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Proceedings of the United States National Museum UNUSUAL FORMS OF FOSSIL CRINOIDS By Frank Springer, Associate in Paleontology, United States National Museum INTRODUCTION It was originally my intention when preparing the monograph on the Crinoidea Flexibilia to follow it up with a similar systematic treatise on the Inadunata. But the time and labor consumed in bringing out such works proved to be too great, and I have been compelled to relinquish the project for the last-mentioned treatise. Many preparatory studies were made, however, with such a work in view, some of which are embraced in the present paper, containing the results of researches extending over many years, which I have long desired to publish, but which have been delayed by the pressure of other matters. The material used is chiefly from my own col- lection, now in the United States National Museum, much of it accumulated with special reference to these researches. Most of the drawings have been made by Mr. Kenneth M. Chapman, of Santa Fe, New Mexico, and some by Miss Francesca Wieser, of the Na- tional Museum. I am much indebted to Dr. R. S. Bassler and Dr. C. E. Resser, of the National Museum, for their assistance in making the photographs upon which the drawings are based. THE USUAL STRUCTURE OF CRINOIDS The generalized picture of a crinoid, as seen in the forms by which the class is most commonly known, is that of a marine organ- ism having a calyx, cup, or theca composed of calcareous plates defi- nitely arranged, inclosing a cavity which contains the visceral organs; attached to the sea bottom or other objects, temporarily or permanently, by a columnar stem formed by a series of centrally perforated segments; and fringed at the opposite end by a ring of movable, food-gathering appendages called arms; these are likewise built up of calcareous segments, but instead of being pierced by a central opening for a tubular canal, as in the stem, they are notched at one side, forming a groove along which the microscopic food is No. 2581.—Proceedings U. S. National Museum, Vol. 67. Art. 9. 23832—26 1 2 PROCEEDINGS OF THE NATIONAL. MUSEUM vol. 67 carried by means of currents produced by minute cilia to the oral center, or mouth. The arms when gathering food are stretched up- ward or outward, but may be folded together when not so employed in such a manner that the vital organs and soft parts are protected by an outer envelope of hard calcareous material. The excrement is discharged through an orifice, either directly piercing the tegmen, or roof, or at the end or side of a projecting tube, or exceptionally through the side wall of the cup. The plates of which the calyx is composed are arranged usually according to a quinqueradiate plan, most plainly developed on the lower side to which the stem is attached, from which the radiation extends upward and continues into the arms. This is called the dorsal side, in contradistinction to that next to which the arms are located, which is the ventral side or tegmen; and the part of the calyx below the origin of the arms is called the dorsal cup, or simply the cup, which incloses or supports the visceral organs. This cup in the usual crinoid is made up of a base, consisting either of a single ring of five, four, three, or two plates (or some of these coalesced into a single undivided disk), called basals; or of this and another alternating ring below it of five or three plates (also sometimes fused into one), called infrabasals; and following the basal ring another set of five alternating plates, called radials, with sometimes a ring of interradials in line with them. The two rings of cup plates—basals and radials—are the funda- mental elements of the adult crinoid structure. They are found throughout all the major divisions of the class, and of the blastoids also. In the typical crinoid these two elements are symmetrically arranged in alternating contact with each other, and the radials are symmetric among themselves. This is the plan in the great majority of fossil, and practically all the existing, crinoids. But to this gen- eralized type there are numerous exceptions. The general tendency of the crinoids as a class, starting in the earliest Paleozoic with the asymmetric form which they inherited from cystidean ancestors, was toward that of complete pentamerous symmetry. This was attained in the Mesozoic and Recent crinoids generally, and in a few genera of the Paleozoic. In most of the Paleozoic crinoids an incomplete pentamerous symmetry was reached, modified by the bilaterial symmetry due to the presence of anal structures. The course of this development was by no means one of continual progress. It was marked by numerous