ig yD
4 . ’ fi . \ , \ FM : wy j 7 i “4 " 1 *
y ‘ ’ . ‘ ' a } ; ve ial : i ‘ na » 7 a» 1 e f ' x
=
“a
: : :
-
SMITHSONIAN INSTITUTION UNITED STATES NATIONAL MUSEUM Bulletin 82
A MONOGRAPH OF THE EXISTING CRINOIDS
BY
AUSTIN HOBART CLARK
Assistant Curator, Division of Marine Invertebrates United States National Museum
VOLUME 1 THE COMATULIDS
PART 1
WASHINGTON GOVERNMENT PRINTING OFFICE 1915
[i “a8 | , see LPNs BULLETIN OF THE UNITED STATES NATIONAL MUSEUM, “i F. IssuED June 10, 1915. ns 7 rhs Tor D 7 4 man . ‘ : | uf is 7 -
ADVERTISEMENT.
The scientific publications of the United States National Museum consist of two series, the Proceedings and the Bulletins.
The Proceedings, the first volume of which was issued in 1878, are intended pri- marily as a medium for the publication of original, and usually brief, papers based on the collections of the National Museum, presenting newly-acquired facts in zoology, geology, and anthropology, including descriptions of new forms of animals, and revisions of limited groups. One or two volumes are issued annually and dis- tributed to libraries and scientific organizations. A limited number of copies of each paper, in pamphlet form, is distributed to specialists and others interested in the different subjects as soon as printed. The date of publication is printed on each paper, and these dates are also recorded in the table of contents of the volumes.
The Bulletins, the first of which was issued in 1875, consist of a series of separate publications comprising chiefly monographs of large zoological groups and other general systematic treatises (occasionally in several volumes), faunal works, reports of expeditions, and catalogues of type-specimens, special collections, etc. The majority of the volumes are octavos, but a quarto size has been adopted in a few instances in which large plates were regarded as indispensable.
Since 1902 a series of octavo volumes containing papers relating to the botanical collections of the Museum, and known as the Contributions from the National Her- barium, has been published as bulletins.
The present work forms No. 82 of the Bulletin series.
RicHarp RaTHBun, Assistant Secretary Smithsonian Institution, In charge of the United States National Museum.
Wasuineton, D. C., April 21, 1915.
HUNRAL ALO
sary An lene iawn fleet (auntlaleess gle blr eave cb yeaa coop Eo bege . 7 aA + Cyit a waa Harney le al sly leghinaeal wad 4 onal litve Temuco jeden ous sige sah) Lewd sxe Jur! slaw mt fetnaiie le Gutfaniy, pits wi a ne i be o? atont denn avi citawri tr aL lati al e iopeerne ulpatiiin Ya Witt ‘imep 10 atbod Jey wee ute eulig ot oli bea
: atte Viiiee ORLA Pepe «ti neue Ww) 2) ONT mclaitre ate VW ehighns hh Gadvite. hohe ‘A AIDA asl ee! batitutemm, tris ure ire Gee tecrpodcr lpia ye Lith Widllawate o byattithin te 5 oF iste) fn hea
efte Ua tiiping ad ud tne Tong Va gtab ult FOF | aie pew « vngif
wurtal'y alt hewiire aaa te te T edt Di Neeser lragmanat dnd a , Le
butte | Wie @ We rani AVE Wi Deaths eal? iste Tee deyttl ont) “ae a id wien ‘iitidte we copa ei he epecetery bree facind ier dley Lie youth Glatolan oy) etait of hina eed ee fh Nhat olf vata adtertyotiow sdpeigfe anes Vaotin yy) Ie tris gerbes brn broken» get nays Ve wAl aiante igpoline imse \ «ah WM ee aiu?,; pit’ mo eae
ihe ye taiay eilbut aw habainet ote « ynalepogral aie f Abas ba dt liat Orbe ot si ial 4 wil CTL es Ai lk meal wo 1d pabel a ws th AWaantiia ld alt an ca rl | tonal whet wih We ey
tin’ ae a Faded ony weve nn
rulball lo Ga 04 ate atv I
whiny Han agrey faci yodiaws pork ta) meget
“| WAL aia Th paws naive)
VAT! «tranoutl ; gi wabwetdewl Myetoa vApleolk lvokheak
ioe WOE: ular aitite. We io val i verona ait
ANE PTS Viney al) Ls
TABLE OF CONTENTS.
PRR CO set oe yar con erate ee 2 io ise AD) eas ae eeee tee: ack Sere ccs dee te eek bal eee History of the work, with an account of the material studied........................------- Generalimethodrot treatments: «425 9-5 stone sce ooo anc cbine 2 51s tees SU
itimbryolopyadevelopmentiand anatomy? —.cse sens a osm ee eet a ses eens Jase Wariantal and aberrants ome. ces ec yee cat actarse dere |= Seto aureys slay mi ee gal eieie stasis Saareoee = Philosophicalliconclusions®.2 ee. = epic ese ens a-ak eee = acess cee ae Relative status ofthe recent crimolds ss s+. seseeer os.c.< cine sects seme tad eases Aree
MU strattiOns: oes taseee Seen ecio ce se eet ose e acai oases ene sees rceccseen eas ssnsee ee Identification of the specimens upon which this work is based. ............-.------------- Individuals and institutions to which the author is indebted.................-..---------- History iomihesubjeet-naseeer ote caen nce Som cece ee ease acessarissecccateues Bee eee eeIs ee (Genera lilintonyseee et ese cine sean asters seats es eine se ee ee Sotelo ue ote =e eer History of the intensive work upon the comatulids...............-.-.--------+--+--+-------- Generallenrveyiouiheynistonyaets s--c2o2e sae seiacioccine comes eine eee ccisecieles seat cise cemiee Glossary of terms used in the description of a comatulid..............--.---.------------------ Hxplanation of symbolase secs ete ks sce o seees Soeericie aes sisie wes Sac selects cia aU a nteeiele curate Weacripwonrola COmAmiiGsen oes yee ee ses case tee nee Sie Sean eee se eee esacss tac meserse aes fidentiicatonobrecent comatn oss 6.22 ceNee mac ee ease ie ane nes eerie eee eee Structure and Anatomy coe srcioe = Sect m sass oc armies mente orarap er ein in alee aes om aise a = siamo HAS Lomsy Ol UN GASEl OJ CC Ose arate efor areca inet taal ese tla Paton sere alas ease aera ore Benes Generalihistory Se est ooo aes Sear ars c coos Sonat Se seae cee See ee se canis t Sons ymoen arse Generalistirveysotthe)histony-seass= sees se ate sae aaa ee ee eee eens sa ast Organizationvorgthe CrinOid saa eere a. 5 ee cele eer ete lar eee aia el a aa ee Generailfremarksscs.55 fue So eee cee en eee soins oe ee as Sa ee ee Se eee ypstandisermenta won cate - n= soe eas eae ag se ae ae on Seen aae aeemeraaaee Developmentronthoe lanvesyce so ster = eote seo ae easiest eee eee ena nae wie iets Hehmodermalsskeleton' 4 sesso soso ees coat eaten cise ome wale oe teem enone see PATHLOLONRY Bare ee = oe te ae tec male St ee ne Re eee oe ee clo oe aries meters ais Orientation and the metameric divisions of the echinoderms.................-.------- Relationships between the digestive tube and asymmetry..............--.-------------
Ones Ol SIMtAT Ake) tal’ POLODCY,sa-ce eee sce no ae en eee ie sis srasite wie eines Rania ere ternalinkelotons-c2. 525 sccsnn cc odes ese ce eee tee erin aan emer oe seinisefac soisiemn Skeleton of the heteroradiate echinoderms................--.--------------------+----
Effect of external mechanics upon the crinoids........-.....---./.---------+---------- HMarliesterinoidss= tote. oc seen sec oa eee nc een no ne ec/ae ee kiss aucinrais cet s ons =
IB Tes COI Bete ee ae ete tare ER ks cee eat ie ee wicca Re INGRVOUR IS VSUG I eee sates siete a actaie tate aa eicte ei ae ee ieee inna eat ae
TE yes ee I wale Sere aetna te ete eles cena ore lay eer cies es into cte Taam
SENET WE bot eae Soec aS pogo Sec oben. Se beanss cecos cb ose cece epaceEEpE eee
IRER@re LOR VO FP ALIN Saisie eae oe =i = Soe eet aia = ere = alate eerie eieie = eter tette mat Ganitaliducistsscsiscnis. oS. am stints asinerseimise a sie oo aaieis ae eee eralaoleint= = mictercte [ate tamale
VI TABLE OF CONTENTS.
Structure and anatomy—Continued.
Organization of the crinoids—Continued. Page. Abies a wesc ee ee a a ea recat ees ae re a ah tS = 190 Promachocrinus and’ ‘Thaumatocrmus::-.. 22 52.0. 2. eee en «secs se eens ee eee 191
Galcarcoun structiras.<.: 2. ei. cece aks ested ete ae ie ao sae derele yore 2 kite he eo 194 Skeleton. asia: whole: <.c625 s2aaesec eco: ates ie Se sete soe nent tos eee eee eee ee 194 Golunin «2:0... 6s oo eee ss SEE Soe SOR POs SE ee Aan ec ce ee ee Ee 198 Gan trodoreal® cae bos a eae eee wea rere citings eS STS SES els hd cote cir ee eee 219 Gime fs oon ose ease Bens RS ws aaa aes BR Ree Te aa = Sn a ee Ae 258 Infrabasalas 2 Coe 68 ong ss Set cee cots salatnctlacnaiwcle ws aoe dl aaw is sate nee ae ees 313 Basals, and structures formed from and associated with them ................-..------ 316 Radianal <s2 v5 Letts cs sonst eae = Je he eae eee eee eee ee ioe oie Dees ere eet 331 Tnitenradials: ansllces << fs ree oe ee ee 335 ‘Perisoniie:interradials ssc. < Peso Ses et ee ee ee 339 Primary, plates of. the disk= 222 2255..2 5. Ste tes Stee eee oe ee ee eee 339 Oral cece aos eae ek cee SUSE SSUES oad SORE O ee Seee ae 340 General proportions of;calyx and ita contents'=>- 3.2.2. ee we ee ee eee 341
Radiale...22< 52: 2esscco eee htt ed Reenter ce ed nee ctad ce oe cae eee ee eee eee 348
Explanation of plates: << <.-)2'22). tose ene ea ee see ae eRe enemas 383
A MONOGRAPH OF THE EXISTING CRINOIDS.
By Austin Hoparr CLark, Assistant Curator, Division of Marine Invertebrates, United States National Museum.
PREFACE. HISTORY OF THE WORK, WITH AN ACCOUNT OF THE MATERIAL STUDIED.
Upon the return of the United States Fisheries steamec Albatross from her cruise in 1906 through the Bering Sea and in Asiatic Russian and Japanese waters, during which I accon:panied her as acting naturalist, the Commissioner of Fisheries, Hon. George M. Bowers, very kindly intrusted to me the work of identifying and describing the Crinoidea which had been collected.
The aim of the work as originally planned was the preparation of a memoir dealing only with the specimens collected on this cruise, but it was later suggested that I include in my study the crinoids from the North Pacific which had previously been collected by the Albatross, and had been deposited in the United States National Museum.
The work proved to be far more of an undertaking than had been anticipated ; so great was the number of new species and so radically did they alter the conception of the recent representatives of the Crinoidea as a whole that I was at last forced to begin at the beginning and to review critically the whole subject.
The two great monographs of Dr. Philip Herbert Carpenter were, of course, the foundation upon which I expected to build; but, with the enormous mass of material at hand, I soon discovered that the subject must be approached along somewhat different lines from those by which it was approached by Carpenter, especially in regard to the comatulids. I therefore laid aside the literature and, with nothing but the specimens before me, attempted to elucidate the systematic problems presented with a mind free from preconceived ideas. The specimens were grouped into species and the species into tentative genera, and these genera again into tentative families, upon characters, both external and internal, which I myself determined; when my ideas had become sufficiently crystallized I again took up the study of the literature and compared my results with those of Carpenter.
Up to this time the work had all been based upon north Pacific species from the Asiatic and American coasts. Radical systematic revision based upon material from a limited district only has seldom proved long lived, and T was therefore extremely anxious to examine additional collections in order to test my conclusions and to investigate further many problems connected with geographic, bathymetric, and
1
2 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
thermal distribution, and with ecology, in which I had become interested largely through my observations while at sea.
Prof. Walter K. Fisher, of Stanford University, California, had been working upon the echinoderms collected by the Albatross among the Hawaiian Islands in 1902; with the greatest liberality he offered me the crinoids of the collections for exami- nation in connection with my other Pacific material.
To Dr. Hubert Lyman Clark, of the Museum of Comparative Zodlogy at Cam- bridge, Massachusetts, had been assigned a large collection of crinoids from Japan and eastern Asia brought together by the Albatross in 1900, and this he most courte- ously offered me to supplement the 1906 collections from the same locality.
The Japanese collections which I had seen up to this time had all been from comparatively deep water, and certain species, long known as inhabitants of the coasts of that country, were conspicuously absent. Mr. Frank Springer, however, realizing the situation, most generously purchased and deposited in the United States National Museum the entire collection made during years of investigation of the marine fauna off southern Japan by Mr. Alan Owston, of Yokohama, in his yacht, the Golden Hind.
Up to now my material had been almost entirely from the North Pacific, and from deep water, although I had examined some of the more common littoral species of Australia and Brazil. The absence of specimens from that great wonder- land of marine zodlogy, the East Indian Archipelago, was keenly felt as a great handicap. But Dr. Theodor Mortensen, of Copenhagen, Denmark, understanding my predicament, with the greatest generosity offered me the entire magnificent collection under his charge, a collection doubly interesting in having been previ- ously examined both by Prof. C. F. Liitken and Dr. P. H. Carpenter. Most of the specimens were from the eastern tropics, many of them having been collected by the Danish consul at Singapore, Mr. Svend Gad; notwithstanding all the Japanese material I had previously studied I found no less than six new species from that country; altogether it formed an invaluable supplement to the Pacific material already at hand.
Shortly after I received the Copenhagen collections, Drs. W. Weltner and R. Hartmeyer, of Berlin, at the instigation of Dr. Th. Studer, of Berne, sent me the collection made by the German steamer Gazelle in northwestern Australia, hitherto an unknown territory so far as regards its crinoid fauna. This collection had been examined by Dr. P. H. Carpenter, and most of the specimens had been tentatively identified, but he had been unable to complete a report upon it before his death.
Mr. Owen Bryant had been conducting dredging operations along the coast of Labrador and had collected some crinoids there, which he very kindly turned over to me.
The great area occupied by the Indian Ocean had hitherto remained almost a blank in so far as our knowledge of its crinoidal inhabitants was concerned; a few specimens had been noted from the Mergui Archipelago, the Andamans, Ceylon, the Red Sea and Mauritius, with one or two, usually more or less doubtful, additional records. I was therefore delighted when Dr. N. Annandale, of the Indian Museum, at the instigation of Dr. F. A. Bather, of the British Museum, offered me for study
MONOGRAPH OF THE EXISTING CRIN OIDS. 3
the entire collection brought together by the Royal Indian Marine Surveying steamer Investigator, as well as the other collections belonging to the Indian Museum, collec- tions remarkable for their unusual completeness.
The large and extensive collections of West Indian crinoids made by the ships of the United States Bureau of Fisheries and deposited in the United States National Museum were now studied in connection with the East Indian material, having been up to this time laid aside awaiting the publication of the report upon the Blake collection of 187879 by Dr. Clemens Hartlaub.
The Berlin Museum, through Drs. W. Weltner and R. Hartmeyer, now submitted to me their entire crinoid collection, an act of courtesy the importance of which to me can only be realized when it is remembered that this collection contains the types of very many of the species described by Prof. Johannes Miller and by Dr. Clemens Hartlaub; and Doctor Mortensen sent me a magnificent collection of Arctic material, undoubtedly the finest in existence, together with the specimens which he himself had collected while in the West Indies. :
At this time the Australian Museum, through Dr. Robert Etheridge, jr., its
curator, sent me for study their entire collection of Australian crinoids, numbering nearly one thousand specimens.
The Albatross was now engaged in an exhaustive survey of the marine resources of the Philippine Islands, and the crinoids which she obtained were, as fast as they accumulated, turned over to me by the Bureau of Fisheries.
Two summers were spent at the Museum of Comparative Zodlogy at Cam- bridge, Massachusetts, working in the library and studying the fine collections of crinoids there, which are especially important in containing a number of species from the Challenger dredgings, named by P. H. Carpenter. Every courtesy Was extended to me, and I was very materially assisted in my work by Mr. Alexander Agassiz, the director of the University Museums, Mr. Samuel Henshaw, the Curator of the Museum of Comparative Zoélogy, and by Dr. Hubert Lyman Clark, the assistant in whose care is the collection of echinoderms. I was also fortunate in having the constant companionship and friendly advice of Prof. Robert Tracy Jackson, of Harvard College, who was at that time engaged in the preparation of his monograph of the palxozoic echinoids.
The collections and library of the Boston Society of Natural History were fre- quently consulted, for which privilege I am indebted to Dr. Glover Morrill Allen and to Mr. Charles W. Johnson. I also visited the Peabody Museum at Yale University, New Haven, Connecticut, where I enjoyed the advantage of reviewing the material with Prof. Addison FE. Verrill; and the museum of the Essex Institute at Salem, Massachusetts, of which Prof. Edward S. Morse is the director.
During the summer of 1910 I spent four months in Europe studying the collec- tions in the various museums, paying particular attention to the types of previous authors; I visited Bergen, Christiania, Stockholm, Copenhagen, London, Leyden, Brussels, Paris, Lyons, Berlin, Hamburg, Dresden, Prague, Vienna, Graz, Monaco, Genoa and Naples.
After my return to Washington the Copenhagen Museum most kindly sent to me the large and important Ingolf collection; the Berlin Museum, through Pro-
4 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
fessors Déderlein and Vanhéffen, sent me the antarctic collections brought together by the Gauss; Prof. F. Doflein, through Prof. Déderlein, sent me his east Asiatic material, and, through the courtesy of Professors Kcehler, Max Weber and Vaney, the Siboga collection of unstalked crinoids was also assigned to me for study. More recently, thanks to the kindness of Prof. Bernard H. Woodward and Mr. Wilfrid B. Alexander, of the Western Australian Museum and Art Gallery, at Perth, I have been enabled to examine the crinoids collected off the coast of southwestern Australia by the Australian steamer Endeavour. ,
Thus in the preparation of this report I have met with the most cordia] coép- eration from all sides. Thanks to the great generosity of all my colleagues I have been enabled to assemble in one place and to compare directly one with another many thousands of specimens of recent crinoids, far more than ever were previously reviewed by any one individual, including examples of practically every known species and a large proportion of the existing types. This material has in many cases been ample for the determination of such questions as the scope of individual and of specific variation, and for the accurate delimitation of species, factors of the greatest importance in the study of all animal groups, but impossible satisfac- ° torily to determine except under the most favorable conditions.
While the present work is a complete monograph of the crinoids living at the present day, based upon the material preserved in practically all of the more impor- tant museums of the world, it is equally a catalogue of the crinoids of the United States National Museum, for my colleagues have been so kind as to permit me to retain duplicates from the collections under their care which I have examined, so that the collection of the United States National Museum now includes, in addition to the very rich material gathered by the vessels of the Bureau of Fisheries, particu- larly by the Albatross and Fish Hawk, and received from other governmental sources, a very large number of specimens, representing numerous species. received as donations from other similar institutions.
GENERAL METHOD OF TREATMENT.
The general method of treatment herein adopted differs in certain important respects from that employed by my distinguished predecessor and by all the other students of this group.
The study of the crinoids heretofore has invariably been approached from the paleontological viewpoint, the recent crinoids being considered as the impoverished and decadent remnants of a once numerous and powerful class, the last forlorn and pitiful exponents of a dwindling phylogenetic strain.
During the 1906 cruise of the Albatross I handled tens of thousands. of speci- mens; several times I saw the forward deck of the steamer literally buried under several tons of individuals belonging to a species exceeding any fossil form in size; everywhere we went we found crinoids; we dredged them at all depths. My ideas of the comparative importance of the recent forms underwent a total change; surely a group so abundant, even though very local and very unevenly distributed over the sea floor, can not be considered as decadent or degenerate. From my
MONOGRAPH OF THE EXISTING CRINOIDS. 5
observations at sea I became convinced that the recent crinoids are in every way as much of a factor in the present day marine biology, and play fully as important a part, as the echinoids, the holothurians, or the asteroids; cecologically they are more interesting than any of these because of their sessile mode of *life and curiously specialized method of procuring food.
I believe that the small importance hitherto attached to the crinoids as recent animals in comparison with the other echinoderms has arisen from three causes: (1) The extraordinary completeness of the palzontological record; this has its origin in the fact that the crinoids exceed almost all other animals in their adapt- ability to fossilization; their organization includes a very large percentage of lime and other inorganic materials, and there are no soft bodied forms among them. it is to be expected, then, that fossil erinoids will be exceedingly numerous, and wil include a far greater variety of diverse types than the fossil representatives of the other echinoderm groups, and therefore will appear greatly to have exceeded in the past in numbers, variety, and general importance the echinoids, asteroids, ophiu- roids or holothurians; while at the same time this splendid palzontological record will tend to blind one to the true importance of the recent representatives and to cause them to appear, in comparison with the recent representatives of the other classes, relatively insignificant; (2) the small number of species hitherto known; the majority of the specimens collected have slipped unheralded into museums; very few investigators have cared to cope with the many difficulties presented by their study, and so the proportionate number of known forms has been allowed to fall far behind those known in the other groups, not because they are really so very much fewer, but because of the much less general interest which they have excited; were the crinoids as enthusiastically studied as the echinoids, ophiuroids, asteroids or holothurians, we should have a wealth of records and of described forms comparing far more favorably with what we find on consulting the literature on those animals; (3) the paucity or absence of accessible species along the shores of the countries where the greatest interest in zoology is taken; one can not expect that a young investigator will devote himself with enthusiasm to the study of a group represented on his shores by one more or less rare or local species as in Europe, or by none at all which are accessible to him as in America, when the representatives of other groups are rich both in number and in species; were the shores of Europe or America as well stocked with littoral crinoids as are those of Borneo or Celebes, I have no doubt that our knowledge of the crinoids would be far in advance of what it is to-day; the semiprofessional zoologist as a rule pursues in foreign lands mainly animals in which he has become interested at home through the study of his own local fauna; animals of classes strange to him, especially if difficult to preserve, are of only incidental interest; therefore he.gen- erally, if he has a leaning toward marine zodlogy, gathers up corals, shells, urchins or starfish, together with the more tenacious ophiuroids, not attempting to save the more brittle species of the latter or the very brittle crinoids.
Firmly believing, therefore, that the recent crinoids are in no way less important than the recent representatives of the echinoids, asteroids, ophiuroids or holothu- rians, and in spite of their remarkably complete paleontological record, I have thought
6 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
it advisable to approach them in a somewhat different way from that which has usually been adopted, in order the more strongly to bring out many points which are obvious enough if the crinoids are considered as recent animals, but which are greatly obscured if one attempts to consider both the recent and the fossil forms together.
This somewhat radical treatment emphasizes some very interesting facts in a way not possible by any other method, and sheds an entirely new light upon many complex problems. Moreover, the results are strictly comparable with the results deduced from the data gathered from a study of other recent groups; a line of investigation may be followed up with the certainty that one is not liable to mistake a very highly specialized for a very primitive structure or type. Comparative anatomy may be employed as an aid in systematic work, so that conclusions do not have to be based upon the skeletal system alone; and, most of important of all, the crinoids in their relations to the other echinoderms and to other marine organisms stand forth in their true light, quite devoid of the false prestige which has hitherto been theirs as a natural result of their magnificent paleontological record, a record which is not surpassed by that of any other marine organisms, and is approached only by one or two restricted groups.
The strongest argument which can be made against this method of treatment is that questions of phylogeny are entirely divorced from any possible solution by the study of chronogenesis, but it seems to me that a phylogeny grafted upon a chronogeny.is a very unsatisfactory structure unless one is certain that the chrono- genesis represents, as of course it should, the true phylogenetic development.
When any group of a class of animals adopts a mode of life entirely different from that of all of the other members of the same class we must be prepared to encounter and to discount extraordinary, sudden, and unexpected changes in the organization which are not connected with the ancestral type of organization by any intermediate stages. Among such animals we almost always find the group char- acters developed in a most erratic manner. Some structures will be very highly specialized, sometime specialized far beyond what is seen in any other member of the class, while others will be in a very rudimentary or primitive state of develop- ment, or perhaps even absent altogether.
The echinoderms differ very abruptly from the crustacean line of descent from which they took their origin, and similarly each of the echinoderm groups differs abruptly from each of the others.
We see in the echinoderms to-day most perplexing combinations of primitive and highly specialized characters associated in all sorts of ways, and this leads naturally to the assumption that there was no definite intergrading form between the echinoderms and the barnacles, which, of all the crustacea, approach them most closely, but that the former sprang from the phylogenetic line, which may by easy stages be traced to the latter, by a broad saltation in which the assumption of the free habit (subsequently modified in the Pelmatozoa) and the correlated assumption of pentamerous symmetry combined to make the existence of intergrading forms impossible, while at the same time it resulted in the formation at the very moment of their origin of two diverse stocks, the heteroradiate (including the Pelmatozoa,
MONOGRAPH OF THE EXISTING CRINOIDS. 7
the Echinoidea, and the Holothuroidea) and the astroradiate (including the Asteroidea and the Ophiuroidea) between which there are, and can be, no interme- diates.
Thus it is evident that we must use the very greatest care in the correlation of the chronogeny and the phylogeny of the echinoderms, and we must be continually on the watch for sudden and aberrant deviations and specializations in the older as well as in the more recent types. A detailed study of the living types will furnish the key to many such deviations, and this subsequently will enable us correctly to interpret the complicated morphology of the extinct species.
As nearly as I can see there is comparatively little of value to be learned in the first instance from the paleontological record of the echinoderms, at least in so far as their comparative morphology and phylogeny is concerned, which can not be learned just as well, or even better, from a study of the recent forms alone, though the fossils furnish invaluable confirmatory evidence of the truth of any conclusions which we may reach.
If we acquire our facts from a study of the comparative anatomy, morphology and development of the recent types and then test them by reference to the extinct series, it seems to me that we can build up eventually a logical phylogenetic sequence of types of progressively increasing specialization and perfection which will be able to withstand all the attacks which may be made upon it.
Of the many and varied recent forms there is abundant material, and this material is always susceptible of detailed study. Furthermore, all of the recent types are interconnected by readily demonstrable phylogenetic lines with all the others.
On the other hand, among the fossils really good and satisfactory specimens are rare, and there are many interesting forms which we are not able, on paleontological evidence alone, to connect in a truly satisfactory manner with related types.
