ea ies eats etste yp ne teed Dias Sat ary) ‘ot ty erie r nipas At by >, ppetierieretrt tthe famieie nana rt esd Prstreuttit Bi fae atte sale tad beets org ry pe rats ete rte ith evokes Digitized by the Internet Archive in 2010 with funding from University of Toronto http://www.archive.org/details/memoirshar20harv - =) MEMOIRS OF THE MUSEUM OF COMPARATIVE ZOOLOGY AT HARVARD COLLEGE VOI XeXe CAMBRIDGE, U.S.A. PRINTED FOR THE MUSEUM. 1897, - ' i Joun Witson anp Son, Camsriper, U.S. A - THE NORTH AMERICAN CRINOIDEA CAMERATA. By CuHartes WacusmutH and CONTENTS. Frank Sprincer. Chapter L-IX. Page 1-359, SHemoirs of the Hluseum of Comparative Zodlogn AT HARVARD COLLEGE. WOT) Sexe THE NORTH AMERICAN CRINOIDEA CAMERATA. By CHARLES WACHSMUTH ann FRANK SPRINGER. IN TWO VOLUMES WITH EIGHTY-THREE PLATES. Worm le CAMBRIDGE, U.S. A.: Printed for the Museum. May, 1897. & Sot a y at ® ae To the Memory . * - LOUIS AGASSIZ, _ TO WHOSE INFLUENCE AS TEACHER, EXPOUNDER, AND INVESTIGATOR, NATURAL on - a HISTORY IN AMERICA IS SO DEEPLY INDEBTED, AND WHO FIRST INSPIRED AND ENCOURAGED US IN OUR EARLIER STUDIES, This Work IS GRATEFULLY DEDICATED BY ' CHARLES WACHSMUTH. 4 FRANK SPRINGER. NOUR. THE Manuscript of this work in its present form was received at Cam- bridge for publication September 1, 1894. Although in press ever since then, no part of it has been published until now, and the date of the work, for bibliographic purposes, will be that which appears on the title-page. During the long time that has been consumed in the printing of the plates and letter-press work since the completion of the text, many new species of Crinoids have been described by American authors, among which some of those herein mentioned as new are included, and thus anticipated. No attempt has been made to cover these cases by modifying the text, nor have any questions arising upon publications appearing subsequent to the above date been considered here. In a work of this size while in press changes could not be made to keep pace with current researches, and the date of delivery of the Manuscript for publication was therefore taken by the authors as final, so far as they were concerned. It is a source of extreme regret that my learned colleague and long-time friend, Dr. Charles Wachsmuth, did not live to see the publication of this Monograph, to which he had devoted so many years of assiduous labor. Never a robust or healthy man, his last few years were almost a continual struggle against disease. His strength gradually failed, and he passed away on February 7, 1896, at the age of sixty-seven years. Dr. Wachsmuth was a native of Hanover, Germany. He came to the United States in 1852, and soon after engaged in mercantile pursuits at Burlington, Iowa, which became his permanent home. Failing health in time compelled him to relinquish business, and for the last thirty years he applied himself to the study of the Crinoids, first as a recreation and to secure outdoor exercise, and afterwards as his life work, with all the ardor of a scientific devotee. His keen powers of observation, sagacious judgment, and indefatigable energy have left their impress upon the works which have been brought out by us. While his death is a loss to Science not easily repaired, it is to none so great—aside from his family — as to the friend with whom he had worked in pleasant collaboration for so many years. It is with a melancholy pleasure that I avail myself of the opportunity afforded by the appearance of his last work to pay this slight tribute to his memory. FRANK SPRINGER. Las Vegas, New Mexico, 1897. Le “i - INTRODUCTION. Tue present work is the outgrowth of studies begun over twenty years ago under the encouragement of Prof. Louis Agassiz, and prosecuted con- tinuously ever since. During that time, we made two very large crincidal collections, of which the original one, in 1873, was secured by Prof. Agassiz, for the Museum of Comparative Zodlogy. Upon this collection one of the writers, while an assistant at the Museum, laid the foundation of the present work. Since 1877 the investigations were conducted by us jointly, and during that time we have built up together the extensive collection which is known as the collection of Wachsmuth and Springer. The advantage of residing, for a time both of us, at Burlington, a locality so well known for the wealth of its crinoidal remains, gave us excellent opportunities to study the Crinoids in all stages of preservation, and being in the field ourselves, we could pick up such material as would help us in the study of minute details. Since the publication of our first paper on the Crinoids, it has been our aim to direct our special attention to studying the morphology of the vari- ous groups as they appeared to us, with a view to future classification, and to revise the work of the previous writers. The various classifications which had been proposed were not based upon strictly morphological principles, and in many cases widely distinct forms were placed together in