recessions and sudden changes. Two of the major divisions of the crinoids, Camerata and Flexi- bilia, and for the most part the third, Inadunata, culminated and were extinguished before the end of the Paleozoic. During this vast stretch of time manv peculiar modifications of the type as we chiefly art 9 UNUSUAL FORMS OF FOSSIL CRINOIDS SPRINGER 3 know it occurred, the causes of which are unknown, but which serve as examples of the infinite variety with which the processes of nature go on. Many of these are already well known, and some others it is the purpose of this paper to illustrate. No general discussion of crinoid morphology is here attempted. For that reference should be had to the works of Bather in part 3 of the Lankester Zoology; of Austin H. Clark on the Existing Crinoids; of Wachsmuth and Springer on the Crinoidea Camerata; of myself on the Crinoidea Flexibilia; and to various papers by Jaekel. I am simply presenting a number of facts, more or less unrelated, bearing upon some of the unusual features and changes above alluded to, by which the general progress of the crinoids in geo- logical time was characterized; together with discussion of some remarkable parallel modifications which I am now able to illustrate more completely than was possible heretofore. THE COILED BILATERAL STEM While the possession of a stem is one of the characters by which the crinoids as a class are best known—so much so that a large part of the early literature treating of them was devoted mainly to discus- sion of the isolated segments of which it is composed—there was in some instances a remarkable tendency to get rid of it. This append- age probably existed in the larval stage of all crinoids, but in by far the greater part of the existing forms, the comatulids, it is cast off at approaching maturity, and the adult crinoid becomes a free floater. In some, the pentacrinites, it attained a great length; and in some (thought to be due to living on reefs in shallow water) the stem is very short, or even disappears, fixation being accomplished through direct attachment by the fused base. In most of the Paleozoic genera the stem was present, sometimes attached or anchored by remarkable specialized structures (Scypho- 1 crlnus, Ancyrocrinus, etc.) ; in many cases only resting in the soft coze of the sea bottom by finely pointed terminals, and often not permanently fixed. 2 The great Paleozoic divisions, Camerata and Flexibilia, and the chiefly Paleozoic Inadunata, had stems with but few exceptions, such as Edriocrinus in which the base was fused and attached directly to other objects or became rounded and free from attach- ment, and Agassizocrinus in which a free floating stage in some of its species was also attained. The same thing is true of the Mesozoic crinoids. 1 Springer, On the crinoid genus Scyphocrinus, Smiths. Misc. Coll. Publication 2440, 1917, pp. 9-12. 2 Dr. Edwin Kirk's instructive paper on Eleutherozoic relmatozoa, Proc. U. S. Nat. Mus., vol. 41, 1911, pp. 1-137. 4 PROCEEDINGS OF THE NATIONAL MUSEUM vol.67 A special form of stem development which I wish to illustrate is that in which the usually cylindrical or pentagonal straight stem loses its characteristic shape, becomes coiled, and takes on a bilateral symmetry, by which for the greater part of its length it is flattened or concave at the inner side, with the columnals elliptic or crescentic in cross section ; and the cirri, instead of occurring in whorls around the column, are borne only in two rows at or near the margins of the flattened or concave side, or sometimes at the back. This structure is correlated—either as a cause or an effect—with a tendency of the crown to bend back upon the stem, and of the stem to coil around it in the opposite direction in such a way that the crown may be tightly enclosed within the coil and completely enveloped by the cirri. The part of the stem proximal to the calyx is circular in section, much re- duced in diameter, relatively short as a rule, but variable in length; and it bears no cirri. The reversed curve is similar to that of a swan's neck, and for convenience the term " neck " may be used for it in the discussions. This character was evidently protective in origin, as the crown is often very small in comparison with the stem; and in some forms, where the cirri are short and closely packed, the crown is so sur- rounded by the stem structures that it can rarely be seen in the fos- sils ; while in others, in which the cirri are more scattered but strong and branching, the neck upon which the crown was borne is long, slender, and exceedingly brittle, not tightly rolled, but evidently loosely enveloped in a fringe formed by the strong cirri.
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