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REATISE ON Z n:;rs:: : -"- ; A TREATISE ON ZOOLOGY A TREATISE ON ZOOLOGY EDITED BY E. RAY LANKESTER M.A., LL.D., F.R.S. HONORARY FELLOW OF EXETER COLLEGE, OXFORD ; CORRESPONDENT OF THE INSTITUTE OF FRANCE ; DIRECTOR OF THE NATDRAL HISTORY DEPARTMENTS OF THE BRITISH MUSEUM PART III THE ECHINODERMA BY F. A. BATHEE, M.A. ASSISTANT IN THE GEOLOGICAL DEPARTMENT OF THE BRITISH MUSEUM ASSISTED BY J. W. GREGORY, D.Sc. LATE ASSISTANT IN THE GEOLOGICAL DEPARTMENT OF THE BRITISH MUSEUM PROFESSOR OF GEOLOGY IN THE UNIVERSITY OF MELBOURNE AND E. S. GOODRICH, M.A. ALDRICHIAN DEMONSTRATOR OF ANATOMY IN THE UNIVERSITY OF OXFORD Reprint A.ASHER & CO. Amsterdam 1964 A TREATISE ON ZOOLOGY EDITED BY E. RAY LANKESTER M.A., LL.D., F.R.S. HONORARY FELLOW OF EXETER COLLEGE, OXFORD ; CORRESPONDENT OF THE INSTITUTE OF FRANCE ', DIRECTOR OF THE NATURAL HISTORY DEPARTMENTS OF THE BRITISH MUSEUM PART III THE ECHINODERMA BY F. A. BATHER, M.A. ASSISTANT IN THE GEOLOGICAL DEPARTMENT OF THE BRITISH MUSEUM ASSISTED BY J. W. GREGORY. D.Sc. LATE ASSISTANT IN THE GEOLOGICAL DEPARTMENT OF THE BRITISH MUSEUM PROFESSOR OF GEOLOGY IN THE UNIVERSITY OF MELBOURNE AND E. S. GOODRICH, M.A. ALDRICHIAN DEMONSTRATOR OF ANATOMY IN THE UNIVERSITY OF OXFORD LONDON ADAM & CHARLES BLACK 1900 Exclusive Agents for U.S.A. STECHERT-HAFNER SERVICE AGENCY, INC. 31 East 10th Street New York, New York 10003 Sole agents for India: Today & Tomorrow' s Book Agency, 22-B/5, Original Road, Karol Bagh, New Del hi-5 PREFACE THE present volume is the " Third Part" in order of a com prehensive treatise on Zoology, which has been for some time in preparation under my editorship. In this treatise each of the larger groups of the Animal Kingdom is to be described by a separate author; whilst, as far as possible, uniformity in method and scope of treatment is aimed at. The authors are, for the most part, graduates 'of the University of Oxford, though it may not be possible to maintain this limitation in future sections of the work. The general aim of the treatise is to give a systematic exposition of the characters of the classes and orders of the Animal Kingdom, with a citation in due place of the families and chief genera included in the groups discussed. The work is addressed to the serious student of Zoology. To a large extent the illustrations are original. A main purpose of the Editor has been that the work shall be an independent and trustworthy presentation, by means of the systematic survey, or taxonomic method, of the main facts and conclusions of Zoology, or, to speak more precisely, of Animal Morphography. The treatise will be completed in ten parts of the size of the present one. It will at once be apparent that this limitation necessitates brevity in treatment which, however, will not, it is believed, be found inconsistent with the fulfil ment of the scope proposed or with the utility of the work vi PREFACE to students. The immediate publication of the following parts may be expected :— Part I. Introduction and the Protozoa. Part II. General Discussion of the Metazoa—The Pori- fera — The Hydromedusae — The Scypho- medusae—The Anthozoa—The Ctenophora. Part III. The Echinoderma (the present volume). Part IV. The Mesozoa—The Platyhelmia—The Nemer- tini. These parts will be issued, without reference to logical sequence, as soon as they are ready for the press. In accord ance with this procedure, which to some extent evades the injustice of making an author, whose work is finished, wait for publication until other more tardy writers have completed their tasks, the present volume, which is Part III, is the first to make its appearance. The following authors have undertaken portions of the work:—Professor Poulton, F.R.S., M.A. Oxon.; Professor Weldon, F.R.S., M.A. Oxon.; Professor Benham, D.Sc, M.A. Oxon.; Mr. G. C. Bourne, M.A. Oxon.; Mr. G. H. Fowler, M.A. Oxon.; Professor Minchin, M.A. Oxon.; Mr. F. A. Bather, M.A.Oxon.; Professor J. W. Gregory, D.Sc; and Mr. E. S. Goodrich, M.A. Oxon. E. RAY LANKESTER. February 1900. CONTENTS CHAPTER VIII GENERAL DESCRIPTION OP THE ECHINODERMA l CHAPTER IX THE PELMATOZOA—CYSTIDEA. 38 CHAPTER X THE PELMATOZOA—BLASTOIDEA 78 CHAPTER XI THE PELMATOZOA—CRINOIDEA 94 CHAPTER XII THE PELMATOZOA—EDRIOASTEROIDEA 205 CHAPTER XIII THE ELEUTHEROZOA—HOLOTHURIOIDEA 217 CHAPTER XIV THE ELEDTHEROZOA—STELLEROIDEA 237 CHAPTER XV THE ELEUTHEROZOA—ECHINOIDEA 282 INDEX TABULAR STATEMENT SHOWING THE SYSTEMATIC POSITION OF THE ECHINODERMA. ANIMALIA. GRADE I. PROTOZOA. „ II. METAZOA. BRANCH A. BRANCH B. PARAZOA. ENTEROZOA. GRADE I. (of the Enterozoa). ENTEROCOELA = COELENTERA. GRADE II. (of the* Enterozoa). COELOMOCOELA = COELOMATA. PHYLA (of the Coelomocoela). ECHINODERMA, MOLLUSCA, APPENDICULATA, PLATYHELMIA, VERTEBRATA, etc. CHAPTER VIII. THE ECHINODERMA.1 PHYLUM ECHINODERMA. GRADE A. PELMATOZOA. CLASS I. CYSTIDEA. „ II. BLASTOIDEA. „ III. CllINOIDEA. „ IV. EDRIOASTEROIDEA. GRADE B. ELEUTHEROZOA. CLASS I. HOLOTHURIOIDEA. „ II. STELLEROIDEA. „ III. ECHINOIDEA. General Features.