the same group. It early became evident to us that we could not hope to gain a correct understanding of the fossil forms except by studying their living represen- tatives. The publication of Carpenter’s two Challenger Reports, and De Loriol’s important Monograph on the Mesozoic and later Crinoids of France, opened to the working paleontologist a new field of research, and enabled him to study the relations between paleozoic and neozoic Crinoids, which had been altogether misunderstood. It had been the general opinion, ever since the time of Johannes Miiller, that all paleeozoic forms were widely distinct from the later ones, a view also held by us until 1890. Before the publication of the first Challenger Report, the attention of paleontologists had been directed almost exclusively to the structure of the dorsal or abactinal side of the calyx; that of the ventral side had been very much neglected, and scarcely any attempt had been made to homologize the 1 2 INTRODUCTION. plates of the tegmen in the different groups. The first attempt in this direction was made by Wachsmuth in 1877, and the subject was taken up again in our Revision of 1879. Dr. P. Herbert Carpenter discussed the question more elaborately in 1884, when it became manifest that our views differed radically upon several important points, and especially as to the identification of the oral plates. The progress of our studies on this and other questions was published from time to time in the Revision, and in short papers. This was done for the double purpose of making known the results of our own studies, and of stimulating inquiry by others upon points that were still obscure. Whatever may be the merits or demerits of the Revision — and that the latter are many and serious none are better aware than we — it accom- plished one of its purposes. It induced research and provoked discussion upon ‘new lines and with an activity unprecedented in this field. The contributions to the knowledge of the subject, resulting directly from these controversies, have been of incalculable value to us, and none the less so because some of our own theories have been from time to time exploded. As the most important result, it has now become clear that the Crinoids were most intimately connected from the Silurian down to the present time, and that only the Camerata—a highly specialized type — became extinct at the close of the Carboniferous. It was not until this fact was realized that the way was opened to a better understanding of the whole Crinoid group, in which, as so often found in Nature, the simpler forms persisted, and led down to present types. Although it seems plain enough now, it was only by slow steps, and after long and patient research, that this result was reached. After a large amount of preliminary work had been done, we proceeded to prepare for publication in permanent form such part of it as we could reasonably hope to finish, and to that end we began the preparation of the illustrations in 1887. The work has grown upon our hands to such an extent that we found it necessary to limit it to the Camerata, the largest and most remarkable group among Palseozoic Crinoids. Thus limited, we could hope to give a reasonably full account of this group, and in connec- tion with discussion of the morphological and systematic relations of the other groups, to give some account of the Crinoids generally. The most of the drawings were made under our personal supervision in our Museum at Burlington; a few were made in Washington. Thirty-five INTRODUCTION. 3 of the plates were drawn by Dr. Charles R. Keyes, the present State geolo- gist of Missouri; thirty-three by Mr. A. M. Westergren, so well known for his drawings for Lovén’s great work on the Echinoids; the remaining twelve by Mr. John R. Ridgway, artist for the United States Geological Survey. The execution of the plates occupied about six years, and we avail ourselves of this opportunity to express our thanks to all of these gentlemen for the fidelity and earnestness with which they performed their work. When the work began to assume a definite shape, Mr. A. Agassiz, on being made acquainted with the extent to which it had progressed, kindly offered to undertake its publication as a part of the Memoirs of the Museum of Comparative Zoélogy at Harvard College. No words of thanks would at all express our sense of the obligation under which this has laid us, not merely for the facility of publication through so desirable a medium, but for the mark of appreciation which this offer implies. If the work shall be found sufficiently useful to science to merit, even in a small degree, the indorsement thus given, we shall deem it the best return we can make. During the studies that led up to this Monograph, we enjoyed the privi- lege of continued communication with our lamented friend, P. Herbert Carpenter, up to the time of his decease. We had some energetic con- troversies in print, and a far greater number in private correspondence that never saw