche de mer (namako), which is greatly appreciated and forms an important staple of export to China.
De Pourtales pointed out in 1870; they consist of remains of deep-sea corals, serpulae, echinoderms and mollusca united Sands, gravels, muds, &c.
In the animal kingdom it occurs as both calcite and aragonite in the tests of the foraminifera, echinoderms, brachiopoda, and mollusca; also in the skeletons of sponges and corals.
The Asterocheridae, which have a good swimming capacity, except in the case of Cancerilla tubulata (Dalyell), lead a semi-parasitic life on echinoderms, sponges, &c., imbibing their food.
Huxley viewed them as equivalent to and on a level with the larvae of Echinoderms, and of such other trocho phore larvae as resem-???,, ally adopted.
Its adhesive foot is paralleled by a cup-shaped ciliated depression, possibly nervous, found in all the larvae cited, except some Echinoderms, and which in Asterids and Crinoids actually serves as an organ of attachment.
Corals, echinoderms, brachiopods and all groups of molluscs abounded.
The shell-bearing forms belonging to this group which were known to Linnaeus were placed by him (in 1748) in the third order of his class Vermes under the name " Testacea," whilst the Echinoderms, Hydroids and Annelids, with the naked Mollusca, formed his second order termed " Zoophyta."
The latter is confined to certain Gastropods which live in Echinoderms and are extremely degenerate in structure.
Corals, both reef-builders and others, flourished in the clearer waters; rugose forms are represented by Amplexoid, Zaphrentid and Cyathophyllid types, and by Lithostrotion and Phillipsastraea; common tabulate forms are Chaetetes, Chladochonus, Michelinia, &c. Amongst the echinoderms crinoids were the most numerous individually, dense submarine thickets of the long-stemmed kinds appear to have flourished in many places where their remains consolidated into thick beds of rock; prominent genera are Cyathocrinus, Woodocrinus, Actinocrinus; sea-urchins, Archaeocidaris, Palaeechinus, &c., were present; while the curious extinct Blastoids, which included the groups of Pentremitidae and Codasteridae, attained their maximum development.
MYZOSTOMIDA, a remarkable group of small parasitic worms which live on crinoid echinoderms; they were first discovered by Leuckart in 1827.
Thus, about 1875, the distinction of Echinoderms from such radiate animals as jelly-fish and corals (see Coelentera), by their possession of a body-cavity ("coelom") distinct from the gut, was fully realized; while their severance from the worms (especially Gephyrea), with which some Echinoderms were long confused, had been necessitated by the recognition in all of a radial symmetry, impressed on the original bilateral symmetry of the larva through the growth of a special division of the coelom, known as the "hydrocoel," and giving rise to a set of water-bearing canals - the watervascular or ambulacral system.
There was also sufficient comprehension of the differences between the main classes of Echinoderms - the sea-urchins or Echinoidea, the starfish or Asteroidea, the brittle-stars and their allies known as Ophiuroidea, the worm-like Holothurians, the feather-stars and sea-lilies called Crinoidea, with their extinct relatives the sac-like Cystidea, the bud-formed Blastoidea, and the flattened Edrioasteroideawhile within the larger of these classes, such as Echinoidea and Crinoidea, fair working classifications had been established.
Jaekel, in addition to valuable studies on crinoids and cystids appearing in the Zeitschrift of the German Geological Society, has published the first volume of Die Stammesgeschichte der Pelmatozoen (Berlin, 1899), a richly suggestive work; the Mesozoic Echinoderms of France, Switzerland and Portugal have been made known by P. de Loriol, G.
The theory which had most influence on the conceptions of Echinoderms in the two concluding decades of the 19th century was that of Lover', elaborated by P. H.
The theory was that all the plates just described, and more particularly those of the cup, which were termed "the calycinal system," could be traced, not merely in all crinoids, but in all Echinoderms, whether fixed forms such as cystids and blastoids, or free forms such as ophiuroids and echinoids, even - with the eye of faith - in holothurians.
- Supposed calycinal systems of free-moving Echinoderms. A, regular sea-urchin (Cidaris); B, sea-urchin with a suranal plate (Salenia); C, developing ophiurid (Amphiura); D, young starfish (Zoroaster).
Those plates are perhaps constant throughout sea-urchins and starfish (though it would puzzle any one to detect them in certain Silurian echinoids), and they may be traced in some of the fixed echinoderms; but there is no proof that they represent the radials of a simple crinoid, and there are certainly many cystids in which no such plates existed.