In treating of the interrelationships of the various echinoderm groups it will be noticed that I have not taken the larvee into consideration. The larvee of the echinoderms are very highly specialized creatures, specialized for a mode of life entirely different from that of the adults, and hence specialized in an entirely different way. To all intents and purposes they are organisms of a different class entirely. Moreover, they are not all specialized in the same direction, and hence are not strictly comparable among themselves. Mechanical considerations of form make comparison between the barrel-shaped larva of Antedon, the bipinnaria of Asterias, the auricularia of Holothuria, and the plutei of Ophiura or of Echinus hazardous and unsatisfactory.
A true comparison between the species of the several echinoderm groups is only possible upon the attainment of the adult form, or at the earliest at the inception of the pentamerous symmetry. However suggestive and instructive the larvee may be, they must be treated quite separately from the adults, as a distinct class of ani- mals, or trouble is sure to result.
In this respect I consider the echinoderms as a whole precisely comparable to those insects and crustaceans which undergo a complete metamorphosis, though in
8 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
the echinoderms the ease is much more complicated than in the insects and crusta- ceans on account of the difference in symmetry between the young and the adults.
Sir Wyville Thomson long ago recognized this fact, that in tracing out the life history of the echinoderms we are apparently dealing with two distinct organisms, each apparently presenting all the essentials of a perfect animal, as had W.9K. Carpenter before him, but succeeding authors have shown a tendency to disregard their warnings.
On account of the curiously aberrant and sudden differentiation of the echino- derms as a whole, and similarly of each of the constituent classes of the group, we can never hope to ascertain the true interrelationships either of the echinoderms and other animals, or of the several constituent classes of the echinoderms, by any direct method of comparison.
The ancestral characters have become so modified by the adoption of radial symmetry, and the bilateral young have become so specialized, that any direct com- parison which is at all conclusive has now become impossible.
We must therefore approach the problem by an indirect method, by the adop- tion of hypotheses which will logically explain all the facts presented and will cover all the data which we are able to accumulate, but which are not primarily the direct and indisputable resultants attained by the correlation of these facts and data.
SYNONYMY.
The synonymy of the recent crinoids was in a decidedly tangled condition, having been only partially elucidated by Carpenter, as he did not discuss in detail any but the species collected by the Challenger. I therefore found it necessary to enter into this phase of the subject somewhat deeply, especially in view of the fact that the group contained a disproportionately large number of floating names— nomina nuda and unidentifiable supposed species—which it was very desirable to allocate if possible. I have attempted to bring together all the references to each species that I could find, in the hope that future workers will be spared the formid- able task of having again to review the enormous mass of literature. The synony- mies given are, I believe, reasonably complete, though numerous notices of species not here included will doubtless come to light in the future. The citations have, with very few exceptions, been personally verified, and may be taken as representing the works consulted in the preparation of this monograph.
SYSTEMATIC TREATMENT.
In the ease of the comatulids it has been found necessary to multiply by about a dozen times the number of genera previously allowed, and to create numerous new families and higher groups. This was the unavoidable result of the discovery of a vast number of new species, throwing a radically different light upon the inter- relationships of the various forms.
The different species of comatulids vary very greatly in the number and obvious- ness of the characters by which they are separable from closely related species; two species, perfectly distinct, may be separable only by a small minority of what are
MONOGRAPH OF THE EXISTING CRINOIDS. 9
commonly considered their specific characters, while two others may have only a small minority in common; and, as in other animals, characters perfectly reliable in one group are more or less unreliable, or even perfectly worthless, in another. Species may be found of all grades of differentiation, from a very small minority of their characters to complete separation, but usually they fall into two classes: (1) those separable from related species by a minority of their characters, the remainder being held in common, and (2) those separable in all their characters. The first division is in reality, of course, arbitrary, for it is undoubtedly true that any two species will be found to be always separable in all their characters, provided we devote a sufficient amount of study to them; it might better be worded ‘‘those separable from related species by a majority of the characters commonly employed in specific diagnosis.”
It is usually found that a number of species differentiated according to the first rule form a circumscribed unit the sum of the diversity of all the characters in which does not overlap the sum of the diversity of all the characters in any other similar unit, the assemblage of forms differentiated under the first rule thus coming as a whole under the second rule. These sharply circumscribed units, as well as species falling within the limits of the second rule, I have considered as representing valid genera, while forms not separated from related forms by the sum of all their characters I have regarded as species. All species agreeing in the majority of their characters as employed in systematic diagnoses I have considered as congeneric.
Now a number of species may, according to this ruling, be strictly congeneric, yet they may be united into several groups by a sharply defined single character which is common to, and exactly similar in, several species, and is not found outside of those species. These groups within the genus I have considered worthy of sub- generic rank. Similarly, subgenera may be differentiated into distinct specific groups, though usually this differentiation is, as would be expected, less apparent. In the separation of the families and of the subfamilies as well as of the higher units the same idea has been followed, but characters of a more fundamental nature, and therefore not sufficiently plastic to be of service in the differentiation of genera and species, have been employed.
As in all other groups of animals the various crinoid species are of very differ- ent relative value. In some (mostly the more highly multibrachiate oligophreate) genera if any one character whereby the species are commonly differentiated be plotted on a species curve, the several species will be found to be indicated not by a series of separate triangles, but by a succession of more or less marked nodes which are united to the mass forming the adjacent nodes by coalesced bases in thickness equal to from 10 to 60 per cent or more of the maximum height of the neighboring nodes. Such variability and lack of absolute fixity in any one character is as a rule reflected in all the characters, and thus there results a species group or genus which may be compared to a small mountain system rising out of a plain, each peak of which represents the separate species.
In such a genus every systematic character varies between two extremes, but there is often no correlation whatever between the different characters. Thus
79146°—Bull. 82—15_—2
10 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
every sort of combination is possible, and a very large variety is found, though the tendency is for the characters to form more or less definitely correlated groupings, and to crystallize into certain definite types.
In other genera (mainly macrophreate) all the characters are more definitely correlated with each other, and then the nodes on the species curve will be found to be very sharp and almost or entirely distinct from each other, the various species indicated exhibiting little or no tendency toward intergradation.
This type of variability is not connected with the geographical origin of the specimens except in a very general way, and therefore the several forms can not be considered as subspecies as that term is commonly understood. It is practically confined to the multibrachiate Oligophreata, and to specimens of oligophreate species from the East Indian region. These same species when extending their range outside of this region gradually become more fixed and definite in their characters, so that individuals from, for example, Madagascar or southern Japan will all be found to be practically uniform in their various features, and to represent the mean of the two extremes seen in a series from the central East Indian region.
The recent representatives of a few families appear to have suddenly deviated from any type which we might reconstruct as the phylogenetic stock whence they had been derived by a process of ‘‘explosion”’ of their characters which have become recombined in a curiously unbalanced manner, exactly as we see to be the case in several fossil groups. A tendency to form an explosive or very aberrant offshoot is more or less evident in every group of animals, but it rarely affects more than a small minority of the genera or of the species.
An earnest effort has been made to avoid the common error of taking into account only obvious differential characters, thereby becoming blinded to the less obvious, but often more reliable, systematic features, by carefully examining every detail of the animal and every point offered by its structure apart from all the others, though in many cases, so far as regards comparative descriptive work, no use has subsequently been found for the data acquired.
Great care has been used in the selection of new generic names, and especially in the selection of the types of new genera; the types are, whenever possible, the first species to have been described, and the commonest species; but in cases where the original description is deficient or the identification doubtful I have taken one of the later species, where circumstances permitted one considered as a synonym of the first. Preference has always been given to species at hand to guard against the possibility of nomenclatorial disturbance through misconception of species not personally known to me.
EMBRYOLOGY, DEVELOPMENT AND ANATOMY.
The systematic study of the comatulids is, no less than that of other groups, based largely upon a knowledge of the development and of the external and internal anatomy; the comatulids, through uniformity of habit, are all built upon the same general plan, and hence the knowledge of their development and anatomy must be comparatively exhaustive in order that the systematic differentiation, at first sight apparently very slight, may properly be appreciated, when it becomes obvious
MONOGRAPH OF THE EXISTING CRINOIDS. 11
that the differences, trifling though they may seem, are really fundamental and valid.
Students of bilaterally symmetrical animals, especially those animals endowed with powers of locomotion, are accustomed to a relatively large coéfficient of specific differentiation; this is true even among other groups of echinoderms in which the individuals lead a more or less bilaterally active life. Also among radially sym- metrical animals which move actively about specific differentiation is usually more marked than among those of sedentary habits.
The difficulty of at first comprehending the comatulid characters is a difficulty of comparative perception, not of fact, and is entirely due to a superficial similarity in the gross anatomy and form.
One can never tell without a most detailed inquiry what are good systematic characters and what are not; the most obscure anatomical features often prove to be of the greatest interest, while in the embryology even such points as the unequal division of the ovum, as well as the absence in certain cases of the anterior tuft of cilia, and the difference in size of the cells at the animal and vegetative poles of the blastosphere, appear to be of specific significance.
It is very important that systematists should consider all these points of apparent difference, especially those which loom up large in the embryo but which disappear more or less in the adults; it is also important that embryologists and anatomists, aroused to a high pitch of enthusiasm over the discovery of certain peculiarities in their material not previously noticed, should not be led either into condemning the work of their predecessors as careless, or into arguing, from a wide anatomical difference between two forms, a correspondingly wide systematic difference.
It is a common fault in works of monographic scope to magnify the systematic side of the subject to the great detriment of the morphological; but a thorough understanding of the anatomy and development of the animals of any group is absolutely essential before the systematic aspect can be intelligently studied. Diverse interpretations of different structures or organs by several authors have often led to corresponding variations in their systematic treatment, variations which have been difficult to appreciate in their true proportions, because of neglect to explain in advance the position taken.
As a general rule systematists are inclined to attach altogether too little impor- tance to anatomical or embryological features, and morphologists altogether too much. For instance, P. H. Carpenter, as a systematist, passed lightly over the peculiarities of the brachial muscles in different forms, while as a morphologist he greatly exaggerated the importance of interradials in the genus Thawmatocrinus.
I have been able to add but little to what has been done by previous workers in the field of development and anatomy; but it is essential that these be explained in some detail before the systematic treatment can be commenced. Instead of giving an account of these phases of the subject taken from a comparative study of the works of others, I have preferred to quote more or less directly from the leading authors on the various points considered, giving full credit to them, and thus mak- ing a far more satisfactory whole, No attempt is herein made to give an exhaustive
12 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
account of the anatomy and embryology of the crinoids, but it is hoped that these points are treated in sufficient detail so that their systematic significance may be appreciated.
The account of the embryology and of the anatomy of the various forms is taken from the latest and most authoritative works, and will be found to be sufficient for the systematic side of the subject; but it must be emphasized that the extracts given are not intended to be, and are not, sufficient to serve as a basis for anatomical or embryological work, and for such work the original papers, which contain much more than the extracts included here, must be used, as especial care has been taken in order that the information herein given shall not detract in any way from the demand for the original papers by providing an easily accessible transcript of the subject matter or of the figures.
Methods of microscopical technique are obviously out of place in a systematic work devoted to animals of comparatively large size, and, therefore, are in all cases omitted; they are, however, given in great detail by the authors cited.
Accounts of obscure anatomical or histological points, or discussions with no systematic bearing, are omitted; this includes the discussion of doubtful structures; information in regard to these may be found in abundance in the original papers. The information here given is, it may be again stated, included for systematic work, and from a systematic point of view, only.
A large amount of exceedingly interesting work has been done on the larval and anatomical homologies of the various echinoderm groups, resulting in a con- siderable diversity of opinion in regard to their interrelationships, and in much speculation as to their common origin and to the original echinodermal prototype. While it is difficult to avoid discussing these matters afresh, it has seemed best to omit reference to them in a work devoted solely to the crinoids, and to only a lim- ited group of the species of that class. The only question that can be of any impor- tance is that of the relationship between the crinoids and the most closely allied classes, and this will be considered at some length.
VARIANTS AND ABERRANTS.
It has long been recognized that a careful study of variants and aberrants often furnishes most instructive data upon which to base a consideration of the origin and phylogenetic significance of the different organs and members, and of an animal as a whole. Inno group is the study of these variants more important than in the echino- derms, and in few groups do they occur with such frequency and along such well- marked lines of progression and retrogression as in the crinoids.
PHILOSOPHICAL CONCLUSIONS,
Many general zodélogical truths are brought out by a study of the crinoids more forcibly than by a study of any other animals, and many others appear in the group in a somewhat new aspect, which sheds a certain very instructive new light upon them.
For instance, in certain genera most of the species will occupy definite and closely circumscribed areas or depths, each different from that inhabited by any of the
MONOGRAPH OF THE EXISTING CRINOIDS. 13
others, the interrelationships being in general accordance with Jordan’s law; but one species, always the most variable and the one occupying the position nearest the center or general mean of the extremes of all the variable specific characters repre- sented in the genus, will be found whose range, both geographical and bathymetrical, is equal to the sum of the ranges of all the other species in the genus.
Again, highly specialized species commonly occupy a specialized and circum- scribed habitat, while generalized species are found among very diverse conditions.
Among the several species in a genus the one occupying the limits of the distri- bution of the genus as a whole is as a rule the most variable in its characters, and similarly in individual species the coéfficient of variation among the individuals increases in proportion to the distance from the center of distribution, primarily as a result of existence under progressively increasing unfavorable or semipathological conditions.
There is a more or less apparent curious and significant exception to this rule, however, for the center of distribution of a large group—and the truth of the obser- vation is, as a rule, greatly increased in proportion to the size and importance of the group—is marked by a most remarkable diversity in the individual, specific, and generic characters of the organisms inhabiting the locality. This is the result of an increase in the number of variants under optimum conditions—a kind of incipient species formation—and has no relation to the more or less pathological type of variation seen along the outer edge of the habitat of a species or of a genus.
Association of species of a single genus or of related genera in pairs, each occu- pying nearly or quite the same geographical and bathymetrical ranges, has fre- quently been reported, cases occurring in most of the animal groups, and instances of it appear among the crinoids. Some of these cases are at once explained by the difference in the breeding seasons of the associated forms which effectually prevents any hybridization; but others are not quite so simple, although they may be accounted for in various other ways.
Not only are the crinoids plant-like in appearance and in the manner of their existence, but some of them have, along with this curious superficial similarity, acquired a more or less close correspondence in the comparative interrelationships of their various systematic characters, just as have many of the arborescent marine organisms.
The degree of stability of the generic and specific characters and of the corre- lation of the characters presented by the several sets of structures and organs among the comatulids is, broadly speaking, inversely proportionate to the fixity of habit of the adults, and therefore in general to the number of arms possessed by the adults. In such groups as the Antedoninz, where the animals are more or less active and are capable of swimming about, the generic and specific characters and the character correlations are, as a rule, strongly marked and readily defined. Such specific or generic intergradation as occurs (and specific and generic intergradation is by no means uncommon) takes the form of a gradual and uniform change in all the char- acters whereby exactly the same balance of correlation is at all times maintained; but in the highly multibrachiate groups in which the musculature in the proximal portion of the arm is greatly reduced, especially in those groups which are highly
14 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
specialized and possess division series of 4(3+4) instead of the more primitive 2, character correlations are unstable and uncertain and are liable to sudden and ex- iraordinary deviations from the normal, resulting in all sorts of grotesque mixtures, not only within a family or genus but even within a group of specimens of the same species from the same locality.
Thus, among the highly multibrachiate comasterids individual specimens of a single species may exhibit, more or less well developed, the essential features of entirely different genera from the one to which they really belong. For instance, examples of Capillaster multiradiata are not uncommon with nearly or quite half of their arms of the type characteristic of the species of Comatella, while examples of Comanthus bennetti are recorded which possess the arm structure of the species of Comanthina and others which possess that of the species of Comantheria; con- versely, specimens of Comanthina schlegelii not infrequently exhibit the arm structure characteristic of Comanthus bennetti.
This shows the necessity for the utmost caution in determining the genus or species of specimens of the highly multibrachiate forms (especially when some or all of the division series are 4[3 +4]), and of specimens of 10-armed forms belonging to highly multibrachiate groups. Each individual must be critically examined not only in respect to the essential features of the group as commonly understood, but also in regard to all of the minor features, for it is sometimes found that the character upon which most stress is ordinarily (and properly) laid is in part or even in its entirety replaced by the character normally diagnostic of an entirely different species or even genus.
The recurrence of nearly or quite identical types of arms, centrodorsals, cirri, pinnules, disks, and other organs in widely different groups raises the question whether such recurrence is really the sporadic reappearance of fixed and definite structural types or whether it may not be merely the result of parallelism.
Now parallelism is the convergence toward a common type of fundamentally different structures or organs. This convergence progresses far enough to satisfy the requirements of the impelling physical, chemical, mechanical, or economic factors, but no further; hence, though two radically different structures or organs may through parallelism be rendered superficially very similar, the modification is never carried far enough entirely to conceal their ultimate diverse origins.
In the comatulids identical types of organs and identical structural types, which, as in the case of the method of arm division, are sometimes quite complex, reappear in widely different groups, in each of which they pass through the same developmental history, but in each of which they are associated with other organs and structures of phylogenetically and developmentally entirely and fundamentally different values which are combined in each case in a radically different way. Such could scarcely be the case were we dealing with structural modifications resulting purely from mechanical, economic, or other exigencies, for we can scarcely imagine parallelisms either to be so erratic in their manifestations and to be in one structure or organ so entirely dissociated from correlated effects upon other structures or organs, or to show, no matter where they appear, the same course of development.
MONOGRAPH OF THE EXISTING CRINOIDS. 15
The general absence of correlation between the several sets of organs and struc- tures which collectively make up the comatulid whole most emphatically negatives the idea that the occurrence of the same structural type in widely different groups is the result of parallelism, and indicates that among the comatulids as a class there is a given number of more or less distinct and independent types of each organ and structure, any one of which may be combined with any one of the types of the other organs and structures.
RELATIVE STATUS OF THE RECENT CRINOIDS.
Since the crinoids are the most nearly strictly sessile of all the animals in the sea, and since their relation to their food supply is always essentially the same no matter what diversity there may be in the chemical and physical nature of their surroundings, the facts furnished by a study of the geographical and bathymetrical distribution of the recent crinoids are of the greatest value in the determination of former land connections, just as the facts brought out by a study of the fossil repre- sentatives of the recent genera and species are of the greatest importance in tracing out the extent and time of existence of the ancient seas.
The remarkable paleontological record of the crinoids, and the abundance of fossil forms closely related to existing genera and species, will allow of an accurate estimate in regard to the geological time when these land connections were estab- lished, and when they became disrupted.
A comparative study of the recent faunas and those of past ages will show at what epoch certain land areas and certain deep channels were formed, as a result of which genera of subsequent origin were unable to spread into territory previously colonized by older forms; while at the same time it will throw much light on the geological age of the components of the deep sea fauna, showing that it is a complex formed of representatives of all the most virile types which have existed in all of the past horizons.
By a careful study of the chemical and physical conditions under which the recent forms live, a determination of their relation to temperature, salinity, light, currents, etc., we shall be able to learn much which will be of the greatest value in ascertaining the exact conditions under which many ancient strata were laid down.
I have reserved the discussion of all these interesting points, as well as of the distribution, ecology, geological history, and the relationships of the recent crinoids to their fossil representatives (including the facts brought out by a comparative study of recent and fossil species belonging to the same genera)—in other words, the bearing of our knowledge of recent crinoids upon the data gathered from a study of paleontology—until the end of this work, not only because the general conclusions find their most logical place after the consideration of all the special features and the complete presentation of all the data, but also for the reason that, as the treatment herein adopted is such a radical departure from any treatment heretofore proposed, and the number of new species is so very large, no general discussion would be of value until after the systematic framework upon which it is of necessity based has been thoroughly elucidated and made easy of comprehension.
16 BULLETIN 52, UNITED STATES NATIONAL MUSEUM.
In the following pages there will be found much speculation in regard to the hypothetical ancestor of the crinoids and of the echinoderms, based upon a study of each of the various systems which, when taken together, make up the ermoid or echinoderm whole, and a figure of the hypothetical ancestor will be found embodying all the data acquired from this study. It is well, perhaps, to emphasize the fact that no claim is made that such a creature ever existed; we see in all the echinoderms to-day most perplexing combinations of primitive and highly special- ized characters, associated in all sorts of different ways, and this leads us naturally, as I have already stated, to the assumption that there was no definite intergrade between the echinoderms and the barnacles, but that the former sprang from the latter (or, more strictly speaking, from the same phylogenetic line which can be traced by easy stages to the latter) by a broad saltation in which the assumption of the free habit and the correlated assumption of the pentaradiate symmetry combined to render the existence of intermediate types impossible, while at the same time it caused the formation by the echinoderms, at the very moment of their origin, of two widely diverse stocks, the heteroradiate, including the Pelmatozoa, the Echinoidea, and the Holothuroidea, and the astroradiate, including the Asteroidea and the Ophiuroidea, between which there are, and can be, no intergrades.
The comatulids must therefore be considered as a biologically extremely com- plex and mixed group in which each organ and structure occurs in a single series all the way from a primitive to a highly specialized type, but in which the various degrees of specialization of each organ or structure, in other words, the progressive steps in the series, as not in any way correlated with species or with genera, or with the comparable degrees of specialization of any other organ or structure.
Thus it is at once evident that there is a most extraordinary uniformity throughout all the comatulid families and genera, and that each is potentially on essentially the same phylogenetic plane as are all of the others.
The comatulids as a group are exactly parallel and comparable to the penta- crinites as a group; they are descended from the same ancestral stock and represent exactly the same phylogenetic stage, but durmg their development they have diverged from their phylogenetic mean in exactly the opposite direction. The pentacrinites have departed widely from their prototypes by enormously increasing the length of the column and at the same time indefinitely reduplicating the cirrifer- ous proximale, a departure which has to a considerable degree lessened the mobility of the crown, this being in part compensated by a corresponding increase in the length of the arms; while the comatulids have departed just as widely by compressing what is virtually the entire column of the pentacrinites within the compass of the single proximale or nodal from which numerous cirri are extruded, fixation by these cirri reducing the possibility of motion by the crown to a minimum so that under ordinary conditions the animals are almost as firmly attached as is Holopus.
As the greater part of the enormously elongated stem of the pentacrinites lies on the sea floor and therefore becomes neutral in its relation to the mechanics of the animals, these forms do not exhibit any very radical departure from a more gener- alized type, such differences as they show being chiefly the result of the very large size of the crown and arms correlated with a reduction in size of the calyx; nor do
MONOGRAPH OF THE EXISTING CRINOIDS. 17
they exhibit any strong tendency toward dissociation of ordinarily correlated char- acters; but the sudden and much more abrupt departure from the normal crinoid habit seen in the comatulids has been accompanied by, or the entirely new conditions under which they live and the consequent extraordinary atrophy of their calyx have induced, the development of all sorts of structural variants and excesses which have not yet had time or, because of the passive part the animals play in their relations to other animals, have not yet been forced, to crystallize to definite types with a definite scheme of correlation.
The morphological difference between the pentacrinites and the comatulids is merely that the weakening of the syzygial union between the first nodal formed and the infranodal just below it in the comatulids leads to its rupture before any additional segments are formed, while in the pentacrinites rupture does not occur until many other columnars have been intercalated between this nodal and the calyx. The pentacrinites thus continue to build a long, many-jointed stem, while the comatulids condense the entire stem within the compass of the first-formed nodal. The morphological difference between the comatulids and the pentacrinites reduced to its lowest terms therefore is merely a slight difference in the develop- ment of the tendency to rupture at the syzygy between the first-formed nodal and the columnar just beneath it.
The comatulids and the pentacrinites occupy a curiously anomalous system- atic position, for both groups are far removed from the direct line representing the progressive phylogenetical development of the class. But both, though widely divergent, agree in differing from all other related types through discarding the proxi- mal portion of the column and in the development of a highly cirriferous proximale, which in the pentacrinites is indefinitely reduplicated.
The genus Thiolliericrinus occupies a position midway between them; species of this genus develop a cirriferous proximale, but retain the larval column; the relation of Thiolliericrinus to the pentacrinites and to the comatulids may roughly be graphically expressed by the followmg formula:
pentacrinites + comatulids 9
= Thiolliericrinus.
Thiolliericrinus, however, is in the direct line representing the progressive phylogenetical development of the class, and approximates very closely, if it does not actually represent, the type from which, by sudden diametrically opposite deviation, both the pentacrinites and the comatulids have been derived.
Systematically the pentacrinites, Thiolliericrinus and the comatulids repre- sent a small group of which Thiolliericrinus is the true phylogenetical exponent, the other two types being aberrant departures from this stock.
Thiollericrinus is fossil only. In the recent seas the comatulids far outnumber all of the other crinoids taken together, at the same time extending through a much wider geographical, bathymetrical and thermal range, while by far the largest of the remaining groups is that of the pentacrinites.
These two highly aberrant types therefore dominate the recent seas, and so pronounced is their dominance that when compared with them all the other types
become relatively insignificant.
18 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
The comatulids alone in their numbers, in the diversity of their habitat, and in the complexity of their systematic interrelationships are in the present seas the strict systematic equivalent of each of the other classes of echinoderms. Of themselves they form what is unmistakably a class, with all the distinctive systematic features of a true class.
Thus the comatulids, in reality only an insignificant and aberrant offshoot from the general phylogenetic crinoidal line, represent in their relationships to the other organisms of the seas of the present day a true class, exhibiting the curious anomaly of a group which, considered from one point of view is a true class, but considered from another point of view does not even rise to the dignity of a subfamily.
ILLUSTRATIONS.
A very considerable amount of time and thought has been expended in attempt-. ing to solve the problem of how best to illustrate the various species of comatulids. These animals differ but slightly in their general build, though very greatly in the finer details of their structure.
In the Challenger monograph the first serious attempt was made to portray the comatulids in a monographic way. Although the figures are exceptionally good, there has always been more or less difficulty in comprehending them, and I experienced a great deal of trouble with them myself. It was not at first evident wherein this difficulty lay. A certain inability of the artist to grasp the significance of such details as the smooth or comparatively rugose appearance of certain speci- mens, details exceptionally difficult of portrayal in a satisfactory manner, account for much of the indefiniteness of certain figures, while the varied position of the arms in the examples given make comparisons between the illustrations exceedingly laborious, and undoubtedly accounts for the rest.
The figures in Hartlaub’s works were drawn by a different artist than were those in the Challenger report; though excellent delineations, a certain personal element has entered into their make-up which makes comparison between them and the Challenger figures more or less unsatisfactory.
No personal element entered into Déderlein’s beautiful photographic repro- ductions; yet they are as difficult to compare with the figures of Carpenter or of Hartlaub as these are with each other. It was therefore evident that I could not hope to produce satisfactory results by placing sole reliance either upon the artist or upon the camera.
A study of Déderlein’s paper side by side with the Challenger report suggested to me that if each species were drawn in detail, and a photographic reproduction of the specimen also given, the former to show the intricate structure and the latter to give the general appearance, a result might be attained which would stand a good chance of being fairly satisfactory.
