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REPORT

ON THE

SCIENTIFIC RESULTS

OF THE

VOYAGE OF H.M.S. CHALLENGER

DURING THE YEARS 1873-76

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.

REGIUS PROFESSOR OF NATURAL HISTORY IN THE UNIVERSITY OF EDINBURGH

DIRECTOR OF THE CIVILIAN SCIENTIFIC STAFF ON BOARD

AND NOW OF

JOHN MURRAY

ONE OF THE NATURALISTS OF THE EXPEDITION

Zoology—Vol. XVIII.

FIRST PART

Published by Order of Her Majesty's Government

PRINTED FOR HER MAJESTY'S STATIONARY OFFICE

AND SOLD BY

LONDON:—EYRE & SPOTTISWOODE, EAST HARDING STREET, FETTER LANE

EDINBURGH:—ADAM & CHARLES BLACK

DUBLIN:—HODGES, FIGGIS, & CO.

1887

Price (in Two Parts, with a Volume of Plates) £5, 10s.

CONTENTS.

Report on the Radiolaria collected by H.M.S. Challenger during the years
1873-1876.

By Ernst Haeckel, M.D., Ph.D., Professor of Zoology in the University of Jena.

FIRST PART.—PORULOSA.

(SPUMELLARIA AND ACANTHARIA.)

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 (from 1834 to 1884) is completely recorded, and the older species (both living and fossil) redescribed, so that the Report is a complete Monograph, which will be an invaluable aid to all future Investigators.

|John Murray.

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 (in the autumn of 1876) 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 (Jenaische Zeitschr. f. Naturw., Bd. xv., 1881). 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 (especially of the smaller ones) 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 (Stations 265 to 274). This remarkable deep-sea mud consists for the greater part of well-preserved siliceous shells of Polycystina (Spumellaria and Nassellaria). Not less important, however, especially for the study of the Acantharia and Phæodaria, are the wonderful preparations stained with carmine and mounted in Canada balsam on the spot by Dr. John Murray. One such preparation (e.g., from Station 271) 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 (compare § 240). 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 (from 1834 to 1884), 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 (an account of which is given in the introduction to my Monograph of the Medusæ), 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 (in the winter of 1866-67) 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 (at the southern extremity of Ceylon) 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 Færöe Islands (on the "Knight Errant" in 1880 and on the "Triton" in 1882). A large number of new Radiolaria were captured in the Færöe 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. Medusæ, Siphonophoræ, Salpæ, 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 (p. 1760). My best thanks are due to Dr. Teuscher for the patient and careful manner in which he discharged these tedious tasks.

The figures of new species of Radiolaria (about 1600 in number) which appear in the atlas of one hundred and forty plates accompanying this Report, were nearly all drawn with the camera lucida, partly by Mr. Adolph Giltsch and partly by myself. The names of the genera which appear at the bottom of the plates have in many cases been changed since they were printed off, as may be seen from the explanations which accompany them. Had it been possible to complete the examination of the material before the plates were commenced this might have been avoided, and in many cases a better selection of figures might have been made. All the drawings have been made upon the stone by the practised hand of Mr. Adolph Giltsch, in his usual masterly manner, and his lithographic work, which has lasted fully ten years, is the more valuable since he has himself microscopically studied the greater part of the species figured. The fact that the atlas presents so full a picture of the marvellous wealth of form of the Radiolaria is especially due to his lively interest in the work, to his unwearying care, and to his morphological acuteness. May it be the means of inducing many naturalists to study more deeply this inexhaustible kingdom of microscopic life, whose endless variety of wonderful forms justifies the saying—Natura in minimis maxima.

CONTENTS.

GENERAL INTRODUCTION.

ANATOMICAL SECTION.

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

Chapter I.—THE UNICELLULAR ORGANISM.

(§§ 1-50.)