—This is one of the best characterised and most distinct Phyla of the Animal Kingdom. Nearly all the living animals included in it, such as the sea-urchin (Echinoid), starfish (Asteroid), brittle-star (Ophiuroid), sea-cucumber (Holothurian), sea-lily (stalked Crinoid), or feather-star (free Crinoid), can readily be distinguished through their possession of a radial symmetry, in which the number five is dominant, of a sub-epidermic skeleton com posed of calcium carbonate, with a characteristic micro-structure resembling trellis-work, and of a system of sacs, canals, and tubes that carry water through the body, especially by means of five radial canals from which small branches called podia are given off to the exterior. The extinct forms known as Blastoidea and Edrio asteroidea appear to have had a similar organisation; and the same statement maybe made of most of the Cystidea, another extinct class. It is true that there are recent forms in which the quinqueradial symmetry or pentamerism is obscure; but, on the whole, it is so marked a feature that the early zoologists, and notably Cuvier, placed the Echinoderma together with the Coelentera in a sub- 1 By F. A. Bather, M.A. ECHINODERMA—GENERAL DESCRIPTION kingdom Radiata. The presence of a gut distinct from the body- cavity (coelom) is alone enough to mark the superiority of Echino- derm organisation, as was first insisted on by Leuckart. The resemblance of certain Holothurians to the Gephyrea is but super ficial and secondary; the above-mentioned characters form sufficient distinction. Examples of the Classes.—Within the Echinoderma is great diversity of organisation. Between the worm-like, semi-gelatinous Holothurian, Synapta, living in the mud of the shore, and the stalked Pentacrinus of the depths of the sea, or the brittle-star of the rock-pools, there might well seem an impassable barrier. Taking typical examples of the various classes, let us note the more obvious differences. In an ordinary Holothurian (e.g. Holo- thuria, Cucumaria, Fig. IV. 4, p. 231) the body is cucumber-shaped, with a mouth at one end and an anus at the other; round the mouth is a ring-canal of the water-vascular system, and from it are given off five radial canals, running below the surface of the flexible integument and sending podia to the exterior; two of the avenues (ambulacra) of podia run along that surface of the body which is away from the ground and may be called " dorsal"; the " ventral" surface, containing the other three ambulacra, is often flattened to form a kind of walking sole. A Holothurian has no arms or projecting rays, but its mouth is surrounded by a circlet of tentacles, often branched, retractile at will, and serving to collect food. A Regular Sea-urchin (e.g. Echinus, Cidaris, Figs. VII., XVII. pp. 290, 303) resembles a Holothurian in being without projecting rays; but it is more spherical in shape, with a rigid test, and moves with its mouth towards the sea-floor, and with its anus at the opposite pole of the body. In a Heart-urchin (e.g. Spatangus, Fig. XLV p. 324), which moves through and swallows mud and sand, the body has become obliquely elongate, i.e. with the long axis at an angle of 36° to the position it occupies in a Holothurian ; the mouth has moved a little forward, and the anus" has moved down from the top of the body to its lower surface, so that both mouth and anus lie on the under surface, at either end of the long axis. In Echmoids, the radial water-vessels are beneath the test ("hypo- thecal") and stretch from the oral to the anal pole. In a Starfish the mouth is in the centre of the under surface, while the anus is almost in the centre of the upper surface, but is absent in a few forms; the body, encased in a yielding theca, is either markedly pentagonal in outline or star-shaped ; in the latter case a central disc maybe distinguished from the "arms." The number of arms varies from five (e.g. Asterias rubens, Figs. L, IV pp. 240,242)to over forty (e.g. Heliaster). The radial water-vessels, one to each arm, lie in a groove on the oral surface (« epithecal") and are fringed by podia, which do not pass on to the aboral surface at all. An ECHINODERMA—GENERAL DESCRIPTION 3 Ophiuroid (Fig. XIII. p. 261) resembles a starfish in which there is a sharp distinction between arms and disc; the mouth is on the under surface, but there is no anus. Whereas the arms of a star fish are merely extensions of the body, containing the generative glands and processes from the stomach, those of an Ophiuroid con tain only blood-vessels, water-vessels, and nerves, and,, being themselves used as locomotor organs, have a stout internal skeleton of separate ossicles, worked on one another by well-developed muscles, but have less developed podia; they are nearly always five, and unbranched except in Astrophytidae (Fig.