After a mature consideration of the matter I decided that, as photographie plates were also to be used, there was no object in burdening the text figures with detail; the simpler they were the more forcibly could the essential differential characters be made to stand out. Moreover, if all the figures were rendered semi-
MONOGRAPH OF THE EXISTING CRINOIDS. 19
diagrammatic by the arbitrary arrangement of the arms in a given position, com- parison of the figures inter se would be greatly facilitated; it would not then be necessary to use the imagination in righting a more or less distorted picture before comparison could be made. with another equally, but differently, distorted.
All the figures included herein have been prepared in line with these ideas, and future workers will be able to determine whether or not they are of any value.
While the portrayal of 5 or 10 armed species which normally carry their arms at more or less of an angle to the surface of the disk is a comparatively simple matter, the question of how to show a multibrachiate or a flattened species without becoming swamped in a multiplicity of detail opened up an additional series of problems. It has seemed to me ample in the case of the flat 10-armed comasterids to show one-fifth of the animal (two arms) in detail, including the centrodorsal and such cirri as may be present on the side opposite the arms as drawn, and to indicate the remaining portions by simple lines; in the case of very many armed forms the sketching in of the arms in the additional four sectors has the effect of diminishing the strength of the detailed sector, as well as by increasing the width of the figure, necessitating a somewhat greater reduction in size than is advisable. Only the centrai portion and one of the so-called “rays” of the multibrachiate species are therefore shown.
In the preparation of the text figures, I was fortunate in securing the codpera- tion of Miss Violet Dandridge, of Shepherdstown, West Virginia, whose experience in preparing figures abounding in detail, especially of shells, fish, and ophiuroids, formed the best possible basis for work upon the crinoids.
The photographs for the plates were made by Mr. T. W. Smillie in the photo- graphic department of the United States National Museum.
IDENTIFICATION OF THE SPECIMENS UPON WHICH THIS WORK IS BASED.
Almost all the specimens which have been examined by the author in the preparation of this report have been marked with a small label stating the fact, and all are herein listed under their respective species, so that any future worker may be able to consult, with the least possible trouble, the material upon which all the statements and deductions herein given have been founded.
The letters following the data for each specimen indicate the collection in which the specimens may be found, as follows:
Amer. M.: American Museum of Natural History, New York. Austr. M.: Australian Museum, Sydney, New South Wales.
B. M.: British Museum.
Berg. M.: Bergen Museum.
Berl. M.: Museum fiir Naturkunde, Berlin.
Bata: Museum of the Boston Society of Natural History. CoM: Zoological Museum, Copenhagen, Denmark.
eM: Dresden Museum.
1 Dare Museum of the Essex Institute, Salem, Massachusetts.
Te: oe: Frank Springer collection.
20 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
F.S. Dep.: Frank Springer deposit collection in the United States Na- tional Museum.
G.M.: Graz University Museum.
I. M.: Indian Museum, Caleutta, India.
K.M.: Christiania Museum.
Mie: Leyden Museum.
soe: Leland Stanford Junior University Museum.
Uys University of Liverpool Museum.
M. C. Z.: Museum of Comparative Zodlogy, Cambridge, Massachusetts.
M. M.: Museum fur Meereskunde, Berlin.
M. O.: Oceanographic Museum, Monaco.
PAY: Museum of the Philadelphia Academy of Natural Sciences.
PeMes Paris Museum.
Be Dades Collection of Prof. Robert Tracy Jackson, of Cambridge, Mas- sachusetts.
Beds Collection of the Naples Station.
WiCks University of California collection.
U.S. B. F., W.: Collection of the United States Bureau of Fisheries, at Woods Hole, Massachusetts.
U.S. B. F., B.: Collection of the United States Bureau of Fisheries, at Beau- fort, South Carolina.
We SoM: United States National Museum collection.
Vis Collection of Prof. Addison E. Verrill, of New Haven, Con- necticut.
W.A.M.: Collection of the Western Australian Museum and Art Gal- lery, at Perth.
Waa Vienna Museum.
Y.M.: Peabody Museum, Yale University, New Haven, Connecticut.
INDIVIDUALS AND INSTITUTIONS TO WHICH THE AUTHOR IS INDEBTED.
There only remains the pleasant duty of expressing my gratitude and offering my most sincere thanks to those who have so kindly aided me in my work with information and with specimens: Mr. Alexander Agassiz, of Cambridge and New- port; Dr. Nelson Annandale, of the Indian Museum, Calcutta; Prof. A. Appellof, of Upsala, Sweden; Dr. F. A. Bather, of the British Museum (Natural History) ; Prof. F. Jeffrey Bell, of the same institution; Mr. Herbert Clifton Chadwick, of Port Erin, Isle of Man; Dr. Hubert Lyman Clark, of the Museum of Comparative Zoélogy, Cambridge, Massachusetts; Prof. Ludwig Déderlein, of Strassburg, Germany; Prof. Franz Doflein, of Munich, Bavaria; Dr. Robert Etheridge, of the Australian Museum, Sydney, New South Wales; Mr. George T. Farran, of Dublin, Ireland; Prof. Walter K. Fisher, of Stanford University, California; Prof. Theodore N. Gill, of Washington; Dr. James A. Grieg, of Bergen, Norway; Prof. Robert Tracy Jack- son, of Cambridge, Massachusetts; Dr. Robert Hartmeyer, of the Museum fir Naturkunde, Berlin; Mr. Samuel Henshaw, of the Museum of Comparative Zoélogy; Prof. W. A. Herdman, of Liverpool, England; Dr. R. Horst, of Leyden, Holland;
MONOGRAPH OF THE EXISTING CRINOIDS. 21
“
Dr. F. A. Jentink, of Leyden; Prof. L. Joubin, of the Natural History Museum, Paris; Prof. René Keehler, of Lyon; Prof. K. Kraepelin, of the Natural History Museum, Hamburg; Prof. Edward L. Mark, of Harvard University; Prof. W. Michaelsen, of the Natural History Museum, Hamburg; Dr. Theodor Mortensen, of the Zodélogical Museum, Copenhagen; Prof. Ed. Perrier, of the Natural History Museum, Paris; Prof. G. Pfeffer, of the Natural History Museum, Hamburg; Mr. Richard Rathbun and Miss Mary J. Rathbun, of Washington; Prof. J. Richard, of Monaco; Prof. William E. Ritter, of the University of California; Dr. Leonhard Stejneger, of Washington; Dr. Charles Wardell Stiles, of Washington; Prof. C. Vaney, of Lyon; Prof. Th. Studer, of Berne, Switzerland; Prof. E. Vanhéffen, of the Museum fir Naturkunde, Berlin; Prof. Addison E. Verrill, of Yale University, New Haven, Connecticut; Prof. Max Weber, of Eerbeek, Holland; Prof. W. Weltner, of the Museum fiir Naturkunde, Berlin; and Prof. Bernard H. Woodward, of Perth, Western Australia.
To Mr. Frank Springer, of Las Vegas, New Mexico, with whom I have been in constant communication since the beginning of the work, and who has assisted me in every possible way, with most valuable information and with specimens, I owe more than I can well express; it is due to his constant encouragement and support that I was at last able to bring my studies to a conclusion.
For their kindness and courtesy in reading the proof of this volume I am deeply indebted to Messrs. Frank Springer, William Patten, and Walter K. Fisher. All three of these gentlemen made numerous suggestions which proved most helpful tome. Itis only fair to them to state, however, that they are not necessarily to be considered as agreeing with all the details of my conclusions.
HISTORY OF THE SUBJECT. GENERAL HISTORY.
The common comatulids of the coasts of Europe (Antedon petasus, A. bifida, A. mediterranea, and A. adriatica) were undoubtedly known, at least to fishermen, long before any record of them appears in literature; so also it is probable that numerous specimens of the large species from the Orient had reached Europe and found their way into the cabinets of collectors soon after the establishment of regular trade between Europe and the East, though they had not aroused sufficient interest to lead to a definite announcement of the fact.
It is in 1592 that we find the first satisfactory reference to a comatulid; its great beauty and delicacy of structure, enhanced, no doubt, by its comparative rarity, led Fabius Columna to treat at some length of the common Mediterranean species (dexadaovaktevoscdijc; Antedon mediterranea), and he even noticed the interesting physiological fact that if a specimen be placed in fresh water its color- ing matter dissolves out, imparting a hue to the water corresponding to the original color of the individual. The remarks of Columna aroused considerable interest, and we find them incorporated, together with a copy of his really excellent figure, in many of the succeeding works on zodlogy.
Fossil erinoids, abundant in many localities, were widely known, and many and curious were the speculations as to their origin; the detached columnals espe-
22 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
cially, on account of their commonly starry shape and delicate markings, had been the objects of much superstitious awe, so that we find them figuring in the early literature, under the names “pentacrinos,” ‘asteroites,’’ or ‘‘entrochos,” much more frequently than the less dreadful but, as we know to-day, far more interesting crowns.
It remained for Edward Llhuyd to first point out (in 1699 and 1703) the con- nection between the fossil crinoids and the recent sea stars, and to go even further and suggest the Rosy Feather Star (‘‘Decempeda cornubiensium,” i. e., Antedon bifida) as the type of sea star to which they were most closely related. Llhuyd really deserves far more credit than is commonly accorded him for dissipating this halo of semi-religious mystery which surrounded the fossil crinoids, and for his great discovery of the relationship between them and the comatulids. His excellent work, which may almost be said to have laid the foundation for the study of the Crinoidea, did not meet with the reception it deserved; his modest, yet convincing, essays could not penetrate the thick wall of popular prejudice, and the comatulids were later again assigned to the place which they had previously occupied.
In 1711 Petiver described and figured the first comatulid known from a locality outside of Europe, calling it the “Stella chinensis perlegens”’ (Capillaster multira- diata). Three years later Barrelier described anew the form first noticed by Columna under the names of barbata and fimbriata.
In 1719 Rosinus, ignorant of the work of Llhuyd, attempted to show the con- nection between the fossil crinoids and the recent sea stars, but he selected the basket stars (Astrophyton, etc.) as the recent forms to which the crinoids are most nearly related, thus not advancing so far as had Llhuyd 16 years before, though in justice to him it must be admitted that he did not have the opportunities for examin- ing the recent comatulids which were enjoyed by Llhuyd.
In his really remarkable work upon the sea stars, published in 1733, John Henry Linck gathered into one volume all of the facts which had been discovered concerning the group. The comatulids he differentiated from the asteroids and from the ophiuroids, placing them in the class ‘Stelle Crinite,’”’ or hair stars, in which he distinguished three genera—dexdxvepoc, with three species, Tpeoxacdexdxvepoc, with one, and Caput- Medusz, with two, as follows:
Class STELLA CRINITA. Genus dexdxvepoc: 4. crocea (founded on the dexadasvaxtevoecdij¢ of Columna)...... Antedon mediterranea. 4. rosacea (founded on the Decempeda cornubiensium of Lihuyd).......2 Antedon bifida. 4. barbata (founded on the Stella fimbriata of Barrelier).......-- Antedon mediterranea. Genus Tpcoxacdexdxvenog (founded on the Stella chinensis perlegens of Petiver.)
Capillaster multiradiata. Genus Caput-Medusex:
CAOMUMMMEN BD: DOVn as dcee ec cee dee onc Omen ene eens ?Comanthus bennetti. C. cinereum, sp. hoy.; according to Miller, Lamprometra palmata, though more likely to be L. protectus, a species which was not differentiated from L. palmata by Miiller.
MONOGRAPH OF THE EXISTING CRINOIDS. 23
The specimens of the two last, which were the only new species described by Linck, were in the collection of Albert Seba and are now probably in the St. Peters- burg museum.
Linck appears to have admitted the close connection shown by Llhuyd between the comatulids and the fossil crimoids; but he had nothing to add to Lilhuyd’s lucid exposition of the facts, so he contented himself with reprinting his dissertation as an appendix.
In spite of the advances which had been made, the next step was a wholesale retrogression and threw the study of the group into utter chaos; for Linné in 1758 placed the comatulids with the starfish and the ophiuroids in the genus Asterias, recognizing only two species, both composites, and neither including any reference to the species represented by the respective type-spectmens.
His first species is:
Asterias pectinata=Antedon bifida+ A. mediterranea + Capillaster multiradiata;
but the type-specimen (at Lund) is not even generically identical with any of these supposed synonyms, being of the species now known as Comatula pectinata; this discrepancy is suggested by the locality given, Indian Seas, whereas Antedon bifida (as known to Linné) is from Cornwall, A. mediterranea from Italy, and Capillaster multiradiata from China. We have to thank Retzius, Miiller, and P. H. Carpenter for redescriptions of the specimen which Linné had in mind when he penned his Asterias pectinata. Linné’s second species is:
Asterias multiradiata=Linck’s Caput-Meduse cinereum+his C. brunnum,
the first of which is undoubtedly a Lamprometra, possibly, as Miller supposed, L. palmata, though more likely L. protectus; the second undoubtedly one of the Comas- teridx, possibly Comanthus bennetti. Retzius and Carpenter have shown, however, as in the case of the preceding, that the type-specimen is generically different from either, and Asterias (Capillaster) multiradiata has been restricted accordingly.
In 1761 the great Dutch collector, Albert Seba, figured and described two multi- brachiate comatulids, one of which was said to have come from Mexico, but both of which probably came from the East Indies.
In the twelfth edition of his work (1767) Linné added to the synonymy of Asterias pectinata Seba’s Stella marinis polyactis, seu Luna marina, said to have come from Mexico (undoubtedly a Himerometra), and his Luna marina altera (which is probably one of the Comasteride), of unknown habitat. In 1758 out of the five references which he cites under Asterias pectinata, four are to 10-armed forms (Antedon) and one to a 13-armed specimen (the Stella chinensis perlegens of Petiver); of the two additional references given in 1767, one of the figures (Stella marinis polyactis) shows 29 arms, the other (Luna marina altera) 37. With this heterogeneous concept in mind it is no wonder that he concludes his discussion of Asterias multiradiata by saying that it is possibly only a variety of A. pectinata.
In 1777 Pennant restricted the Linnean Asterias pectinata by describing his Asterias bifida and A. decacnemus, both of which, however, represent the same
24 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
species, the Decempeda cornubiensium of Lihuyd, or the Antedon bifida as now known; and in 1783 Retzius carefully redescribed the Linnean types of Asterias pectinata and A. multiradiata, at the same time adding to science a new species from the American side of the Atlantic, Asterias tenella, the Hathrometra tenella of to-day.
Brugiére, in the ‘‘Encyclopedie méthodique” (1792), republished the figures of comatulids given by previous authors.
Toward the end of the eighteenth century, Pennant, Forster, and Latham and Davis, in the various editions of the ‘‘Faunula Indica,” included both the Linnean species as given by that author and on his authority, bemg able to add no original matter of their own.
Speaking solely with reference to the Linnean system of nomenclature, de Fréminville took the first step in the recognition of the comatulids as a group dis- tinct from the other sea stars; in a short paper published in 1811 he proposed the genus Antedon for the common west European species (A. bifida), a specimen of which he had found in a dry dock at Havre, adhering to the growth on a ship’s bottom. He made no attempt to elucidate the two Linnean species, or any others previously known, in connection with the new one he described (A. gorgonia), nor did he go further than to show in what way it differed from the ophiuroids.
Simultaneously Lamarck had become dissatisfied with the heterogeneous character of the Linnean genus Asterias, and in the following year (1812), in the second volume preliminary to his great work on the invertebrates, he suggested the vernacular name ‘‘Comatule” (though without diagnosis) for the comatulids, which he latinized and formally described in 1816 as Comatula, assigning to his new genus eight species, seven of them new, and overlooking the Asterias tenella of Retzius.
But in the meantime (1815) William Elford Leach had slipped in with his new genus Alecto, covering the same ground as Lamarck’s Comatula, to which he assigned three species, all of which, as well as the genus itself, were very poorly diagnosed. Leach’s new species were based upon specimens then in the British Museum; he made no reference to any other worker and, as his types have since been lost, we do not know for certain (except in one case by a fairly reasonable inference) what his species were. As given by himself the three species are:
Alecto NOTUAG © o.oo fe sa 5 cals oS ee Sot ee ee Eee eee rn Alecto Cur pea. oe sas Soa ete ROS ce ae ORs Antedon bifida. Alecto) CONNALGE 3. S326 55524 -F= n> ee eee eee eee (most probably) Tropiometra, sp.
It is important to scrutinize carefully Leach’s arrangement in order to determine the availability of Alecto as a generic name. All subsequent authors, for instance Schweigger in 1819 and Miller in 1840, have accepted Alecto horrida as the repre- sentative species of the genus. Alecto europza is the same as the Antedon gorgonia of De Fréminville, and is therefore the type of Antedon, 1811; moreover, it is also the same thing as the Ganymeda pulchella of J. E. Gray, 1834, which is the type of the genus Ganymeda. Alecto carinata is possibly the same as the Comatula carinata of Lamarck, 1816, which is the type of the genus Tropiometra, 1907; this process of elimination thus leaving Alecto horrida as the type of Alecto. Alecto horrida is quite unidentifiable, and therefore Alecto is unavailable as a generic name among the
MONOGRAPH OF THE EXISTING CRINOIDS. 25
comatulids, which is rather fortunate in view of the fact that a subsequently estab- lished Alecto has been widely used as a generic name among the Bryozoa.
Schweigger attempted to make Alecto horrida a synonym of the Linnean Asterias multiradiata; but we can not attach any importance to this, as it was cus- tomary until a much later date to consider all multibrachiate comatulids as belong- ing to the species ‘‘ multiradiata,” as was done, for instance, by Audouin and Leuck- art, through ignorance of the real generic and specific, as well as of the family, characters of the animals.
The comatulids mentioned and described by Lamarck in the year following Leach’s description of his three new species of Alecto are:
Comatula solaris, sp. NOV-.-----------------------2-+200+- Comatula solaris. Capillaster sentosa. Comatula multiradiata.......--------+--------+-------++---- {come bennetti. % Comaster multifida. Comatula rotalaria, sp. NOV.-------------------------------- Comatula rotalaria. Comatula fimbriata, sp. NOV ...---------------------+-+----- Capillaster multiradiata. Comatula carinata, sp. NOV...-.----------------------------- Tropiometra carinata. Comatula mediterranea, 8p. NOV ..--.------------------------ Antedon mediterranea. Comatula adeonx, sp. NOV-..---..-------------------+--+---- Oligometrides adeonz. Comatula brachiolata, sp. NOV .------------------------------ Comatulella brachiolata.
The determination of the type of the genus Comatula is a matter of consider- able importance in crinoid nomenclature ; succeeding authors have either accepted it in the sense of Lamarck to cover all comatulids, or have dropped it altogether ; the genus has never been properly revised. Now Lamarck’s generic diagnosis is quite explicit; it reads, ‘‘bouche inférieur, centrale, isolée, membraneuse, tubu- leuse, saillante;” this obviously refers to the anal tube which was mistaken by Lamarck for the mouth, and shows that when it was written he had in mind an exocyclic form, or a member of the family Comasteridx, thus eliminating from consideration the species adeonz, carinata, fimbriata (which has a central or sub- central mouth, though belonging to the Comasteride), and mediterranea, and leaving solaris, brachiolata, rotalaria, and multiradiata, the last having been subsequently eliminated by L. Agassiz, who made it the type of his new genus Comaster in 1836. Rotalaria was designated as the type of Comanthus in 1907, thus leaving the two species solaris and brachiolata as possible types of Comatula; of the two solaris agrees best with the generic description which, moreover, could not by any chance have been based upon brachiolata, as the two specimens of that form known to Lamarck are both very small, and have the arms folded in such a way as to conceal the disk. Thus we find that solaris must be taken as the type of the genus Comatula.
Lamarck had undoubtedly originated the name Comatula or, in its French form, “Comatule” long before he published it, and before either Antedon or Alecto were published, and, as priority of publication was not such a vital matter in those days as it is now, he was unwilling to relinquish it in favor of either of the earlier names, the more so as both of these were ill-defined and covered the ground only in a rudimentary way; his reputation was so great that practically all succeeding authors followed him, only a very few resurrecting Leach’s name Alecto, while Antedon was completely buried.
79146°—Bull. 82—15 3
26 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
Prof. Johannes Miller was largely responsible for the later disregard of the generic name Comatula in favor of Alecto, rehabilitated, and Actinometra, newly coined; for he employed Comatula as a term to include all comatulids, and expressed the finer divisions by Alecto and Actinometra, used in a subgeneric sense. Dr. P. H. Carpenter followed Miiller in this perversion of Comatula, and used the name only in a sort of vernacular way, much as we now use the word ‘‘comatulid.” In speaking exactly he always used Eudiocrinus, Antedon, Actinometra, Atelecrinus, etc., but when he merely wished to differentiate the free from the stalked forms he always spoke of the former, or of any one of them (most commonly Antedon bifida, which he regarded as the type of the group), as ‘‘Comatula.”
Lamarck entirely failed to recognize the affinities of the comatulids, and placed them with the starfishes, though in a separate genus, as other post-Linnean authors had dote.
In the year following the appearance of Lamarck’s treatise on the comatulids the portion of Savigny’s description of Egypt dealing with the echinoderms was published; in it were figured two comatulids from the Red Sea, one of which was designated (the identifications being by Audouin) as ‘‘Comatula sp.,” the other as “Comatula multiradiata.”’ There is no further reference to the first of these figures, which represents the local species of Tropiometra; but in 1836 de Blainyille copied the second in the atlas to his ‘‘Manuel d’Actinologie;”’ in doing this he made a curious mistake, for the plate is lettered ‘‘Comatula adeonex,” though in the text the description of Comatula adeone is taken from Lamarck, and the species is correctly said to have 10 arms. In the year following the ‘‘Penny Encyclopedia” copied de Blainville’s account of Comatula adeonx, multiradiate figure and all, and the same slip was made by Knight in his ‘‘Natural History,” published in 1867.
In 1819 Schweigger figured various parts of a species which he determined, without doubt incorrectly, as ‘‘Comatula multiradiata;” he further identified this with Leach’s Alecto horrida.
J.S. Miller, in his epoch making monograph published in 1821, again raised the comatulids to a position next to the fossil crinoids, and thus brought the conception of the group as a whole to the same level at which it had been left by Llhuyd 120 years before. Miller proposed the name Crinoidea for the class, but he only mentioned one comatulid, the Rosy Feather Star (the only one with which he was personally acquainted), which he had found at Milford Haven. He was unable to place this species in reference to those described by Lamarck, and therefore tentatively described it as new under the name of Comatula fimbriata, which name Miiller in 1841 changed to milleri owing to the conflict with the Lamarckian Comatula fimbriata which is quite a different thing. Lamarck’s Comatula fimbriata is the species now known as Capillaster multiradiata, a species belonging to the Comasteridx, while Miller’s Comatula fimbriata is the common Antedon bifida, a species belonging to the Antedonide.
In 1822 we find the first reference to a comatulid in American zodlogical litera- ture, Prof. S. L. Mitchill recording two specimens, which he did not identify, from Gaspar Strait. In 1825 Mr. Titian Peale found on the beach at Great Egg Harbor, New Jersey, a specimen which he sent to the Museum of the Philadelphia Academy;
MONOGRAPH OF THE EXISTING CRINOIDS. 21
there it was studied by Thomas Say who, however, could not identify it with any of the species then known, so he described it as new, calling it (emending Leach’s generic name) Alectro dentata. Say’s species has never been properly understood; it has been very generally confused with Alecto sarsii, later described, and with the Asterias tenella of Retzius which also came from America, but from farther north, although it is in reality perfectly distinct from both. It is probable that up to the present time no one has been able to make direct comparisons between these three forms, for certainly Carpenter, had he done so, could never, as he did, have called them identical.
About this time (but just when I have been unable to ascertain) W. E. Leach described the common and magnificent arctic species, from specimens brought from Spitzbergen, as Alecto (i. e., Heliometra) glacialis.
In 1826 Risso published his Comatula coralina and C. annulata (both synonyms of Lamarck’s Comatula mediterranea), basing them upon specimens obtained at Nice; and in the same year J. E. Gray published a paper on the digestive system of the comatulids in which he proposed uniting them with the so-called Crinoidea of Miller under the family name of Encrinitidx; in other words proposing Encrinitide (or Encrinidz) as a synonym of Miller’s Crinoidea.
The year 1827 was a memorable one in the history of the comatulids, for in that year Dr. John Vaughan Thompson discovered in the Cove of Cork in Ireland a small organism which he at once recognized as a crinoid and described in detail in his classical memoir on the ‘‘Pentacrinus europeus.’’ In the following year Fleming became impressed with the differences between this small species and the larger pentacrinites, and proposed for it the new generic name Hibernula, this being rejected two years later by de Blainville who, considering that the names of all stalked crinoids should end in ‘‘-crinus,” rechristened it Phytocrinus. But Thompson had not been satisfied with the mere discovery of this interesting animal ; he made it the object of careful study, and in 1835 he announced that it was nothing more nor less than the young of the common comatulid, Antedon bifida.
Fleming in 1828 suggested the recognition of two species of British comatu- lids, as had been done by Pennant, but for them he resurrected the long-forgotten names of Linck, calling them Comatula rosacea and C. barbata. The former was quickly adopted, both because of its eminent appropriateness and because of the great and deserved prestige of its author, and had become firmly fixed in the nomenclature before growmg sentiment in favor of a more stringent adherence to the principle of adopting the works of Linné as the starting point in all zoélog- ical nomenclature finally dislodged it. Some sacrifice must of necessity be made to secure nomenclatorial uniformity, but we can not help regretting the rejection of the appropriate names conferred upon the sea stars by such a master of the subject as Linck in favor of the attenuated and often questionable nomenclatorial resultants obtained by the analysis of the unwieldy composites created by his less discriminating successor. At the same time Fleming proposed the family Comatulade for the comatulids, together with the Pentacrinus europzus of Thomp- son, and he suggested a division of the family, one part to contain certain forms having the digestive apparatus with two apertures (as Gray had shown to be the
28 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
case in Antedon bifida), the other to contain those like Pentacrinus ewropeus in which the digestive system was supposed to have but a single opening.
In 1831 Ferussac recorded that M. Lemare-Piquot brought back many coma- tulids collected on his voyage to the East Indies and South Africa.
Georg August Goldfuss in 1832 published a description and a figure of a speci- men which he had found at Bonn, which he referred to ‘‘ Comatula multiradiata” ; the species represented is the Alecto bennetti subsequently described by Miller. At the same time Goldfuss gave a good comparative account of the common Mediter- ranean Antedon for comparison with the fossil species with which he was mainly concerned,
Riippel, in the course of his travels, found in the Red Sea an interesting multi- radiate comatulid upon which he bestowed the manuscript name of ‘‘ Comatula leucomelas,”’ but he does not appear to have mentioned it anywhere in his works. In 1833 Leuckart came across his specimens in the Senckenberg Museum at Frank- fort and published the name together with the locality, though without any diag- nosis. Recently Hartlaub has reéxamined the specimens, and has found them to be examples of the Alecto palmata later described by Miller.