the light. To his incisive and suggestive mind is due the over- throw of more than one promising but untenable theory; and we take a melancholy pleasure in recording here our appreciation of his high attain- ments, and our sense of the great loss which Science has suffered through his untimely death. It has been our purpose to give descriptions of all American species of the Camerata known up to this date, and those that could be recognized have been described anew, with the aids derived from the material brought to light since the original descriptions were made. Many of the species were defined from very imperfect specimens, and often without illustrations. In the latter cases we have, when practicable, figured the type specimen, and when necessary and possible have given figures of additional specimens. During the preparation of the work we have had access to most of the type specimens in the United States and Canada, which were placed in our hands for comparison, study, and illustration. A few only of Prof. Hall’s types in the New York State Cabinet of Natural History at Albany, and some of §S. A. Miller's later species, we were unable to procure. £ INTRODUCTION. Not the least of the pleasure we find in bringing our work to a conclusion, is the opportunity it affords us of acknowledging our obligations to the men of science and collectors of America, for their liberality and personal con- fidence shown to us, by placing in our hands — often for indefinite periods — original, unique, and priceless collections, without the use of which this work would have been impossible. It would be difficult to express in fitting terms of acknowledgment the full measure of our indebtedness to them, and we can only venture the hope that they may find in the work itself some small return for the valuable contributions they have made toward it. To Mr, Agassiz we owe a lasting debt of gratitude: first of all for his personal encouragement and valuable counsel, and next for the use of the magnificent collection of the Museum of Comparative Zoélogy. This has been placed at our disposal without restriction, not only for examination at Cambridge, but for removal to Burlington of all specimens we desired, with liberty to use them as if they were our own. Only those who are acquainted with the character and value of this unrivalled collection can appreciate our obligation for such a use of it. It contains the original collection of De Koninck, of the Belgian Carboniferous Crinoids, and the Schultze collection from the Devonian of the Eifel, —by far the finest collections that have ever been made of the rare Crinoids of those interesting localities. There are also the collections made by Hon. B. J. Hall, Prof. W. H. Barris, and the original collection of Wachsmuth, all from the Burlington limestone, which include the types of a large number of the species described by Hall, White, and Meek and Worthen. In addition to these is the fine collection made by C. B. Dyer, from the Hudson River group, of Cincinnati, con- taining many types of species described by Meek in the Ohio report, besides most excellent material from Waldron, Crawfordsville, and other celebrated localities of the West; also the Walcott collection from New York. A full account of the various collections made use of by us would exceed the limits of a preface, but we cannot refrain from making par- ticular mention of some of them: — The collections in the American Museum of Natural History at New York, containing many of the type specimens of the New York Paleonto- logical Reports, have been at all times accessible to us through the courtesy of Prof. R. P. Whitfield, who has been prompt to send us such specimens as we needed for illustration or comparison, and to give us any desired infor- mation obtainable from the extensive material under his charge. INTRODUCTION. 5 During the lifetime of Prof. Worthen, the eminent Director of the I]linois Geological Survey, and afterwards under the administration of his accomplished successor, Dr. Josua Lindahl, we enjoyed the privilege of unrestricted facilities in the use of the type and other specimens in the State Museum of Natural History at Springfield. The private collection of Worthen, containing a large number of the types of the earlier species described in Hall’s Iowa Reports, was packed up and inaccessible while he held the position of State Geologist; but after his death, when the col- lection was acquired by the State of Illinois and incorporated in the State Museum, we were permitted through the courtesy of Dr. Lindahl to examine it, and were given full use of the valuable type specimens. As a mark of our personal esteem, and in justice to the memory of this pioneer collector and geologist, we have inserted the name of Worthen in the notation of such of his type specimens as are now in the State Collection. These types are of great value, as they are the only types of the early Bur- lington and Keokuk species still in existence, so far as we know, with