a p- | tr . r- ! un ; r-q I i-n i CD I CD □ m CD = CD fc THE VOYAGE OF H.M.S. CHALLENGER. ZOOLOGY-VOL. XXX. TEXT. - REPORT ON THE SCIENTIFIC RESULTS OF THE VOYAGE OF H.M.S. CHALLENGER DURING THE YEARS i873-76 UNDER THE COMMAND OF Captain GEORGE S. NARES, R.N., F.R.S. AND THE LATE Captain FRANK TOURLE THOMSON, R.N. PREPARED UNDER THE SUPERINTENDENCE OF THE LATE Sir C. WYVILLE THOMSON, Knt, F.R.S., &c. ^5!lIfo REGIUS PROFESSOR OF NATURAL HISTORY IN THE UNIVERSITY OF EDINBURGH DIRECTOR OF THE CIVILIAN SCIENTIFIC STAFF ON BOARD AND NOW OF 7 JH Y\ %S W JOHN MURRAY, LL.D., Ph.D., &c. ONE OF THE NATURALISTS OF THE EXPEDITION Zoology— Vol. XXX. TEXT MARINE BIOLOGICAL LABORATORY LIBRARY WOODS HOLE, :, W. H. 0. I. IPubltsfoeo bp ©rHer of l£er iflajestp's (Sobmunent PRINTED FOR HER MAJESTY'S STATIONERY OFFICE AND SOLD BY LONDON:— EYRE & SPOTTISWOODE, EAST HARDING STREET, FETTER LANE EDINBURGH:— ADAM & CHARLES BLACK DUBLIN:— HODGES, FIGGIS, & CO. 1889 Price (with Volume of Plates) £4, 10s. PRINTED BY BALLANTYNE, HANSON AND CO., EDINBURGH AND LONDON, FOR HER MAJESTY'S STATIONERY OFFICE. x rts CONTENTS. Report on the Asteroidea collected by H.M.S. Challenger during the years 1873-1876. By W. Percy Sladen, F.L.S., &c. (The Manuscript was received in Instalments betiveen 15th March 188G and 21st January 1889.) MARINE BIOLOGICAL LABORATORY LIBRARY WOODS HOLE, I VV. H. C I. ERRATA. Page 27, line 2 from below, for " Expedition " read "Expedition." „ 28, „ 8 from below, for " l°\t C." read "- 1"-1 C." „ 29, „ 4, for "1M C." read "-1°-1 C." „ 75, ,, 8 from below, for "magniying" read "magnifying." ., 87, „ 9 from below, for "0°-8 C." read "-0"-8 C"; and line 7 from below, for "0'-6 C," read «-0°-6 C." „ 110, in table, for " 49 " read " 85." „ 124, line 23, for "Billeton" read "Billiton." „ 174, „ 15, for "Buono" read "Bueno." ,, 209, ,, 12 from below, for " echinulata" read " echimdatus." „ 238, ,, 15, for "pumilis " read " pumilus" „ 248, „ 8, for "115 "read "150." 253, „ 17, delete " manuscript," and delete lines 26 and 27. (For Luidia chefuensis, Grabe, see Jahres-Ber. Schles. Getellsch. f. voted. Cultur, 1876, p. 28.)] „ 254, „ 7, for "Delia" read " Delle." „ 261, „ " 23, for " childreni " read " eqitestris." „ 287, „ 16 from below, add " PI. XLVIII. figs. 5 and 6." „ 295, last line, for " west " read " east." ,, 309, line 16, for " western " read " eastern." „ 317, „ 10 from below, for " 78° 0 " read " 78°-8." „ 321, „ 11 from below, for "tuberculatus" read " tuberculosus." ,, 32G, ,, 3 from below, for "piercei" read " peircei." „ 344, ., 6 from below, for "latter" read "former." „ 402, ,, 17, for "pustidata" read " pusUdafus." „ 411, „ 3 from below, after " Mag." read " Nat:' „ 448, „ 3 from below, for "E<3r " read " E >3 ;•." „ 551, „ 5, add "PI. LXILfig. 1." „ 553, lines 2 and 9 from below, and p. 554, line 2, for "fallax" read " purpureus." „ 739, line 12, for "row" read "non." EDITOKIAL NOTE. The collection of Asteroidea is one of the most important made during the voyage of the Challenger. The late Sir C. Wyville Thomson paid con- siderable attention to these organisms, and at one time had the intention of describing the new species discovered during the Expedition ; with this end in view, indeed, he had a few of the specimens figured and litho- graphed. Eventually, however, the whole collections were handed over to Mr. W. Percy Sladen for examination and description. The result of Mr. Sladen's work, extending over many years, is now presented in this magnificent Report, which forms Part LI. and Volume XXX. of the Zoological series of Reports, the text and plates being bound up in separate parts. Not only on account of the large number of forms, but also on account of their representative character, the Challenger collection is unquestion- ably the most important contribution to this group of animals that has ever been made. In the Report, supplementary to the treatment of the Challenger material, a compact account of the whole group has been aimed at. Thus (1) under each family an account of all the genera included therein, whether Challenger or not, has been given, and where necessary the grounds on which the classification has been adopted are discussed ; (2) under each genus there is a statement of all the known species which are recognised as belonging to that genus, with an account of their geo- graphical distribution : this presents a compact idea of the composition and distribution of each genus ; (3) a complete list of all the known species of THE VOYAGE OF H.M.S. CHALLENGER. recent Asteroidea, -with their geographical and bathymetrical distribution, and a full account of all synonyms and changes of name are given ; every known starfish is believed to be included, and the recognition of " good species " and " synonyms " is based on personal observation (except in the case of some American types and a few recently described from the French dredgings). The Report may be considered a monograph on the group, though of course it need hardly be said that previously described and recognised species are not re-described. An entirely new classification of the Aster- oidea has been given — it was very unsatisfactory and quite unworkable previously. All naturalists who appreciate careful and accurate work will join with me in congratulating Mr. Sladen on the completion of his valuable memoir, which extends to 935 pages, and is illustrated by 118 lithographic plates and a map. John Murray. Challenger Office, 32 Queen Street, Edinburgh, January 23, 1889. THE VOYAGE OF H.M.S. CHALLENGER. ZOOLOGY. REPORT on the Asteroidea collected during the Voyage of H.M.S. Challenger during the Years 1873-76. By W. Percy Sladen, F.L.S., F.G.S., &c, Secretary of the Linnean Society. PREFACE. The preparation of the Eeport on the Asteroidea obtained by the Challenger Expedition was entrusted to me by the late Sir C. Wyville Thomson, and I received the collection in the early part of the year 1881. Dr. John Murray subsequently placed in my hands the starfishes collected during the cruises of the "Lightning," "Porcupine," "Knight Errant," and " Triton ; " and an account of these is included in the present Report, which has thus become a statement of all the Asteroidea obtained by British deep-sea dredging Expeditions up to the present date. In the following pages there are described 33 genera, 2 subgenera, 196 species, and 15 varieties first discovered by the Expeditions above named ; and 9 other new genera and 3 subgenera, based on forms previously known or not obtained by these Expeditions, are established. Summarising each Expedition independently : in the Challenger collection there are 84 genera, 5 subgenera, 268 species, and 13 varieties; of which 34 genera, 4 subgenera, 184 species, and 12 varieties are new (five of the new genera and two subgenera being based on types of which one or more representatives have been previously known). In the "Porcupine" collections of 1869 and 1870 there are 25 genera, 1 subgenus, 29 species, and 1 variety ; of which 7 genera, 1 subgenus, 10 species, and 1 variety are new (two of the genera being based on types of which representatives were previously known). In the "Lightning" collection are 3 genera and 3 species; of which 1 genus and 1 species are new. (In the case of the "Lightning" and " Porcupine " Expeditions it is (ZOOL. CHALL. EXP. — PART LI. — 1888.) a 11 THE VOYAGE OF H.M.S. CHALLENGER. probable that a greater number of starfishes were obtained than are here recorded ; those enumerated, however, appear to be all the authenticated examples now preserved.) In the " Knight Errant " collection are 9 genera, 1 subgenus, 10 species, and 2 varieties ; of which 4 genera, 2 species, and 2 varieties are new (two of the new genera being based on types of which representatives were previously known). In the "Triton" collection are 11 genera, 11 species, and 2 varieties; of which 5 genera, 4 species, and 1 variety are new, and two of the new genera are established on types of which representatives were previously known. In the latest summary of the Asteroidea, published by M. Perrier1 in 187S, 454 species are enumerated, representing 52 genera. Three of the genera, however, included by M. Perrier are invalid or synonymous, the abstraction of which reduces the number recognised by him to 49. Representatives of 38 of these genera were obtaiued by the Challenger ; that is to say, representatives of more than three-fourths of the previously known types. The remaining 11 genera are either extremely rare or of very local occurrence. These figures indicate in a striking; manner that the collection made by the Challenger affords a fair representation of the general character of the Asterid fauna of the globe, so far as known. The proportionally