Leuckart was the first to describe the curious parasitic worms belonging to the genus Myzostoma with which crinoids are usually infested, his attention having been first called to them by mistaking one for a madreporic plate. In discussing the genus Jfyzostoma he mentions a multiradiate comatulid from the Red Sea which, following Audouin, he identifies as “‘Comatula multiradiata,’ but which von Graff, acting on the advice of P. H. Carpenter, has suggested was probably an example of Heterometra savignii, the species to which Audouin’s Comatula multiradiata has always been referred.
In 1834 Dr. J. E. Gray found upon the coast of Kent a peculiar organism which he was unable to place, and he therefore described it as new under the name of Ganymeda pulchella. Later it was discovered that his supposedly anomalous creature was merely the detached centrodorsal of the common Antedon bifida.
In 1835 the first mention of a recent crinoid occurs in Australian zodlogical literature; in that year the Rey. C. Pleydell N. Wilton described, under the name of Enerinus australis, what he supposed to be a new species, but which has since proved not to be a crinoid at all. Ten years later his paper was in part translated into French and reprinted, the author's name being incorrectly given as ‘Rev. C. Pleydell.’’
In the year 1836 de Blainville published a valuable summary of the knowledge which had been acquired in regard to the comatulids; his account of them is prac- tically the same as that contained in the later editions of the work of Lamarck. de Blainville had previously published two less extended treatises on the group in the well known “‘Dictionaire d’histoire naturelle,’ one in volume 10 (1818), the other in volume 60 (1830).
Prof. Louis Agassiz in 1836 founded his genus Comaster, based upon the Comat- ula multiradiata of Lamarck, which unfortunately is not the same as the Asterias multiradiata of Linné and of Retzius. Agassiz employed as the differential char- acter for his new genus the excess of the numbor of arms over the 10 found in
MONOGRAPH OF THE EXISTING CRINOIDS. 29
“
Comatula as he restricted it, a character which we now know to be of very uncertain value.
In the ‘Inconographie du Regne Animal” published by Guérin-Ménéville during the years from 1828 to 1837 there are two figures supposed to represent the species described as Comatula carinata from Mauritius; possibly the first (2) does represent this-species, though it looks more like some species of Antedon; but the second (2a) appears to be a species of Amphimetra, and agrees fairly well with A. discoidea from northern Australia and the East Indies. There is a specimen of Amphimetra discoidea (labeled by P. H. Carpenter Antedon milberti var. dibra- chiata) in the Paris Museum from which I suspect this figure was drawn.
In the course of his studies on the echinoderms Prof. Johannes Miller had become interested in the comatulids, and in 1841 he published a paper upon the group in which he described the new genus Actinometra. The type of his new genus was the new species Actinometra impertalis, founded upon a magnificent speci- men two feet in expanse which he had found in the Vienna Museum labeled ‘ Coma- tula solaris.’ In addition to Actinometra imperialis Miller described as new the following species, all of which he referred to Leach’s genus Alecto:
Alecto milleri (new name for Comatula jimbriata Miller, not
Comatula fimbriata Lamarck)........--------------------4 Antedon bifida. Alecto phalangium.......-----------+-+----+- 2222-22222 rete Leptometra phalangium. AllectO: €8CRTUCKL = 9) = 21 = «=m iniei= == infec = ale min me wn = Heliometra glacialis.
Alecto echinoptera
Ey es ers are oinias et Neteller tai Comactinia echinoptera. Alecto rosea
oe ey oie ae et stS eee ee cleepaie ET Comatula brachiolata.
AEG TGUIG TR aoe ceacre ce anaacscopeusee oun dns sous - saree Amphimetra tessellata. Alecto polyarthra........-----------------+--------+-----+-=- (Not identifiable.) Alecto multifida (see below)..---...------------++------------- Comaster multifida. Allecto 80VUQIIN. 22-10 0 - oan = = a oa meses ei ain in Heterometra savignit.
| Lamprometra palmata.
Alecto palmata ecto p |Lamprometra protectus.
Alecto parvicirra. .....-------+-+----- +--+ 2-222 eee etree eee Comanthus parvicirra, PAllechONLUTILONeNSIS sane eens so sila sate le epee ae etal ote na aii Comanthus parvicirra. Alecto japonicd.......-----+--+-+-------- 2-2 - == 2-222 eee eee Comanthus japonica.
WAlectoNfiag CUlatas ee. on sae ne ee le aaa inl aeae = = Dichrometra flagellata. Alecto novxe-guine®. ..---------- +++ +--+ 2-2-2 - 22 e rere Comaster novxeguinex. Alecto elongata........--------------------------+-----+---- Dichrometra flagellata. AUD GDB acaba ceessecnneetod sho seocus Jecooceueadsopenese Comanthus bennetti.
Miiller found in the literature three species which bore the name multiradiata, in addition to the so-called ‘‘multiradiatas” of Leuckart, Audouin, and Schweigger, which he seems to have correctly considered wrongly so called; one of these had been described by Linné (Asterias multiradiata) and later redescribed by Retzius, another had been described by Lamarck (Comatula multiradiata), while a third had been described and beautifully figured by Goldfuss (Comatula multiradiata). Both Lamarck and Goldfuss had been under the impression that the species they had in hand was the one originally diagnosed by Linné. Miller took the ground that the name should hold for the species or form which was best described, and he treated the Asterias multiradiata of Linné and the Comatula multiradiata of Lamarck as being quite unrecognizable from the published descriptions, and therefore not ten- able. The Comatula multiradiata of Goldfuss, well described and illustrated with
30 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
an excellent figure, he considered as the true multiradiata, and when he came to examine Troschel’s notes upon the Lamarckian types at Paris he never thought of restoring to them the name multiradiata, but renamed one of the two forms repre- sented among them (Alecto) multifida, at the same time describing it in detail.
This action of Miller in describing anew the Comatula multiradiata of Lamarck, hitherto unidentifiable, resulted in the positive identification of that species, and with it, the genus of which it is the type, the Comaster of L. Agassiz. The type of this genus now becomes Alecto multifida Miller—Comatula multiradiata Lamarék reidentified. Concurrently with his perversion of the specific name multiradiata, Miiller shifted the genus Comaster of Agassiz to cover the species described and figured by Goldfuss, in spite of Agassiz’ statement that the multiradiata of Lamarck was the fype.
Although P. H. Carpenter in his earlier work partially rectified this error, he later accepted Miiller’s views in regard to Comaster, and thus failed to recognize its rightful place in nomenclature.
In the year in which Miller published his first paper on the comatulids (1841) Delle Chiaie described his Comatula bicolor, which seems to have attracted little attention, as it was generally recognized as merely a synonym of Lamarck’s Comatula mediterranea.
Miller went to Sweden and examined at Lund the Linnean types, publishing in 1843 a redescription of both Asterias multiradiata and A. pectinata, but he curi- ously overlooked the type of Retzius’ Asterias tenella. At the same time he de- scribed two new species, Alecto purpurea, which he found in the Berlin Museum, and Alecto wahlbergii, which he found in the Stockholm Museum. Both of these species have since been strangely neglected, the former being incorrectly treated as a synonym of the Linnean Asterias pectinata, and the latter as a synonym of Miiller’s earlier Alecto parvicirra.
Michelin in 1845 noted the occurrence of Comatula carinata (Tropiometra carinata) at Mauritius.
In 1846 Diiben and Koren announced the discovery on the coasts of Scandinavia of two species which they were unable to identify with any of the previously de- scribed forms; they accordingly proposed for them the names Alecto petasus and Alecto sarsii, following Miller in the use of Leach’s name Alecto. The first of these species had been reported from the Scandinavian coast by Prof. Michael Sars in 1835 under the name of Comatula mediterranea, but his notice of its occurrence does not seem to have attracted much attention.
In 1846 Miiller deseribed four additional species (Comatula macronema, C. jacquinoti, OC. trichoptera, and C. reynaudii) which he found in the Paris Museum, and in 1849 he published his very important memoir on the genus Comatula and its species, the first really adequate work on the subject, in which he treated of all the forms then known. His genus Actinometra had given him considerable trouble, for in many cases he had been unable to determine whether a specimen should be referred to that genus or to Alecto (as understood by him), and in specimens in which the disk was lost or concealed, as he knew of no other differences than those afforded by the,arrangement of the ambulacra, he was, of course, quite at a loss. He there- fore reduced Actinometra and Alecto to subgeneric rank under Comatula, which he
MONOGRAPH OF THE EXISTING CRINOIDS. 31
employed as a general term to cover all species; if he could make out with cer- tainty the arrangement of the ambulacra, he inserted Alecto or Actinometra, as the case happened to be, between Comatula and the specific name; if he could not, he omitted the subgeneric designation and referred the species unqualifiedly to Comatula. Some idea of the difficulties which he encountered (undoubtedly largely through differences in the state ot preservation and consequent different degrees of distortion of the soft parts of the specimens examined by him) may be gathered from the fact that he placed a single species, Comanthus parvicirra, both in Actinometra (twice) and in Alecto, and also in the incerte sedis under Comatula.
In the course of his studies Miller had discovered that his Actinometra impe- rialis was identical with Lamarck’s Comatula solaris; but while he dropped the spe- cific name imperialis he still clung to his Actinometra, not relinquishing it, as he should have done, in favor of Comatula. In this, as in other things, he was followed by P. H. Carpenter.
Miiller’s final arrangement of the comatulids was as follows:
Comatula (Actinometra) solaris..........--------------------Comatula solaris. (GE(CActimometra) rota lamiasme sae eae se eas ae rea Comatula rotalaria.
Ga eAchinometi@) MUGlt DERG Vcc etree se ae ne nee oll Comanthus wahlbergii. ( (Aldecta eChinio Pten Omen ees arte lateral Comactinia echinoptera. (GE (Allecto) Wrrred tern O10 (eet eet eee oe ei Antedon mediterranea. (Oa CAlecto caries eee ee eee ee eae tee Tropiometra carinata. CR (Alecto) mail bertyaeeen eran oie ae a ea Amphimetra milberti. CUCAllecto) phalanguinis esses ee ee ee eee Leptometra phalangium. (GR(CAllecto) pelos is aceite ere a ete ale aera er ei oi Antedon petasus. Ga(Allecto) sareiisen se eee eee ee eee Hathrometra sarsii.
(GA (Allecto es chirich titan semen em esto alata ee Heliometra glacialis.
Ga (Al ecto) sagittis ees a eee ola Heterometra savignii.
OS (CAllecio) jumbria asec = eee eee ee ele eerie ee Capillaster multiradiata. C. (Alecto) reynaudii.........-------------------++=-------=- Heterometra reynaudii. (GN(Allecto)) DOTViCir TG eae = ae alae a ale aoe eer = Comanthus parvicirra.
Lamprometra protectus. G, (Alecto) palmata......---.-------=-9>-<---------=- ee Sche Lainproniesea boatnintan
er Capillaster sentosa. C. (Alecto) multiradiata......-...----+----+-+--------+------- | Capiiasten enaiinesats C. (Alecto) articulata, sp. nOV...----------------------------- Liparometra articulata. GGTACRO Latte eee eae © = aaa ee ae nine Comatula brachiolata. Guniiilenttease eee oe cn cen = oe oe oa ie ae ee a eae ee Antedon bifida. (GS OSEO ee eos sane n= aslo sie ee aes Pelee nal Comatula brachiolata. Gado eke eae sss eS Acie ees aoe eee Oligometrides adeonx. C. cumingii, sp. NOV.-.---.--------+---+---+2 202s rere eee ee Comatula pectinata. C. elongata.....-.------2-----+ 022222 cere tenner e serene Dichrometra flagellata. C. trichopterd...-...---------- +--+ 22-22-2222 reer t etree Comanthus trichoptera. G@. macronema.....=-..-----=-+-+-2-2-- 22-225 - + ester ess Ptilometra macronema. C. philiberti......-.--------++---02- 22222 e eee eter e estes Amphimetra philiberti. C. japonica: .......---.--------------+---=+----- See ees Comanthus japonica. G, multifida.....-.-2.-2------22 + 2-222 2c e nese e eset esse eee Comaster multifida. GEALUT ONENESS Seer iors Ge = ie wi aia mass sie ein Tans = cine Comanthus parvicirra. C. flagellata....--..----------+--+ +++ +2222 e eee terete Dichrometra flagellata. C. nove-quinex....--.-----------+----- Dee ape a nate nya tiete ...-Comaster novxguinex. Galben el iis eee ane = cide nia = Sales siele erin Seiei=ieleitae sie erate Comanthus bennetti. C.. jacquinoti......--2-----------0- 222222 cee neers eee Amphimetra jacquinoti.
Gin festaiiataMee eee cess. c1- owe niod nine hiss Ass Satan eas ia ar Amphimetra (?) tessellata.
82 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
It will be noticed that there is no mention in this list of the Asterias tenella of Retzius, the Alectro dentata of Say, or of the Alecto glacialis of Leach.
While working up the natural history of Chile for inclusion in his monographie account of that country, Gay found in the Paris Museum a number of comatulids which had been labeled by Valenciennes Comatula picta, and which were said to have been obtained in Chile. In the eighth volume of his work (1854) Gay described these under Valenciennes’ name of Comatula picta; but he makes no further men- tion of their occurrence in that country. The specimens belong to the common Brazilian species of Tropiometra, and could not have come from Chile; they prob- ably came to France on a ship from Chile which had touched at some Brazilian port on the way.
In 1857 Barrett discovered in the Sound of Skye a curious species which he named Comatula woodwardii (Leptometra celtica), but which he renamed (jointly with McAndrew) in the following year Comatula celtica, to avoid conflict with a previously described fossil Comatula woodwardii.
Dujardin and Hupé in 1862 published their great work on the so-called zo06- phytes, the former being responsible for that part which dealt with the comatulids. These authors followed Miller closely, but corrected many of his mistakes, while making some additional errors of their own. They recognized three genera of recent comatulids which they called Actinometra, Comatula, and Comaster, the last being based upon Goldfuss’, Comatula multiradiata and used, therefore, in the same sense in which it was understood by Miiller. Actinometra as described by them has a central anus, the brachial ambulacra leading to a horseshoe-shaped peripheral furrow; Comatula included the forms in which the mouth is central and forms the converging point of five equal radiating ambulacra on the disk. Actino- metra imperialis, which Miller himself had shown to be but a synonym of Comatula solaris, they reinstated as a valid species, even going so far as to consider it generically different from C. solaris.
Their arrangement of the yarious species is:
Comatula medterrdnens. os sess oa oe ek aoe tae ee eee Antedon mediterranea. Comimtina phalanguivimys sso. c56 os eecae sh eRe eee Leptometra phalangium. Comatula petasus. <2 <2222-s05ssvecenes oese de <5>- 52 —Antedon petasus: COMUAtULE RORBE Se ssa es oda 4 Se GA Ee RR eee Hathrometra sarsit. Comatula eschrichtii.............-..-- ae Aen ee ane eee ..-Heliometra glacialis. Comatila carinata® Sas. 22222 22 5s oe eee eee 3 Sades Soe Tropiometra carinata. Comatilar ddhonige Noe oss Rec ee Oligometrides adeonz. Comatula trichoplenam 428. % 5.) bu SUR eye cee eo Comanthus trichoptera. Comatitlals eynatuiy ake 8 32228 = ots 9-7 Boers Sa ieeranrn Heterometra reynaudii. Comatilareolaris mare pecan cos tc eas ce en ee Comatula solaris. COMABIG UTRCNOUAG. Ae lycn 2c an oe Bone oe renee Comatulella brachiolata. Comatila'echnnoplerd.. 22... 25 scat cce- fee Comactinia echinoptera. Conuriitlaposedi ss soc 20 knee Sot SS a ee ec ee eee Comatulella brachiolata. Conant a tease tata ee is mene See hee tn ge rote Amphimetra (?) tessellata. CRITE DURES sheet a SUS ee eats ee Comatula purpurea. Comatnlen philber tie. 202585 orc ooceh vines comet ances cue sen See Amphimetra philiberti. COMMER NAWE Coser es one woe ph Lean See tLe Seok Cer ee Amphimetra milberti.
COMMU A FORRMO = aos one Ee eae See eee Amphimetra jacquinoti.
MONOGRAPH OF THE EXISTING CRINOIDS. 33
Comatula macronema...---------+++++-++-- +22 25rrett tt Ptilometra macronema. Comatula savignyi ....---------++---+---2220e cette ttre Heterometra savignit. Comatula rotalaria.......--------------------+-22 crt Comatula rotalaria. Comatula fimbriata....-.------------------+++-2222e etree Capillaster multiradiata. Comatula elongata....-------------------------72007 0700 Dichrometra flagellata. Comatula parvicirra. .....----------+-----2222020 05 tr rte Comanthus parvicirra. Comatula japonica. .-..-----------------+-+-222222 700 or ee Comanthus japonica. Comatula flagellata.......--------------++-+-------77 77707 Dichrometra flagellata. Comatula timorensis...-.-.---------------------- +t ttre Comanthus parvicura. Comatula articulata. ....------------+----------- 22000 ttt ce Liparometra articulata. Comatula multifida......--------------------220222 2200 Comaster multifida. Comatula novx-guinex.-..------------------+222227 000 0t Comaster novxguiner. Comatula bennetti.....-.------------------- 2220 ttre Comanthus bennettt. Actinometra imperialis......----------------+-++77277 00077 Comatula solaris. Actinometra pectinata...--.--------------------702 tr ttttt Comatula pectinata. Actinometra multiradiata...-.---------+--------------7777- lon ae Sem Capillaster multiradiata. Actinometra wahlbergii....-------------------7----70 ttt? Comanthus wahlbergit. Comaster multiradiatus.....---------------+----+2-27-07777> Comanthus bennetti.
In addition to these described forms they gave a list of undescribed species, taking the names from labeled specimens in the Paris Museum.
While we are not at present directly concerned except with the systematic history of the comatulids, it would be impossible to appreciate this properly without some idea of the relative progress made along other lines of study, and it is therefore fitting that some mention be made of the new era in the elucidation of the structure and development of the group which began in the year 1863.
Adams in 1800 had called attention to the two apertures on the comatulid disk, while in the years 1823-1826 Péron, Gray, Leuckart, Meckel and Heusinger independently demonstrated, in varying degrees of completeness, the existence of a coiled digestive tract. In 1835 Dujardin showed that the eggs of the comatulids are borne externally on the pinnules and are not internal as in the other echinoderms, while in the same year J. V. Thompson demonstrated the stalked condition of the young. In 1843 Miller made a valuable contribution to the knowledge of the struc- ture of the comatulids in his classical memoir on the structure of Pentacrinus caput- meduse. (Isocrinus asteria) ; but the true understanding of the comatulid embryology, development and structure may be justly said to date from the epoch-making memoirs of Prof. George J. Allman, 1863 (‘prebrachial” larval stage), Prof. Sir C. Wyville Thomson, 1865 (early development), and especially of Dr. William Ben- jamin Carpenter, 1866 (ater development, history and structure).
Canon Alfred Merle Norman in 1865 published the results of his researches on British echinoderms, in which he followed Gray (1848) in the use of Antedon in preference to Comatula, at the same time changing the family name to Antedonide. He described no new species, but he recognized, as Pennant and Fleming had done, two British species of the A. bifida type, Antedon rosacea (following Fleming in the use of Linck’s name) and A. milleri, which latter he included on the authority of Sir Wyville Thomson.
In the same year Mr. Alexander Agassiz and Mrs. Elizabeth Cary Agassiz definitely made known the first species of the family Comasteride, Comatula
34 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
meridionalis (Comactinia meridionalis), from the American coast of the Atlantic, though it has subsequently been found that Miiller’s Alecto echinoptera is also an American form.
In 1866 Wilhelm Bohlsche described as new a curious little comatulid from the coast of Brazil which he had been unable to identify with any known form. He called it, in compliment to the justly famous Norwegian naturalist of that name, Antedon diibenii. This species has been the cause of considerable confusion; P. H. Carpenter identified with it a specimen which the Challenger dredged at Bahia, and figured both this specimen and the type in the Challenger report on the “Comatulx.” The Challenger specimen is a young example of Tropiometra picta, but the type specimen obviously belongs to the Antedonide, and to the genus Antedon. It is only within the past year that this species has been rediscovered, the second known specimen having been collected on the island of St. Thomas.
It was in 1866 also that Prof. Sven Lovén instituted the new genus Phanogenia for the reception of a curious exocyclic comatulid from Singapore which differed from all the other species then known in having the centrodorsal very much reduced, in fact merely a small stellate plate, and quite without cirri. This form he called Phanogenia typica (Comaster typica).
Two years afterwards (1868) Professor Lovén announced the startling discovery of a recent cystid at Cape York, Australia, which subsequently proved to be nothing but the detached disk of one of the Zygometride. This so-called Hyponome sarsii of Lovén was the first zygometrid known; but in the same year Prof. Carl Semper introduced to science a second, the peculiar Ophiocrinus (Eudiocrinus) indivisus, remarkable in possessing but 5 arms, whereas all the other comatulids then known had at least 10.
The United States Coast Survey had been for some time engaged in a systematic study of the marine conditions off the coast of the southern United States, and Count L. F. de Pourtalés was thus enabled in 1868 to make known the interesting Comatula brevipinna (Crinometra brevipinna, the first known species of the Charito- metride) and Comatula hagenii (Coccometra hagenii), the first comatulids definitely known from the West Indies, C. brevipinna being, moreover, the first species known with “plated ambulacra”’ like those of the pentacrinites, though their existence in this form was not demonstrated until many years later.
In the following year Kuhl and van Hasselt gave colored figures of two large comatulids, one of which was described as new under the name of Comatula (Acti- nometra) hamata (Comatula solaris), and Pourtalés added to the known fauna of the West Indies his Antedon armata (Analcidometra armata), A. cubensis (Antedon cubensis and Atelecrinus balanoides), and A. rubiginosa (Comactinia meridionalis). At the same time Prof. E. von Martens recorded from the Red Sea the Alecto palmata of Miller, which had originally been described from the Red Sea, and, erro- neously, India, and recorded Comatula solaris (based on a specimen of Tropiometra carinata) from Zanzibar.
Dr. C. F. Liitken had become interested in the comatulids, and had discovered that in the exocyclie species the oral pinnules are furnished with a peculiar terminal comb; he retained Actinometra for the exocyclic forms and used Antedon or Alecto
MONOGRAPH OF THE EXISTING CRINOIDS. 35
for the endocyclic. Unfortunately he never published any detailed account of his studies himself, but he gave to Dr. P. H. Carpenter the results of his researches, by whom they were published, together with his own observations, 10 or 12 years later (1879). Doctor Liitken had, however, in 1866, 1869, 1874, 1877 (two), and 1879, published lists of the comatulids in the collection of the Godeffroy Museum at Hamburg, which clearly show that his conception of the generic limits of ‘“‘ Ante- don” and “ Actinometra” at that time was the same as that elaborated by P. H. Carpenter in 1879 and in 1888. The names used by Liitken were all nomina nuda, but all have since been identified.
In the United States Prof. Addison E. Verrill had taken up the study of the echinoderms and, beginning in 1866, he published various papers in which he brought up to date the somewhat scanty knowledge of the comatulids of North and South America.
Sir C. Wyville Thomson, in his prelimimary report upon the crinoids collected by the Porcupine expedition (1872) and in his semipopular work “The Depths of the Sea,” published in 1873, as well as in “The Atlantic,”’ published in 1877, brought out many new facts concerning the crinoid fauna of the north Atlantic and of the Mediterranean.
In 1875 Grube described three new comatulids from Borneo, Comatula levis- sima (Amphimetra levissima +Amphimetra malberti), Comatula (Actinometra) borneensis (Capillaster multiradiata) and Comatula mertensi (Comanthus parvicirra), reverting to the classification of Miller which had been abandoned by Verrill and Pourtalés, these authors placing all their species in the genus Antedon, following Norman and Gray.
In the year 1877 Prof. E. P. Wright described a supposed new genus and species of sponge from Australia, which he called Kallispongia archeri. Mr. S. O. Ridley, in reviewing the paper for the “Zodlogical Record,” at once noticed the similarity of the animal to the stalked larva of Antedon, and expressed his doubts as to whether it really was a sponge. Subsequent study has shown that Kallispongia archeri is in reality the stalked larva of two Australian crinoids, Ptilometra miilleri and (probably) Compsometra loveni. Were it not that the figure of the pentacrinoid of Ptilometra miilleri is given as a “variety” of the supposed species, Kallispongia would have to be used instead of Ptilometra.
At the same time the Rev. T. R. R. Stebbing, who had been interested in the then current speculation in regard to the origin of the generic name Antedon, pub- lished a short note stating that ’4v@jdov was the name of anymph mentioned by Pausanias, and that the name would be more correct if spelled “ Anthedon.”” This emendation has not, however, been adopted by any one except Minckert, who employed it in one of his papers published in 1905.
Mention should here be made of the monograph published in 1877 by Prof. Ludwig von Graff on the myzostomes, a group of curious ‘‘worms”’ until recently known only as parasites upon the crinoids. In the preparation of this monograph Professor von Graff received many specimens taken from crinoids bearing unpub- lished museum names and from crinoids taken in localities not previously known to support a crinoid fauna. Later Professor von Graff studied the myzostomes from
36 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
the Challenger erinoids, and from those collected by the Blake in the West Indies. The names included in these later reports (two on the Challenger and one on the Blake material) were furnished by P. H. Carpenter; but some of them were subse- quently dropped by Carpenter, and others were never mentioned by him at all, so that they now stand in von Graff’s works as nomina nuda.
In 1878 Pourtalés, continuing his studies, described Antedon alata (Neocomatella alata), A. pulchella (Neocomatella alata), and A. granulifera (Crinometra granulifera).
Owing to the great difficulty which he must have had in comprehending the vague descriptions of the early authors, and to a lack of the true appreciation of the somewhat intricate differential specific characters of the group, as well as to the almost complete absence of material with which to make comparisons, we find the diagnoses of Pourtalés somewhat difficult to comprehend, the more so as many of them are short and indefinite; the absence of authentic type-specimens, and a trans- ference of certain of his original labels to species not agreeing with his diagnoses have added to the confusion. Carpenter attempted to straighten matters out in 1881, but in some ways made things rather worse. Antedon granulifera Carpenter at first decided was an ‘‘ Actinometra”’; later (1888) he shifted the name to a species (Crinometra imbricata) resembling Crinometra brevipinna but entirely lacking the peculiar granulated ornamentation which induced Pourtalés to bestow the name granulifera upon it, and renamed Antedon pourtalesii what is most probably the type of granulifera. Carpenter’s action in regard to Antedon alata and A. pulchella was extremely arbitrary; he saw that the two were synonyms, but, instead of choosing the first name given (alata), he chose the later (pulchella) as being more appropriate.