the exception of a few in the Shumard collection. We have been unable to obtain any information as to the types of Owen and Shumard’s descriptions in the Report for Iowa, Wisconsin, and Minnesota, in 1852, — the first Sub- carboniferous Crinoids described from the West. A considerable part of the collections made during the first Iowa Geological Survey are said to have been destroyed by fire, either at Burlington or Keokuk, and it is supposed that a number of type specimens were lost in this way. McChesney’s types were all lost in the great Chicago fire. The collections in the Canada Survey Museum at Ottawa, containing the types of all of E. Billings’s Lower Silurian species, and the later ones of Whiteaves, have been freely open to us under the authority of Sir Alfred Selwyn, and through the unremitting courtesy of Prof. J. F. Whiteaves. Through the attention of Dr. C. A. White and Prof. C. D. Walcott, we obtained the use of the types of some of Meek’s descriptions in the National Museum at Washington. Prof. §. H. Williams of Ithaca, New York, had the goodness to furnish us for examination the types of species described by him, from the Museum of Cornell University, and some of the types from the Colonel Jewett collection. Through Prof. A. H. Winchell we had the use of the specimens in the collection made by Dr. White, now in the University Museum at Ann Arbor, containing the types of a number of well known Subcarboniferous species. (=p) INTRODUCTION. To Dr. G. Hambach of St. Louis we owe the facility of examining the type specimens in the Shumard collection at the Washington University. We are under special obligations to Prof. Borden, of Borden Institute at New Providence, Indiana, for the opportunity of examining the original collection of Dr. Knapp, of Louisville, from the now exhausted Bear Grass locality near Louisville, containing some of the types of species described by Lyon, Shumard, and Yandell, which now form a part of the Museum of the Borden Institute. To Prof. S. Calvin we are indebted for the loan of fine specimens from the Hamilton of Iowa and New York, from which we made descriptions of several species. Our thanks are also due to Prof. W. H. Barris, of Davenport, Ia., who gave us the use of his type specimens and other valuable material from the Hamilton group of Iowa and Michigan, which were under his charge in the Museum of the Davenport Academy of Science. We also avail ourselves of this opportunity of expressing our high appre- ciation of the favors extended to us by Dr. G. Lindstrém, of the National Museum of Sweden at Stockholm, in which are deposited the magnificent collections of Crinoids from the Upper Silurian of Gotland that formed the basis of Angelin’s descriptions. Not only has he at all times allowed us the privilege of having special drawings made from unique specimens in the Museum, but on one occasion, on learning of the difficulty under which we labored from want of adequate material to study the genus Crotalocrinus, he sent us, without solicitation, a series of specimens, including some of Angelin’s originals, with liberty to retain them as long as might be necessary for the examination we desired to make. We have also to acknowledge our indebtedness to Mr. Walter R. Billings, of Ottawa, Canada, for the loan of types of Trenton species in his own col- lection, and also for his good offices in securing for our use the collections of Messrs. Stewart and I. F. Sowter. Besides this, Mr. Billings has from time to time furnished us valuable notes in relation to many rare and inter- esting forms, often illustrated by exquisite drawings from his own hand. We extend our thanks to Mr. John Stewart and Mr. I. F. Sowter, of Ottawa, Canada, for the use of their specimens, — Mr. Stewart having at one time sent us his whole collection for study. To the owners of private collections in the United States our obligations are so numerous and varied that we cannot attempt to express in proper terms of appreciation our indebtedness to each one. INTRODUCTION. 7 It is especially difficult for us to express our obligations to Mr. Victor W. Lyon, of Jeffersonville, Ind., who with the utmost liberality placed his own collection at our disposal, and also that of his father, the late Major Sidney S. Lyon, through which we secured the use of all the types of the species described by Major Lyon himself, and by Lyon and Casseday. Mr. Lisbon A. Cox, of Keokuk, Ia., gave us access to his extensive and unique collection from the Keokuk limestone, containing the types of a large number of species described by Worthen in Vol. VII. of the Mlinois Reports. Mrs. Yandell, of Louisville, Kentucky, has sent us for examination some rare types in the collection of the late Dr. L. P. Yandell. We tender our special thanks to this venerable lady for the efforts she made to serve us. To the naturalists and collectors of Cincinnati and vicinity we are indebted for great facilities in studying the crinoidal fauna of the Lower Silurian of that