large number of new species is not surprising, when it is borne in mind that a considerable portion of the track of the Challenger passed through regions which had hitherto been wholly unexplored, or in which the dredge had never previously been employed. The long list of Asterids inhabiting abyssal depths brought to light by the Challenger and the other Expeditions herein discussed may be said to have opened a new chapter in the history of the Asteroidea. The summary of these forms, nearly all of which belong to previously unknown types, given in the Tables appended to the Report affords the most striking evidence of the important contribution made by the Challenger to our knowledge of the group. Reference to these Tables will render any words of comment by me unnecessary here. The archaic characters of a number of the deep-sea forms are highly remarkable, and furnish not only a confirmation of the validity of the classification now adopted for the Asteroidea, but also give an important clue to the systematic position of many of the extinct members of the Class. The large accession of new forms has greatly extended our knowledge of the morpho- logical plasticity of many types, and has thrown important light on the constitution and relations of groups and families. In all cases the descriptions take cognisance of external and general morphological structure, and are not confined merely to details of specific difference, from the systematist's point of view. All the systems of classification previously employed by writers on the Asteroidea are, in 1 Nouv. Archives Mus. Hist. Nat., 2e S£rie, t. i. p. 73. REPORT ON THE ASTEROIDEA. iii my opinion, inadequate and artificial ; and I have been led to formulate a classification more in accordance with morphological characters as at present understood. I venture to believe that this will be found more convenient for general workers, as well as more natural. In dealing with the synonymy of a species I have in each case given all the different name-combinations under which the form has been recognised and the author by whom each was originated, but I have not given a citation of all subsequent writers by whom the terms have been copied or used. By this means brevity has been gained, without sacrificing anything pertinent to the history of the name. In order to add to the completeness of the work, I have given under each genus a notice of all authentic species belonging thereto, and their geographical distribution. I have also added at the end of the Eeport a synoptic list of the known species of recent Asteroidea, with particulars of their geographical and bathyrnetrical distribution, the changes of name to which they have been subjected, and their synonyms, which I hope will increase the utility of the Report. A few species which have been inadvertently omitted in the chorological statement of the genera will be found duly recorded in this list. 137 genera and 810 species are here enumerated. The urgent need of a critical examination by one person of the large number of types distributed throughout the various Continental collections has long been felt by every worker at the group, as many of the types in question have been very imperfectly known, and, owing to insufficient description, numberless false determinations and a confusing multiplication of synonymous terms have arisen. To eliminate as far as possible this element of perplexity, I have visited the chief Continental museums and have personally studied the rich collections at Christiania, Stockholm, Lund, Copenhagen, Leyden, Paris, Berlin, Kiel, Hamburg, Leipzig, Breslau, and Vienna. I venture to believe that the labour thus expended will place the list now given on a more reliable basis than was pre- viously possible. I desire to acknowledge gratefully the privileges afforded me in this undertaking, and to express my hearty thanks to the Heads of the various universities and museums, who, without exception, placed the collections under their charge unreservedly at my disposal. To the many eminent naturalists who have given me valuable assistance and information I desire to tender my sincere thanks. Some obligations are ackoowdedged in the text, and in addition to those I cannot refrain from naming here the late Prof. M. Esmark and Dr. Robert Collett of Christiania ; Prof. S. Loven of Stockholm ; Dr. Hjalmar Theel of Upsala ; Dr. C. D. E. Roth of Lund ; Prof. J. J. S. Steenstrup and Prof. Chr. F. Lutken of Copenhagen ; the late Prof. H. Schlegel and Dr. F. A. Jentink of Leyden ; Prof. Edmond Perrier and Ms J. Poirier of Paris ; the late Prof. W. Peters, Prof. Ed. von Martens and Dr. F. Hilgendorff of Berlin ; Prof. Karl Mobius of Kiel ; Prof. H. A. Pagenstecher and Dr. G. Pfeffer of Hamburg ; Prof. R. Leuckart and Prof. J. V. Carus of Leipzig ; Prof. Anton Schneider of Breslau ; Dr. F. Steindachner and Dr. Emil von iv THE VOYAGE OF H.M.S. CHALLENGER. Marenzeller of Vienna. To each and to all I desire to express my hearty appreciation of the courtesy and friendly kindness I received at their hands. My warmest thanks are also due to Dr. A. C. L. G. Giinther, the keeper of the Zoolo- gical Department of the British Museum, for the many facilities afforded to me during the progress of my work ; and especially to my friend, Prof. F. Jeffrey Bell, whose constant and ever-ready assistance has placed me under an indebtedness greater than I can adequately express in words. In like manner, my cordial thanks are given to my friend, Dr. P. Herbert Carpenter, whose critical advice and wide knowledge of Eckinoderm literature have always been at my disposal. In conclusion, I beg to thank most heartily Dr. John Murray for many acts of kindness and assistance during the progress of my Report, and for the friendly willingness he has always shown to meet my views. My thanks are also due to Mr. Wm. E. Hoyle for the attention and trouble he bestowed on my proof-sheets whilst on the Editorial Staff; and to Mr. James Chumley for similar good offices. To the memory of the late Sir Wyville Thomson, under whose direction this work was commenced, I would pay the tribute of sincere and personal esteem, repeating — not inaptly as it seems to me — the words used by the Danish naturalist, Abildgaard, in writing of his friend, the first great dredger, Otho Frederick Miiller, " Loquuntur Eius scripta cum mundo peritura." CONTENTS. I. CLASSIFICATION. PAGE I. History of the Classification of the Asteroidea ..... xvii II. Keasons for considering previous Classifications Invalid .... xxii III. A Classification of the Asteroidea based on Factors of Morphological Importance xxiii Synopsis of the Orders and Families of the Sub-class Euasteroidea . . . xl II. DESCRIPTION OF GENERA AND SPECIES. Order PHANEROZONIA . Family Archasterid.s . Subfamily Pararchasterin/E Genus Pararcliaster, Sladen . Pararchaster semisquamatus, n. sp. semisquamatus, var. occidentalis, nov. antarcticus, n. sp. spinosissimus, n. sp. pedicifer, n. sp. . armatus, n. sp. Genus Pontaster, Sladen Pontaster tenuispinus, Diiben and Koren tenuispinus, var. plati/nofa, nov. planeta, n. sp. hebitus, n. sp. limbatus, n. sp. oxyacanthus, n. sp. teres, n. sp. forcipatus, n. sp. . forcipatus, var. eclrinata, nov, mimicus, n. sp. pristinus, n. sp. venustus, n. sp. venustus, var. robusia, nov trullipes, n. sp. subtubercidatus, n. sp. Subfamily Plutonasterin^e Genus Dytaster, Sladen Dytaster spinosus, n. sp. exilis, n. sp. exilis, var. gracilis, nov. exilis, var. carinata, nov sp. 1 1 4 4 7 10 11 12 15 19 23 28 29 30 33 35 38 41 43 47 48 50 52 55 55 58 60 60 63 65 68 69 \'l THE VOYAGE OF H.M.S. CHALLENGER. Dijtaster madreporifer, n. sp. nobilis, n. sp. xquivocus, n. sp. . biserialis, n. sp. . inermis, n. sp. Genus Plutonaster, Sladen P/uto7iaster bifrons, Wyville Thomson, sp. bifrons, (?) juv. . marginatus, n. sp. rigidus, n. sp. rigidus, var. semiarmafa, nov. ambiguus, n. sp. . notatus, n. sp. abbreviates, n. sp. Subgenus Tethyaster, nov. . Plutonaster {Tethyaster) subinermis, Philippi, sp. ptarelii, Diiben and Koren, sp, Genus Lonchotaster, Sladen Lonchotaster tartareus, n. sp. forcipifer, n. sp. . Subfamily Pseudarchasterinj: . Genus Pseudarchaster, Sladen . Pseudarchaster discus, n. sp. tessellatus, n. sp. . intermedins, n. sp. Genus Aphrodit aster, Sladen . Apihroditaster gracilis, n. sp. Subfamily Archasterin.e Genus Archaster (Muller and Troschel), emend. Archaster typicus, Muller and Troschel Family Porcellanasteridje Subfamily Porcellanasterinj-: . Genus Porcellanaster, 'Wyville Thomson Porcellanaster cxruleus, Wyville Thomson caulifer, Sladen . tuberosus, Sladen . crassus, Sladen gracilis, Sladen . eremicus, n. sp. . Genus Styracaster, Sladen Styracasfer liorridus, Sladen armatus, Sladen . Genus Hyphcdaster, Sladen Hyi^lialaster hyalinus, Sladen diadematus, Sladen inermis, Sladen . planus, Sladen Genus Tlioracaster, Sladen . Thoracaster cylindratus, Sladen REPORT ON THE ASTEROIDEA. vu Subfamily Ctenodiscin.