In 1879 Dr. Edgar A. Smith described in great detail a new comatulid from the island of Rodriguez, which remains to-day the only crinoid known from that locality; he called it Comatula indica (Stephanometra indica) and it was the first species to be discovered belonging to the family Stephanometride. In the same year Dr. Richard Rathbun published the results of his study of the Brazilian comatulids, carefully comparing Brazilian and African specimens of the corresponding species of Tropio- metra, and describing in detail, though conscientiously refraining from naming, another species from Brazil which has since proved to be the interesting Nemaster lineata.
The year 1879 marked the beginning of a new epoch in the study of the comatu- lids, for in that year was published Philip Herbert Carpenter’s masterly monograph on the genus “‘ Actinometra,”’ which is, in many ways, the best work he ever did, and which is free from a number of the more serious errors which mar the Chal- lenger report published nine years later. In this work he reviews the whole subject of the comatulids and gives a detailed account of the comparative structure of such species as were available. One new species, Actinometra polymorpha, is described, which, however, he soon found to be the same as the Alecto parvicirra of Miller.
In the same year Carpenter published a preliminary account of the comatulids which had been collected by the Challenger, in which he diagnosed the remarkable new genus Promachocrinus which has 10 radials instead of the usual 5.
In 1881 Carpenter followed this with a similar report on the collections of the United States Coast Survey steamer Blake, in which he gave us an idea of the fauna
MONOGRAPH OF THE EXISTING CRINOIDS. 37
of the Caribbean Sea, and made known the remarkable new genus Atelecrinus, assigning to it three species, Atelecrinus balanoides (sp. nov.), A. cubensis (Antedon cubensis Pourtalés, in part; immature A. balanoides), and A. sp. (Atelecrinus wyvillii). In addition he described the interesting Antedon spinifera (Stylometra spinifera), and first recorded (in that species) the presence of a complete ambulacral plating in a comatulid comparable to that found in the pentacrinites, while he also figured, without giving a formal description, the extraordinary form which he called Antedon columnaris (Zenometra columnaris). In the same year he reported upon the rich comatulid collection of the Leyden Museum (which had previously been studied by Miller), and laid the foundation for knowledge of the remarkable comatulid fauna of the East Indies. The species which he discussed in this paper were:
Tropiometra encrinus.
Antedon carinata...-.-.------------=+-2-2-2---272777 27° Tropiometra carinata. Tropiometra picta. Antedon serripinna, sp. NOV----------------+---72 705700 Oligometra serripinna. Antedon pinniformis, sp. DOV. .--------------727 7777777774 Amphimetra pinniformis. Antedon perspinosa, Sp. NOV----------+-------7 725522777 Colobometra perspinosa. Antedon spicata, sp. DOV----------------+---7 2770000077" Stephanometra spicata. Antedon levicirra, 8p. DOV. .------------------7270rstt tte Lamprometra protectus. Antedon flagellata......-------------0+7 277752 crtttttt Dichrometra flagellata. Antedon bimaculata, sp. NOV.------------+------277707 7777 Dichrometra bimaculata. Writedon: elongates = ee ana aoe an cin Dichrometra flagellata. Actinometra typica...-----------------2227 ror Comaster typica. Actinometra japonica. .-.-----------+2222-7r rrr rrrr ttt Comanthus japonica. Actinometra schlegelii, sp. NOV. ------------------7777 777777 Comanthina schlegelit. Actinometra nove-guine®....-------+---2252 cer sttttrt Comaster noveguinex. Actinometra robustipinna, sp. DOV. -------------227777 777" Himerometra robustipinna. Actinometra alternans, sp. DOV--------++------772777 77777 Comantheria allernans. Actinometra parvicirra ...---------------27 5572 trct tr Comanthus parvicirra. (Alecto timorensis)-..------=---+--2- =r aceon c recs t neo Comanthus parvicirra. (Comatula simplex) ..--------2-+---2000 borne Comanthus parvicirra. Actinometra peronii, Sp. DOV--------------27 7720007 T Comanthus bennett. Actinometra bennetti....----------=22- 2277-57250 Comanthus bennetti.
In 1882 he further elucidated the East Indian fauna in a similar paper on the comatulids of the Hamburg Museum, in which he also takes up the peculiar genus Ophiocrinus (Semper, 1868), changing the name to Eudiocrinus (Ophiocrinus being preoccupied), and describing some additional species, which have recently been shown to have only a very remote relation to the original Ophiocrinus indivisus.
The comatulids considered in this paper are:
Atelecrinus balanoides...--.-----<--=2 =~ - 9925223 Atelecrinus balanoides. eelecrinus: CUDENSIS = = <== 2 Atelecrinus balanoides. Atelecrinus wyvillii, sp. NOV.------------77 225-02 r Atelecrinus wyvillit. Eudiocrinus indivisus.--.--------------07 ct tc rr Eudiocrinus indivisus. Eudiocrinus varians, sp. DOV---------------7557 7775050 Pentametrocrinus varians. Eudiocrinus semperi, Sp. ROV--------------7277 7770777777 Pentametrocrinus sempert.
Budiocrinus japonicus, 8p. DOV----------+-22--7 77777550 Pentametrocrinus japonicus.
38 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
(Specimens in the Hamburg Museum.)
Tropiometra andouini. Tropiometra encrinus. Tropiometra carinata.
Tropiometra picta.
Avitedon COANOIG. 2 see ce eee eae cee elem
Antedon levipinna, sp. NOV ....---.---------------------- Lamprometra protectus. Antedon xquipinna, sp. NOV.. ...-.---------------------- Lamprometra protectus. Antedon imparipinna, sp. NOV.....--.---------------------Lamprometra protectus. Antedon variipinna, sp. NOV..-.....---------------------- Amphimetra variipinna. Antedon crenulata, sp. NOV.........-----------+-+--------- Amphimetra crenulata. Antedon acuticirra, 8p. NOV. ....-...-.----..--.------------ Craspedometra acuticirra. Antedony LUdOMICL, BPs) DOVs eos ee ane alas a= ael= onelnle mini l= Craspedometra acuticirra. Antedon bipartipinna, sp. NOV...--------------------------Craspedometra acuticirra. (Antedon australis, nom. nov.) -...---.------------------- Craspedometra acuticirra, SACI ITLOENA GO Lane sa aan erene ae aie aa eie i= ee rem ie Comatula solaris. Actinometra robusta, 8p. NOV......---------+---+---+----- Comatula solaris. PA CIUTOM EUR UNC NO noe eee ols aia nia are ain a = aie inelm miele a= Comanthus parvicirra. Actinometra multiradiata. ©... <2. 5... 2c eceen + sce sees ses ee ee aged: : Capillaster multiradiata. Actinometra grandicalyx, sp. NOV..------------------------ Comantheria grandicalyx. Actinometra meyeri, Sp. NOV...----- Ba) Sees iets miners Comanthus annulata. SACHINOMERE DENNEN comm ee ecole tne Selo sia =~ Selenite sie Comanthus bennetti.
In the same year Prof. F. Jeffrey Bell invented a very ingenious, but unfortu- nately impracticable, scheme for the expression of the specific characters of the comatulids by means of so-called “specific formule,” and gave a list of all the species known to him with their specific formule attached; in this list he inserted the names of some undescribed species which had been obtained by the Alert in Australia, and he added an appendix describing Actinometra annulata (Comanthus annulata) from Cape York. Later in the same year he very briefly diagnosed a new form from the Straits of Magellan, Antedon magellanica (Florometra magell- anica), treating it as a variety of the arctic Heliometra glacialis.
It was in 1882 also that Greeff reported the occurrence at the island of Rolas in the Gulf of Guinea, near Sio Thomé, of a comatulid which he called Antedon rosa- cea, but which is probably the same as the species afterwards named by Hartlaub, from specimens obtained on the Ivory Coast, Antedon hupferi. This curious species is the west African representative of the Brazilian Antedon diibenii and of the European Antedon bifida. ;
Early in the following year Carpenter reviewed Bell’s system of formulation, pointed out numerous errors, and gave a revised list of all the species which he could determine; and Prof. Edmond Perrier diagnosed a new species of Eudiocrinus, E. atlanticus (Pentametrocrinus atlanticus). The genus Eudiocrinus was hitherto supposed to be peculiar to the Pacifie—being in fact named for the Pacific Ocean— and the discovery of a species in the Bay of Biscay was an occurrence of more than ordinary interest.
In 1883 also Prof. Percival de Loriol discussed the echinoderms of Mauritius, noting the occurrence there of Tropiometra carinata.
The report on the collections made by H. M. S. Alert in Australian and East African waters was published by Bell in 1884. In it certain species, badly in need
MONOGRAPH OF THE EXISTING CRINOIDS. 39
of redescription, were recorded with no data but the localities, others were given erroneous and misleading diagnoses, the species briareus was again, as in 1882, referred to “Antedon” instead of to ‘ Actinometra”’ where it belonged, and some of the names conferred in 1882 were shifted about and applied to quite different species.
The Australian species included in the Alert report are:
PAW LEO TEU ICOTIE ae ee eens eee aaa sao eal { Tropto TREO BE ON Oligometrides adeonzx.
Amphimetra milberti.
Antedon milberti.......-------2------- 2-2-2 e errr |smeknar discoidea. Oligometra carpentert.
Antedon pinniformis...-.--------++-------+22222 corr Oligometrides adeonz. Antedon carpenteri, 8p. MOV..--------------+222722220 ttt Oligometra carpentert.
Antedon pumila, sp. NOV..-------------+-+-2222 272000 tt tro (=Antedon loveni, 1882)-.---.---------+++222-22500000t7 Antedon bidens, sp. NOV. .-------------------+++ 222200000 -Oligometrides adeonz. Antedon loveni, sp. NOV..----------+-++---+-- 202000 (=Antedon insignis, 1882)..-----------+++--------7 20707777
\ Compsometra loveni.
}Colobometra perspinosa.
Antedon decipiens, sp. NOV. ----------------+-----7 7777 5r tt Amphimetra crenulata. Antedon reging, 8p. NOV. .----------------++----22 705575000 Lamprometra gyges. Antedon articulata.......---------+-------+2-25520e0r tert Liparometra articulata. Antedon gyges, SP. DOV- --------------- +++ +2222 2 2c Lamprometra gyges. Antedon irregularis, sp. NOV.---------------+--+---777770 7007 _Amphimetra crenulata. Antedon elegans, 8p. NOV.---------------+-------7700tr ttt Zygometra elegans. Antedon briareus, 8p. NOV. .-------------------22- 2050 r rte Comantheria briareus. Antedon microdiscus, 8p. NOV..---------------+-2-205 ttt ttre Zygometra microdiscus. Actinometra solaris...--.------------------+20 corr Comaiula solaris. Actinometra albonotata, sp. NOV. .----------------------77777> -Comatula solaris. Actinometra intermedia, sp. NOV. ---------------------77 77 77> Comatula solaris. Actinometra robusta...----------+------+ +020 r rrr Comatula solaris. Actinometra strota....------------------++--7 2222220 t Comatula solaris. Actinometra cumingii....-------+--+-+--+--27tr rr etrr Comanthus parvicirra. Actinometra coppingeri, 8p. NOV------------------- 7257057 Capillaster multiradiata. Actinometra jukesi...--------------7 70222 et trc rt Comatula rotalaria. Actinometra parvicirra..-..------------2+077-0orr nett Comanthus parvicirra. Actinometra alternans....------------------27070ccrt te Comantheria alternans. Actinometra paucicirra, 8p. DOV----------------2+520050 rrr Comatula rotalaria Actinometra multifida.....------------+-+72 cre ct rcrtn { Gominates WUE ee Comanthina schlegelit. 5; aie Comaster typica. Actinometra variabilis, sp. DOV. .-----------+++---77000 07> heal y ltifida, Actinometra, sp. jUV------------------ 7200 t tr Comatula pectinata.
The east African species included in the Alert report is: Actinomelnas Sp-s-/2= 2-2-2 s2s- 222 ot ee ae Comissia ignota.
In 1884 also P. H. Carpenter diagnosed his remarkable new genus Thaumato- crinus, which recently has been shown to be only the young of a species belonging to one section of his genus Promachocrinus, the section which was included by Minckert in 1905 in his new genus Decametrocrinus, over which name Thawmatocri- nus has, of course, priority.
In the same year that the Alert report was published P. H. Carpenter also pub- lished an account of the crinoids occurring between the Faeroe Islands and Gib-
40 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
raltar, mainly based upon the results of the work of the Porcupine expedition, and he also finished the monograph on the stalked crinoids which had been obtained by the Challenger. This latter is much more comprehensive in scope than is indicated by its title, for it includes a valuable discussion of the morphology both of the coma- tulids and of the stalked species.
In 1885 Bell published a list of the Australian crinoids which had been sent to the International Fisheries Exhibition in London. This list was published in New South Wales, and was the first contribution to the study of the recent crinoids, properly identified as such, to be printed in Australia.
Bell in 1887 reported upon a collection of echinoderms from the Andaman Islands, which included a single crinoid; this he refrained from naming, as the same species alsa occurred in a collection from the Mergui Archipelago that had been assigned to Carpenter for report. Carpenter later called it Antedon andersoni (Pontiometra andersoni).
In the year 1888 the great Challenger report was published, which, though based upon the Challenger collections, amounts to a complete and thorough mono- graph of the group; this work brought the knowledge of the comatulids up to date, and has ever since served as a foundation upon which authors have built.
The following classification of the comatulids was adopted by Carpenter in this volume:
“J. Crinoids with the calyx closed below by the enlarged top joint of the larval stem, which develops cirriand generally separates from the stem joints below it, so that the calyx is free. The basals may form a more or less complete ring on the exterior of the calyx, or be only represented by an internal rosette. Five or ten rays, either simple or more or less divided. The first axillary is the
second, or (very rarely) the first, joint above the calyx-radials. Definite interradial plates usually absent. The mouth central, except in one genus.
Family Comatulidx d’Orbigny. A. Centro-dorsal has no articular facet on its lower surface. a. Five rays. i. Mouth central or subcentral. Oral pinnules have no comb. Radials separated by interradials....................-..-..--- 1. Thaumatocrinus. Radials united laterally. 1. Basals persist as a closed ring. No pinnules on lower
prachiala 32.5 [5k se. eee ese sos ewan iae se ee te ACERT 2. Basal ring incomplete or invisible externally. Ee ive anma only. 5 «<<< 165 <isin ede ola dome osialosmeiele 3. Budiocrinus. EDS Tenvarms $3.55 oss Vessac tees cectececee aeeaee sae 4, Antedon. ii. Mouth excentric or marginal. Oral pinnules havea terminalcomb. 5. Actinometra. Dy SLOMTRYBr conccocccescsic ea ceaeere Beet ciaaee webingas sale cectceeeese cece 6. Promachocrinus.
B. Centro-dorsal has an articular facet below...................-.---+------- 7. Thiolliericrinus.”’
These genera contained in all 188 recent species, divided among them as follows:
PETAR ULLO PANS 36 os or i ois seg BE Le oe ele pT a el a a uy SRIBIECTETEUBS oa So 2s pines te So ioc Ape eee Cp oe eC Be oS a eee 3 TOLER OUINTNAS once owner ole oe ee rae ORL TER OE CREE Eee ee 5 PUN TL Na eee St Sle ay IRE SS reg eh ait 8: A ns SE oy 2 Bi te Ve 122 BCHNOMEING Bind o's ele - Bois Geen SS wole se he Ree Re 54 PrOovunchocranuss. 2. 2.25 de axe shee ase habee ee ae. ee 3
MONOGRAPH OF THE EXISTING CRINOIDS. Al
Carpenter did not subdivide the genera Antedon and Actinometra, but he arranged the species in a number of more or less well defined groups for the better appreciation of their differential specific characters (but not of their specific inter- relationships), as follows:
ANTEDON.
Series I. The two outer radials united by syzygy. (This includes only the “Elegans group”; Carpenter did not employ this name in the Challenger report, introducing it for the first time in his report on the comatulids of the Mergui Archipelago, 1889.)
Series II. The two outer radials articulated; 10 arms.
The radials and lower brachials have flattened sides; pinnule ambulacra
generally plated cat 2s\< - omenasios sme Sac ee seis weicioe sm paec c/cisas 1. “Basicurva group.” The rays not flattened laterally. Pinnule-ambulacra well plated....... 2. ‘‘Acela group.” The first two or three pairs of pinnules long and flagellate, with numerous
BOD Gt UO ay Oj OLIM GS mere eereeta tate aye teste tattet eter tsar) ay atestarerats clare 3. ““Eschrichti group.”’ The joints of the lowest pinnules, which are often long and slender, are
longer than wide, frequently very much so......-........-.-------- 4. “Tenella group.”
The first pair of pinnules is comparatively small, and their joints but little longer than wide; one or more of the second, third, and fourth pairs are longer and more massive, with stouter joints than their suc- (CEO ere ete te tet tate ete te olan lala teeter te et 5. ‘‘ Milberti group.”’
There are in addition six 10-armed species which Carpenter does not assign to any of the preceding groups.
Series III. Two articulated distichals. Bidistichate species with the radial axillaries and some of the following joints more or less wall-sided, and a well marked ambulacral skeleton onthe pinnules= see sero nemesis eae wee eles nie'= wtelats stars elereeerelste 6. “‘Spinifera group.” Bidistichate species with an unplated disk and no definite ambulacral skeleton. The sides of the lower brachials are scarcely, if at all, flattened. The first pinnule smaller than its successors.........-.-.- 7. ‘‘Palmata group.”’ Series IV. Three distichals, the first two articulated, the third axillary with a syzysy- Tridistichate species with plated ambulacra and the lower parts of the TN let HOMO LOL Ye ete apatite ote 8. “Granulifera group.” Tridistichate species with an unplated disk and no definite ambulacral skeleton; the bases of the raysare not flattened laterally............. 9. ‘“Savignyi group.”’
ACTINOMETRA.
Series I. The two outer radials and the two first brachials respectively united
by syzygy. BST NEAT Se at a aoa ott ree 1. “Solaris group.” Two distichals, united by syzygy.-.----------------------------------. 2. “Paucicirra group.”’ Three distichals, the axillary a syzygy-..-.------------.---.----.--.-- 3. “‘Typica group.” Series II. The two outer radials articulated; 10 arms...........--- ee ---- 4. “Echinoptera group.”
Series III. Two articulated distichals. Two articulated distichals. The palmars and subsequent series, when present, are of the same character; but the first two brachials are TINIE VARY ZY BY ee ee 3 ale = tae la tole e oti telat la laine ete 5. ee group.” Two articulated distichals; the first arm syzygy in the third baiecbaiee “‘Valida group.”’
79146°—Bull. 82—15——4
42 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
Series IV. Three distichals, the first two articulated, and the third axillary with a syzygy. Tridistichate species with a pinnule on the first brachial and a syzygy in
the second. The palmar and post-palmar series, when present, con-
sist of two joints, the first bearing a pinnule, and the second axillary
WAHL ABV RYE Vins aceon ee tee mete mee tee mater te oon ion imi=in 7. ‘‘Fimbriata group.” Tridistichate species, with a pinnule on the second brachial and a syzygy TTT PNG GIR 2 fata eee en oite siete eee here ni wate = Blom intel meal aie 8. ‘‘Parvicirra group.”
The following species of comatulids were considered as valid by Carpenter, and were included by him in the Challenger report:
THAUMATOCRINUS. Thaumatocrinus renovatus........--------------+--+---- Thaumatocrinus renovatus. ATELECRINUS. Atelacrinus OLanOVdes = 2 ext aalmntm ale sie wim nto farnn afm == alae Atelecrinus balanoides. ALelerranitie AYN oe acne eters oat toe nie) emia etalvia Atelecrinus wyvillii. VA TELECYATAUG CUDENIBES tet ete sin een ae oe ie isos cece Atelecrinus balanoides. EUDIOCRINUS. FAUOCHINUS TNOWISUS ws ao 5,5 a ace ome aioe ionic es mas oe Budiocrinus indivisus. POUMO CHINAS VATIONS «ca daa scan cae rscis sisiele'winte's nas ee Se Pentametrocrinus varians. TRACROCUTIUS RENUDEN «oo nino crete sain eee nee ie ae nae iain Pentametrocrinus semperi. Thin pon Gos Cee japonicus. alles ea i i a a Be Pentametrocrinus tuberculatus. OUCROCTAYUUG CULQVLRCIES crater a) aie mye feel ale stem ieee es Pentametrocrinus atlanticus. ANTEDON. Sertes I. AntedonsUChlans- 5-05 one ioe acne chen ceeeconces Zygometra elegans. AMtEdON TUL AGI AO. ote ewe erate fois eteie le eiae aja Zygometra microdiscus. ATited ON TRACT OUNSCIRS ae Schon ate alae ee ele ee eer Zygometra microdiscus. Series II. “ Basicurva group.” ANLEAON: LONGICNTO aaa causa laiat Ase Eoin om sia aR ee Asterometra longicirra. Antedon Malis arent coee ais caisinntoie Samer ee eee ee Aglaometra valida. ANTEAON WENO an 6 Ses a we soos ww ce a elem Binley eet Nore Aglaometra incerta. ANLEAON GTACHIS® aos fais ee Vere scien se ate nee ech Thalassometra pergracilis PA niledomy) ULSWANACTE = foro wloe v wre os cee eee eee Thalassometra lusitanica. PANLEGON UT CUITUMU ooo oc concn kno os Sos Ae Stiremetra breviradia. PAOLO BIRT T naan ania ee oe ee eee eae Stiremetra spinicirra. PATITENON (CUETO. 2 os camiee to ocean thee ae eee Stiremetra acutiradia. Antedon Giepinosa os . Se os ck tee oe a Thalassometra bispinosa. Antedon: ANP UINE = 3. ane Fo Sew ae aos Bae ee Lee Thalassometra latipinna. Aritedon Mulan: . 2c coos seemed <2. pees yee Thalassometra multispina. Antedon échinatac = set oes Sai aes 232 ee ne Thalassometra echinata. Antedon basicurva eels data a aonk ae mE Oey asic ane See eee Charitometra basicurva. WA SILECOVRMAONS Cc 2-228 )icheee ys eee E CIN WS StS eae Charitometra incisa.
ATBOOT MUDPT ORG 2 occ 8 7 Ate Ne sae Pu ee saeneeete Glyptometra tuberosa.
MONOGRAPH OF THE EXISTING
Antedon parvipinna Antedon flewilis......------------+-++2+-2222--2-22---- TA TiEEdOTNACILLEA LES amet awioin = oeiaie ee lelaleiei= = ==) =I Antedon denticulata Antedon pusilla
WAri ted OTC Lat eee ale oie le os tee oa areal =i == Antedon discoidea
“ Eschrichti group.”
Antedon eschrichti Antedon antarctica Antedon australis Antedon rhomboidea Antedon quadrata Antedon magellanica
{
“ Tenella group.”
Antedon phalangium
Antedon hystrix Antedon prolixa
{
|
Antedon tenella
Antedon exigua
Antedon alternata
Antedon rosacea
Antedon petasus Antedon diibeni
Antedon lineata Antedon remota Antedon longipinna Antedon tenuicirra Antedon lxvis Antedon hirsuta Antedon angustipinna Antedon abyssorum. . --------++---+++02rrtetttrt tree
Antedon abyssicola....---------++++++++ 222220007 707+ {
|
“ Milberti group.”
Antedon pinniformis Antedon serripinna Antedon carpentert
CRINOIDS. 438 Strotometra parvipinna.
Pachylometra flexilis.
Chlorometra aculeata.
Pecilometra acela. Calometra discoidea.
Heliometra glacialis. Solanometra antarctica. Solanometra antarctica. Florometra magellanica. Heliometra glacialis. Florometra magellanica.
Leptometra phalangium. Leptometra celtica. Hathrometra prolixa. Hathrometra prolixa. (All the smaller species belonging to the genus Hathrometra.) Hathrometra exigua. Thaumatometra alternata. Thaumatometra cypris. Trichometra persina. Antedon bifida. Antedon moroccana. Antedon hupferi. Antedon mediterranea. Antedon adriatica. Antedon petasus. Antedon diibenii. Tropiometra picta. Tsometra angustipinna, Thaumatometra remota. Thaumatometra longipinna. Thysanometra tenuicirra. Thaumatometra levis. Eumorphometra hirsuta. Tsometra angustipinna. Thaumatometra abyssorum. Bathymetra abyssicola.
Bathymetra carpenteri.
Amphimetra pinniformis. Oligometra serripinna. Oligometra carpenteri.
44 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
PA TALON PUMA. son coe emeee eam ote annie ce =e eee =a Compsometra loveni. Amphimetra milberti. Antaion milbertis..- csi eee nee =e. shee erat eeeee Amphimetra mélleri. Amphimetra discoidea. Antedon levissima 2.52 see he tee e none 5042 bane Amphimetra levissima. PAMIIACON: LESROL UAT = enema neee one a eel ee eee ee eee (?) Antadore perepinos@ ofan <= oe 2 ss aia tee ee Colobometra perspinosa. PAIGLOQOYY ONCE 208 ere emo eteene ercle oiet s ee Amphimetra anceps. Antedon VAVWPINNls sce s< coe ses om chine lima = sine <i cieee Amphimetra crenulata.
Tropiometra encrinus. Tropiometra carinata. ANLEAON: COMMA == os oo Se eee Hee a= eels eas saa Tropiometra indica. Tropiometra audouini. Tropiometra picta.
AnledOn: POTUCOUTE =< ae elms aoe = eee sess Jes ane sees Tridometra parvicirra. Antedon <n formas. 2s-5. de team scl-eese cis sehen ce ere Decametra informis. Antedon lovent 5s. et eee ee as oni s eee ese ESE Colobometra perspinosa.
The six following 10-armed species do not fall into any of the groups previously given:
Antedon! dens 722 oe ccs ste ae ie epee so ssa soe see Oligometrides adeonx. Antedon: CdeOn tren. ante cees oh nana wN oetiea nose Oligometrides adeonz. Antecon jist inirl see a ne ees ee te Amphimetra milberti. Arledon Galenouiee en sos toet Nees ee Se ae eens Balanometra balanoides. AUGON Ae feciemra = ser a ane ae ene sne eee Hypalometra defecta. MntedonAmPVn nace ae cio venetian as (?)
Six other 10-armed species are mentioned by name, but are not discussed; these are:
ARLEN OTMOAUI saat cen tees Yee alee eee Kee Analcidometra armata. Antadon: breviminnds nosaee eee 5.5 ee ne tes cesee a (?)
PANLEAOM COULNNONAE = mn Ske bye 5 sacs, 5. one see coes Zenometra columnaris. sAmtedoT CUben site eo Sees 8 ais so eee caine seein Antedon cubensis. Anitedon du plern® sextet ea Ae oo i Sas cea elena bes nie Horxometra duplex. pAntelOn uLgenN eet ~ bitte so ctoe Usa. ee eee) Coccometra hagenii.
Serigs III. “Spinifera group.”
anny macronema. Ptilometra miilleri.