The calycinal theory is not merely an assertion of certain homologies, a few of which might be disputed without affecting the rest: it governs our whole conception of the echinoderms, because it implies their descent from a calyculate ancestor - not a "crinoidphantom," that bogey of the Sarasins, but a form with definite plates subject to a quinqueradiate arrangement, with which its internal organs must likewise have been correlated.
The absence of an apical system of plates; the fact that radial symmetry has not affected the generative organs, as it has in all other recent classes; the welldeveloped muscles of the body-wall, supposed to be directly inherited from some worm-like ancestor; the presence on the inner walls of the body in the family Synaptidae of ciliated funnels, which have been rashly compared to the excretory organs (nephridia) of many worms; the outgrowth from the rectum in other genera of caeca (Cuvierian organs and respiratory trees), which recall the anal glands of the Gephyrean worms; the absence of podia (tube-feet) in many genera, and even of the radial water-vessels in Synaptidae; the absence of that peculiar structure known in other echinoderms by the names "axial organ," "ovoid gland," &c; the simpler form of the larva - all these features have, for good reason or bad, been regarded as primitive.
Semon called this stage the Pentactula, and supposed that, in its early history, the class had passed through a similar stage, which he called the Pentactaea, and regarded as the ancestor of all Echinoderms. It has since been proved that the five tentacles with their canals are interradial, so that one can scarcely look on the Pentactula as a primitive stage, while the apparent simplicity of the Synaptidae, at least as compared with other holothurians, is now believed to be the result of regressive vlu.
Study of the earliest larval stages has always led to the conclusion that the Echinoderms must have descended from some freely-moving form with a bilateral symmetry, and, connecting this with the ideas just mentioned, we reach the conception that this supposed bilateral ancestor (or Dipleurula) may have become fixed, and may have gradually acquired a radial symmetry in consequence of its sedentary mode of life.
The tracing of this history, and the explanation of the general characters of Echinoderms and of the differentiating features of the classes in accordance therewith, constitutes the Pelmatozoic theory.
The Pelmatozoic theory thus regards the Pelmatozoa as the more ancestral forms, and the Pelmatozoan stage as one that must have been passed through by all Echinoderms during their evolution from the Dipleurula.
Although, in the extreme correlation of the radial food-grooves, nerves, watervessels, and so forth, with a radiate symmetry of the theca, such a type differs from the Cystidea, while in the possession of jointed processes from the radial plates, bearing the grooves and the various body-systems outwards from the theca, it differs from all other Echinoderms, nevertheless ancient forms are known which, if they are not themselves the actual links, suggest how the crinoid type may have been evolved from some of the more regular cystids.
Instead of discussing all these questions separately, with the details necessary for an adequate presentation of the argument, we shall now sketch the history of the Echinoderms in accordance with the Pelmatozoic theory.
But only slight modifications are required to produce the Tornaria larva of the Enteropneusta and other larvae, including the special type that is inferred from the Dipleurula larval stages of recent forms to have characterized the ancestor of the Echinoderms. We cannot enter here into all the details of comparison between these larval forms; amid much that is hypothetical a few homologies are widely accepted, and the preceding account will show the kind of relation that the Echinoderms bear to other animals, including what are now usually regarded as the ancestors of the Chordata (to which back-boned animals belong), as well as the nature of the evidence that their study has been, or may be, made to yield.
Which of the coelomic cavities this last is connected with is uncertain, for there is considerable doubt as to the origin of the genital glands in the embryonic development of recent echinoderms. It seems clear, however, that there was but a single duct and a single bunch of reproductive cells, as in the holothurians, though perhaps bifurcate, as in some of those animals.
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 freeswimming, 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: Cystidea Edrioasteroidea Pelmatozoa Holothurioidea Crinoidea Blastoidea Eleutherozoa Stelliformia Echinoidea Brief systematic accounts of these classes follow: Grade A.
It is true that some specialized forms, such as the Brisingidae among starfish, A strophiura and Ophioteresis among ophiurans, contravene the usual diagnoses; but this neither obscures their systematic position, nor does it alter the fact that since early Palaeozoic times these two great groups of stellate echinoderms have evolved along separate lines.
If then we place these groups in a single class, it is not on account of a few anomalous genera, but because the characters set forth above sharply distinguish them from all other echinoderms, and because we have good reason to believe that the ophiurans did not arise independently but have descended from primitive starfish.
Bell, Catalogue of the British Echinoderms in the British Museum (London, 1892); P. H.
P. Sladen, "Homologies of the Primary Larval Plates in the Test of Brachiate Echinoderms," Quart.