£ Genus Ctenodiscus, Miiller and Troschel Ctenodiscus corniculatus (Linck), Perrier australis, Lutken procurator, n. sp. Family Astropectinid.-e .... Subfamily Astbopectinin^: Genus Craspidaster, n. gen. Craspidaster hesperus, Miiller and Troschel, sp Genus Leptopty 'chaster, Smith . Leptoptychaster herguelenensis, Smith . arcticus, Sars, sp. arcticus, var. elongata, nov. antardicus, n. sp. Genus Moiraster, n. gen. Moiraster magnificus, Bell, sp. . Genus Astropecten, Linck Astropecten brasiliensis, Miiller and Troschel breoispinus, Sladen polyacanthus, Miiller and Troschel pectinatus, Sladen acanthifer, Sladen japonicus, Miiller and Troschel imbellis, Sladen . liermatophilus, Sladen irregularis, Linck pontopormis, Sladen zebra, Sladen zebra, var. rosea, Sladen . velitaris, von Martens granulatus, Miiller and Troschel . monacanthus, Sladen cingulatus, Sladen mesactus, Sladen Genus Psilaster, Sladen Psilaster cmdromeda, Miiller and Troschel, sp. acuminatus, n. sp. ■ cassiope, n. sp. . gracilis, n. sp. . patagiatus, n. sp. Genus Phozaster, Sladen Phoxaster pumilus, n. sp. Genus Bathyldaster, Danielssen and Koren . Bathybiaster loripes, n. sp. loripes, var. obesa, nov. . vexillifer, Wyville Thomson, sp. . Subfamily Luidiin^e .... Genus Luidia, Forbes Luidia aspera, n. sp. . PAGE 170 170 171 172 173 174 175 17.5 177 182 184 1S9 189 190 192 193 193 198 198 201 202 203 205 206 207 209 210 212 214 214 215 216 218 219 221 224 225 228 230 232 234 236 239 240 242 243 244 244 248 Vlll THE VOYAGE OF H.M.S. CHALLENGER. Luidia altcrnata (Say), Liitken limbata, n. sp. claihrata (Say), Liitken . ciliaris (Philippi), Gray . longispina, n. sp. africana, n. sp. . sarsii, Diiben and Koren forficifer, n. sp. . Family Pentagon asteridse Subfamily Pextagonastbrin^ . Genus Pentagonaster, Linck . Pentagonaster semilunatus, Linck granulans, Retzius, sp. . astrologorum (Muller and Troschel), Perrier patagonicus, n. sp. japonicus, n. sp. . lepidus, n. sp. arcuatus, n. sp. . Genus Calliaster, Gray Calliaster baccatm, n. sp. Genus Chitonaster, Sladen CJiitonaster cataphradus, n. sp. Genus Gnathaster, n. gen. Gnathaster meridionals, Smith, sp. elongatus, n. sp. . pilulatus, n. sp. . Genus Nyrnphaster, Sladen . Nyrnphaster symbolicus, n. sp. symbolicus, var. breviradiata, nov, bipunctus, n. sp. . protentus, n. sp. . albidus, n. sp. basUicus, n. sp. . Genus Paragonaster, Sladen . Paragonaster ctenipes, n. sp. ■ cylindratus, n. sp. Genus Ncctria, Gray . Nedria ocdlifera (Lamarck), Gray Subfamily Goniodiscin^! Genus Stellaster, Gray Stdlaster incei, Gray . princeps, n. sp. . Genus Leptogonaster, Sladen . Leptogonaster cristatus, n. sp. Subfamily Misiasterinj: Genus Mimaster, Sladen Mimasier tizardi, Sladen cognatus, n. sp. . KEPORT ON THE ASTEROIDEA. IX Family Antheneid^e .... Genus Anthenea, Gray Anthenca acuta, Perrier tuberculosa, Gray tuberculosa, (?) juv. Genus Hippasteria, Gray Hippasteria plana (Linck), Gray Family PentacerotidsE .... Genus Pentaceros, Linck Pentaceros dorsatus (Linne-), Perrier . turritus, Linck . productus, var. tuberata, nov. callimorphus, n. sp. Genus Pentacerop>sis, n. gen. . Petitaceropsis obtusatus, Bory de St. Vincent, s Genus Culcita, Agassiz Culcita novx-guinese, Miiller and Troschel Genus Asterodiscus, Gray Asterodiscus elegans, Gray Genus Choriaster, Liitken Clwriaster granulatus, Liitken . Family Gymnasteriidm .... Genus Gymnasteria, Gray Gymnasteria carinifera (Lamarck), von Martens Genus Porania, Gray .... Porania pidvillus (0. F. Miiller), Norman antarctica, Smith glaber, n. sp. spiculata, n. sp. . magellanica, Studer Genus Marginaster, Perrier . Marginaster fimbriatus, n. sp. . Genus Rhegaster, Sladen Rhegaster murrayi, Sladen Genus Lasiaster, n. gen. Lasiaster villosus, n. sp. Family Asterinw/E .... Subfamily Ganeriin.e .... Genus Cycethra, Bell .... Cycethra electUis, n. sp. nitida, n. sp. . pinguis, n. sp. . Genus Ganeria, Gray Ganeria falklandica, Gray Subfamily Asterininje .... Genus Patiria, Gray .... Patiria bellida, n. sp. . Genus Nepanthia, Gray Nepanthia brevis, Perrier, sp. . (ZOOL. OHALL. EXP. — PART LI. — 1888.) PAGE 338 339 340 340 340 341 341 342 343 346 346 347 347 350 351 351 352 353 353 354 354 355 356 357 358 360 360 360 362 363 364 365 367 368 371 372 374 376 376 377 379 380 382 383 384 384 385 386 387 THE VOYAGE OF H.M.S. CHALLENGER. Nepanthia maculata, Gray Genus Asterina, Nardo Asterina regularis, Verrill exigua (Lamarck), Perrier gunnii, Gray pectini/era (Miiller and Troschel), von Martens folium (Llitken), Agassiz.. cepheus (Miiller and Troschel), von Martens penicillaris (Lamarck), von Martens Subfamily Palmipedin^e . . Genus Palmipes, Linck . , Palmipes membranaeeus, Linck diaphanus, n. sp. . Order CRYPTOZONIA Family Linckiidm Subfamily Ch.etasterin2E Genus Chxtaster, Miiller and Troschel Chsdaster longipes (Retzius), Sars Subfamily Linckiin^; Genus Fromia, Gray . Fromia milleporella (Lamarck), Gray Genus Ophidiaster, Agassiz . Ophidiaster attenuatus, Gray . ophidianus (Lamarck), Agassiz cylindricus (Lamarck), Miiller and Troschel tuberi/er, n. sp. . . helicostichus, n. sp. . Genus Leiaster, Peters . ." Leiaster speciosus, von Martens Genus Linckia, Nardo . . Linckia guildingii, Gray — miliaris (Linck), von Martens pacifica, var. diplax (Miiller and Troschel) Genus Nardoa, Gray . Nardoa tuberculata, Gray Genus Narcissia, Gray Narcissia canariensis, d'Orbigny, sp. trigonaria, n. sp. Subfamily Metrodirin^; Genus Metrodira, Gray Metrodira subulata, Gray Family Zoroasterid/e Genus Zoroaster, Wyville Thomson . Zoroaster fulgens, Wyville Thomson — tenuis, n. sp. Genus Cnemidaster, n. gen. Cnernidaster wyvillii, n. sp. KEPORT ON THE ASTEROIDEA. XI Genus Pholidaster, Sladcn .. .. . Phulidaster squamaius, n. sp. . distincius, n. sp. . Family Stichasterid/e , Genus Stichaster, M tiller and Troschel Stichaster aurantiacus (Meyen), Verrill roseus (0. F. Miiller), Sars albulus (Stimpson), Verrill pohjplax, Miiller and Troschel, sp. felipes, n. sp. . . i polygrammus, n. sp. . . Genus Neomorphaster, n. gen. . ■. Neomorphaster etistichus, n. sp. Genus Tarsaster, n. gen. . . i Tarsaster stoichodes, n. sp. Family Solasterid.& . Subfamily Solasterin* . . < Genus Crossaster, Miiller and Troschel Crossaster papposus (Linck), Miiller and Troschel papposus, var. septentrionalis, nov. penicillatus, n. sp. Genus Rhipidaster, n. gen. Rhipidaster vannipes, n. sp. Genus Solaster, Forbes Solaster endeca (Retzius), Forbes paxillatus, n. sp. . — — regularis, n. sp. . subarcuatus, n. sp. . . tondatus, n. sp. . . .• Genus Lophaster, Verrill Lophaster furcifer (Diiben and Koren), Verrill ■ stellans, n. sp. . Subfamily KoRETHRASTERINiE Genus Korethraster, Wyville Thomson Korethraster hispidus, Wyville Thomson Genus Peribolaster, Sladen Peribolaster folliculatus, n. sp. . Family Pterasterid& .... Subfamily PterasterinjE ... Genus Pteraster, Miiller and Troschel Pteraster rnilitaris (0. F. Miiller), Miiller and Troschel rnilitaris, var. prolata, nov-, affinis, Smith rugatus, Sladen . stellifer, Sladen . semireticulatus, Sladen Genus Retaster, Perrier Retaster verrucosus, Sladen ^— peregrinator, Sladen PAGE 426 427 429 430 430 431 432 432 432 433 434 436 438 439 440 442 442 442 444 444 446 447 448 450 452 452 454 455 457 458 459 460 462 462 464 464 465 468 470 470 471 472 473 473 474 475 477 478 479 Xll THE VOYAGE OF H.M.S. CHALLENGER. Retaster gibber, Sladen insignis, Sladen . Genus Marsipaster, Sladen Marsipaster spinosissimus, Sladen hirsutus, Sladen . Genus Calyptraster, Sladen Calyptraster coa, Sladen Genus Hymenaster, Wyville Thomson Hymenaster nobilis, Wyville Thomson ■ formosus, Sladen . pergamentaceus, Sladen . sacculatus, Sladen echinulatus, Sladen carnosus, Sladen . glaucus, Sladen . vicarius, Sladen . pellucidus, Wyville Thomson ■ infernalis, Sladen ceelahis, Sladen . crucifer, Sladen . anomalus, Sladen latebrosus, Sladen ^ porosissinius, Sladen graniferus, Sladen geometricus, Sladen pullatus, Sladen . membranaceus, Wyville Thomson ■ — — ■ coecincdus, Sladen prsecoquis, Sladen . Genus Bentliaster, Sladen Bentliaster icyville-thomsoni, Sladen penicillatus, Sladen Subfamily PYTHOXASTERiNiE Genus Pythoaaster, Sladen Pythonaster murrayi, n. sp. Family EchinasteriDsE . Subfamily Acanthasterin^e Genus Acanthaster, Gervais . Acanthaster eehinites (Ellis and Solander), Liitken Subfamily Mithrodiixj: Genus Mifhrodia, Gray Mithrodia clavigera (Lamarck), Perrier Subfamily Echinasterin^e Genus Cribrella (Agassiz), Forbes Cribrella oculata (Linck), Forbes ornata, Perrier compada, n. sp. . obesa, n. sp. prxstans, n. sp. . REPORT ON THE