Antedon quinquecostata.<. 2 ~..<i2-.0220000050ckesesecss Stenometra quinquecostata. PANLEROTY BINNS er One n sees ee ae 6 sas Sosa: oe eee Stylometra spinifera.
pA ntelon i diip les ieee, US Vee Nee 2 Le Se ee Oe ce aS Horxometra duplex. Anitetory iuetaritca Me ene Gah BI ee Thalassometra lusitanica. ATUELON fERblie 3-2 Se OTM Bete ONE Pachylometra flerilis. ANtedon pibisite. 223" - tee eee ce. een Pachylometra patula. PATRON TOUUBA Choco 2 are es oe, eae Pachylometra robusta. PATIBAON DON UAEM os. oo. 5 ee Crinometra granulifera. Antmion bre niving: <6, ibe een occ.) Crinometra brevipinna.
(eae compressa. Parametra granulifera.
MONOGRAPH OF THE EXISTING CRINOIDS. 45
“ Palmata group.”
PANLEAONINOANCH AS Pena ere efe ene ais inane 2 aie e as ate Cyllometra manca. PAM LECOTL CLS CU ONTIUS -teteteetatate) =e miele ee eae 2 ae Cyllometra disciformis. Antedon clemeénsmree mae saean nas ase dene a= se aetoa= = =a Amphimetra anceps. PAT LECOTU MON GULL eee ae oat o eee reesei ee omnis Stephanometra marginata. Anitedon tubercular ces Ss 2 Sa sae sinc dsc cies eee Stephanometra tuberculata. SANULEMO SDVCL LM ret see nein ini == ela ie= ee eels =a = ae ie Stephanometra spicata. PA TELECON TAU ae reso a= eran 2 Soci ae ae che sie eerie = Stephanometra indica. BA NtedOTePROCE Cla pho mera. 3\ op aia;n (o's xjaiaeeis wratat ae Nataia efela(as ==) Lamprometra protectus. CA mteloT CONG UT ENS a wear a = ola soo) ato ea Ee 1c eae Lamprometra protectus. PANLELOTNE GUY PUN etn sata l= a eos ae este aisle Lamprometra protectus. PAM LEUOTY LEWICUTO =< a12 15 hae aicioerate asta eae eee Lamprometra protectus. PAM TEMOTALIN DOTS PANG nee areola eee ere oe Lamprometra protectus. PAN TECON RENIN omer cnysie ise ees Saas Sei = Lamprometra gyges. Antedon gyges....--- a eR Seren Stee ete or Lamprometra gyges. PATULCOOTO TO UITUO LO aa meee ice totstaerotte eat tetas eto at Lamprometra palmate. PAN LEAON) DV EVUCUM EAL = aie erate ia rnise seas = meee sale sine = Lamprometra protectus. JU OUD SH ON Oeste trem arin a eadenabosadcnsscicoDeOenne Lamprometra similis. Antedon occultans. 25-82 e ae sere ce asec senes- se Lamprometra protectus. SAM LEMONY OUEICIIOLO ee ee ena oe eee eee ean a Inparometra articulata. WA TLLECOTTEG OLS aap ee eee ear ere relate im aletaier Iiparometra regalis. eAnitedon Clon gatas seq =) \asaerne = om aia aie )saseie oases ime > a Dichrometra flagellata. SA LedON LAG ella erasers eee iale neato tenia = Dichrometra flagellata. Antedonibimaculaton scp. 22 sao s ss cee ese senee + Dichrometra bimaculata. Series IV.
“Granulifera group.”
PATIL CAOTOT GO USCUCULY Ie a ena eels see ieee eae ae Pachylometra angusticalyz. PA TERCCOTU TILL QUES see teetal atelier ea Pachylometra inxqualis. PATILCAOTGR ONAL Yer. ae eee ee eerste ieee er Crinometra imbricata. PAntedons GStin Cle aere encase ea ee ieee ieee a= Pachylometra distincta. VAT LEAO TU NILLETS T7170 eae eet ate ea etal Thalassometra multispina. VAN ECON DONT ECLOa ets eee ee tater eee etre Crotalometra porrecta.
“ Savignyi group.”
PATHELON QTUGUSEN COLO == eas sate ae ee ee ea Adelometra angustiradia. PATE ONU TEU ITAU Cena tal- ta oleae eee ae ae ree Heterometra reynaudit.
PA TILEROTUEGWIGNY Uaee seers nee aie =n ie sisal Heterometra savignit. VAT LEMOTNCTICE DS se ee ios hola teresa ieler ata ale alia Amphimetra anceps.
a phimetra crenulata.
Antedon variipinna.........---+----+--+-+-+-++----- Amphimetra variipinna,
Antedon quinduplicava.......---.-.------------------ Heterometra quinduplicave.
PAN teHOT UCULUCUT Waa malate = ai lm =m alee =)= li Craspedometra acuticirra.
PAMTECOTIIULEO UIC Ieee eas ne ea = ieee ee aie ati Craspedometra acuticirra.
Antedon philiberti..........-.---------+--+--------=-- Amphimetra philiberti.
Antedon bipartipinna.......-.------------+-----++---- Craspedometra acuticirra. ACTINOMETRA.
“ Solaris group.”
Comatula purpurea. Comatula pectinata. VACHNOMEIRC SOLOING. smc sc eo cence eee See ile Canalo ow Comatula solaris. Actinometra brachiolata.......--------+--+--++---+---+> Comatulella brachiolata.
Actinometra pectinata.......-----+-------+-+-+---++--- {
46
BULLETIN 82, UNITED STATES NATIONAL MUSEUM,
“ Paucicirra group.”
Actinometra PQucicirrd.......sccceeeeeeeeeee sere en eeee= Comatula rotalaria.
Actinometra distincla: <<... - 20. 255. D2 geen cece nce tces Comaster distincta.
ActenO metry Cy Diu ee ote ole te sta mito etetw nc = ona Comaster typica. .
Actinometra NOv#-GuUine#.......2. 20-2002 e cence eee e ees Comaster novxguinex.
Actinometra multibrachiata..........--- nicl aroin Ao kere attenie Comaster multibrachiata. Series II.
“ Echinoptera group.”’
Actinometra echinoptera.............---+--+--+-+-+-------Comactinia echinoptera. Actrriometra pulchella=s.-9 eee sae oc <8 eax wa sonic ies Comactinia echinoptera. Actinometra blakei (nomen nudum).....-...--.--------- (?) Actinometra meridionalis..........-..------------------Comactinia meridionalis.
Series III.
“ Stelligera group.”’
Neocomatella alata. Aolinowietereppdcatin oe a ae eee ene Neocomatella atlantica.
Neocomatella europxa.
Palxocomatella difficilis. ACHNOMENG MACUL so62 3 foto sass eee Seen cee Comatella maculata. Achnometa slelligera-s 225-22 = sss Dae eee cence cee tee Comatella stelligera. Achinomet a NigTi=s.-S25. «0a o cede Seip en coee ok Comatella nigra.
“ Valida group.” ACtiNOMERGBLONG CED a0 in a aioe 3 ax Foo siamese eine wisi Comanthus parvicirra. ACUMOMEIN SUNDICL eis PE oon cing om on eee aeons ieee Comanthus parvicirra. ALCURTLOMEL OT OMIUIING arate nctere oaravoscinie 2 /-foinin c= ve CS Comanthus parvicirra. Achinometra MONAG enn kes ane sats oe soe yates eels Comanthus annulata. Series IV. “ Fimbriata group.”
Achnometa jUnUTIalas oo eae saa: else oes aie ee ee Capillaster multiradiata. ALENOMEECODDINGEN «= oe ctia = eee ee e OE Capillaster multiradiata. Actinometra borneensis........--..- nie ett Eni eer Capillaster multiradiata.
Capillaster marizx. Actinometra multiradiata..........----.200200002e000- Capillaster multiradiata.
Capillaster coccodistoma. Actinometra Senvond ccc: soc 2 eos SEN eee Capillaster sentosa. Arhnomnstra Winbatis os. 2% sons Ses oan oa ae eee ee Nemaster lineata. Actinometra discoidea (nomen nudum)...........-.-.--- (?)
“ Parvicirra group.”
Comanthus annulata. Achmomettn partici Tayssneacs 2c Sencar ons eee Comanthus Pees Comanthus parvicirra. : Comaster distincta. Actinometra quadrata..........- SaaS ate eee Comanthus parvicirra.
MONOGRAPH OF THE EXISTING CRINOIDS. 47
rActinometn a tmicho pled sary nae se) eye Cee see Se Comanthus trichoptera.
PACHINOMEN A JAPOTICI Ee nee eee ene an = es Comanthus japonica.
PACH NOME OG MUU lddeeereeeeeaee see ses see = ee acl Comaster multifida.
PA CHINO MEM ALUOTIC DUIS eee eeee ate yniy ae cio eie =e Jomaster multifida.
Actinometra grandicalyx.............-----------.-------Comantheria grandicalyz.
Actinometra alternans........-.---.-.---------.--------Comantheria alternans.
PA CLIN ONLEETOAU MATEUS nae Aas se es eree elaine eiresee elaine Comantheria briareus.
PACH NO MEL OOVVETROOO 2 mains aise wee = oes sing els = = Comantheria briareus.
Actinometia;MAQnificd= 2 = = ne = ae eee a= = fepisee Comantheria magnifica.
EA CLIN OMELTONOCLL Dano Netto arse Seta eee ee we Comaster belli.
A ChiNOMENG CUPLED =a a 22s aa ss = 311s 2,2) oso y=) sions se 3 5 Comanthina schlegelii.
VA CHINOMEM A MOUWIS se 2 ean ager etree iets Comanthina schlegelit.
Actinometra robustipinna..........-----.--------------Himerometra robustipinna.
Actinometra UiHtOnGlise. -seee oe cee ace sess oe Comanthus annulata.
Actinometraregauiss~sscent. eins seen = tie = See - esi Comanthina schlegelii.
Actinonelta Schiegelncus. 02-5 eaneas Jae sees oa 3 Comanthina schlegelii.
PA CEIMO MEN GD ENON sata mata mane eerie mame a) =intaie ial Comanthus bennett.
FACEINOMEL CI UCTEREC se rate Sat santa) erat aerate srerciel sl Comanthus bennetti. PROMACHOCRINUS.
Promachocrinus kerguelensis......-....---------------- Promachocrinus kerguelensis.
‘Promachocrinusiabyssorwin= 3. = - = - 2 eas oe eee = Thaumatocrinus renovatus.
IPT OM OCKOCRUTULS TILL CR tater eer te ete eee erase Thaumatocrinus nares.
Besides the systematic account of the various species, the Challenger report contains a vast amount of information on the morphology of ecrinoids, and an exhaustive discussion of the relation between the recent and the fossil species. Most of this, however, is included in the volume on the stalked crinoids published in 1884.
The myzostomes found upon the crinoids which were studied by Carpenter were, as previously noted, sent to Prof. Ludwig von Graff, who reported upon them in four papers (1877, 1883, 1884, and 1887) in which he included many manuscript names which had been furnished him by Carpenter and by Semper.
In the same year that the Challenger report was published Bell reported upon a small collection of crinoids which had been sent him by Mr. J. Bracebridge Wilson from Port Phillip, Victoria; among them were two forms which he described as new, under the names of Antedon wilsoni (Ptilometra macronema, juv.), and A. incommoda ( Compsometra incommoda).
In 1889 Professor Bell reported upon a collection of echinoderms made at Tuticorin, in the Madras presidency, by Mr. Edgar Thurston, and also upon some echinoderms obtained off the southwest coast of Ireland. Mr. James A. Grieg also recorded some crinoids which had been dredged in Vestlandske Fjord.
Professor Bell had received some additional examples of the species which he had described in the Alert report as Antedon pumila, and had discovered that the first pinnule was the longest, and not short as he had stated, he having been misled by the broken condition of the original specimens. His Antedon incommoda was supposed to differ from the earlier A. pumila through the greater length of the first pinnule, but this difference being now shown to be nonexistent, he now relegated the former to the synonymy of the latter, though, curiously enough, the two are well differentiated on other characters.
48 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
The chief paper of the year was Carpenter's account of the comatulids of the Mergui Archipelago, based upon a collection made by Mr. John Anderson. In this paper the following comatulids are noticed:
ATLAAOT CLI OMS ae te eraee ata ee een aa este arete pinta ania) ate aVarole\e wiele = ete Zygometra comata, Antedon andéreonicBPRUOVeee: oc. ec seems tic ei. so nanisateclseieers Pontiometra andersoni. Antec logis eet ee ancien oa cele isis a ninis nis aeiee ete ates Amphimetra milberti. ANLECOTE SOUCOLO are en iw aiarsols Soicte swine = ets (eer Stephanometra spicata. ANLedOM CONFUNG ENS a ane lam a into aloe elrninle oe = ine esetent~ Lamprometra protectus. eA Chinometra OLA Ape NOV =e socom <= (sie ams ae Saigo = miele Comatella stelligera.
The difficulties attending the use of the various specific groups instituted by Carpenter were first brought to notice by this article, for he referred Actinometra notata to the ‘ Paucicirra group” in which he described it as a new species near Act. paucicirra; it really belongs in the “Stelligera group,” and had Carpenter placed it here he would have seen at once that it is the same as the Actinometra stelligera described at great length in the Challenger report. It is in this paper that Carpenter gives to the Series I of Antedon the name of “‘ Elegans group;”’ at first he had considered the single species represented in the collection as new, and when he sent some myzostomes which he found upon it to Professor von Graff he gave him the name of Antedon comata for it. Later he decided that it was the same as the Australian species described by Bell, and suppressed the name. It has been recently shown, however, that his first decision was correct.
Aside from some papers of purely local interest, the chief contribution in 1890 was the preliminary paper by Dr. Clemens Hartlaub describing a large number of new forms from the Indian Ocean. The complete work on the littoral comatulid fauna of the “Indian Archipelago” appeared in 1891; it is exhaustive in its treat- ment, and, besides most excellent descriptions (accompanied by figures) of all the new species, includes redescriptions of many imperfectly known forms, taken from the types. During the preparation of this work Hartlaub was in constant com- munication with Dr. P. H. Carpenter, to whom he referred several of the more difficult problems; it thus comes to have an additional authoritativeness, as it embodies to a certain extent conclusions reached by Carpenter from a study of material upon which he never published. Hartlaub identified many of Liitken’s nomina nuda, placing them correctly in the synonymy.
The species considered by Hartlaub are:
AMIedOn: DENGUIENSS |AD (DOW ashe). eee ee = eee eae Heterometra bengalensis. ANLEdON UITENAT RD TOV sate ek: one Semen oe ee Himerometra martensi. BATION Kr Ae DEANAS BD. NOV ieaeiecoc ee ene oe te ee ee Himerometra robustipinna. Antedon Grockst, Bp: NOV: 5254-5 0 NN Ce ee es Amphimetra variipinna. SANLECON A AiVAS; (SP. NOV eo oe OS eee. See ee Heterometra affinis.
Antedonmematodon: ap! mov 825.255: Sco. ado once eee Amphimetra nematodon.
Antedon ludovici [ee ee amboinx. Craspedometra acuticirra.
ANUOGON CTASMPINNG, BP. NOV... ---6-- 1-52... nn ee Himerometra robustipivna. ANILEGOW CUD IC Bp: NOV: oe cue eee ee Petasometra clarx. Antedow bella, Spi novils. ios.) ak J 23-bit Cenometra bella.
Antedon bella, var. brunnea, var. nov ...................-- Cenometra brunnea.
Antedon kiinsingert, Bp: NOV... .c sss: 0se- us case eee Lamprometra palmata.
MONOGRAPH OF THE EXISTING CRINOIDS. 49
PAmtedon finschtixSpsnON eee as = css -istaetahias Aaisicis sae caclae seni Oxymetra finschii. Antedon palmata......----.------ SEU E SRO CBP ABER ESaeeerisse lead UE Lamprometra protectus. PAMLEHON CRINALEGNAD DOVE Rime ets aoe eC eas en's oeiccce ae ee oe Oxymetra erinacea. Antedon tenuipinna, sp. noy ...---.---- ee eel ee Stephanometra tenuipinna. ‘Antedon oxyacantna,| Sp; NOV)..--2 22.00) 02 02sec eee e esses Stephanometra oxyacantha. Antedon monacantha, sp. Nov ...-.-....---------------------Stephanometra monacantha. PANLCAONSPANIMINTA; BP. WOVe ae a aise aia) = lacie = ot Stephanometra spinipinna. SAALELOTUUME OTE UT IU wares afer a Sle ae aici aera) = oS ania Lamprometra protectus. TAmicdonitenera:; BD. NOV = -5- sas = 5 ais ce sam mimes 4s slomis ease Lamprometra gyges. Amt edOnHOTEVtCUNENLD)e aire ots eee ee steee = sais =e 2 .---Lamprometra protectus. PAMEEMOTN CLOTHS OL atom sew =e tna een er aS ate elmo eee ten : tAmtedon flagellata® 2 =tese. cout ch oe soe a toss eee ae | Dichrometra Jegeliate. PATCH OTN CON EN OAS): hLOVianinne neem ee rn SP arse els ola) state te Cosmiometra conifera. PAM LEAOTITACT OTLEN ate aaa ata ae ent aa are ee a Ptilometra miilleri. PANTEAOTY OTULCT BONY oso wiac om Salem eee = eae Soe ee ae Pontiometra andersoni. Amphimetra molleri. pAntedon milberty. 2c eto ee ctte alent San dese sacle se 3 [mph milberti. Amphimetra discoidea. PAMECAOMSCLIADITULnememem eee eo sere eerie eee ee sete Oligometra serripinna. PAriLedOn GAMONACH, BPs) MOV gars clases wana mia nies 2 =| t= =f /eim ae ay Oligometra japonica. : Colobometra vepretum. Antadon perspinnen cu 4222-10: Ub satd0-b4 2..gaese 2a «262 Colokontid eeaees PATtEdONI OTA VSD a NOVmes ent eee en a aeiaaae ne eee eee ent Tropiometra afra. Antedon hupfert, 8p. NOVse22-so2s2-0- 20 ta oe sass ceeseee Antedon hupferi. Amtedon Nand (Sp ONp ssn eree ts seni 2s secs nes See ee eee Iridometra nana. PA CEI OTIVEL TO CVU CTCL eee ene ee oo it eal Comantheria briareus.
FA CEULOMLELT Oy POTN CUT Oalctnrs states clatca ciao aaain' ws oie ae erere te eos Comanthus samoana.
| Comanthina schlegelit. Comanthus parvicirra.
PA CLIMOMEN.O TEQUILA tet eee ae See oh) tae eee eee Comanthina schlegelit. Actinometra coppingeri......------------------------------- Capillaster multiradiata. Actinometra macrobrachius, sp. NOV ....--.--.------------- Capillaster macrobrachius. Actinomelra fim Urniatare= seco cere ae ee ees Capillaster multiradiata. ‘Actinometra muliwadiatGecees == tee re nee ane Capillaster multiradiata. PA CENTIO NICU O) SLELING ET (ne aaa ea ea Comatella stelligera.
BA CEYTVOTIVEUT CTU CTLL LU fo tere ao tate eee ol ee Comatella maculata. CALL UTIOTIVEET CU PTEL LCI Be UL Cee ele ee aoa ete lel ene Comatella maculata.
PA CEETOMELT A SOLONLS meas ele ee ae ee eee ler Comatula solaris. Actinometra pectinata....- - See eee eee eee eee Comatula pectinata.
PA CHINOMEMUUTACRIO LALO = aa a ayaa niete ieleee ee es ee es Comatulella brachiolata. PA CINTIGTICEET OEY PUI a= meme ote eae ea altel tated rat Comaster typica. Actimometra gracilis, Sp. NOY ---------------------2e+-e5e- Comaster gracilis.
In addition to the new species indicated above, Hartlaub described in the pre- liminary paper Antedon lepida, A. protecta, and A. amboinensis, which he later re- ferred to Antedon palmata, A. imparipinna, and A. brevicuneata, respectively; all three of them are synonyms of Lamprometra protectus.
Dr. P. H. Carpenter in 1891 published a paper on a small collection of crinoids from Madeira, in which he discussed the vexed question of the synonymy of the common European species, combining as a single form all the species which are now understood as constituting the genus Antedon; and Canon Norman wrote a short note in which he called attention to the fact that Actinometra, as used by Carpenter
50 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
in the Challenger report, is clearly preoccupied by the Comaster of Agassiz. The perversion by Miiller of this latter name is explained, and for Comaster, as used by Miller (that is, with the type Comatula multiradiata Goldfuss, not Lamarck—Alecto bennetti Miller), he suggested the term Goldfussia, which, however, was promptly shown by Dr. F. A. Bather to be preoccupied and therefore unavailable.
The work of the two French steamers, the 7’ravailleur and the Talisman, had resulted in the discovery of many interesting crinoids off the coast of southern Europe and northwestern Africa. Scattered references to these are found in the writings of E. Perrier, Captain Parfait, de Folin, and of the Marquis de F ilhol, but they are mostly very indefinite and unsatisfactory. Interest in these crinoids appears to have soon died out, and no detailed report upon them has as yet been published.
In 1892 Professor Bell recorded some crinoids which had been dredged off the west coast of Ireland, and described a new species from Mauritius, Antedon emen- datriz (Cenometra emendatriz) which is difficult to understand owing to the inade- quateness of the description and to the lack of correlation between the description and the figures. In the same year he published a useful epitome of the knowledge in regard to the British comatulids. The account of the comatulids which had been collected by the Norwegian North Atlantic Expedition, by Prof. D. C. Dan- ielssen, also appeared in this year, as well as a list of Norwegian species, by Miss L. Buckley, from the dredgings of the steam yacht Argo.
In 1893 Professor Bell reported upon a small collection of crinoids from the Sahul Bank, north of Australia, describing one new species, Antedon wood-masoni (Cosmiometra woodmasoni).
In 1894 de Loriol again recorded Tropiometra carinata from Mauritius; Prof. Georg Pfeffer recorded some species from east Spitzbergen; Mr. Edgar Thurston recorded a number of forms from various localities in southeastern India, the identifications having been furnished by Professor Bell, and Professor Bell published an account of the crinoids of Macclesfield Bank, near the Philippines, adding to it lists of the species known from northwestern Australia and from the Arafura and Banda Seas. The crinoids he gives are:
MaAccLEsFIELD BANK.
Eudiocrinus granulatus, sp. nov.................-.------- Budiocrinus indivisus. ANIACON: COMNOLG 1 oa wetcle eh oe ins cee see eee ee Oligometra serripinna. ANFEU OT REDMON x see oe oni od 2 oly Sete ee eee Stephanometra tuberculata. AMLEdOn: MOT MALT, AP: NOVoc sea cae... eee See ea ee Himerometra robustipinna. Antedon bassett-smithi, sp, NOV........--..--2..22-22+---- Comatella stelligera. Antedon vicaria, sp. nov........ So aentcta ete alate Ie eT RE Mariametra vicaria. Antedon brevicirra, sp. nov......- Pain c tear Saree ...-Comaster distincta. Antedon flavomaculata, sp. nOV.........2.0+-ec---eeeceess Stephanometra monacantha. ANTEAOTE MOO HD eNO. << ons He inate ee cl oot es ee Lamprometra protectus. PANLOGOT I ieLOt, BD. NOY 22), civ noose oes ete eee (?) OLE LOCOS DUO «<5 oud cae Le ean ko ee ee Mariametra vicaria. wcesnometrafimbriata: 2A the eee er S20 Capillaster multiradiata. ACHMNOM CH POTTIONT GW 32250 SIE ee ONS 5. ce Comanthus parvicirra.
LACUMMON Et U DOTNEacinc sk Ee ee eee. Ls ee Comanthus bennetti.
MONOGRAPH OF THE EXISTING CRINOIDS. 51
Actinometra simplex.--.-.--------+---+------- ttt tree Comatella maculata. Actinometra duplex..---------------------2 2000 e ttre Comanthina schlegelii. Actinometra maculata.....--------------+--+-+-200cr ctr Comatella stelligera. Actinometra rotalaria...-------------------++---22tttt Comanthus parvicirra. Actinometra regalis....---.----------+---+-2+7227 77700777 Comaster multibrachiata. Actinometra peregrina, 8p. NOV-------------------- +77 777- Comissia peregrina.
Nortuwest AUSTRALIA. er Amphi iscoidea. Antedon milberti { caropliemiclra Cae
Oligometra carpenteri.
Antedon serripinna...---------+--------2222-02 rrr Oligometra carpenteri. Antedon variipinna...--------------------+22-rtt rrr Amphimetra crenulata. Antedon, sp. (‘‘near macronema”) ..-------++++-----277777> Cenometra cornuta. : 2 Comat ti : Actinometra pectinata......-----+-------+222207 rrr { Cmte peas Comatula purpurea. Actinometra nobilis.......------------------2-007 ttt Comaster belli. Actinometra paucicirra...-.------------+-+2 20222 t rrr Comatula rotalaria. Actinometra parvicirra {eee Ree: eee ace eam Lee aT Sy a Comantheria briareus. Actinometra variabilis......---------------++++20ctrttctte Comanthus parvicirra. A : Comaster typica. Actinometra multifida.....--------------++---002 terre { : ‘ jnometra mullyfi Comanthina belli. Actinometra multiradiata...---.---------------++---7-7- .. Capillaster multiradiata. ARAFURA AND BANDA SEAS. Actinometra maculata...-------------+------+22 220 cr ttre Comatella maculata.
In 1895 Dr. Clemens Hartlaub published a paper on some comatulids from the Bay of Panama, the first definitely known from the eastern Pacific, announcing the important discovery of Florometra (Carpenter’s ““Eschrichti group” of Ante- don) within the tropics, and extending the known range of one species of that genus (Florometra magellanica) from the Straits of Magellan to Panama; at the same time he described a new species of Florometra from Panama, and three spe- cies of other genera from the Galapagos Islands; in an appendix he described a new Lamprometra from Gaspard Strait, between Banka and Billeton. The species mentioned by him are:
Antedon agassizii, sp. NOV---------+-+--+22222 22205 r rrr Thalassometra agassizit. Antedon rhomboidea..-------------------- 77 t2 ttt Florometra magellanica. Antedon tanneri, sp. DOV..----------------22-72 0 rrr Florometra tanneri. Antedon parvula, sp. NOV.---------++----222552ttrt rts Thaumatometra parvula. Antedon bigradata, Sp. DOV.-------+---------27 222555 t ttt Psathyrometra bigradata. Vinedon Spi ie = eee 2a ae acer ae es ee eo ae Trichometra, sp. Wraicdon BD meee sectese sence er = ccc reeecr cenaas Psathyrometra, sp. Antedon subtilis, sp. DOV. ---------------+2---2000r tr Lamprometra subtilis.
In this paper Hartlaub suggests the following arrangement of the comatulids:
I. Series I. Species with plated ambulacra: (a) The two outer radials articulated.
Menara eens ewes = hee sewer cen scare cia stan le CU Oes OUP: Acela group.
tcaaayG Hietaralir lS Ueto eeeee rae so be don uc ome pag OCe EC Ses ““Spinifera group.”
