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REPORT on the RADIOLARIA collected by H.M.S. CHALLENGER during the years 1873-1876.

FIRST PART.--PORULOSA.

EDITORIAL NOTES.

The Report on the RADIOLARIA by Professor Ernst Haeckel of Jena occupies the whole of the present Volume, the text being bound up in Two Separate Parts and the Plates in a Third Part. The Report forms Part XL. of the Zoological Series of Reports on the Scientific Results of the Expedition, and is the largest single Report of the series which has up to this time been published.

The Manuscript of the Systematic Part was written by Professor Haeckel in the English language, and was received by me in instalments on the 12th August 1884, 13th July and 4th December 1885, and 3rd June 1886. The Introduction was written in German and was translated into the English language by Mr. W. E. Hoyle of the Challenger Editorial Staff; the German text being received in instalments between the 15th July 1886, and the 25th January 1887.

The Challenger Naturalists found the representatives of this group of animals to be universally distributed throughout ocean waters, and their dead remains to be nearly equally widely distributed over the floor of the ocean, the relative abundance and the species differing, however, with change of locality, and their abundance or variety being intimately connected with some of the most interesting and intricate problems of general oceanography.

It was a fortunate circumstance that so distinguished a Naturalist, with such an intimate knowledge of the RADIOLARIA, should have been willing to undertake the laborious examination and description of the extensive collections made during the Expedition. Professor Haeckel has devoted ten years of his life to this work, and this Report sets forth the results of his labours, on the conclusion of which he will be congratulated by all Naturalists. The entire literature of the RADIOLARIA is completely recorded, and the older species redescribed, so that the Report is a complete Monograph, which will be an invaluable aid to all future Investigators.

THE

VOYAGE OF H.M.S. CHALLENGER.

#ZOOLOGY.#

PREFACE

The significance of the Radiolaria in regard to the relations of life in the ocean has been increased in a most unexpected manner by the discoveries of the Challenger. Large swarms of these delicate Rhizopoda were found not only at the surface of the open ocean but also in its different bathymetrical zones. Thousands of new species make up the wonderful Radiolarian ooze, which covers large areas of the deep-sea bed, and was brought up from abysses of from 2000 to 4000 fathoms by the sounding machine of the Challenger. They open a new world to morphological investigation.

When ten years ago I accepted the enticing invitation of Sir Wyville Thomson to undertake the investigation of these microscopic creatures, I hoped to be able to accomplish the task with some degree of completeness within a period of from three to five years, but the further my investigations proceeded the more immeasurable seemed the range of forms, like the boundless firmament of stars. I soon found myself compelled to decide between making a detailed study of a selection of special forms or giving as complete a survey as possible of the varied forms of the whole class; and I decided upon the latter course, having regard both to the general plan of the Challenger Reports, and to the interests of our acquaintance with the class as a whole. I must, however, confess at the close of my work that my original intention is far from having been fulfilled. The extraordinary extent and varied difficulties of the undertaking must excuse the many deficiencies.

The special examination of the Challenger collection was for the most part completed in the summer of 1881; I collected its results in my Entwurf eines Radiolarien-Systems auf Grund von Studien der Challenger-Radiolarien . Since the manuscript of this preliminary communication was completed only a few days before my departure for Ceylon, and since I was unable to correct the proofs myself, several errors have crept into the Prodromus Systematis Radiolarium included in it. These have been corrected in the following more extensive working out of it. Even at that time I had distinguished 630 genera and more than 2000 species; but on the revision of these, which I undertook immediately on my return from India, this number was considerably increased. The total number of forms here described amounts to 739 genera and 4318 species; of these 3508 are new, as against 810 previously described. In spite of this large number, however, and in spite of the astonishing variety of the new and marvellous forms, the riches of the Challenger collection are by no means exhausted. A careful and patient worker who would devote a second decade to the work, would probably increase the number of new forms by more than a thousand; but for a really complete examination, the lifetime of one man would not suffice.