1. Definition of the Radiolaria.—Radiolaria are marine Rhizopoda, whose unicellular body always consists of two main portions, separated by a membrane; an inner Central capsule (with one or more nuclei) and an Extracapsulum (the external calymma, which has no nucleus, and the pseudopodia); the endoplasm of the former and the exoplasm of the latter are connected by openings in the capsule-membrane. The central capsule is partly the general central organ of the Radiolarian cell, partly the special organ of reproduction, since its intracapsular protoplasm, along with the nuclei embedded in it, serves for the formation of flagellate spores. The extracapsulum is partly the general organ for intercourse with the outer world (by means of the pseudopodia), partly the special organ of protection (calymma) and nutrition (sarcomatrix). The majority of Radiolaria develop also a skeleton for support and protection, which presents the utmost variety of form, and is generally composed of silica, sometimes of an organic substance (acanthin). The Radiolarian cell usually leads an isolated existence (Monozoa vel Monocyttaria); only in a small minority (of one legion) are the unicellular organisms united in colonies or cœnobia (Polyzoa vel Polycyttaria).

2. The Two Subclasses of the Radiolaria.—The systematic catalogue of the Radiolaria, which forms the second part of this Report, and is brought up to the year {ii}1884, contains 20 orders, 85 families, 739 genera, and 4318 species. The consideration that but a small proportion of the ocean his yet been investigated renders it likely, however, that even this large number does not include the half of the recent species. The great progress which our knowledge of the organisation of the Radiolaria has made, by means of comparative study, renders it possible to arrange this enormous mass of forms in four main divisions or legions, and these are again related in pairs, so that two divisions of the highest rank or subclasses are constituted, the Porulosa (or Holotrypasta) and Osculosa (or Merotrypasta).

3. Porulosa or Holotrypasta.—The subclass Porulosa or Holotrypasta includes the two legions, Peripylea or Spumellaria, and Actipylea or Acantharia, which agree in the following constant and important characters:—(1) The Central Capsule is primitively a sphere, and retains this homaxon form in the majority of the species. (2) The Membrane of the central capsule is everywhere perforated by very numerous minute pores, but possesses no larger principal aperture (osculum). (3) The Pseudopodia radiate in all directions and in great numbers from the central capsule, passing through its pores. (4) The Equilibrium of the floating unicellular body is in most Porulosa pantostatic (indifferent) or polystatic (plural-stable), since a vertical axis is either absent, or, if present, has its two poles similarly constituted. (5) The Ground-forms of the skeleton are therefore almost always either spherotypic or isopolar-monaxon, very rarely zygotypic. The two legions of the Porulosa are distinguished mainly by the skeleton of the Spumellaria (or Peripylea) being siliceous, never centrogenous, nor composed of acanthin, whilst in the Acantharia (or Actipylea) it is always centrogenous and made up of acanthin; hence in the former the nucleus is always central, in the latter always excentric.

4. Osculosa or Merotrypasta.—The subclass Osculosa or Merotrypasta includes the two legions Monopylea or Nassellaria, and Cannopylea or Phæodaria, which agree in the following constant and important characters:—(1) The Central Capsule is originally monaxon (ovoid or spheroidal) and retains this ground-form in most of the species. (2) The Membrane of the central capsule possesses a single large principal aperture (osculum) at the basal pole of the vertical main axis. (3) The Pseudopodia radiate from a stream of sarcode which passes out from the central capsule only on one side, namely, through the principal aperture. (4) The Equilibrium of the floating body is {iii}monostatic or unistable, since the two poles of the principal axis are always more or less different from each other. (5) The Ground-forms of the skeleton are, therefore, for the most part grammotypic (centraxon) or zygotypic (centroplan), rarely spherotypic. The two legions of the Osculosa are distinguished chiefly by the principal opening (osculum) being closed by a porous plate (porochora with its podoconus) in the Nassellaria (or Monopylea), and by a radiate cover (operculum with its astropyle) in the Phæodaria (or Cannopylea).