Mhredilistichals:.-.-+.<-s-----s0-=-ene~ oma 2 2s 2 ooo Snes “Granulifera group.”
(6) The two outer radials united by 8yZygy--------------+---2°+2-7-7> “ Blegans group.”
52 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
II. Series II. Species with unplated ambulacra: “ Eschrichti group.”
Went gums? sss. soe ee eee ee eo ae se encore a eee “ Milberti group.” ‘ Tenella group.”
Two Gistichals-seenes sae ee > oman ome nee ate winston“ eae ‘* Palmata group.”
Three distichale.tc<2 he - so -cs- > See ae eae - ~ ne cena elie nis' a eelasle = “Savignyt group.”
In this year also Hara described Antedon macrodiscus (Tropiometra macrodiscus) from Japan, at the same time mentioning the fact that Comanthus japonica is abundant at Misaki. Prof. E. von Marenzeller gave a detailed account of the occurrence of comatulids in the eastern Mediterranean; and Prof. René Keehler described a new form, Antedon flava (Crotalometra flava) from the dredgings of the French steamer Caudan, and in addition recorded a number of species from Amboina. In this last paper he records Comanthus bennetti under the name of Actinometra robustipinna, being unaware that the type-specimen of the latter is an endocyclic example, representing a species in the “‘Savignyi group” of Carpenter.
In 1898 Prof. Ludwig Déderlein published the results of his study of a small collection of comatulids from Amboina and Thursday Island; the species noted by him were the following:
ANLEMON ELEQUNS = coe neice ee new csioe oe eee seer ee Zygometra elegans. VANTECOTS INNCTOCIECUS i= ate tie ne ao ctears ea wee sinew eine or Zygometra microdiscus. BAMLENOTL OLGEIS hae oie eee aie eS ee ee eee Oligometrides adeonz. PAN tedON LAOUsCi sme ene te one fo a eae neem laintatar Craspedometra acuticirra. VANTECOTE AT DORN onns =e oer eee e eee Lamprometra protectus. VA CHET ONIVEERG SPECTROM tam aan 3 wolfe eee ie eel Comatula pectinata, ACHNOMEHCSOUMNE nec = oman a<,~ 32 252 eee see eee ee a Comatula solaris. SACEETLOMIERNEL PROVCUON Tila ae naam ae ae tale ale a alsie = nieai i Comatula rotalaria. itinitie ODER eo ee 55 oh Sten Ab he Neon rane. Comaster belli.
, as Comanthus annulata, DA CONTEOILOCINL PION URRY Teeter siete re oe ree | Comanthus parvicirra. SACRO TEQAUS = 2 cone ce eosin esis he ~ see eee ee sce Comanthina schlegelit.
In the following year Prof. Hubert Ludwig discussed the crinoid fauna of Zan- zibar, adding to the species already known from the region Antedon flagellata (Dich- rometra flagellata, var. afra), and recapitulating the previous records of others. At the same time he published a paper on the crinoids of the Magellanic region, taking the opportunity to compare the arctic and the antarctic faunas. Professor Bell in the same year recorded the echinoderms which had been obtained by Mr. J. Stan- ley Gardiner at Rotuma and Funafuti; there was only one comatulid (Comatella maculata) among them. He also published a list of the species which were obtained by Dr. Arthur Willey during his expedition to the Pacific in search of the eggs of the pearly nautilus. The species mentioned in this latter paper are:
EANIEPL OTE NOLO» oes fimo ee eee ee oR ee ee CEE Lamprometra protectus. Antedon tuberculata.............--.----- So ahe Pb eee Stephanometra tuberculata. PALA ONLEH MOT ONGICN i =) 2 <5 ose an es stay oe te eee Comanthus bennetti. PAPEMLEN TONIC OM UIRS « oa 3 tata en, aaa, pee
Comaster gracilis.
PLCEMLONIORNE UCTHAREG oo oe Sa Se Bn Sc Comanthus bennetti. PLCUEOTCN ONTRE ACU TO) << occ cise oes nics le eee Comanthus parvicirra,
MONOGRAPH OF THE EXISTING CRINOIDS. 53
Prof. Georg Pfeffer in 1900 published a list of the comatulids which had been obtained at Ternate by Prof. W. Kiikenthal, and Prof. Percival de Loriol described Antedon déderleini (Dichrometra déderleini) from Japan.
It was in 1900 also that Prof. Carl Chun brought out his interesting semi- popular account of the cruise of the German steamer Valdivia, in which he figures a new species of Eudiocrinus (Pentametrocrinus) which was dredged off the coast of Somaliland, thus extending the known range of the genus in the Indian Ocean from the eastern portion of the Bay of Bengal, whence a specimen had been recorded, without a specific name, by Wood-Mason and Alcock in 1891.
The only paper of general interest in 1901 was Prof. Hubert Lyman Clark’s memoir on the echinoderms which had been collected by the Bureau of Fisheries steamer Fish Hawk about the shores of Porto Rico. In this paper he mentions the following species:
PAMLEMON NAGEN TD x02 tae nla niai= clea ewe o= ena ores Sone ln ce ore Coccometra nigrolineata. PA CLUTLONLELN, Cn TILEDULO TUL LLB S eratctalas ae etalaratte set atel et aie eenet = t=)n le ate) ators oe Comactinia echinoptera. PA CHINOTEN A TU DIQUIL OSC tare atatata stata siaiele alosiai=ieisin cise esi se Comactinia echinoptera.
In the year 1904 Mr. Frank Springer described Actinometra iowensis (Nemaster dowensis) which had been obtained in three feet of water on the Florida reefs; and for the first time described covering plates, comparable to those seen in many of the endocyclic forms, in a comasterid. In the same year Professor Bell published a list of the comatulids which had been collected by Mr. J. Stanley Gardiner in the Maldive and Laccadive archipelagoes, noting the following:
Amphimetra producta.
FANUEAON LE VSSUMN Dana cses cee s ace ceecaaeciece sane esac ese = Amphimetra molleri. Decametra taprobanes.
Decametra mobiusi.
PANLECONNMLL DET Ela eee eee eies a sene eee see ete esses Amphimetra producta. -| Amphimetra molleri.
PANLELON PUL ans sense ccm seiaceee ncn son eae sents ee seelele Himerometra sol. LANLECONANAICHS sara) sans ee sa Sees Se sine eee ee Comaster gracilis. VATHECOTO UAIUPINNA. Seen seers ae eee ae ee see Stee aera ? PA CHUNONTEN Qily DUCH = Sekpay isms SUP foes =) oats eee te eke Comanthina schlegelit. PACH NOMEN CifUIMNOTUALG sass o= we wile eels pale iia sea eer Capillaster multiradiata. Actinometra multiradiata........-..--.-.-------- “ise ose ts Capillaster multiradiata. JOA OCI ARONA MIO Win ccacesbagsasdeds ssesaeesddkooseoseacoke Capillaster sentosa. PACEUTLOMVELT CL TIVACULOL Ga oe ean ine oan nee ieee ee eee Stephanometra indica.
In a paper on the echinoderms of East Greenland, published in 1904, Dr. Theodor Mortensen calls attention to the presence of covering plates along the ambulacra of Antedon eschrichtii (Heliometra glacialis), and suggests that ae systematic characters may be found in the structure of the outer pinnules of the comatulid arms, which have hitherto been quite neglected from a systematic stand- point. In the same year Mr. Herbert Clifton Chadwick published a list of the comatulids which had been collected by Prof. W. A. Herdman at Ceylon during his investigations of the pearl oyster fisheries about that island. The species recorded by Chadwick are:
DATULE CLOTS CNTY TIVE et erate aol a= a ole aati ai Oligometra serripinna. VAG ITECE TE IUNCOET Leta te = aaa alae alo ete eae alas shelton fa teed Amphimetra milberti.
54 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
AN LAROTT COTANGUD = 0 las ete wos Ss toile ho Sine setae ale Tropiometra indica. Aritedon marqvnatd..<2..-- a. 29-2 wane eee eee elm Stephanometra marginata. SANIOUON SIUNCL: «<< tos cena Seals a aoe eee eine pial Stephanometra indica. PATTEM OGL = woe nin «amen eee ae ee eee ee ei i= Cenometra herdmani. Antedon okellt, Bp NOV) << = <0 62 = sie ne oe alee ernie Lamprometra protectus. Am edON FEY NGUGI ace nee mee ioe Sern aie eee tama sie eer es Heterometra reynaudii. ANEMONE GNCEDE: fon on mw snide o> - rin tla el Heterometra bengalensis. AntedOn VOTUPUNNG= «monn oem cnn == = nwo oem onan Heterometra reynaudii. SACEUNOMAING MOTAUE alana eee ole einle a= laatete ental Comatella stelligera. Actinometra multwradiata:-.. =~ - ------ 20+ - 5-9-2 2-2 Capillaster multiradiata. Comanthus annulata. AchinOmie ia pane Nernst leas sm stale aie oe eae tea tates eet Comanthus parvicirra.
Comissia chadwickt. In 1905 Professor Bell recorded four species of comatulids from South Africa,
three of which he described as new, all in ‘‘groups’’ widely different from those in which they belong; the four species are:
Antedon capensts, SP. DOV <---..cc0- oe nono - nano - ans =e Tropiometra carinata. PAM TOOONNECIOIEN BDO DOM ese ee sre a eee se Pachylometra sclateri. SATILEQOTS INOAGTICU TAs) KPa OV mniatnlnie ain oa a = feminine Crotalometra magnicirra. PA CIMLONELT Os DON TACIT seen oie iettel eiste ee epee eee nice Comanthus wahlbergii.
In 1905 also was published Wilhelm Minckert’s important and instructive treatise on autotomy and arm regeneration, with especial reference to the syzygy; in this he proposed a new ‘‘group,” the “Brevipinna group,” to receive species from the “Basicurva,”’ “Spinifera,” and “Granulifera”’ groups of Carpenter in which the IlBr series are either 2 or 4 (3+4) indiscriminately; but he evidently had a very hazy idea of the specific interrelationships of the forms within the group, as his group type comprises at least four distinct species. In another paper published at the same time he very rightly splits Carpenter’s genus Promachocrinus into two components (Promachocrinus and Decametrocrinus), but very illogically creates the family Decametrocrinide for their reception, or as an equivalent to the old genus Promachocrinus, his final arrangement being little, if any, in advance of that of Carpenter; a new species of Promachocrinus as restricted (P. vanhéffenianus; a synonym of P. kerguelensis) is described, and the suggestion is made that the comatulids be recognized as a distinct order under the name of Eleutherocrinoidea (having nothing to do with the pentremite genus Eleutherocrinus), the stalked crinoids to be considered as representing another order, the Stylocrinoidea.
In 1907 Dr. Hubert Lyman Clark recorded two comatulids which had been obtained by Mr. Alan Owston off southern Japan and given by him to Mr. Thomas Barbour; these were: Tropiometra macrodiscus and Cyllometra manca (C. albo- purpurea).
In 1908 Mr. Chadwick published an account of a collection of comatulids brought together by Mr. Cyril Crossland during Professor Herdman’s biological survey of the Sudanese Red Sea; in this paper six species are listed, as follows:
Antedon serri pinna ea as ee Ete ne et aha ed ef Prometra chadwicki . PANMBQOTL ANTONIN. «Sosa cee gee is ee ee Tridometra xgyptica. Antedon MOLINA ss oa ona cose sane elm inn eee ew an ce eelate ?Stephanometra marginata. Antedon imparipinna........ pale a ae a lala eC ne Lamprometra palmata.
PATS ORLON, TRELLIS «rch sao Ove) See hte cine oe eS OE Lamprometra palmata.
LANOR OD EOVIONY tec tc nk Se ee, Anne, Heterometra savignii.
MONOGRAPH OF THE EXISTING CRINOIDS. 55
Professor Bell in 1909 reported upon a collection of echinoderms made by the Perey Sladen Trust expedition under the direction of Prof. J. Stanley Gardiner; the species he records are:
PA CELLOMEL.G) LULU LOGO td eae nee mera rye tence sens select salts Comatella maculata.
PA itedOny COU Usama ataretss see elleetonlsaeieice sic oes cnc nees ?Cosmiometra gardineri. AMCEAOT PAUL eos a eralatalatsizixt= nis (elsia}=ieinis,=\=|s\cisieleicie <s osiesine o Stephanometra indica. PAM LEAO TY S DUCA ettate o ela stotere le iatnieine eile aeioc te eases aciaseree Cenometra emendatriz.
In 1909, also, Professor Kcehler summarized, in a magnificent monograph, the results of the researches of the Princesse-Alice; in this eight comatulids are included, as follows:
PAMLCAOTESCHTICN UA Stas ie steer a hy eee ee ee ae Heliometra glacialis. A OBIRT OTR E Des een as oseleeiaoe asco Sccue cue ee aecs Thalassometra lusitanica. PAN LEdON OMUSEC: BD NOV aaeane eee Sete tne Smee ise ee Thalassometra omissa. Antedonphalanguunys aoa et see se ose ae eee ee Leptometra celiicn. ‘ Leptometra phalangium. CANLCAONDTOULE me peee caciae cl eee eine aa ecn neces oer Hathrometra prolixa, Amtedory MOSGCER: tasraras-faiarasta Seta Ae se etee lela o= seins crays Scie Antedon bifi da. Antedon mediterranea. Antedon tenella. -- 22-2 an = ee ene ae Hathrometra, sp.
PEO UOCKUNILS CLLR TULLCTES Seen te ae ae Pentametrocrinus atlanticus.
In 1910 Professor Kehler and M. C. Vaney published a preliminary note upon the crinoids collected by the French steamers Travailleur and Talisman, and M. Vaney described a new species of Promachocrinus (P. joubini) from the collections of the Pourquoi Pas? under Dr. Jean Charcot.
Beginning in 1907 the present author published a number of papers on the Cri- noidea, describing new forms, suggesting new interpretations for various morpho- logical and anatomical structures, and developing an entirely new scheme of classi- fication which it was believed would be more satisfactory than any of the schemes previously employed. These papers are all preliminary and more or less incomplete expositions of the matter presented in the present memoir, and it has, therefore, not seemed necessary to review them in this connection; but an account of the development by the author of each of the systematic units herein used, showing the steps by which it has been brought into its present form, is included under each of the systematic headings.
A study of these preliminary papers’ shows numerous misconceptions of sys- tematic and morphological affinities and errors of other kinds, especially among the earlier ones. These were chiefly the result of lack of material and necessary dependence upon insufficiently detailed descriptions and figures. It is easy for the man who does nothing to avoid making errors; but activity of any kind necessi- tates occasional mistakes. No thorough revision or comprehensive work of any kind was ever done without a similar history, and the author feels confident that his errors will be found to be no more numerous nor more serious than those of his predecessors
56 BULLETIN 82, UNITED STATES NATIONAL MUSEUM. HISTORY OF THE INTENSIVE WORK UPON THE COMATULIDS.
The preceding sketch shows the gradual development of the systematic side of the study of the comatulids from the first beginnings up to the present day; but beside this constructive work a very considerable amount of intensive work has been done. This intensive work, whereby our knowledge of single species, but not of the group as a whole, has been advanced, has been mainly confined to mul- tiplying records of locality within restricted areas.
As might be expected, Antedon bifida is the chief species concerned; but it is rather strange that out of the very numerous records published of the capture of this form, by far the greater part are in English journals. Antedon petasus has also come in for a fair share of attention, but we are rather surprised at the lack of in- terest which has been displayed in regard to A. mediterranea. Known from the vicinity of Naples so long ago as 1592, it has been repeatedly recorded from that district, although other locality records are very few; we do not understand it nearly so well as we do Antedon bifida in spite of the fact that we have known it for more more than 100 years longer. Antedon adriatica, although reported as abundant in the Adriatic Sea, by Olivi, as far back as 1792, has been so neglected that it was not even differentiated as a valid species until the past year.
The echinoderm fauna surrounding the coasts of Great Britain is now, thanks to the early and enthusiastic interest shown by the British naturalists in dredging, fairly well understood; and since the first discovery of Antedon petasus in 1835 and of Hathrometra sarsii in 1844, but especially since the discovery of Rhizocrinus lofo- tensis in 1864, the Norwegian naturalists, particularly M. Sars, Danielssen, Koren, and J. A. Grieg, have greatly developed the echinoderm fauna of the rich Nor- wegian coast, and we now have at hand a large mass of data concerning these species.
There has been only a slight and transient interest shown in the comatulids of the corresponding portion of North America. Retzius described Hathrometra tenella from “St. Croix” in 1783, and Say described H. dentata from New Jersey in 1825; since then a number of records of their capture in the early explorations by the ships of the United States Fish Commission (in which, however, both are given under the same name) have been published by Prof. Addison E. Verrill, but prac- tically nothing by anybody else, or in recent years.
The western coast of North America remained absolutely a terra incognita so far as its crinoids were concerned until 1907, in which year many species were described from the region.
Chiefly within comparatively recent years a notable advance has been made in the intensive study of the crinoids inhabiting the coasts of Australia. The first local record, published in Tasmania in 1835 by Wilton, proves to have been based on some organism not a crinoid. There is the same difficulty with the second record, published by Sir Richard Owen in 1862. The third record is scarcely more fortunate, for here a portion of a comatulid is described as a cystidean. Nine years after this we find described and figured two comatulid pentacrinoid larve, but they are given a place in the Porifera instead of in the Echinodermata. Except for these records and notices of Australian species inserted in comprehensive works, Bell’s
MONOGRAPH OF THE EXISTING CRINOIDS. 57
list in the Alert report (1884) is the foundation upon which the knowledge of the crinoid fauna of Australia must be built up. This was followed in the year suc- ceeding by a list published at Sydney, and in 1888, 1889 and 1890 by lists and dis- cussions of Australian species published both in England and in Australia, of which the most important are the records of Mr. Thomas Whitelegge and of Prof. E. P. Ramsay (Sydney) and of the Port Phillip biological survey (Melbourne). In 1894 the foundation was laid for the intensive study of the crinoids of the west coast of Australia, while within recent years the work of the Hamburg west Australian expedition and of the local surveying steamers Thetis and Endeavour has done much to give us a clear idea of the Australian fauna.
The gradual development of knowledge in regard to arctic comatulids must be considered quite apart from the development of the subject as a whole, for the arctic regions have been made the scene of a vast amount of detailed investiga- tion, far exceeding that bestowed upon any other area of equal importance, and the abundance of reliable records from the seas north of America, Europe, and Asia finds no counterpart in any other district.
About 40 workers have assisted in the elucidation of the arctic comatulids, the majority taking little or no interest in those of other regions.
So long ago as 1770 comatulids were found in abundance in the Arctic Ocean and we find many references to them in the writings of the old explorers, more es- pecially those of Phipps, Scoresby and Dewhurst. Dr. W. E. Leach applied the name glacialis to the largest, most characteristic, and most abundant of the Arctic species some time before 1830, Professor Miiller, ignorant of Leach’s work, rechristening it in 1841. In 1859 Edward Forbes remarked upon the enormous abundance of this form at Spitzbergen in moderate depths, and since then there has been a continuous accumulation of data regarding this and other arctic species, at first more or less unsatisfactory but soon becoming definite and exact, so that now we know more about the arctic species and the bathymetric, thermal, and cecological conditions under which they live than we do about any one of the species of Antedon occurring along the European coasts, or about any other crinoid.
A detailed history of all this Arctic research would be in effect a history of but a single species, and is therefore reserved until the consideration of Heliometra glacialis; but it would be an injustice not to mention the investigators by whom this history has been mainly written. Beginning with Wright (1866), Wyville Thomson (1872), Nordenskjold (1876), Sladen (1877) and Stuxberg (1878), who were the first to present really satisfactory data, we meet with the writings of Litken, d’Urban, von Marenzeller, Hoffman, Verrill, Fischer, P. H. Carpenter, Ganong, Levinsen, Danielssen, Pfeffer, Drygalski, Schaudinn, the Prince of Monaco, Déderlein, Hartlaub, Richard, Keehler, Kolthoff, Rankin, Michailovskij, Mortensen, Schmidt, Grieg and Derjugin. Almost all of these gentlemen published at least two papers on the subject, and some of them quite a number. Déderlein’s contribu- tion to the ‘‘Fauna Arctica”’ is especially noteworthy in giving a valuable summary of the records of all previous authors. ~
79146°—Bull. 82—15——5
58 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
Professor von Marenzeller was the first to indicate that, so far as its crinoids are concerned, the fauna of the western part of the Sea of Japan is in reality the same as that of the Arctic Ocean north of Europe.
GENERAL SURVEY OF THE HISTORY.
The history of the development of the study of the comatulids is strangely short when compared to the corresponding history of other groups of marine inverte- brates. There has been a curious reluctance among investigators in regard to attempting work upon these animals. But on the whole this is probably a fortunate circumstance, for few organisms are so baffling and so difficult of systematic analysis, and few have so well resisted the efforts of able zodlogists properly to understand them.
The four works which may justly be considered as marking the four epochs in the study of the comatulids are those of Linck (1733), Lamarck (1816), J. Miller (1849), and P. H. Carpenter (1888), and about these four works the work of all the other authors may be said to have centered, with a remarkably close corre- spondence to the model. There has been an absence of originality and of attempts at revision which is especially striking when we compare the history of the coma- tulids with that of the stalked crinoids.
Although many serious errors have been made, and many wholly illogical methods of systematic treatment proposed, it is perhaps remarkable that the mis- takes have been so few. One can not help commenting upon the fact that the study of the comatulids has been followed by so many of the greatest zodlogists of the past two centuries, and how few are the names of men who have not attained to the highest eminence along other lines.
At the present day the study of the comatulids is in its infancy; nothing more than a beginning has been made, even in the systematic aspect, the phase of the study of every group which commonly first appeals to the novice. One of the chief aims of the present contribution is to demonstrate how woeful is our lack of definite information in regard to even the commonest species, of their systematic interre- lationships, their habitat, their habits, their life history, their anatomy, and of their geological significance, not to mention their relations to temperature, depth, pressure, light, salinity, and in general to all the physics and chemistry of their environment, and to the other animals and the plants surrounded by which they live. It is greatly to be hoped that the present memoir will call attention to these animals in a way that will result in a great increase in the amount of work upon them, and will serve as a stimulus and suggestive guide to young investigators looking for an uncrowded and promising field in which to prosecute their labors, so that we may, in the not far distant future, appreciate the general truths in regard to their “natural history,’”” whereby we may, as we can through no other animals so well, arrive at a clear understanding of many problems in marine biology and in geology.
MONOGRAPH OF THE EXISTING CRINOIDS. 59
GLOSSARY OF TERMS USED IN THE DESCRIPTION OF A COMATULID. Vale
Aboral.—The surface opposite to that which includes the mouth and the anal tube; the dorsal surface. In life this is the lower surface under normal conditions (see figs. 77, p. 130, 78, p. 131, 79, p. 132, 80, p. 133, 81, p. 134, 82, p. 135, 101, p. 163, 107, p. 173, 114, p. 181, 160-162, p. 223, and 163, p. 225).
Adambulacral.—Bordering the ambulacral grooves.
Adapical.—Aboral or dorsal.
Adolescent autotomy.—See under Autotomy 2.
Adoral.—The surface upon which is situated the mouth and the anal tube; the ventral surface. In life this surface is uppermost under normal conditions (see fig. 117, p. 183, and p. 110 [7]).
Ambulacra.—(1) Shallow grooves running along the ventral (adoral) surface of the pinnules and arms and traversing the disk, converging at the mouth; they serve to convey food to the mouth (see figs. 15-19, p. 67, 22-27, p. 69, 45a, p. 79, and 117, p. 183).
(2) This term as used by Guilding is equivalent to cirri.
Ambulacral grooves.—See Ambulacra (1).
Ambulacral lappets.—Small epidermal folds which border the ambulacral grooves on either side, giving their margins a scalloped appearance.
Ambulacral plates.—Small plates developed in two rows (more rarely im a single row) along either side of the ambulacral grooves; the Side and Covering plates taken together (see figs. 7, p. 63, 18, 19, p. 67, and 55, p. 81).
Ambulacral structures.—(1) All the structures, both calcareous and noncalcareous, internal and external, associated with the ambulacra.
(2) The structures in the radial, as opposed to the interradial, portion of the animal.
Anal appendage.—See Anal process.
Anal area.—The interambulacral area at or near the center of which is situated the anal tube (see figs. 15-19, p. 67, and pp. 110 [7], 111).
Anal funnel.—See Anal tube.
Anal interradial.—The interradial situated on the margin of the ana: area.
In cases where there is only one interradial present it is invariably the anal interradial, and this is then known simply as Anal a.
In recent species if the anal interradial is present, all the other interradials are also present (see figs. 115, 117, p. 183, and pp. 335-339).
Anal plate-—See(1) Radianal and (2) Anal x.
Anal process.—The name given to a short segmented process borne on the posterior interradial (anal z) in the so-called Thawmatocrinus renovatus. Thaumato- crinus renovatus is the young of the species later described as Promachocrinus abyssorum, and the anal process is the rudiment of the first of the interradial arms to be formed. Similar processes, each developing into an interradial arm, subsequently appear on all the other interradial plates (see figs. 115-117, p. 183, and pp. 335-339).
60 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
Anal tube.-—A fleshy conical tube, usually of considerable height, situated in one of the interradial areas of the disk (the anal area) and bearing at its summit the anal opening (see figs. 15-19, p. 67, and pp. 110 [7], 111).
Anal x.—The interradial situated between the two posterior radials, distal to and to the left of the radianal if that plate is present. In all the recent forms anal «
ORAL PINNULES Br (PRIMIBRACHS)
CENTRO-DORSAL
CIRRUS SOCKETS
CIRRI
FiG. 1.—LATERAL VIEW OF A SPECIMEN OF ANTEDON ADRIATICA FROM TRIESTE; FOR THE SAKE OF SIMPLICITY THE FOUR ARMS ON THE SIDE OPPOSITE THAT FIGURED ARE OMITTED.
is exactly like the other four interradials, and these are always present if anal z is present. In the recent crinoids anal z, if persistent, gives rise to a post- radial series exactly resembling those on the radials, becoming itself transformed into a plate indistinguisable from a true radial. This is the cause of the forma- tion of 6-rayed variants, the sixth ray being situated between the two posterior
MONOGRAPH OF THE EXISTING CRINOIDS.
ID, SA SWS Y WS Nees ate: tT NE i
BRACHIALS
BASALS (B)
Fia.2.—LATERAL VIEW OF THE TYPE SPECIMEN OF PHRYNOCRINUS NUDUS FROM “ALBATROSS”? STATION 4971; A PORTION OF THE COLUMN AND MOST OF THE ARMS ARE OMITTED. THE CALYX, CONSISTING OF THE BASALS
AND THE RADIALS, IS HEAVILY OUTLINED (DRAWING BY THE AUTHOR).