region. Mr. I. H. Harris, of Waynesville, Ohio, placed at our disposal the species of his magnificent collection of Hudson River Cri- noids. Mr. 8. A. Miller favored us with the loan of his valuable types of Lower Silurian species. Mr. E. O. Ulrich, of Newport, Ky., has sent us for examination the types of his species, and besides other instructive speci- mens. He also used his influence in our behalf with Messrs. Oeh and Vaupel, who placed some of their finest specimens in our hands. Dr. E. N. 8. Ringueberg, of Lockport, N. Y., has sent us the types of his species of the Niagara group of Western New York, with liberty to use them as we might find desirable. To Prof. J. M. Clarke, of Albany, N. Y., we are indebted for the use of valuable type specimens from the Hamilton group of New York, then in his private collection, but since passed into the New York State Cabinet. Mr. Thomas A. Greene, of Milwaukee, Wis., placed in our hands a large collection of natural casts from the Niagara group, including types of the Waukegan species; and Mr. W. C. Egan, of Chicago, a similar collection from near Chicago, containing the types of the species described from that locality. Mr. F. A. Sampson, of Sedalia, Mo., gave us the use of his collection, containing the types of a large number of species described by S. A. Miller in the Missouri and Indiana Reports; and Prof. R. R. Rowley, of Louisiana, Mo., furnished us the types of his species. We are also indebted for the use of specimens and friendly acts in vari- 8 INTRODUCTION. ous ways to Mr. Asa S. Tiffany, of Davenport, Dr. C. C. Washburn, of Wal- dron, Ind., Dr. Moses Elrod, of Hartsville, Ind., Rev. H. Herzer, of Berea, Ohio, Rey. John Davis, of Louisiana, Mo., Mr. D. H. Todd, of Kansas City, Mo., Mr. G. M. Nickels, of Sparta, Ill, Mr. E. Brown, of Belfast, N. Y., and others. To Dr. Horace G. Griffith, formerly of Burlington, now of Philadelphia, we express our grateful acknowledgments for his intelligent and unremitting efforts to aid us in the prosecution of this work, and for his steadfast devo- tion to our interests manifested upon every occasion. We also bear in kindly remembrance our former townsman, Mr, James Love, whose fine collection was always at our disposal, and which, together with one made by Mr. J. W. Giles, afterwards passed into our hands. Dr. Charles R. Keyes has at all times exhibited a lively interest in the progress of our work, and we owe to him not only the procurement of some valuable specimens, but other friendly offices. Nor do we forget our good friend, Orestes St. John, whose keen eye and rare judgment, and no less his skilful pencil, have always been at our ser- vice. We have from him some unsurpassed structural drawings, and he presented us several unique Crinoids from the Coal Measures of Kansas. Our thanks are due to Mr. Wm. F. E. Gurley, of Danville, Ill., for the use of specimens from Waldron, and to Mr. A. C. Benedict, of Indianapolis, for the use of specimens obtained by him at St. Paul, Ind. In addition to the facilities above mentioned, we have had during the preparation of this work our own collection, which contains authentic speci- mens of nine-tenths of the species of Crinoids described from the United States, and two-thirds of all the European species. From many of the typical localities we have been able to obtain, either by purchase of local collections, or by personal exertions, large series of specimens, by means of which it has been possible to study in many cases, and among different genera, the indi- vidual variation existing in the limits of a species, and the modifications due to growth. In looking over the descriptions it will probably surprise some of the authors to find so many of their species placed in the synonym lists, but we were obliged to do so after careful study and comparison with authentic specimens. CHARLES WACHSMUTH. FRANK SPRINGER. Burtinecton, Towa, May 1, 1894. Received at Cambridge, September 1, 1894. ALEXANDER: AGASSIZ. TABLE OF CONTENTS. INTRODUCTORY PART. INTRODUCTION HisToRIcAL TERMINOLOGY . . - + «+ « MORPHOLOGICAL PART. PRIMARY AND SUPPLEMENTARY PLATES 38 Tuer PLATES OF THE ABACTINAL SysTEM 38-88 The Stem and its Appendages 38-52 Basals and Infrabasals . 52-73 The Radials . . : 68-73 The Arms and Pinules . 73-88 Tur PLATES OF THE ACTINAL SystEM 88-104 The Orals . F 88-89 Mouth and pean rie 99 Tue SUPPLEMENTARY PLATES The distribution of the Plates and their relations to the different SYSTEMATIC PART. CLASSIFICATION . ‘ 144-172 Definition of the Crinoidea and their Primary Subdivisions . 169 Analysis of the Families 170 Geological and Geographical Distri- bution of the Camerata 172 DESCRIPTIVE. RETEOCRINIDAE . . 173-187 Analysis of the Genera . 173 Geological and Geographical Dis- tribution . 173 Reteocrinus 176 Xenocrinus « 182 Tanaocrinus : 185 THYSANOCRINIDAE . 188-214 Analysis of the Genera . 188 Geological and Geographical Distri- bution 188 Thysanocrinus . 190 Ptychocrinus 197 Hyptiocrinus 200 Idiocrinus 202 Lampterocrinus 207 Siphonocrinus . 