ASTEROIDEA. Xlll Cribrella simplex, n. sp. simplex, var. granulosa, nov. sufflata, n. sp. , Genus Perknaster, n. gen. Perlmaster fuscus, n. sp. densus, n. sp. Genus Echinaster, Miiller and Troschel Echinaster spitwsus (Retzius), Miiller and Troschel — — ■ eridanella (Val. ), Miiller and Troschel spimdifer, Smith . Family^ Heliasterid.z: .... Genus Heliaster, Gray . . , Heliaster heliantlms (Lamarck), Dujardin and Hupe Family Pedicellasterid.e Genus Pedicellaster, Sars Pedicellaster scaler, Smitli hypernotius, n. sp. Family^ A steri/d^: .... Genus Asterias, Linne ( Asterias vesiculosa, n. sp. meridiondlis, Perrier perrieri, Smith . torquata, n. sp. . glomerata, n. sp. . rubens, Linne versicolor, n. sp. . amurensis, Liitken cunningliami, Perrier Subgenus Cosmasterias, nov. Asterias (Cosmasterias) tomidata, n. sp. sulcifera, Perrier, sp. . Subgenus Smilasterias, nov. . . Asterias (Smilasterias) scalprifera, n. sp. triremis, n. sp. Subgenus Hydrasterias, nov. Asterias (Hydrasterias) opliidion, n. sp. Subgenus Leptasterias (Verrill) Asterias (Leptasterias) miilleri, Sars, sp. ■ compta, Stimpson, sp. Subgenus Stolasterias, nov. Asterias (Stolasterias) gemmifera (Val.), Perrier, sp. tenuispina, Lamarck, sp. calamaria, Gray, sp. . volsellata, n. sp. stichantha, n. sp. ■ eustyla, n. sp. . ■ glacialis, O. F. Miiller, sp. africana, Miiller and Troschel, sp, TAGE 547 548 549 550 551 552 553 554 555 555 555 555 556 556 557 558 558 560 560 568 569 569 570 571 572 573 575 576 576 576 578 578 578 579 5S1 581 582 582 583 583 583 583 583 584 586 587 588 589 XIV THE VOYAGE OF H.M.S. CHALLENGER. Genus Calvaslerias, Perrier . Calvasterias dolidota, n. sp. Family Brisingwje Genus Labidiaster, Liitken . Labidiaster radiosus (Lov6n), annulatus, n. sp. Liitken Genus Odinia, Perrier Odinia pandina, n. sp. , Genus Brisiwja, Asbj0rnsen . Brisinga endecacnemos, Asbj0rnsen coronata, Sars vcrtirillata, n. sp. ericophora, n. sp. armillafa, n. sp. . membranacea, n. sp. discincta, n. sp. Genus Freyella, Perrier Freyella pennata, n. sp. poly enema, n, sp. echinata, n. sp. . fragilissima, n. sp. bracteata, n. sp. , dimorpha, n. sp. . remex, n, sp. tuberculata, n. sp. benthophila, n. sp. heroina, n. sp. attenuate/,, n, sp. . ^Genus Colpaster, n. gen. Colpaster scutigerida, sp. PAGE 589 590 592 594 595 595 597 598 601 603 604 604 606 608 611 613 614 618 621 623 626 629 632 635 638 641 643 645 647 648 III. CHOEOLOGY OF THE SPECIES. A. List of Stations and the Geographical Positions at which Asteroidea were Collected, with the Enumeration of the Species obtained at each Station. I. Challenger Expedition . . . . ." . . . .651 II. " Porcupine " Expeditions— i. 1869 Cruise ........... 675 ii. 1870 Cruise .......... 679 III. "Knight Errant" Expedition ......... 680 IV. " Triton " Expedition .......... 681 V. " Lightning " Expedition ......... 682 Table of the Distribution of the Genera of Asteroidea collected by the Challenger in the great Oceanic Areas, showing the number of Species by which each is known to be represented and the number of Species which are common to two or more Oceans ...... 683 REPORT ON THE ASTEROIDEA. xv B. Bathymetrical Range of the Species. I. Challenger Expedition — (1.) Littoral Zone ...'.*..... 687 (2.) Continental Zone . . . ". '. '. . . .090 (3.) Abyssal Zone ......... 692 II. "Porcupine" Expeditions — (1.) Littoral Zone ......... 695 (2.) Continental Zone ......... 695 (3.) Abyssal Zone ......... 696 III. " Knight Errant" Expedition— (1.) Littoral Zone ......... 697 (2.) Continental Zone ......... 697 (3.) Abyssal Zone ......... 697 IV. " Triton" Expedition— (1.) Littoral Zone ......... 698 (2.) Continental Zone ......... 698 (3.) Abyssal Zone ......... 698 Tablejshowing the known Bathymetrical Range of the Genera of Asteroidea collected by the Challenger in each of the great Oceanic Areas ........ 699 C. Nature of the Sea-bottom on which the Asteroidea collected by the Challenger were FOUND, WITH AN ENUMERATION OF THE SPECIES OCCURRING ON EACH DEPOSIT. I. Terrigenous Deposits — a. Gravel, Stones b. Coarse Gravel . c. Sand . d. Sand, Gravel e. Sand and Shells f. Rock . ti- Hard Ground, "Gravel ll . Hard Ground i. Mud . 3- Blue Mud k. Green Mud 1. Green Sand m. Red Mud n. Volcanic Mud 0. Greenish Volcanic Mud P- Volcanic Sand 1- Coral Mud r. Coral Reefs 702 702 703 704 704 704 705 705 705 706 707 709 709 709 710 711 711 712 XVI THE VOYAGE OE H.M.S. CHALLENGER. II. Abyssal Deposits — a. Globigerina Ooze b. Pteropod Ooze c. Diatom Ooze . d. Radiolarian Ooze c. Red Clay IV. LIST OF THE KNOWN SPECIES OF RECENT ASTEROIDEA V. SYSTEMATIC INDEX VI. INDEX TO AUTHORS ...... VII. EXPLANATIONS OF PLATES. VIII. MAP. PAGE 713 714 714 715 715 716 839 888 CLASSIFICATION. CONTENTS. I. History of the Classification of the Asteroidea. II. Seasons for Considering Previous Classifications Invalid. III. A Classification of the Asteroidea based on Factors of Morphological Importance. I. History of the Classification of the Asteroidea. The earliest attempt to classify the Asteroidea was made by Linck1 in 17.33, who published the first monograph devoted to starfishes, entitled : De Stellis marinis liber singularis. It is a work worthy of honour, and one which I hold in reverential estima- tion. Linck divided the "Stellas marinas" into two sections, named " Stellas fssas" and " Stellas integral " — the former characterised by the presence of an open channel or anibu- lacral furrow along the actinal surface of each of the lobes or rays of the body, the latter by the absence of these open channels. The Stellas fissas correspond to the Asteroidea of the present nomenclature ; the Stellas integras comprise the Ophiuroidea and the Comatulse. Linck divided the Stellas fissas, or section equivalent to the Asteroidea, into three classes, which were based on the number of the rays ; the first class embraced forms with less than five rays (all, by the way, abnormal or monstrous forms), the second forms with five rays, and the third all those with more than five rays. The classes were subdivided into genera, those included in the first and third being distinguished by the number of their rays, jg In the second class, however, the genera were characterised by the general form. This class contained seven genera, four of which are still maintained, viz., Pentagonaster, Pentaceros, Astropecten, and Palmvpes. It will be seen from the above that the basis of Linck's classification is purely artificial, the number of rays being the determinating factor. In 1766 Linne,2 in the 12th edition of his Systema Naturse, grouped all the Asteroidea, Ophiuroidea, and Comatulse known to him in one genus, Asterias, which he 1 De Stellis marinis liber singularis, Lipsiae, 1733. 2 Systema Nature?, Editio Duodeeima, Reformats, Holmia?, 1766. (ZOOL. CH-ALL. EXP. — part li. — 1888.) c XVlll THE VOYAGE OF H.M.S. CHALLENGER. divided into three sections, named Integra, Stellatss, and Radiatw. The representative of the first of these sections cannot now be recognised ; the second section is equivalent to the Asteroid ea, and comprised nine species ; the third section embraced the Ophiuroidea and Comatulse. Lamarck,1 in 1816, divided the Asterias of Linne (which he regarded as a family under the name of " Les Stellerides ") into four genera, which he named Comatula, Euryale, Ophiura, and Asterias, the last being equivalent to the section " Stellatse " of Linne, and to the Asteroidea of the present day. He divided the genus Asterias into two sections: (i.) "Asteries scutellees," and (ii.) " Asteries rayonne'es," the former embracing the species in which the length of the ray does not exceed that of the diameter of the disk, and the latter those in which the length of the ray is greater than the diameter of the disk. In 1830 de Blainville 2 proposed to divide the Asteroidea (which he ranked as a family, " Asteridea ") into five sections (" genres ou sous-genres "), which were characterised by the general form, and by the number of the rays. The composition of the different groups was more or less heterogeneous, and none of them have been maintained. In 1834 Nardo3 grouped a number of species known to him into five genera, two of which are maintained, viz., Aster ina and Linchia. In 1835 Agassiz4 divided the Asterias of Lamarck (which he ranked as a family in the order " Les Stellerides ") into nine genera, including fossil forms. Of the eight genera in which recent forms were included, two are maintained in statu quo (Ophidiaster and Cidcita), and the name of a third (Goniaster) is used in a restricted sense. The remaining genera correspond to groups named by Linck or Nardo. This partition of genera was based on a number of structural characters, and was the first approach towards a morphological classification of the group. In April 1840 Midler and Troschel5 published a preliminary classification of the Asteroidea, which they divided into three unnamed families, characterised by the presence or absence of an anal aperture, and by the quadriserial or biserial arrangement of the arnbu- lacral tube-feet. Fourteen genera were defined by means of structural characters, and representative species were cited. Two years later this work was expanded into a com- plete monograph on the group, entitled: System der Asteriden (Braunschweig, 1842), which has formed the basis of all subsequent systematic work on the Asteroidea. In the monograph the three families above indicated comprised eighteen genera, which included 140 species. 