The richest source of the Challenger material is the Radiolarian ooze of the central Pacific Ocean . This remarkable deep-sea mud consists for the greater part of well-preserved siliceous shells of Polycystina . Not less important, however, especially for the study of the ACANTHARIA and PHAEODARIA, are the wonderful preparations stained with carmine and mounted in Canada balsam on the spot by Dr. John Murray. One such preparation often contains twenty or thirty, sometimes even fifty new species. In many of these preparations the individual parts of the unicellular organism are so well preserved that they show clearly the characteristic peculiarities of the legions and orders. Since the material for these preparations was taken with the tow-net, not only from the surface of the sea but also from different bathymetrical zones, it furnishes valuable conclusions regarding the chorology, as well as the physiology and morphology of the group. For many new discoveries I am indebted to the study of such preparations, of which I have examined about a thousand from 168 different Stations . In addition to these about 100 bottles were handed to me, containing partly bottom-deposits, partly tow-net gatherings.

Sir Wyville Thomson, who directed the investigations of the Challenger with so much devotion, and only partly saw its results, has laid me under a deep debt of obligation; not less is this the case, however, with his successor, Dr. John Murray. I am especially indebted to both gentlemen for the freedom they have allowed me in the carrying out of my work, and especially for the permission to include a description of all known Radiolaria in the Challenger Report, which has thus become a second edition many times enlarged of my Monograph published in 1862. Since all previous literature of the subject has been consulted and critically revised, it is hoped that this Report will form a useful foundation for future investigations. All names of sufficiently described Radiolaria published during the first half century of our knowledge of the class , are inserted in alphabetical order in the index at the end of this work.

In addition to the treasures of the Challenger, my own collection of Radiolaria has yielded many new forms whose description is here included. On my journeys to the Mediterranean , I have given special attention to these delicate microscopic organisms for more than thirty years. Besides the various points on the Mediterranean, the Atlantic Ocean at the Canaries yielded many interesting new forms; whilst my voyage across the Indian Ocean, from Aden to Bombay, in November 1881, thence to Ceylon and back by Socotra in March 1882, was still more productive. In particular, some extended excursions which I had the opportunity of making from Belligemma and Matura gave me an insight into the rich treasures of the Indian Ocean.

Most important, however, as regards the knowledge of the Indian Radiolaria, are the collections which Captain Heinrich Rabbe of Bremen has so beautifully preserved during his many voyages through that region. In the neighbourhood of Madagascar and the Cocos Islands more especially, and also in the Sunda Archipelago, he met with large swarms of Radiolaria, among which were many new and remarkable forms. These were of special value for completing the chorology, and the more so since the course of the Challenger in the Indian Ocean lay very far to the southwards. I will therefore take this opportunity of repeating my best thanks to Captain Rabbe for the friendly donation of his valuable collection.

The Radiolarian fauna of the North Atlantic Ocean, which was previously but little known and only slightly increased by the investigations of the Challenger, received a valuable increase from the interesting collections made by Dr. John Murray on various expeditions to the Faer?e Islands . A large number of new Radiolaria were captured in the Faer?e Channel, partly at the surface of the Gulf Stream, partly at various depths, and the proof was thus furnished that at certain points in the North Atlantic Ocean Radiolaria are very richly developed. I am further indebted to Dr. John Murray for the free use of this important material as well as for much other assistance in the carrying out of my work. Another rich source of Radiolaria I found in the alimentary canal of pelagic animals from all seas. Medusae, Siphonophorae, Salpae, Pteropoda, Heteropoda, Crustacea, &c., which live partly at the surface of the sea and partly at various depths, and swallow large masses of Radiolaria, often contain numbers of their shells well-preserved in their intestine. The alimentary canal of Fishes and Cephalopods too, which live upon these pelagic animal frequently contains considerable quantities of siliceous shells; and another newly discovered source has been found in the coprolites of the Jurassic period, which consist largely of Radiolarian skeletons.

In the investigation of this complicated system of organisms, I have endeavoured on the one hand to give accurately the forms and dimensions of the species observed, and on the other hand to present a survey of the relationships of the different genera and families; and in this I have striven especially to combine the phylogenetic aims of the natural system with the essentially artificial divisions of a practical classification. Being, however, a conscientious supporter of the theory of descent, I can of course lay no stress upon the value of the categories, which are here distinguished as Legions, Orders, Families, Genera, &c. All these artificial systematic grades I regard as of merely relative value; and from the same cause I attach no importance to the distinction of all the species here described; many of them are probably only developmental stages, and like my predecessors I have determined their boundaries on subjective grounds. In the systematic working out of so much material one always runs the risk of doing either too much or too little in the way of creating species; but in the light of the theory of descent this danger is of no consequence.