5. The four Legions of Radiolaria.—The four principal groups of Radiolaria, to which we have given the name "legions," are natural units, since the most important peculiarities in the structure of the central capsule are quite constant within the limits of the same legion, and since all the forms in the same legion may be traced without violence to the same phylogenetic stem. The four legions are, however, related to each other, in so far as they all exhibit those characters which distinguish the Radiolaria from other Protista. The two which compose the Porulosa (§ 3) seem somewhat more nearly related to each other than to the two which make up the Osculosa (§ 4). When, however, the attempt is made to bring them all into a phylogenetic relationship, it undoubtedly appears that the Spumellaria (or Peripylea) are the primitive stem, out of which the other three have been developed as independent branches. All three have been derived, probably independently, from the most ancient stem-form of the Spumellaria, the spherical Actissa.

6. Peripylea or Spumellaria.—Those Radiolaria which we call "Peripylea" on account of the constitution of their central capsule, or "Spumellaria" on account of the nature of their skeleton, are separated from the other three legions of the class by the combination of the following constant characters:—(1) The Membrane of the central capsule is single and evenly perforated all over by innumerable fine pore-canals, but without any larger principal opening (osculum). (2) The Nucleus always lies centrally in the Spumellaria monozoa and is serotinous, for it divides only at a later period into the nuclei of the spores; in the Spumellaria polyzoa it is precocious, and divides early into many small nuclei. (3) The Pseudopodia are exceedingly numerous and distributed evenly over the whole surface of the central capsule. (4) The Calymma contains no phæodium. (5) The Skeleton is seldom wanting, is never centrogenous, and is always siliceous. (6) The Ground-form of the central capsule is originally spherical (often modified); that of the skeleton is also spherical or, in the majority of cases, derived in different ways from the sphere.

7. Actipylea or Acantharia.—These Radiolaria which we call "Actipylea" on account of the constitution of their central capsule, or "Acantharia" from the formation of their skeleton, are separated from the other three legions by the combination of the following constant characters:—(1) The Membrane of the central capsule is single and {iv}perforated by numerous fine pore-canals, which are regularly distributed in series or groups, but without a larger principal opening (osculum). (2) The Nucleus is always excentric and generally precocious, since it divides early by a peculiar process of budding into numerous small nuclei. (3) The Pseudopodia are very numerous and distributed regularly in groups (or series united into a network). (4) The Calymma contains no phæodium. (5) The Skeleton is generally present, always centrogenous, and composed of acanthin. (6) The Ground-form of the central capsule is originally spherical (often modified), that of the skeleton polyaxon (often modified).

8. Monopylea or Nassellaria.—Those Radiolaria which we call "Monopylea" from the formation of their central capsule, or "Nassellaria" from the nature of their skeleton, are distinguished from the other three legions of the class by the combination of the following constant characters:—(1) The Membrane of the central capsule is single, and has only one large principal opening (osculum) at the basal pole of the vertical main axis; this osculum is closed by a perforated lid (porochora or operculum porosum) from which there arises within the central capsule a peculiar cone of threads or pseudopodia (podoconus). (2) The Nucleus is usually excentric and is always serotinous, since it only divides at a comparatively late period into spore-nuclei. (3) The Pseudopodia are not very numerous and arise by division of a single stem or bundle of threads of sarcode, which issues from the porochora. (4) The Calymma contains no phæodium. (5) The Skeleton (very rarely absent) is never centrogenous, but always extracapsular and siliceous. (6) The Ground-form of the central capsule is always monaxon (with a vertical allopolar main axis), originally ovoid, often modified; that of the skeleton is also generally monaxon, often modified (triradial or bilateral).

9. Cannopylea or Phæodaria.—Those Radiolaria which we call "Cannopylea" from the constitution of their central capsule, or "Phæodaria" on account of their peculiar phæodium, are distinguished from the other three legions by the combination of the following characters:—(1) The Membrane of the central capsule is double, consisting of a strong outer and delicate inner capsule, and has only one principal opening (osculum) at the basal pole of the vertical main axis; this osculum is closed by a radiate cover (astropyle or operculum radiatum), from the centre of which arises an external tubular spout (proboscis). Occasionally a few small accessory openings (parapylæ) are present besides the principal opening. (2) The Nucleus lies centrally or subcentrally in the capsule (in the vertical main axis), and is serotinous, inasmuch as it only divides at a late period into spore-nuclei. (3) The Pseudopodia are usually very numerous and arise from a thick sarcomatrix, formed by the spreading out of a thick stem of sarcode, which issues from the astropyle. (4) The Calymma always contains a phæodium or peculiar voluminous excentric mass of pigment. (5) The Skeleton (very rarely absent) is never centrogenous, always extracapsular and formed of a silicate of carbon. (6) The {v}Ground-form of the central capsule is always monaxon (with a vertical allopolar main axis) and generally spheroidal; that of the skeleton is very varied.