209 RHODOCRINIDAE . . 215-263 Analysis of the Genera . 215 Geological and Geographical Distri- bution .. . 216 Rhodocrinns . ..- +--+ - 218 Gilbertsocrinus . . . . . - 233 Pace 1-10 11-31 32-37 105 groups - - 105-123 The Anal Plates and the Anus. 124-139 INTERNAL Cavity or THE Catyx 140-142 The Chambered Organ and the Axial Canals 140 The Conyoluted Organ . 145 Thylacocrinus . 248 Diabolocrinus . 249 Archeocrinus . 253 Rhaphanocrinus 258 Lyriocrinus 5 261 MELOCRINIDAE . 264-329 Analysis of the Gener oars . 264 Geological and Geographical Distr bution 265 Melocrinites 267 Glyptocrinus 267 Periglyptocrinus 277 Stelidioerinus . 279 Mariacrinus 281 Macrostylocrinus . 285 Melocrinus . 292 Dolatocrinites 304 Technocrinus 304 * Allocrinus 806 Centrocrinus 308 Dolatocrinus 310 Stereocrinus 324 Hadrocrinus a. On CALYPTOCRINIDAE . 830-359 Analysis of the (Gener a. 330 Geological and Geographical Distri- bution 330 Eucaly ptocrinus 332 Callicrinus . . + + + « « « 803 - > a => e ‘= ‘ 4 +. = . ‘ i > " > y & . he i} THE CRINOIDEA CAMERATA OF NORTH AMERICA. INTRODUCTORY PART. I. HISTORICAL. Tue first reference to Fossil Crinoids, according to De Koninck, was made by Agricola in the second half of the sixteenth century. He distin- guished between Trochites, Entrochus, and Encrinus. The former name he applied to all detached stem-joints; Hntrochus to a series of joints, and Encrinus to the calyx of Enerinus liliformis, at that time the only Crinoid in which a crown had been found in connection with the stem. As early as the seventeenth and eighteenth centuries the crinoidal remains received the attention of a large number of writers, some of whom regarded them as plants, others as animals. Rosinus, who lived at the beginning of the eighteenth century, was the first writer to show that the Crimoids were not plants, as before then gene- rally supposed, but were closely related to the Asterids, and especially to the group which afterwards received the name Euryale. He also supposed that the Trochites and Entrochites were parts of Znerinus, and not inde- pendent bodies. An important advance in the knowledge of the Crinoids was made by Guettard,* who described the first recent Stalked Crinoid that ever came to Europe. He gave this species, which was afterwards known ‘as Pentacrinus caput-meduse Lamk., the popular name ‘‘ Palmier marin,” and took it to be the type of all fossil Crinoids with pentagonal stem, as opposed to those with a round stem, of which he thought the living type had not been dis- covered. He gave a moderately fair description of its structure; but added * Mémoire sur les Encrinites et les pitrres étoilées, dans lequel on traitera aussi des Entroques. (Mém. de l Acad. Roy. Soc. de Paris, 1755 (published 1761), pp. 224-318. 12 THE CRINOIDEA CAMERATA OF NORTH AMERICA. little as to the systematic position of the Crinoids generally, stating, how- ever, that they were neither Polyps nor Starfishes. Linné, throughout all the editions of his “Systema Naturae,” placed the Crinoids among the corals. Blumenbach* has the credit of having been the first writer who ranked them with the Asteroids and Ophiurids among the order “ Vermes crustacei,” which corresponds approximately to our pres- ent Echinoderms. Lamarck, in the first edition of his “Systéme des Ani- maux sans vertébres,” published in 1801, ranged them among the “ Polypes a rayons coralligénes,” along with Gorgonia, Umbellula, and Pennatula; but he afterwards modified this opinion, and in 18127 referred the Crinoids to the “ Polypes flottants,’ which he arranged next to the Radiata. In 18164 he placed the Encrinites (Stalked Crinoids) among the Polyps, but the Comatulz (Free-floaters) among the Echinoderms. Schweigger § directed attention to the close resemblance that he found to exist between the arm structure of stalked Crinoids and Comatule, and he considered the two forms to be closely related. Cuvier in 1817, || and again in 1830, placed the Crinoids among the Echinoderms. The name “Crinoidea,” with the rank of a family, was proposed in 1821 by J. S. Miller, for the lily-shaped, radiate animals which theretofore had been known as Hncrinites and Pentacrinites. He restricted the group to the Brachiate forms, and to those provided with a stem, as appears by the follow- ing definition: ** “An animal with a round, oval or angular column, com- posed of numerous articulating joints forming a cup-like body containing the viscera, from whose upper rim proceed five articulated fingers.” This description includes neither Blastoids nor Cystids, which were placed by Miller’s successors as subordinate groups under the Crinoids. It also ex- cludes the Comatule and the genus Marsupites, which have no stem, and which probably for this reason were referred by him to the “ Stelleride.” Among the latter he recognized four divisions: “Comatule, Euryale, Ophiura, and Asteria,” and he placed Marsupites in the same group with Euryale. Miller knew little of the structure of the Comatule, but enough * Handbuch der Naturgeschichte, 1780. + Extrait du cours de Zodlogie du Muséum d’histoire naturelle sur les Animaux sans vertébres, ete., 1812. { Histoire naturelle des Animaux sans Vertébres, ete., 1815-1822. § Handb. der skeletlosen, ungegliederten Thiere, Leipzig, 1820, p. 528. || Le Régne animal, 1817 (1), Vol. IV., p. 12. {| Op. cit. (Bd. of 1830.) ** A Natural History of the Crinoidea, Bristol, 1821, p. 7. HISTORICAL. 