In the carrying out of this extensive task the friendly aid of Dr. Reinhold Teuscher of Jena was of the greatest benefit to me; at my request he was at the trouble of making a large number of accurate drawings with the camera lucida, and he also undertook a long series, amounting to some 8000, accurate micrometric measurements, which were of the greatest value in the attempt to settle the important question of the constancy of the various species; I have alluded to this in a note at the conclusion of the Report . My best thanks are due to Dr. Teuscher for the patient and careful manner in which he discharged these tedious tasks.

FIRST PART.

GENERAL INTRODUCTION-- PAGE

SYSTEMATIC PART, 1

SECOND PART.

Note on the Dimensions and Measurements, 1760

ADDENDA, 1761

ERRATA, 1763

INDEX, 1765

GENERAL INTRODUCTION.

ANATOMICAL SECTION.

A SKETCH OF OUR KNOWLEDGE OF THE ORGANISATION OF THE RADIOLARIA IN THE YEAR 1884.

The extent of the Radiolaria, as limited by the above definition, which I have made as compact as possible, differs in several important respects from that allowed to the group by all previous diagnoses. The shortest expression of its scope might perhaps be:--Rhizopoda with central capsule and calymma; for the most important character of the Radiolaria, and that by which they are distinguished from all other Rhizopoda, is the differentiation of the unicellular body into two principal parts of equal importance and their separation by a constant capsule-membrane.

The division of the Radiolaria into two subclasses and four legions , I sought to establish in 1883 in a communication on the Orders of the Radiolaria . As a believer in the theory of descent, I regard all the systematic arrangements of specialists as artificial, and all their divisions as subjective abstractions, and hence I shall be guided in the establishment of such groups as subclasses, legions, orders, &c., by purely practical considerations, especially by the desire to give as ready a survey as possible of the complex multitude of forms .

In most of the more modern delineations of the Radiolaria the central capsule is regarded as the "cell proper" and its membrane as the "cell-wall." The following facts are opposed to the correctness of this interpretation:--1. In most Radiolaria the exoplasm is clearly different from the endoplasm, and the former is more voluminous than the latter. 2. In all Radiolaria the division of labour is so carried out between the central capsule and the extracapsulum, that the physiological significance and independence of both principal parts of the cell is almost equally great. 3. It is only in the ACANTHARIA that the formation of the skeleton takes place within the central capsule; in all the other three legions it is quite independent of it.

Regarding the general system of ground-forms compare my Generelle Morphologie . The ground-forms there proposed and systematically defined have, however, found but little acceptance ; but having now, twenty years after their publication, anew carefully revised and critically studied them, I can find no sufficient reason for abandoning the principles there adopted. On the contrary the study of the Challenger Radiolaria during the last ten years, with its incomparable wealth of forms, has only confirmed the accuracy of my system of ground-forms. The customary treatment of these in zoological and botanical handbooks is quite insufficient.

For a complete system of the geometrical ground-forms and their relation to promorphological classification, see Gener. Morphol., Bd. i. pp. 555-558.

"What are these?" Mrs. Harbinger asked.

He shook his head, as if he had given up all hope of understanding anything.

"The butler put them in my hands," he murmured.

"Upon my word, Mrs. Harbinger," spoke up Mrs. Croydon, seeming more offended than there was any apparent reason for her to be, "you have the most extraordinary butler that ever existed."

Mrs. Harbinger threw out her hands in a gesture by which she evidently disclaimed all responsibility for Graham and his doings.

"Extraordinary! Why, he makes my life a burden. There is no mistake he cannot make, and he invents fresh ones every day. Really, I know of no reason why the creature is tolerated in the house except that he makes a cocktail to suit Tom."

"Dat ees ver' greet veertue," Count Shimbowski commented genially.

"I do not agree with you, Count," Miss Wentstile responded stiffly.

The spinster had been hovering about the Count ever since his accident with the teacup, apparently seeking an opportunity of snubbing him.

"Oh, but I die but eef Mees Wentsteele agree of me!" the Count declared with his hand on his heart.

Mrs. Croydon in the meanwhile had taken the letters from the hand of Barnstable, and was looking at them with a scrutiny perhaps closer than was exactly compatible with strict good-breeding.

"Why, here is a letter that has never been posted," she said.

Mr. Harbinger took the whole bundle from her hand.

"I dare say," was his remark, "that any letter that's been given to Graham to mail in the last week is there. Why, this letter is addressed to Christopher Calumus."

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