10. Synopsis of the Subclasses and Legions:—

11. Individuality of the Radiolaria.—Like other Protozoa the Radiolaria are unicellular organisms, the whole fully developed organisation of which falls under the category of a single cell, both morphologically and physiologically. Since this view is based upon the composition of the individual body out of two different morphological elements, nucleus and protoplasm, it is at once justified in the case of the majority of Radiolaria, in which the plasmatic body encloses only a single nucleus (the so-called "Binnen-Bläschen"); such is the case in all the Spumellaria monozoa, Nassellaria and Phæodaria. This aspect of the case might appear doubtful in those Radiolaria in which the simple primary cell-nucleus divides early into numerous small secondary nuclei, as is the case in the Spumellaria polyzoa and most Acantharia. Strictly speaking, the multinucleate central capsule should in such cases be regarded as a syncytium; but since the individual unity of the unicellular organism is as clearly defined in these precocious multinuclear Radiolaria as in the ordinary serotinous forms, the former must be considered unicellular Rhizopods just as are the latter. This mode of regarding {vi}the case is the more necessary, inasmuch as the early division of the nucleus has no further influence upon the organisation. Just as in many other classes of the Protista there are monozootic (solitary) and polyzootic (social) forms, so also in the Radiolaria there are in addition to the ordinary monozootic or monobious forms certain families in which colonies or cœnobia are formed by the association of individuals; this distinction may be expressed by the terms "Monocyttaria" and "Polycyttaria."

12. Morphological Individuality.—From the morphological standpoint the individuality of the unicellular elementary organism is obvious in the ordinary solitary Radiolaria (Monobia), and is to be so regarded that the whole body with all its constituent parts, and not merely the central capsule, is to be regarded as a cell. Naturally the xanthellæ or yellow cells (§§ 76, 90), which as independent algæ live in symbiosis with many Radiolaria, must be excluded. The unicellular organisation of the Radiolaria is further to be distinguished from that of the other Protista, inasmuch as an internal membrane (capsule-membrane) separates the central (medullary) from the peripheral (cortical) portion. In the cœnobia of the social Radiolaria (or Polycyttaria), the morphological individuality persists only as regards the medullary portions of the aggregated cells (the individual central capsules), while the cortical portions fuse completely to form a common extracapsulum. Hence in these Spumellaria polyzoa two different stages of morphological individuality must be distinguished, the Cell as a Morphon of the first stage, and the Cœnobium as a Morphon of the second stage.

13. Physiological Individuality.—From the physiological standpoint also the individuality of the unicellular organism is immediately obvious in the case of the ordinary solitary Radiolaria (Monobia); as in other Protista it fulfils all the functions of life by itself alone. This physiological individuality of the monobious Radiolarian cell is furthermore not influenced by the xanthellæ, which live as independent algæ in symbiosis with many Radiolaria; even though these often by the production of starch assist in the {vii}nourishment of the Radiolaria, yet they are by no means indispensable to them. On the other hand, the physiological individuality offers more complicated relations in the social Radiolaria (Polycyttaria) which live united in colonies or cœnobia. Here the actual Bion (or the fully developed physiological individual) is not represented by the individual cells, but by the whole multicellular cœnobium, which in each species has a definite form and size. In these cœnobia, which are usually spherical or cylindrical jelly-like masses, several millimeters in diameter, numerous cells are so intimately united that only their medullary portions (the central capsule with the endoplasm) remain independent; the cortical portions (calymma and exoplasm) on the contrary uniting into a common extracapsulum. This discharges, as a whole, the functions of locomotion, sensation, and inception of nutriment, while the separate central capsules act in the main only as reproductive organs (forming spores) and partly also as the central organs of metastasis (digestion). Each cœnobium may also be regarded as a polycyttarium, i.e., a "multicellular Radiolarian," whose numerous central capsules represent so many sporangia or spore-capsules.