1 Hist. Nat. Anim. s. vert., t. ii. p. 530, 1816. 2 Diet. Sci. Nat., art. "Zoophyte," t. lx. p. 216, 1830; Manuel d'Actinologie ou de Zoophytologie, 1834, p. 235. 3 De Asteriis, Oken's Isis, 1834, Heft vii. p. 716. 4 Prodrome d'une Monographic des Radiaires ou Echinodermes, Mem. Soc. Set. Nat. Neuchatel, t. i. 1835, p. 190. 5 Monatsber. d. k. preuss. Akad. d. Wins. Berlin, April 1840, p. 102. REPORT ON THE ASTEROIDEA. xix In November and December 1840, and consequently only a few months later than Midler and Troscbel's preliminary paper above referred to, Gray1 published a synopsis of the genera and species of starfish. He divided the Asteroidea into two unnamed sections, one characterised by havinsr four rows of ambulacral tube-feet, the other having two rows. The first section contained one family, named " Asteriadae ; " the second section three families, named Astropectinidee, Pentacerotidae, and Asterinidae. These four families comprised forty-five genera and nine subgenera, which were briefly denned by structural characters. This classification was republished as a separate work2 in 18G6, with a few alterations and eight additional genera which had been described in the interim. In 1869 Perrier3 published a valuable memoir on the pedicellariae of Asterids and Echinids. In 1875 Perrier4 pointed out that the characters upon which the sectional divisions or families made by Mtiller and Troschel and by Gray were based were insufficient and unsatisfactory. He insisted upon the importance of the pedicellariaa as a classificatory factor, and he divided the group into two unnamed divisions by means of the character of the pedicellariae and of the disposition of the ambulacral tube-feet. The first division was characterised by the presence of pedunculate forficiform or forcipiform pedicellariae, and a usually quadriserial arrangement of the ambulacral tube-feet ; the second division by sessile, pincer-formed, or valvate pedicellariae, and a usually biserial arrangement of the ambulacral tube-feet. The first division contained only one family, the "Asteriadae;" the second division sis families, the Echinasteridaa, the " Linckiadaj," the Goniasteridae, the Asterinida3, the Astropectinidae, and the Pterasteridae. These seven families embraced forty-five genera (forty-two recognised in the text), a number of Gray's genera being absorbed or altered. Both families and genera were characterised by the structure of the skeleton. This work contains a careful revision of the species contained in the Museum d'Histoire Naturelle of Paris, and supplemented by those in the British Museum. In 1878 Viguier5 published an elaborate investigation on the Asterid skeleton, and propounded his discovery that in the mouth-ring of some starfishes the ambulacral plates were predominant, whereas in others the adambulacral plates were predominant. He also showed that a great range of variation occurred in the form of a plate which he called the " odontophore " (the basal interbrachial plate), as well as in the character of the mouth-plates, which he called teeth. 1 A Synopsis of the Genera and Species of the Class Hypostoma (Asterias, Linnaeus), Ann. and Mug. Nat. Hist, 1840, vol. vi. p. 175. 2 Synopsis of the Species of Starfish in the British Museum, London, 1866. 3 Recherches sur les PMicellaires et les Ambulacres des Asteries et des Oursins, Ann. Sci. Nat., 5e Sene, t. xii. 1869, p. 197. 4 Revision de la Collection de Stellerides du Museum d'Histoire Naturelle de Paris {Archives de Zool. exper., t. iv. 1875, pp. 265-450 ; t. v. 1876, pp. 1-104, 209-304). 5 Anatomie compared du Squelette des Stellerides (Archives de Zool. exper., t. vii. 1878, pp. 33-250, pis. v.-xvi.) XX THE VOYAGE OF H.M.S. CHALLENGER. On the basis of these researches Viguier proposed an amended classification of the Asteroidea. He divided the class into two sub-classes, named " Astdries ambulacraires " and " Asteries adambulacraires." The first characterised by the predominance of the ambulacra! plates in the mouth-ring, the presence of pedunculate forficiform or forcipiform pedicellarise, and the usually quadriserial arrangement of the ambulacral tube-feet ; the second sub-class characterised by the predominance of the adambulacral plates in the mouth-ring, by the presence of sessile, pincer-like, or valvulate pedicellarise, and by the almost constant biserial arrangement of the ambulacral tube-feet. The first sub-class was divided into three families, the " Asteriadae," Heliasteridse, and Brisingidaa. The second sub-class was divided into seven families, the Echinasteridas, " Linckiadaj," Goniasteridae, Asterinida?, Pterasterida?, Astropectinidse, and Archasteridse. The three families, Heliasterida?, Brisingidoe, and Archasterida? are additions to those given in Perrier's list. Perrier, however, included the family Brisingidse in his remarks, but did not include it in his synoptical table. The genera recognised by Perrier are accepted by Viguier, Metrodira, Nepanthia, and Brisinga being the only additions on his list. Viguier, however, defined the genera chiefly on the basis of his own investiga- tions on the skeleton, the characters of the odontophore and the mouth-plates being used by him as important factors in the classification. On these grounds several of the genera are placed by Viguier in different families and associations from those to which they were referred by Perrier. In 1879 Zittel1 published a classification having special reference to fossil forms. He divided the class Asteroidea into two orders — (1) the Ophiuridse, which was again divided into two sub-orders, the Euryalese and the Ophiurese ; and (2) the Stelleridas, which was subdivided into two sub-orders, the Encrinasteriad and the Asterias verse. The latter group — which comprises all the recent forms — was simply divided into two unnamed sections, characterised by the quadriserial and the biserial arrangement of the ambulacral tube-feet. In 1884 Perrier2 again discussed the question of classification, and pointed out that the structure of the mouth determined by Viguier, and the character of the ambulacral furrow, are morphologically correlated, the one dependent on the other, and that the modifications in the form of the pedicellarise stand in no definite connection with the structure of the ambulacral furrow and the mouth. From this he was led to discuss the relative taxo- nomic values of the structure of the ambulacral furrow and mouth on the one hand, and of the form of the pedicellariaa on the other. He decided in favour of the pedicel- larise, on the ground that they are in his opinion aborted rudiments of ancestral organs 1 llantlbuch der Pakeontologie, Miinchen, 1879, Band i. p. 437. 2 Memoire sur les Etoiles de mer recueillies dans la mer des Antilles et le Golfe de Mexique durant les expeditions de dragage faites sous la direction de M. Alexandre Agassiz (Nouv. Archives Mus. Hist, Nat., 2e Serie, t. vi. (1884) p. 134). REPORT ON THE ASTEROIDEA. xxi which were originally more important than at present. As the outcome of these consi- derations, Perrier relinquished the division of the Asteroidea into two sections or sub- divisions, and divided the class into four orders, which he considers may be characterised by the form of the pedicellariae. The following is Perrier's1 summary of his classification, given verbatim : — 1. FORCIPULAT.E. Pedicellaires pedoncules, droits ou croise's. Families : BeISINGID^E, PEDICELLASTERIDiE, ASTERIAD^E, HELIASTEMDiE. 2. SPINULOSES. Pedicellaires en pince, resultant d'une modification des piquants. Families : EchinasteriDyE, Pterasterid.e, AsterinidyE. 3. VALVAT^E. Pedicellaires valvulaires ou en saliere. Families : Linckiad/E, GoniasteridyE, AsteropsidyE. 