Francis Arthur Bather

“ The starfish are the scavengers of the sea, but unfortunately do not confine their attentions to decaying matter; they eat oysters, clams, mussels, barnacles, sea-snails, worms, Crustacea and even smaller starfish. There is constant war between oyster-fishers and starfish; no less than 42,000 bushels of starfish were removed from the oyster-beds of Connecticut in a single year, but not till they had worked damage to the amount of $631,500. The simplest way in which a starfish eats is by taking small bits of food into the stomach, and ejecting the refuse again through the mouth. ”

“ Thus, the mouth is central and turned away from the sea-floor; the animal does not seize its food by tentacles, limbs or jaws, neither does it move in search of it, but a series of ciliated grooves which radiate from the mouth sweep along currents of water, in the eddies of which minute food-particles are caught up and carried down into the gullet; the undigested food is driven out through an anus which is on the upper or oral side of the theca, but as far distant as practicable from the mouth and ciliated grooves. ”

“ But the scattered details of comparative anatomy are capable of manifold arrangement, while the palimpsest of individual development is not merely fragmentary, but often has the fragments misplaced. The morphologist may propose classifications, and the embryologist may erect genealogical trees, but all schemes which do not agree with the direct evidence of fossils must be abandoned ”

“ In the middle of the body is a small anal opening, and near the angle between two rays is a furrowed plate pierced by many minute pores and called the madreporite. The under surface of the body has the mouth in the centre, and from it deep grooves radiate to the ends of the arms. At the bottom of each groove is a water-vessel, which gives off branches to the podia or sucking-feet on each side of it. ”

“ Starfish begin to eat voraciously when quite young; one less than 3/8th in. across has been observed to eat over fifty young clams of half that length in six days. The more a starfish has to eat the quicker it grows, and it may become sexually mature in less than a year, then producing many thousands of young. Fortunately the increase is kept in check by many causes. The young, while still in the stage of free-swimming larvae, are swallowed in millions by various fish. When they settle down on seaweed their bright colours attract eels and many small fishes. ”

“ Although, in the extreme correlation of the radial food-grooves, nerves, water-vessels, 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. ”

“ Evolution of the Echinoderms.—It is reasonable to suppose that the Coelomata—animals in which the body-cavity is divided into a gut passing from mouth to anus and a hollow (coelom) surrounding it—were derived from the simpler Coelentera, in which the primitive body-cavity (archenteron) is not so divided, and has only one aperture serving as both mouth and anus. ”

“ At a later stage tangles of hemp or cotton waste may, be dragged over the oyster-beds, when the starfish will cling to them by their pedicellariae. They make excellent manure, but are of ho further service to man. Fishermen who catch them in their nets or on their lines often tear them in half and throw them back into the sea. Some of these mutilated animals may, however, grow fresh rays, and thus one may find a starfish consisting of one large ray and four quite small ones, the whole shaped like a comet. ”

“ There is no reason to suppose that the central apical plate of certain free-swimming crinoids has any more to do with the distal foot-plate of the larval Antedon stem than has the so-called centro-dorsal of Antedon itself, which is nothing but the compressed proximal end of the stem. As for the supposed basals of Echinoidea, Asteroidea and Ophiuroidea, they are scarcely to be distinguished among the ten or more small plates that surround the anus of Bothriocidaris, which is the oldest and probably the most ancestral of fossil sea-urchins (fig. 5) . ”

“ But since the mouth is quite small and the food often large, the starfish finds it more convenient to turn its stomach inside out and to wrap it around the animal to be eaten, which is then digested quietly and the stomach withdrawn again. ”

“ 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 Edrioasteroidea—while within the larger of these classes, such as Echinoidea and Crinoidea, fair working classifications had been established. ”

“ The stomach is not a long coil, but a simple sac with branched blind tubes extending into each ray. A generative gland also passes down the side of each ray, and emits the milt or eggs when ripe through a pore near the body. Spawning takes place in spring or early summer. ”

“ 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. ”

“ Probably the best way in which man can keep down the numbers of starfish is by dredging the seaweed in the latter half of July when it is covered with young; a single cartload thrown on shore would capture many millions. ”