138 to be struck by the resemblance they bear to the crown of Pentacrinus, and he pointed out that the pentagonal plate at the base of the subglobose body of the Comatulz occupies the position of the first column joint of the “ Crinoidea.” Miller subdivided the Crinoidea into four groups: the ArTICULATA, to which he referred the genera “ Apiocrinites, Encrinites and Pentacriniles ;” the SEMIARTICULATA with “ Poferiocrinites;” the INARTICULATA with “ Cyathocri- nites, Actinocrinites, Rhodocrinites, and Platyerinites ;”’ and the Coapunata with “ Eugeniacrinites.” His primary groups were based upon the mode of union between the stem and calyx, and between the latter and the arms; his genera upon the number and arrangement of the plates in the dorsal cup. Considering that in 1821 only about twenty-five species of Stalked Crinoids, recent and fossil, were known, and many of them only from fragmentary specimens, we cannot help admiring the genius of Miller, who brought order out of chaos, and laid the foundation of the present classification of the Crinoidea. His genera have been generally accepted, and are now rec- ognized as the types of well-marked families, Miller introduced an elaborate terminology, but unfortunately did not always apply his terms to the same parts. In some of his genera he gave the term “pelvis” to the proximal ring of the plates witliin the calyx, in others to the plates of the ring above. In Apiocrinus and Actinocrinus he called the radials “ first costals;” the succeeding ones “ second costals,” and the first axillaries “scapule.” In Platycrinus, however, and in Poteriocrinus and Cyathocrinus, the radials are his scapule, and are followed by arm plates. Platycrinus, according to Miller, has no costals at all; but in the dicyclic Cyathocrinus and Poteriocrinus costals are said to be represented by the plates of the interradial basal ring. A year after the appearance of Miller's work, Schlotheim published the first part of the Supplement to his Petrefactenkunde,* reproducing therein Miller's figures together with his own, and adopting his generic and specific names. A year later, however, in the second part of that work, he withdrew Miller’s generic names, and referred all Stalked Crinoids back to Enerinus and Pentacrinus respectively. Goldfuss in his great work + adopted Miller’s classification and termi- nology. Cumberland ¢ did not consider Miller’s name “ Crinoidea” appro- * Nachtrige zur Petrefactenkunde, 1822-1823 (2 Bande, mit 37 Kupfer-Platten), + Petrefacta Germania, Diisseldorf, 1826-1833. } Reliquie conservate, Bristol, 1826. 14 THE CRINOIDEA CAMERATA OF NORTH AMERICA. priate, as not a single Hnerinus or Pentacrinus resembled in the smallest degree a lily, either in stem, root, flower, or bud. Nor did he think it absolutely proved that they were animals instead of coraline sensitive plants. In 1825, Say * described three new species under the genus Pentremites, which he made the type of a new family of the Crinoidea, and proposed for it the name Blastoidea. He also described the genus Caryocrinites, which he took to be intermediate between Cyathocrinus and Actinocrinus. In the same year two additional species of Pentremiles were described by G. B. Sowerby. In the years following up to 1840, a number of new species of Crinoids were described by Mantell (1822), Pander (1830), Steininger (1831, 1837, and 1838), Goldfuss (1832 and 1838), Zenker (1833), Phillips (1835-1836), F. A. Roemer (1836 and 1839), Heisinger (1857), Sedgwick and Murchison (1837), D’Orbigny (1837), Miinster (1858-1846), and others; but they added little to the general knowledge of the Crinoids. L. Agassiz, in his Prodréme d’une Monographie des Radiaires ou Echino- dermes,t referred the Crinoids to the “order” Séellerides, together with the “genera” Comatula and Marsupites, which, as he stated, differ from the Crinoids only in not having a stem. J. V. Thompson, in 1836, discovered { that the small species, which he had described in 1827 as Pentacrinus europaeus, loses its stem at a more advanced stage of growth, and changes into a free-floating Comatula. Thompson also discovered the ovaries along the pinnules. Other important discoveries in relation to the anatomy and development of recent Crinoids were made by Adams, Heusinger, Savigny, Delle Chiaje, Blainville, and Dujardin. D’Orbigny in 1859 described the remarkable recent genus LZolopus,§ a Crinoid not attached by a jointed stem, but by the lower end of the calyx. In 1840 appeared the classical work of Johannes Miiller, “ Ueber den Bau des Pentacrinus caput-medusx,” || which marked a new era in the history of the Crinoidea, and threw a flood of light upon the whole group. Miiller in this work discussed the relation between the Pentacrinites and Comatule, and pointed out the anatomical differences in the structure of Crinoids and * Journ. Acad. Nat. Sci. Phila., Vol. 1V., pp. 292-296. + Mémoires de la Société des Sciences Naturelles de Neufchatel, 1835, Tom. I, p. 168. { Memoir on the Starfish of the genus Comatula (Edinburgh New Philos. Journ., Vol. 20, p. 295. § Wiegmann’s Archiv fiir Naturgeschichte, I, p. 185, Taf. 5, Figs. 2-7. || Read before the Berlin Akademie der Wissenschaften, April 30, 1840. HISTORICAL. 