4. PAXILLOS^E. Pedicellaires formes par un ossicule squelettique et les piquants qui le recouvrent. Families : Archasteridje, Astropectinhxe. On subsequent pages of the same memoir the names " Spinulosaj" and "Valvatae" are changed to " Echinulatae " z and " Valvulatae " 3 respectively, and the following additions are made to the families mentioned : — In the order Spinulosse, the families " Mithrodime " and Solasteridse are included,4 and in the order Valvatae (or Valvulatoe) the families " Gymnasteriadae," Pentacerotidae, Antheneidae, and Pentagonasteridae take the place5 of the Goniasteridae and Asteropsidas of the above list. Three pages further on another family, Goniopectinidae,0 is also referred to this order, Valvulatae ; but in the descriptive part of the memoir, the type (and only) genus, Goniopecten, is placed7 under the family Archas- teridae in the order Paxillosae. 1 Op. tit., p. 154. 2 Op. tit., pp. 167, 20G. 3 Op. tit., pp. 1G4, 1G7, 221. 4 Op. cit., p. 164. 5 Op. cit., p. 165. G Op. cit, p. 168. 7 Op. cit., p. 249. XXU THE VOYAGE OF H.M.S. CHALLENGER. II. Reasons for considering previous Classifications invalid. As the foregoing is the latest scheme of classification, and is, in fact, the embodiment of all that have gone before, I propose to examine briefly the fundamental characters upon which it is based. Perrier's classification is based upon the character of the pedicellarise. He considers that the pedicellarise furnish characters of the highest taxonomic value ; in other words, he regards them as class characters, upon the modifications of which divisions of ordinal rank may be made. He assigns to the pedicellarise this importance on the ground that they are the degenerated rudiments of organs whose functions were more important in the ancestral forms than those which pedicellarise now perform. Perrier states that the pedi- cellarise appear earlier in the embryo of Echinids than the spines. He regards them as more ancient organs. He rejects the view that they are modified forms of spines, as sug- gested by A. Agassiz. He considers that they furnish positive ordinal characters in the Echinoidea. He asserts that these statements are even more clearly applicable to the Asteroidea than to the Echinoidea. He regards the more complex forms of pedicellarise as older than the simpler forms, and believes that the forcipiform pedicellarise are older and more typical than the more simple forficiform pedicellarise. Confining my remarks to the Asteroidea, I venture to think that facts do not support any one of these statements, so far as that class is concerned. (1.) Eespecting the priority of appearance in the pedicellarise and spines, I may say that in no starfish embryo which I have examined have I found anything to warrant the assumption that pedicellarias appear before spines ; in fact, my observations indicate unequivocally that the spines are formed before the pedicellarias. In Asterias, a form which is crowded with pedicellarise when adult, and is one grouped by Perrier amongst what he considers (erroneously in my opinion) the oldest forms of Asteroidea, this is certainly the case. Neither has any other observer who has written upon the development of starfishes recorded, so far as I am aware, the appearance of pedicellarise before spines. (2.) As to whether pedicellarise are modified forms of spinelets, and as to whether the older forms of the organ are simpler or more complex than the more recent, I consider that those Asterids, which I believe to represent the most archaic forms, distinctly support the views, (i.) that the pedicellarias are modified spinelets, and (ii.) that the older forms of the organ were simpler and less complex than the more recent. As to the mode in which the more complex forms may have been evolved it is unnecessary to speculate here. The further outcome of the argument that the most complex form of pedicellarias indicates the most ancient organism would logically lead to the conclusion — although such an opinion is not definitely expressed by Perrier — that the Echinoidea are phylogenetically older than the Asteroidea, for I imagine that it will be generally admitted that the pedicellarias of Echinoidea are more complex than those of Asteroidea. Such a conclusion, I venture to REPORT ON THE ASTEROIDEA. xxiii believe, would not meet with general acceptance on other grounds, as the Asteroidea are by- many considered to represent a more archaic type than the Echinoidea. (3.) As to whether the pedicellariae furnish characters by which the four "orders" indicated by Perrier may be distinguished, I consider that they are insufficient and unsatisfactory ; and I would venture to say that in my opinion two of the orders in question would be more correctly described as defined by the character of their spinulation. I refer to the Spinuloses and the Paxillosae. In his diagnosis of the order Spinuloses (Echinulatae), Perrier distinctly states that the pedicellariae are simply formed of modified spinelets (op. cit., p. 206), and in that of the order Paxillosae (op. cit., p. 249) no mention whatever is made of pedicellariae ; in the abridged synopsis of the orders, however, given above, the Paxillosae are defined as characterised by pedicellariae, formed of an ossicle of the skeleton and the spinelets which cover it (op. cit., p. 154). The statements in the case of these two " orders " would seem to negative Perrier 's argument that pedicellariae are not modified spinelets and that they have nothing to do with those appendages. Furthermore, I fail to see that the characters invoked from the modifications in the form of the pedi- cellariae are of sufficient importance to indicate differences of an ordinal degree. Apart from the above considerations, which negative the view that the pedicellariae in the Asteroidea afford characters by which orders may be distinguished, I make bold to say that I am unable to regard either pedicellariae, or spines, or any other mere tegu- mentary appendages as furnishing characters of sufficient importance to warrant their employment as taxonomic factors of ordinal rank. Though I admit that pedicellariae do possess characters of a certain taxonomic value, I cannot regard them as characters either of primary or even of secondary importance in the great question of the division of a class. I may remark in passing that I do not consider the plate to which Viguier has unfor- tunately given the inappropriate name of " odontophore" to merit the importance which he has placed upon it. The plate in question, which is the basal plate of the interbrachial system, is pushed by development upon the first pair of adambulacral plates, or mouth- plates, and is moulded into form to a certain extent by these plates, its shape being largely dependent on the number of the rays and the character of the adambulacral plates. So far, however, as my own observations go, I am inclined to think that in not a few cases greater differences may be found to exist between the "odontophores," or, as I should prefer to call them, "basal interbrachials," of congeneric species, than between those of species of distinctly separate genera. The form of the plate appears to be extremely variable, and not to present characters of very great taxonomic value. III. A Classification of the Asteroidea based on Factors of Morphological Importance. Passing in review the various morphological features or fundamental points of struc- ture which are common to the whole class, the following appear to me to present char- xxiv THE VOYAGE OF H.M.S. CHALLENGER. acters whose differences indicate changes in structural detail resulting from modification and development, and show a relationship of subordination which warrant their being ranked as ordinal factors : — 1. The adaptation of the organism to subserve the functions of respiration and excretion. 2. The character of the ambulacral skeleton. 3. The character of the ambital skeleton. I will remark briefly on each of these topics, confining, however, my observations on the present occasion to the sub-class Euasteroidea, as the other constituents of the Asteroidea are fossil forms whose classification it is not my intention to discuss in this place. 