“ For, whereas to one brilliant suggestion of far-reaching homology another can always be opposed, by the detailed comparison of individual growth-stages in carefully selected series of fossils, and by the minute application to these of the principle that individual history repeats race history, it actually is possible to unfold lines of descent that do not admit of doubt. ”

“ Sand-stars and brittle-stars are found in all seas, usually occurring in quantities, but are most abundant in the rock-pools of the tropics. By constantly sweeping their arms over the sea bottom, they gather food consisting of minute animals. ”

“ Each plate of the upper circlet supports an arm, and is called a “radial”; the plates of the lower circlet, the “basals,” rest on the stem and alternate with those of the upper circlet, i.e. are interradial in position. Some crinoids have yet another circlet below these, the constituent plates of which are called “infrabasals,” and are situated radially. The tegmen in most primitive forms, as well as in the embryonic stages of the living Antedon (fig. 2) , consists of five large triangular plates, alternating with the radials, and called “orals,” because they roof over the mouth. ”

“ In the sand-stars the rays are comparatively short, with their spines closely pressed to their sides, so that they look like lizards’ tails; in the brittle-stars the rays are much longer and more flexible, with the spines standing out, so that they look like wriggling centipedes attached round a little sea-urchin. The brittle-stars are more active than the sand-stars, and can go more than two yards in a minute ”

“ 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 axial ossicles, which correspond to the plates flooring the arm-groove in a starfish, resemble vertebrae connected by pairs of straight muscular bundles, and articulated by tenon-and-mortise joints, according to whose degree of development the arms vary in their power of coiling. ”

“ The food-grooves are exothecal, i.e. are stretched out from the theca on jointed skeletal processes (brachioles) . These either are close to the mouth or are removed from it upon a series of ambulacral or sub-ambulacral plates not derived immediately from thecal plates, or are separated from the oral centre by hypothecal passages passing beneath terminal plates. The stereom and stroma become arranged in folds and strands at right angles to the sutures of the thecal plates; in higher forms the stereom-folds are in part specialized as pectini-rhombs. Families: Echinosphaeridae ”

“ Pentactaea Theory.—In opposition to the calycinal theory has been the Pentactaea theory of R. Semon. There have always been many zoologists prepared to ascribe an ancestral character to the holothurians. 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 well-developed muscles of the body-wall, supposed to be directly inherited from some worm-like ancestor ”

“ The two former, which may often be found hiding under the rocks, or in the seaweed, or in pools at low tide, resemble the ordinary starfish in having five distinct arms. These, however, as shown in fig. 3, are long and serpent-like, and are attached to a relatively small body or disk. The digestive and generative systems do not extend to the rays but are confined to the body. The arms are cylindrical and have no groove on the under side such as exists in starfish ”

“ The thecal plates, however irregular in some species, always show defined basals and a distinct plate (“radial”) at the end of each ambulacrum; they are in all cases so far affected by pentamerous symmetry that their sutures never cross the ambulacra. ”

“ The evidence for the Pelmatozoic theory is supplied by palaeontology, embryology, the comparative anatomy of the classes, and a consideration of other phyla. Palaeontology, so far as it goes, is a sure guide, but some of the oldest fossiliferous rocks yield remains of distinctly differentiated crinoids, asteroids and echinoids, so that the problem is not solved merely by collecting fossils. ”

“ 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 ”

“ 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. ”

“ Cryptozonia, with marginals inconspicuous and somewhat atrophied in the adult, the supero-marginals separated from the infero-marginals by intercalated plates, with papulae distributed over the whole body, with ambulacrals crowded and narrow, either ambulacrals or adambulacrals prominent in the mouth-skeleton, with pedicellariae stalked or sessile. ”

“ Those that were able to accommodate themselves to this topsy-turvy 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. ”

“ 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. ”

“ 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 apical plates by a stretch of dorsal integument containing skeletal elements ”

“ Rays simple or branched, capable of coiling, since the vertebrae articulate by surfaces of hour-glass shape; ventral arm-plates, and often the others, much reduced; spines reduced or absent. Families: Euryalidae, Gorgonocephalidae, Astrochelidae, Astroschemidae, Astronycidae.
The Silurian genera Eucladia and Euthemon have the rays greatly reduced and merged in the disk, so that the ambulacrals are unseen. There are a few large dorsal, lateral and ventral arm-plates, and at the angles of the latter emerge huge podia with a granular or plated skin. ”