.DORSO-CENTRAL (TERMINAL STEM PLATE)
61
62 BULLETIN 82, UNITED STATES NATIONAL MUSEUM,
rays and receiving its ambulacra from the ray to its left. In the genera Promachocrinus and Thaumatocrinus anal x and all the other interradials give rise to additional (interradial) post-radial series so that a normally 10-rayed animal results (see figs. 113, 114, p. 181, 115-117, p. 183, and 122, p. 191). Anambulacral.—Bordering the ambulacral grooves. Angles of the calyx.—A term sometimes enployed to designate the points of union between the interradial sutures and the suture between the centrodorsal and
CROWN,
+1 Brs/(PRIMIBRACHIAL AXILLARY) 4.
1 Bf, (FIRST PRIMIBRACH) IALS
s
Fic. 3.—LATERAL VIEW OF A SPECIMEN OF ILYCRINUS COMPLANATUS FROM “ALBATROSS”? STATION 3783; THE MAJOR PART OF THE COLUMN AND FOUR OF THE ARMS ARE OMITTED. THE CALYX, CONSISTING OF THE BASALS AND THE RADIALS, IS HEAVILY OUTLINED (DRAWING BY THE AUTHOR).
the radial circlet in the comatulids. It is here that the outer ends of the basal rays appear (see fig. 415, p. 319).
Antepenultimate segment.—Of the cirri; the segment immediately preceding the penultimate (see figs. 314-317, p. 273, and pp. 278-283).
Anterior arm.—The arm situated directly opposite the anal area; in the endocyclic species the ambulacrum leading from this arm across the disk would, if con- tinued beyond the mouth, pass through the anal tube; in the exocyclic species
MONOGRAPH OF THE EXISTING CRINOIDS. 63
CIRRALS
X RADICULAR CIRRI
TERMINAL CLAW
BASAL SEGMENTS Fic. 5.
COVERING PLATES PERLSOMIC PLATES SIDE PLATES
PINNULARS
. PINNULARS
FIG. 7.
Fia. 8.
Fies. 4-8.—4, LATERAL VIEW OF A DORSAL CIRRUS FROM A SPECIMEN OF PARAMETRA ORION FROM SOUTHERN Taran. 5, LATERAL VIEW OF THE ROOT OF A SPECIMEN OF BATHYCRINUS PACIFICUS FROM SOUTHERN JAPAN, SHOWING STUMPS OF RADICULAR CIRRI(DRAWING BY THE AUTHOR). 6, ‘THE OUTER SIDE OF THE PROXIMAL PORTION OF A FREE UNDIVIDED ARM FROM A SPECI- MEN OF STEPHANOMETRA MONACANTHA FROM Ful. 7, PORTION OF A DISTAL PINNULE FROM A SPECIMEN OF PCCILOMETRA ACCELA FROM NEAR THE MEANGIS ISLANDS (ADAPTED FROM P. H. CARPENTER). 8, GENITAL (OR MIDDLE) PINNULES FROM A SPECIMEN OF PCECILOMETRA ACOELA FROM NEAR THE MEANGIS ISLANDS, SHOWING THE EXPANSION AND THE VENTRAL
PLATING (ADAPTED FROM P. H. CARPENTER).
64 BULLETIN 82, UNITED STATES NATIONAL MUSEUM,
this ambulacrum usually, at the base of the arm, makes a more or less abrupt turn to the right to reach the interradial mouth, which is situated between the bases of the anterior and the right anterior arms (see figs. 22-27, p. 69, and 117, p. 183, and p. 110 [6]); (see Avis and Orientation).
Anterior radii.—(1) The radius in which the anterior arm is situated is commonly distinguished as the anterior radius (see fig. 22, p. 69).
(2) It is sometimes convenient to differentiate the radii on either side of the anal area from the three others, in which case there are distinguished 2 posterior and 3 anterior radii.
(3) In certain of the Comasteride, where the left posterior radius is curiously modified, this is often referred to as the posterior radius, the remaining four being collectively termed anterior radii (see fig. 27, p. 69, and p. 111).
Apical.—(1) Aboral or dorsal.
(2) Applied to the centrodorsal (or cirri), situated at or near the dorsal pole (see fig. 310, p. 269, and pp. 304-306).
Apical plate.—The hypothetical plate covering the center of the dorsal side of the primitive crinoid (compare fig. 71, p. 127, and see pp. 198-200).
Appendicular skeleton.—The skeleton of the division series and arms; the skeleton of the post-radial series.
Arm bases.—The proximal brachials; this term is commonly employed to distin- guish the more or less oblong earlier brachials as distinct from the triangular brachials beyond them (see figs. 30, p. 71, 6la-c, p. 87, 79, p.132, 94, p. 155, 109, p. 175, and 110 p. 176).
Arm pair.—Any two free undivided arms which arise from the same axillary. This term is rarely met with except in reference to 10-armed species, in which each of the post-radial series is sometimes referred to as an arm pair-
Arms.—(1) Strictly speaking, the series of ossicles subsequent to the last straight muscular articulation; or the series of ossicles beginning with the one imme- diately preceding the last synarthry; thus in the Pentametrocrinide the arms begin with the first ossicle beyond the radials; in the Uintacrinide they begin with the third ossicle beyond the [Br (costal) axillary; in the remaining coma- tulid families they ordinarily begin with the first segment after the last axillary, except in the genus Eudiocrinus, in which the third segment beyond the radials is the first arm ossicle. In the recent comatulids the true arms never divide (see figs. 6la—c, p. 87, and pp. 109 [5], 110 [6]).
(2) While the preceding definition delimits morphologically homologous arms, it is more convenient for practical descriptive purposes to consider the arms as including the entire undivided series of ossicles beyond the last axillary, or beyond the radials in the Pentametrocrinide and in the genus Eudiocrinus (see figs. 1, p. 60, and 2, p. 61, and p. 110 [6]).
(3) Several authors have considered all the ossicles beyond the radials, no matter how many divisions there may be, and without regard for the type of division, as morphologically comparable arms; this view is inadmissible, for the reason that the radial is an integral part of the series of ossicles following, and is not properly a calyx plate at all.
MONOGRAPH OF THE EXISTING CRINOIDS. 65
CENTRAL CANAL
MUSCULAR FOSS
INTERARTICULAR LIGAMENT FOSSAZ
=
a BASAL RAYS TRANSVERSE RIDGE
|=) RADIAL RIDGES 2y DORSAL LIGAMENT FOSSA
ie - CIRRUS SOCKETS VW CRA centro-porsat
=
< INTERRADIAL RIDGES CIRRUS SOCKETS
Ss Fia. 10. ¢ DORSAL POLE
Fia. 9.
¢ % A “%, Ue oa) UN Wo ip ENTRAL CAVITY % Se % ci SYZYGIES S %, NO T Br, (FIRST PRIMIBRACH <7) , pees es Fig. 13. ri ( aT es Ss WATER PORES BASALS
CENTRO-DORSAL Fia. 14.
Figs. 9-14.—9, LATERAL VIEW OF THE CENTRODORSAL AND RADIALS OF A SPECIMEN OF STENOMETRA QUINQUE COSTATA FROM THE Ky ISLANDS (ADAPTED FROM P. H. CARPENTER). 10, LATERAL VIEW OF THE CENTRODORSAL AND RADIALS OF A SPECI MEN OF HIMEROMETRA MARTENSI FROM SINGAPORE (DRAWING BY THE AUTHOR). 11, VENTRAL VIEW OF THE RADIAL PENTA- GON OF A SPECIMEN OF HIMEROMETRA MARTENSI FROM SINGAPORE (DRAWING BY THE AUTHOR). 12, DORSAL VIEW OF THE RADIAL PENTAGON OF A SPECIMEN OF PTILOMETRA MULLERI FROM SYDNEY, NEW SouTH WALES (DRAWING BY THE AUTHOR). 13, VENTRAL VIEW OF THE CENTRODORSAL OF A SPECIMEN OF PEROMETRA DIOMEDEX FROM SOUTHERN JAPAN (DRAWING BY THE AUTHOR). 14, DIAGRAMMATIC LATERAL VIEW OF THE PROXIMAL PORTION OF A SPECIMEN OF ATELECRINUS CONIFER FROM THE HAWAIIAN ISLANDS (DRAWING BY THE AUTHOR).
66 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
Articular faces.—The apposed surfaces of two segments united by articulation, as opposed to suture (see figs. 31-34, p. 71, and 36-40, p. 75, and pp. 113, 376); (see Articulations).
Articular facets—See Articular faces.
Articulations—The unions between adjacent ossicles when composed of ligament bundles or of muscles, or of a combination of both (see Suture); articulations are of two types, each type being subdivided into two subtypes, as follows:
A. Muscular articulations —The apposed articular faces are marked by an approximately hemispherical pit lodging the dorsal ligament, anterior (ventral) to which is a strong transverse ridge; slightly anterior to the center of this ridge is the central canal through which passes the axial cord of the dorsal nervous system; just anterior to the transverse ridge lies a pair of interarticular ligament fosse, one on either side of the central canal; these interarticular ligament fossee are bounded anteriorly by strong oblique ridges which separate them from the pair of muscular fossx (see figs. 31, 32, p. 71, 431, 432, p. 349 and pp. 114, 376).
a. Straight muscular articulation (often known simply as Muscular articulation).—A type of muscular articulation in which the transverse ridge is perpendicular to the dorsoventral axis of the joint face, and the dorsoventral axis divides the joint face into two equivalent and similar halves (see figs. 31, p. 71, 431, 432, p. 349, and pp. 114, 376).
b. Oblique muscular articulation.—A type of muscular articulation in which the transverse ridge is strongly oblique in reference to the dorsoventral axis of the joint face (typically making with it an angle of 45°) and the interarticular and muscular fosse of the two sides are more or less unequal (see figs. 6, p. 63, and 30, 32, p.71).
B. Nonmuscular articulations—Articulations in which muscles are absent, the union being effected solely by ligaments (see figs. 33, 34, p. 71, 36-40, p. 75, and p. 113).
a. Synarthry.—A type of non-muscular articulation in which the apposed articular faces show two hemispherical fosse for the recep- tion of a pair of ligament bundles, separated by a strong ridge running in the direction of the dorsoventral axis of the joint face, which is pierced in the center by the central canal (see figs. 6, p. 63, 14, p. 65, 30, 33, p. 71).
b. Syzygy—A type of nonmuscular articulation in which the apposed surfaces are flat, and are marked by fine low radiating ridges (see figs. 2, p. 61, 6, p. 63, 14, p. 65, 34, p. 71, and 35, p. 73).
(See also Cryptosynarthry and Pseudosyzyqy.)
Asteriz.—Same as Pentacrini.
Autotomy.—(1) A process by which a comatulid inflicts self-mutilation, usually by breaking off a part or all of an arm; this usually occurs at either a syzygy or at asynarthry. This process of autotomy in the crinoids has commonly been supposed to be voluntary, but is in reality the result of a state of panic which causes a total relaxation of the muscles (see pp. 140-142).
MONOGRAPH OF THE EXISTING CRINOIDS. 67
AMBULA GROOV INTERAMBULACRAL (INTERPALMAR) ees 7 a
uu ® ; SACCULL 8 3 x ° = é ANUS 2 ANAL TUBE MOUTH J ~ a = -<
(YVWIVduaLNI) TVaOVINaWVYaLNI
Fia. 16.
AMBULACRAL GROOVES
(\ (iereRAMBULACRAL (INTERPALMAR) AREAS
INTERAMBULACRAL (INTERPALMAR) AREAS
PERISOMIC PLATES
SE oe Si ANS NOY LES PAS x
Q Sah \ Y X oO \Y Fe
y=
AMBULACRAL GROOVES
MOUTH
ANAL TUBE’ anus ‘ANAL AREA ©
Fia. 18. ANAL AREA ANAL AREA
Fie. 19.
Figs. 15-19.—15, THE NAKED ENTIRE DISK OF A SPECIMEN OF TROPIOMETRA PICTA FROM RIO DE JANEIRO. 16, THE NAKED INCISED DISK OF A SPECIMEN OF CENOMETRA BELLA FROM THE CHINA SEA. 17, THE DEEPLY INCISED DISK OF A SPECIMEN OF MARIAMETRA DELICATISSIMA FROM SOUTHWESTERN JAPAN. 18, THE PARTIALLY PLATED, SLIGHTLY INCISED DISK OF A SPECIMEN OF PARAMETRA ORION FROM SOUTHERN JAPAN. 19, THE COMPLETELY PLATED ENTIRE DISK OF A SPECIMEN OF
NEOMETRA MULTICOLOR FROM SOUTHERN JAPAN.
68 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
(2) In the comatulids this process is always invoked to produce a greater number of arms than 10; the young animal always has 10 arms until a con- siderable size is reached, when the arms are broken off either at the first syzygy or at the first synarthry, and from the stump an axillary is regenerated bearing two or more arms in the places of the one lost; this is known as Adolescent autotomy.
Autotomy at any other place than the first syzygy or the first synarthry always results in the regeneration of a single arm similar to the one lost, though with a longer and more irregular intersyzygial interval (see Regeneration).
Adolescent autotomy is caused by natural growth changes in the arms, and is not in any way subject to the will of the animal (see pp. 140-142).
Axial cavity—The small hole left in the dorsal pole of the centrodorsal after the loss of the larval stem. It is almost immediately closed by a deposition of calcareous matter (see fig. 594, pl. 16, and pp. 228, 229).
Azial cord—(1) The large nerve cord which runs along the arm in the canal (the central canal) just anterior to the transverse ridge seen on the joint faces (see figs. 31-34, p. 71, 63, 64, p. 89, and 65, p. 91 and pp. 350-354).
(2) This term is sometimes used to include all the nerves belonging to the dorsal nervous system.
Arial interradial canals —The more or less complete canals in the interior of the radial pentagon which lie on the sutures between the radials.
They inclose branches from the water vascular system (see pp. 375, 376).
Axial interradial furrow.—The furrows seen on the inner side of the radial pentagon which coincide in position with the sutures between the radials; when bridged over by calcareous deposit they form the axial interradial canals (see pp. 375,376).
Arial nerve cord. See Axial cord.
Axial prolongation—A prolongation of the radial canals of the water vascular system whereby they come to end upon the ventral surface of the centrodorsal, or even to extend outward between the centrodorsal and the radial pentagon (see figs. 252-255, p. 253, 256-261, p. 255, 468-470, p. 359, 471-476, p. 361, 477, p. 363, and 508, p. 371, and pp. 374, 375).
Arial radial canals —The radial canals of the water vascular system, when more or less surrounded by calcareous deposit.
Avial radial furrows.—The furrows on the interior surface of the radial pentagon which when bridged by calcareous deposit form the axial radial canals.
Arial skeleton —The Radial skeleton.
Arillary—An ossicle at which the arms divide; a single ossicle which bears distally two similar series of ossicles arising from a pair of similar muscular articulations (see figs. 1, p. 60, 3, p. 62, 14, p. 65, 30, p. 71, and 61 a-c, p. 87, and pp. 358- 360).
Arvis.—The axes commonly considered in the description of the comatulids are:
(1) Anteroposterior axis ——This axis divides the animal into two bilaterally similar halves; it is found in two positions, a (1) primary and
a (2) secondary.
MONOGRAPH OF THE EXISTING CRINOIDS. 69
Fia. 20. Ria 22.
Fia. 26. Fie. 27. Fie. 28.
‘
Figs. 20-28.—20, THE DIGESTIVE TUBE AND DISK AMBULACRA OF ANTEDON BIFIDA, ILLUSTRATING A COMATULID WITH AN ENDO- CYCLIC MOUTH (ADAPTED FROM P. H,. CARPENTER). 21, THE DIGESTIVE TUBE AND DISK AMBULACRA OF ONE OF THE SPECIES OF THE FAMILY COMASTERID#, ILLUSTRATING A COMATULID WITH AN EXOCYCLIC MOUTH (ADAPTED FROM P. H. CARPENTER). 22, DIAGRAM SHOWING THE COMPARATIVE RELATIONSHIPS BETWEEN THE AMBULACRA, ANAL TUBE, AND ARMS IN A FIVE- ARMED ENDOCYCLIC COMATULID; THE AXIS d-@ IS THE PRIMARY ANTEROPOSTERIOR AXIS. 23, DIAGRAM SHOWING THE COM- PARATIVE RELATIQNSHIPS BETWEEN THE AMBULACRA, ANAL TUBE, AND ARMS IN A TEN-ARMED ENDOCYCLIC COMATULID; THE AXIS a-@ IS THE PRIMARY ANTEROPOSTERIOR AXIS, 24, DIAGRAM SHOWING THE COMPARATIVE RELATIONSHIPS BETWEEN THE AMBULACRA, ANAL TUBE, AND ARMS IN A TWENTY-ARMED ENDOCYCLIC COMATULID; THE AXIS @-a IS THE PRIMARY ANTERO- POSTERIOR AXIS. 25, DIAGRAM SHOWING THE COMPARATIVE RELATIONSHIPS BETWEEN THE AMBULACRA, ANAL TUBE, AND ARMS IN A TEN-ARMED EXOCYCLIC COMATULID, OR COMASTERID, IN WHICH ALL OF THE ARMS ARE PROVIDED WITH AMBULACRAL GROOVES, AND IN WHICH THE MOUTH IS RADIALIN POSITION; THE AXIS a-@ IS THE PRIMARY ANTEROPOSTERIOR AXIS. 26, DIA- GRAM SHOWING THE COMPARATIVE RELATIONSHIPS BETWEEN THE AMBULACRA, ANAL TUBE, AND ARMS IN A TEN-ARMED EXOCYCLIC COMATULID, OR COMASTERID, IN WHICH ALL OF THE ARMS ARE PROVIDED WITH AMBULACRAL GROOVES, AND IN WHICH THE MOUTH IS INTERRADIALIN POSITION; THE AXIS a-@ IS THE PRIMARY ANTEROPOSTERIOR, THE AXIS 0-b THE SEC- ONDARY ANTEROPOSTERIOR. 27, DIAGRAM SHOWING THE COMPARATIVE RELATIONSHIPS BETWEEN THE AMBULACRA, ANAL TUBE, AND ARMS IN A TEN-ARMED EXOCYCLIC COMATULID, OR COMASTERID, IN WHICH EIGHT OF THE ARMS ARE PROVIDED WITH AMBULACRAL GROOVES AND TWO ARE UNGROOVED, AND IN WHICH THE MOUTH IS INTERRADIAL IN POSITION; THE AXIS a-a IS THE PRIMARY ANTEROPOSTERIOR, THE AXIS b-b THE SECONDARY ANTEROPOSTERIOR. 28, DIAGRAM SHOWING THE COMPARATIVE RELATIONSHIPS BETWEEN THE AMBULACRA, ANAL TUBE, AND ARMS IN A TEN-ARMED EXOCYCLIC COMATULID, OR COMASTERID, IN WHICH FOUR OF THE ARMS ARE PROVIDED WITH AMBULACRAL GROOVES AND SIX ARE UNGROOVED; THE AXIS a-2 IS THE PRIMARY ANTEROPOSTERIOR, THE AXIS b-b THE SECONDARY ANTEROPOSTERIOR.
70 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
a. Primary anteroposterior avis—In the endocyclie comatulids the axis passing along the anterior arm and continued through the mouth and anal tube, leaving the animal in the center of the posterior border of the anal area, divides it into two exactly similar halves (see figs. 22-28, p. 69).
b. Secondary anteroposterior axis—In such of the exocyclic comatulids as have an interradial mouth, situated on the edge of the disk between the bases of the anterior and right anterior rays the anteroposterior axis which divides the animal into two bilaterally equal halves passes through the middle of the interambulacral area between the anterior and right anterior arms, through the mouth, through the anal tube, and along the median line of the left posterior ray (see figs. 26-28, p. 69, and pp. 152-161).
(2) Dorsoventral axis —This axis passes through the dorsal pole and through the center of the disk, being at right angles to the plane in which the arms lie when extended horizontally.
(3) Longitudinal axis—In speaking of the arms individually this axis refers to the mid line of the arms; it is occasionally used in reference to the pinnules or to the cirri.
Azygous tentacle—The median tentacle of a tentacle group; usually the term refers to the first tentacle which is formed in the larva (see fig. 543, pl. 4.)
B.
Basal.—See Basals.
Basal bridge-—A narrow rounded ridge or rod connecting the inner ends of the basal rays; the five basal bridges form a pentagon within which is seen the rosette (see figs. 424-426, p. 321, 447-449, p. 353, 454, p. 355, 459-463, p. 357, and 479, 480, p. 363, and pp. 324, 335).
Basal cirrals.—The one, two, three, or four very short cirrus segments immediately adjacent to the centrodorsal (see fig. 4, p. 63, and p. 276). :
Basal fold.—The incurved edge of the basal grooves, which is applied to the basal ray.
Basal grooves.—The grooves on the dorsal surface of the radial pentagon which lodge the basal rays; they occur on the lines of suture between the radials (see figs. 229-233, p.247, 236-242, p. 249, 243-249, p. 251, 256-258, p. 255, and pp. 236-238, 370).
Basal pentagon.—The Radial pentagon.
Basal rays.—Prismatic calcareous rods of secondary origin developed in the basal grooves between the radial pentagon and the centrodorsal; their inner ends are usually connected with the rosette, and by basal bridges with the inner ends of the adjacent basal rays (see figs. 9-12, p. 65, 97, p. 159, 208-215, p. 241, 227, p. 245, 229-233, p. 247, 416-427, p. 321, and 447-451, p. 353, and pp. 326-330).
Basal ring.—A structure formed by anchylosed basals which show no trace of the interbasal sutures (see figs. 3, p. 62, and 134, p. 203).
Basal star.—The five basal rays, plus the five connecting basal bridges (see figs. 447-451, p. 353, and pp. 324, 325).
MONOGRAPH OF THE EXISTING CRINOIDS. 71
. BRACHIALS
“I1l Br (TERTIBRACHS) 1 Br; (FIRST PRIMIBRACH)
Il Br (SECUNDIBRACHS) I Br (PRIMIBRACHS) CENTRO-DORSAL
Fia. 29.
MUSCULAR FOSS SEPTUM
INTERARTICULAR LIGAMENT FOSSA= LONGITUDINAL RIDGE CENTRAL CANAL CENTRAL CANAL TRANSVERSE RIDGE
LIGAMENT PIT Fia. 33. DORSAL LIGAMENT FOSSA
Fia. 31.
MUSCULAR FOSS: SEPTUM {NTERARTICULAR LIGAMENT FOSS RADIAL RIDGES CENTRAL CANAL TRANSVERSE RIDGE CENTRAL CANAL LIGAMENT PIT
DORSAL LIGAMENT FOSSA Fia. 34. Fig. 32.
Fics. 29-34.—29, DIAGRAM OF THE CENTRAL STRUCTURES AND ARM BASES OF A SPECIMEN OF 4 SPECIES OF COMANTHUS WITH THE CIRRI REMOVED (DRAWING BY THE AUTHOR). 30, THE CENTRAL STRUCTURES AND PART OF A POSTRADIAL SERIES OF A SPECI- MEN OF THAUMATOMETRA TENUIS FROM THE WESTERN PART OF THE SEA OF JAPAN (DRAWING BY THE AUTHOR). 31, D1A- GRAM OF A STRAIGHT MUSCULAR ARTICULATION (DRAWING BY THE AUTHOR). 32, DIAGRAM OF AN OBLIQUE MUSCULAR ARTICULATION (DRAWING BY THE AUTHOR). 33, DIAGRAM OF A SYNARTHRY (DRAWING BY THE AUTHOR). 34, DIAGRAM OF A SYZYGY (DRAWING BY THE AUTHOR).
72 BULLETIN 82, UNITED STATES NATIONAL MUSEUM.
Basals (BB) —The plates which collectively form a circlet just below the radials; they are usually five in number and alternate in position with the radials, but many forms possess only three; they may be entirely distinct, with the suture lines easily visible between them, or they may be solidly anchylosed, forming a solid ring or funnel (see figs. 2, p. 61, 3, p. 62, 14, p. 65, 115-118, p. 183, 122, p. 191, 130-134, p. 203, 144, p. 207, 145, p. 209, and 407-413, p. 317, and pp. 316-331).
In the recent comatulids the basals, at first forming an essential part of the calyx wall, become in early life metamorphosed into the rosette and there- fore disappear from external view, except in the family Atelecrinide where they are almost always to be seen forming a narrow ring between the centro- dorsal and the radials (see figs. 123, p. 192, 124, 125, p. 193, 414, p. 319, and 430," p. 321, and pl. 8, figs. 573-575, and pp. 318-320).
Many recent comatulids have, just above the centrodorsal in the interradial angles, more or less pronounced tubercules which are often so large as to appear as true basals; these are, however, Basal rays of secondary origin, and have no connection with the larval basals (see figs. 415, p. 319, and 416-427, p. 321, and pp. 326-330).
In the recent crinoids the infrabasals, when present, form a circlet within the basals and are entirely concealed by them; in the comatulids they fuse with the uppermost columnal in early life to form the centrodorsal, or are entirely absent (see figs. 565-572, pl. 7, and pp. 313-316).
The basals are the equivalent of the genitals in the echinoids. »
Basal surface.—Of the centrodorsal; the dorsal pole.
Base.—(1) Of the calyx; the Radial pentagon;
(2) Of the centrodorsal, the surface which is applied to the radials (see figs. 229, 230, 232-234, p. 247, 235-242, p. 249, and p. 232).
Bifascial articulation—Same as Synarthry.
Bilateral symmetry.—See Symmetry and Azis.
Bivium.—A term used to designate the posterior pair of arms, or rays, when these differ from the three anterior in being short, ungrooved, and nontentaculiferous, as in many of the Comasteride (see figs. 45a—b, p. 79, and pp. 110, 111).
Bourgueticrinoid stem.—A stem or column of the type found in the species of the genus Bourgueticrinus. This type of stem is characteristic of the young of the comatulids and of the pentacrinites (see figs. 135-139, 141-143, p. 205, 518-524, 526, pl. 1, and 527, pl. 2, and pp. 208-210).
Brachial ambulacra.—The ambulacra on the ventral surface of the arms and of the division series (in contrast to those of the disk and the pinnule ambulacra) (see fig. 45a, p. 79, and pp. 110, 111).
Brachial axillary.—A term used by some authors for any of the axillaries except the first, which is differentiated as the IBr, primibrachial, radial, or costal axillary.
Brachial perisome.—The perisome upon the ventral surface of the arms, beyond the second brachial.
Brachials (Br).—The calcareous segments or ossicles of which the arms are composed ; many authors have used this term for all the ossicles beyond the radials, but it is more properly used, as herein, for the ossicles beyond the last division series only (see fig. 1, p. 60, fig. 2, p. 61, and fig. 6, p. 63).
MONOGRAPH OF THE EXISTING CRINOIDS. 73
0.
Calyx.—The base of a crinoid; that is, the part remaining after the stem (or centro- dorsal) and postradial structures have been removed; it includes the infra- basals (when present), the basals and the radials, with any supplementary plates such as<