1. The organs which Stimpson ' first named " papulae " (abactinal or dorsal water- tubes, A. Agassiz ; Hautkiemeu, Ludwig ; respiratory processes, Carpenter ; Kiemen- blaschen, Hamann ; tubules, Vogt and Yung ; brauchies lymphatiques, Cudnot ; dermal branchiae, Durham), which puncture the body-wall in the form of delicate transparent mem- branous cceca, permit an exchange by osmosis of fresh oxygenated fluid from without, and of the effete or carbonised fluid from within the body-cavity. According to Mr. Durham,2 they would also seem to permit of the passage of "scavenging amoeboid cells" and more or less solid particles. The papulae may be distributed over the whole body, or may be confined to a limited area. By means of their mode of occurrence, the Euasteroidea may be divided into two groups : in one the papulae are confined to the abactinal surface, and never pass beyond the boundary of the supero-marginal plates, and consequently do not occur in the lateral walls or on the actinal surface ; in the other group the papulae extend beyond the boundary of the supero-marginal plates, and occur in the lateral walls and on the actinal surface. The former of these groups may be called the Stenopneusia, the latter the Adetopneusia. I regard the first group (the Stenopneusia) as the older, and as indi- cating a simpler or less complex stage of organisation for the performance of the functions in question. Embryology supports this view, for at an early stage in the life history of an Adetopneusate Asterid no papulae are present; those first formed areconfined to the abactinal surface, and the earliest to appear are situated near the base of the ray. It is only at a later stage of growth that the papulae invade the lateral walls and the actinal surface. In other words, the members of the more highly developed group (the Adeto- pneusia) pass in the course of their development through a stage which represents the characters of the adult condition of the more primitive group (the Stenopneusia). 2. The ambulacral skeleton — by which I understand not only the ambulacral plates and their associated adambulacral plates, but also the correlated series of tube-feet — exhibits, 1 Proc Boston Soc. Nat. Hist., vol. viii. (1862), p. 2G1. 2 Proa. Roy. Soc, 1888, vol. xliii. p. 329. REPORT ON THE ASTEROIDEA. • XXV when the sub-class is viewed as a whole, two modes of growth ; in the one the production of parts is accelerated in relation to the growth of the starfish, in the other the production of parts is retarded, or proceeds pari passu with the general development of the skeleton. In the first group, the production of the ambulacral elements is so rapid and prolific in relation to the development of the starfish and the length of the ray, that the plate3 become crowded and crushed together, and their individual development, though not essen- tially reduced, is greatest in a plane at right angles to the axis of the ray, the plates beino- very short in the longitudinal direction of the ray, and crushed together in consequence of the action of growth-pressure. The segments may become so numerous, and the growth- pressure so great, that the normally single series of ambulacral tube-feet on each side of the median line may be crushed into a zigzag or alternating double series of tube-feet, thus pro- ducing a quadriserial arrangement of the tube-feet in each furrow. This character is secon- dary, however, in my opinion, and results from the degree to which the crushing is carried. I do not therefore consider the quadriserial disposition of the tube-feet to be of sufficient importance to define the primary divisions of a class as previous syste- matists have done. A further result of this growth-movement and crushing of parts in the ambulacral skeleton is found in the form and posture of the adambulacral plates, which assume a more or less crowded character, and their position becomes transverse. A secondary effect of this action on the adambulacral plates is the influence on the form of the mouth-plates, which is especially conspicuous in the actinostomial ring, the ambulacral elements being rendered the most prominent and define the mouth character, whilst the adambulacral elements are retarded, and have a relatively insignificant character. In consequence of the mode of its formation, I consider the form of the mouth a secondary character. I do not consider it to be of such importance as Viguier does, for although it is true that all Asterids in which the ambulacral skeleton is of the retarded type, or with its growth concurrent with the general development, have an adambulacral mouth, the occurrence of the ambulacral mouth is simply dependent on the degree to which the growth- pressure is extended, as well as the influence of other factors which determine the form of the adambulacral plates. In the second group, in which the production of parts in the ambulacral skeleton is retarded, or proceeds pari passu with the general development of the skeleton, there is little or no action of growth-pressure, the ambulacral and adambulacral plates are not diminished in the longitudinal direction of the ray by crushing, the ambulacral tube-feet are biserial in arrangement, and the adambulacral plates are most prominent in the actino- stomial ring, and define the character of the mouth. These two groups may be called the Leptostroteria and the Eurystroteria respectively. I consider the last mentioned of these two groups (the Eurystroteria) to be the older. Embryology supports this view, for at an early stage of a Leptostroterate Asterid (in which when adult the ambulacral tube-feet are quadriserial and the mouth ambulacral), the ambulacral skeleton shows no effects of growth- (ZOOL. CHALL. EXP. — PART LI. 1888.) d xxvi THE VOYAGE OF ELKS. CHALLENGER. pressure, the plates are long in the longitudinal direction of the ray, the ambulacral tube- feet are in regular biserial arrangement, and the mouth is defined by adambulacral plates ; it is not until a later stage that the plates become crowded by growth-pressure, that the ambulacral tube-feet are crushed into zigzag or alternating series, forming a quadriserial arrangement, and the mouth-ring is defined by the prominence of the ambulacral plates. It will thus be seen that the members of the more highly developed group (the Leptostroteria) pass in the course of their development through a stage which represents the character of the adult condition of the more primitive group (the Eurystroteria). It is of interest to note that this occurrence and action of growth-pressure in the ambulacral region of Asterids strikingly recalls the growth-pressure shown by Loven1 and by Duncan2 to occur in the ambulacra of the poly geminate forms of regular Echini. 3. The ambital skeleton is formed by the marginal plates and their supplementaries when present. I consider them to be one of the most important systems of plates in the body as determining form and superficial character. Two distinct modes of growth may be defined in the marginal plates of the Euasteroidea, by which the sub-class may be divided into two groups. In the first the marginal plates develop rapidly, and continue to increase in size throughout the life of the starfish, being usually in the adult state the most conspicuous series of plates present, and forming a prominent and massive marginal border. In the second group the marginal plates do not increase in size, or only to an insignificant extent ; in fact, their relative size in relation to the body is conspicuously diminished in the adult stage as compared with their proportions in the young stage. They are in con- sequence often quite inconspicuous in the adult starfish, and can only be discovered by means of dissection, and even then frequently with difficulty. The first of these modes of growth is characterised by the increscent or accelerated development of the marginal plates, and the second by the decrescent or retarded growth of the mar- ginal plates. The former group I call Phanerozonia, in consequence of the prominent and conspicuous character of the marginal plates ; the latter group Cryptozonia, in conse- quence of the insignificant and often hidden character of the marginal plates. I consider that the Phanerozonate group is older than the Cryptozonate, and that the latter indicates a divergence in character and a modification in form correlated with the development of other organic systems. Embryology supports this view of the relative age of the two groups, as in the early stages of the life history of a Cryptozonate Asterid, the marginal plates are large and conspicuous, forming a prominent phanerozonoid margin ; it is not until a later stage that the growth of the marginal plates is retarded, and that with the increase in size in the other plates of the test they then appear to be relatively decrescent, 1 Etudes sur les Echino'idees. K. SvensA: Vet.-Almd. Handl, Ed. xi. No. 7, pp. 23 et seq., 1874. 