15 Asterids. He also introduced a more rational terminology for the plates of the calyx, which is still used by Zoblogists and Palzontologists. He proposed the name “ Basala” for the pelvis plates of Miller, and “ 2adialia” for Mil- ler’s scapule and costals, including the first axillary. When the rays are free above the first radial, the axillary supports the arms; but when that plate forms a part of the calyx, it is followed by the “ Distichalia,” and these by the “ Palmaria”’ For the supplementary plates he introduced the terms “ Interradialia, Interdistichalia and Interpalmaria.” Miiller divided the Crinoids into three great groups: the “ Crinoidea Arti- culata,’ the “ Crinoidea Tessellata” and the “ Crinoidea Costata,’ which he defined, and of which he gave a list of the principal genera. Of the Articulata, to which Miiller referred Pentacrinus, Apiocrinus, Encrinus, and the Comatule, he said that the rays develop directly from the base of the calyx, and the. lower ray plates are united laterally by a skin, which is either naked or paved with irregular plates; that this skin is continued to the ven- tral disk, closing the ventral side of calyx and arms; and that the radials consist of three successive plates, of which the first and second, and the axillary and the first arm-joint, are united by muscles. Miiller’s Tessellata combine Miller’s “ Semiarticulata,’ and “ Inarticu- lata,”.and include the Blastoidea, Cystidea and the Cretaceous genus Mw- supites. The calyx is composed of 3, 4 or 5 basals, which are sometimes separated from the radials by a ring of “ parabasals.” Between the radials there may be “ interradials,” and between the distichals and palmars, “in- terdistichals,” and “interpalmars.” The “Scheitel” (ventral disk) covers the whole ventral surface; it is constructed of solid plates, united at their edges. The Tessellata were subdivided by Miiller into two groups: Crinoidea with arms, and Crinoidea without arms. To the former he referred all true Crinoids and the Cystid genus Caryocrinus, forms having no separate anal opening and no “ Tentakelfurchen ” (food grooves) upon the disk, and none probably upon the arms. The armless Crinoids comprise the “ Pentremites” (Blastoidea) and “ Sphaeronites ” (Cystidea), forms with separate mouth and anus. Miiller’s classification, although a great advance upon that of Miller, was not accepted by the French and English writers succeeding him; but it was revived later on by Ferd. Roemer and von Zittel. In 1842 a classification was proposed by Thos. Austin and Thos. Austin 16 THE CRINOIDEA CAMERATA OF NORTH AMERICA. Jr.,* who made the Crinoidea in the widest sense a class of the Echinoder- mata, to which they gave the name “ Pinnastella.” Among the latter they placed as orders : — I. CIONACINETI (Stalked Crinoids). Il. LIBERIA (Free-floaters). Families: Apiocrinoidea. Gnuathocrinoidea. Poteriocrinoidea. Astracrinoidea. Encrinoidea. Comastella. Pentacrinoidea. Marspiocrinoidea. Platycrinoidea. Actinocrinoidea. Dimerocrinoidea. Only the Platycrinoidea and Poteriocrinoidea ¢ were defined. To the former they referred the genera: Platycrinus, Dichocrinus, Hexa- crinus, Caryocrinus, and Cyathocrinus ; to the latter: Poteriocrinus, Symbatho- erinus, Extracrinus, and Pentacrinus. Their Encrinoidea include: Enerinites, Eucalyptocrinites, Cupressocrinites, and Euryocrinites ; their Marsupiocrinoidea : Marsupiocrinites and Crotalocrintes ; the Actinocrinoidea : Actinocrinites, Rho- docrinites, Melocrinites and Tetracrinites. 'The Austins placed the Blastoidea with the Sphaeroidocrinoidea, and the Periechinidze under the Columnida. D’Orbigny in 18524 undertook to subdivide the Crinoids (including Blastoids and Cystids) into twelve families, which contain most heterogene- ous elements. For description he divided the plates of the calyx into zones, without reference to their radial or interradial position. The next classification was that of Ferd. Roemer, who wrote in 1855,§ and divided the Crinoidea into three great groups : — I. Actinoidea, or true Crinoids, having large, pinnule-bearing arms. Il. Cystidea, Crinoids in which the arms are feebly developed or wanting, and mouth and anus are separate. III. Blastoidea, Crinoids without arms, the soft parts of the animal enclosed within a calyx, which is closed from all sides, leaving only a few openings. The