2 On the Anatomy of the Ambulacra of Recent Diadematidse, Journ. Linn. Soe. Lond. (Zool.), vol. xix. p. 95 ; On the Structure of the Ambulacra of some Fossil Genera and Species of Regular Echinoidea, Quart. Journ. Geol. Soc. Lond., 1885, vol. xli, p. 419. REPORT ON THE ASTEROIDEA. xxvii and at the adult stage have become more or less insignificant. It may hence be stated briefly that the members of the more highly developed group (the Cryptozonia) pass in the course of their development through a stage which represents the characters of the adult condition of the more primitive group (the Phanerozonia). Of the development of intermediate plates between the supero-marginal and infero- marginal series, and of the relative posture of the plates in these two series in the Crypto- zonate Asterids, it is unnecessary here to remark. From the foregoing considerations it will be seen that three of the most important class characters of the Asteroidea exhibit two modes of presentment, and that by means of any one of the three the Euasteroidea may be divided into two groups, one of which in each case is apparently older than the other. Now the divisions thus made are essentially equivalent in each of the three categories ; that is to say, the older group as determined by one of the characters corresponds to the older group determined by the other two, and similarly with the newer and more highlv developed group. The corresponding divisions in each of the three categories are not, however, exactly coequal when regarded independently, nor is this to be expected ; but the slight overlap or extension of one or other of the characters in the case of transitional groups is comparatively insignificant : in fact, the result is a "dove-tailing" or interlocking of the groups, which is perfectly natural and in accordance with what is found to occur in other branches of the animal kingdom. We have thus a natural morphological division of the sub-class into two sections or groups on the basis of three independent and im- portant structural factors, one group being more archaic or more primitive than the other. These groups I regard as true natural orders in the strictest sense of the term. To the first or older group I have given the name Phanerozonia, to the second and more highly developed group the name of Cryptozonia. I have selected the marginal plates for the name character of the orders in consequence of their importance in the Asterid skeleton as well as for the compara- tive ease with which their character may be observed in the superficial aspect of a starfish. 1 shall now proceed to define the families comprised in the two orders, in accordance with my views as to their morphological relations, and shall cite under each the consti- tuent genera. I have been led on structural grounds to establish subfamilies where the divergence of form indicated by the included genera demanded in my opinion this expression of their affinities. In many cases this presentment of the constitution of a group will be found to give a solidarity to the family and an intelligent view of its com- position, as well as to indicate the relationships of genera which otherwise would appear isolated and unnaturally placed. As an evolutionist, however, I regard the ideas of species, genera, families, and other divisions, as merely abstract particulars which con- stitute a synopsis or working key to our views of the affinities of organic forms — the XXVm THE VOYAGE OF H.M.S. CHALLENGER. details in an outline sketch of our attempt to put together the scattered evidences and to trace the lost records of the pedigree of life. The following is a summary of the classification I have adopted : — Class ASTEROIDEA. Sub-class EUASTEROIDEA, Sladen, 1886.1 Order I. PHANEROZONIA, Sladen, 1886. [Stenopneusia : Eurystroteria.] Euasteroidea with marginal plates large and highly developed in the adult. The supero-marginal and the infero-marginal plates contingent and with their axes usually in parallel planes. Papulas restricted to the abactinal area, circumscribed by the supero- marginal plates. Ambulacral plates well spaced and usually broad, the development of the ambulacral skeleton being retarded, or concurrent with that of the test generally. Actinostomial ring Avith the adambulacral plates prominent. Pedicellarise valvate, fora- minate, or excavate. Family I. ARCHASTERiDiE (Viguier, 1878), emend. Sladen, 1886. Phanerozonate Asterids with thick and massive marginal plates bearing spines or spini- form papillae. Abactinal skeleton with simple spiniform spicules, with pseudo-paxillse, or with true paxillae. Actinal interradial areas with plates bearing spinelets. Adam- bulacral plates large and not compressed. Proctuchous. Pedicellariae frequently present. No superambulacral plates. Subfamily 1. Paearchasteein^:, Sladen, 1886. Archasteridae with the papulae confined to a limited area at the base of the ray. Marginal plates more or less alternate. Actinal intermediate plates absent or very few in number. Genus 1. Pararchaster, Sladen. Genus 2. Pontaster, Sladen. Subfamily 2. Plutonasterisle, Sladen, 1886. Archasteridoe with the papulae distributed over the whole abactinal area. Marginal plates opposite. Actinal intermediate plates well developed. No definite medio-radial line of abactinal plates. Genus 1. Dy taster, Sladen. Genus 2. Plutonaster, Sladen. Genus 3. Lonchotaster, Sladen. 1 Date when the corresponding sheet of the descriptive portion of this work left my hands for press. KErORT ON THE ASTEROIDEA. xxix Subfamily 3. Pseudarchasterix.e, Slaclen, 1886. Archasteridse with a definite medio-radial line of abactinal plates. Abactinal plates arranged in longitudinal series parallel to the median series. Actiual intermediate plates well developed. Abactinal plates devoid of internal imbricating ridges. No pedi- cellariiE. Genus 1. Pseudarchastcr, Sladen. Genus 2. Aphroditaster, Sladen. Subfamily 4. Archasterin^e, Sladen, 18S6. Archasteridse with a definite medio-radial line of abactinal plates. Abactinal plates in oblique transverse rows on each side of the median series. Abactinal plates with special internal imbricating ridges. Actinal intermediate plates aborted, or very few in number. Pedicellariae present. Genus 1. Archaster (Miiller and Troschel), emend. Sladen. For a Synopsis of the Subfamilies and Genera included in the Family Archasteridse, see p. 2. Archasterid^; incertse sedis. The following recently described genera appear to me to belong to the family Archas- terinse, but the descriptions published do not enable me to indicate their position more definitely, and in some cases the reference to the family is doubtful. Genus Benthopecten, Verrill. Genus Blahiaster, Perrier. Genus Cheiraster, Studer. Genus Crenaster, Perrier. Genus Goniopecten, Perrier. Genus Pectinaster, Perrier. Genus Luidiaster, Studer. ? Genus Odontaster, Verrill. Remarks. — Judging from the description alone, I have great hesitation in placing Odontaster with the Archasteridse ; it may possibly prove to be more closely allied to the Pentagonasteridse : in any case it appears to be an annectant genus between the two groups. Family II. Porcellanasterid.-e, Sladen (1883), emend. 1886. Phanerozonate Asterids with thin and lamelliform marginal plates, naked or covered only with membrane. Abactinal area covered with membrane, beset with simple spini- xxx THE- VOYAGE OF H.M.S. CHALLENGER. ferous spicules or pseudo-paxilla3. A central epiproctal prominence, more or less defined, and frequently developed into an elongate tubular prolongation. Actinal interradial areas more or less extensive, paved with squamiform intermediate plates, covered with delicate membrane. Cribriform organs present. Adambulacral plates large, with a simple marginal armature, uniserially disposed. Subfamily 1. PoRCELLANASTERiNyE, Sladen, 1883. Porcellanasteridaa with cribriform organs highly developed ; localised. Actinal inter- radial areas, with squamiform plates covered with a simple membrane, and not traversed with fimbriated channels. Genus 1. Porcellanastcr, Wyville Thomson. Genus 2. Styracaster, Sladen. Genus 3. Hyphalaster, Sladen. Genus 4. Thoracaster, Sladen. Genus 5. Pseudaster, Perrier. Subfamily 2. Ctenodiscin^e, Sladen, 1886. Porcellanasteridse with a simplified form of cribriform organ on the margins of each pair of marginal plates. Actinal interradial areas traversed by fimbriated channels, in con- tinuation of the fasciolar or cribriform channels between the marginal plates. Genus 1. Ctenodiscus, Midler and Troschel. For a Synopsis of the Subfamilies and Genera included in the Family Porcellanas- teridaj, see p. 126. Family III. Astropectinid-E (Gray, 1840). emend. Phanerozonate Asterids with large marginal plates bearing spines or spiniform papillge. Abactinal skeleton with true columnar paxilla?. Actinal interradial areas small, inter- mediate plates when present spinose. Adambulacral plates short and more or less com- pressed. Superambulacral plates present. Aproctuchous. Pedicellarias rarely present. Subfamily 1. Astropectinin^e, Sladen, 1887. Astropectinidre with adambulacral plates touching the infero-marginal plates along the ray. Marginal and adambulacral plates not correspondent in length and number. Supero-marginal plates more or less well developed. Genus 1. Craspidaster, n. gen. Genus 2. Leptopty chaster, Smith. REPORT ON THE ASTEROIDEA. XXXI Genus 3. Moiraster, n.