14. Monocyttaria and Polycyttaria.—In the majority of the Radiolaria each unicellular organism passes its individual life in an isolated condition (as a Monocyttarium). Only in a part of the Spumellaria numerous unicellular individuals are united into societies which are regarded as cœnobia or colonies (Polycyttaria). This is the case in three different families belonging to the Peripylea, in the Collozoida (without a skeleton, Pl. 3), the Sphærozoida (with a Beloid skeleton, Pl. 4), and the Collosphærida (with a Sphæroid skeleton, Pls. 5-8). All three families of Polycyttaria (or social Radiolaria), agree in their mode of forming colonies, since the central capsules of the social individuals remain separate and lie in a common jelly-like mass, which is formed by the fusion of their extracapsulum. The chief part of the voluminous colonies, which attain a diameter of several millimetres (sometimes more than 1 cm.), and are generally spherical, ellipsoidal or cylindrical, consists therefore of the jelly-like calymma, and this is penetrated by a sarcoplegma, to whose meshes all the individual organisms contribute by means of the pseudopodia, which radiate from their sarcomatrix. A further peculiarity in which the social Spumellaria differ from the solitary consists in the fact that the former are precocious and the latter serotinous in the division of the nucleus (§ 64). Whilst in the solitary or monozootic Spumellaria the middle of the central capsule is occupied by the simple nucleus, and this divides only at a late period (immediately before the formation of spores) into the numerous spore nuclei, in the colonial or polyzootic Spumellaria this division takes place very early, and the middle of each central capsule is usually occupied by an oil-globule.

15. The Central Capsule and Extracapsulum.—The special peculiarity of the unicellular Radiolarian organism, by which it is clearly distinguished from all other Rhizopoda (and indeed from most other Protista), is its differentiation into two separate chief constituents, the central capsule and extracapsulum, and the formation of a special membrane which separates them. This, the capsule-membrane, is not to be compared with an ordinary cell-membrane, as an external layer, but rather to be regarded as an internal differentiated product. The extracapsulum or external (cortical) portion of the body is in most Radiolaria more voluminous than the central capsule or inner (medullary) portion. The exoplasm of the former (the cortical or extracapsular protoplasm) is emphatically different from the endoplasm of the latter (the medullary or intracapsular protoplasm). Besides the most important vital processes are distributed by division of labour so completely between them that they appear most distinctly co-ordinated. The central capsule is on the one hand the general central organ of the "cell-soul" for the discharge of its sensory and motor functions (comparable to a ganglion-cell), on the other hand the special organ of reproduction (sporangium). The extracapsulum, also, is not less significant, since on the one hand its calymma acts as a protecting envelope to the central capsule, as a support to the pseudopodia, and a foundation for the skeleton or a matrix for the development of the shell, and on the other hand its pseudopodia are of the utmost importance as peripheral organs of movement and sensation as well as of nutrition and respiration. The central capsule and the extracapsulum are therefore to be regarded both morphologically and physiologically as the two characteristic co-ordinated principal parts of the unicellular Radiolarian organism.

16. The Malacoma and Skeleton.—Whilst the division of the unicellular organism into central capsule and extracapsulum is undoubtedly the most important character of the Radiolarian organism, the development of a skeleton of peculiar and most varied form is of very striking significance. This skeleton is always a secondary product of the cell, but is always anatomically so independent, and so clearly marked off from the soft parts or malacoma, that it seems advisable to regard both separately in a general morphological survey. The skeleton stands in a different relation to each of the two principal constituents of the malacoma. Only in the Acantharia is it centrogenous and developed from the central capsule outwards. In the other three legions the skeleton never arises in the centre of the capsule; in the Nassellaria and Phæodaria it is always extracapsular; in the Spumellaria it is also outside the central capsule originally, but afterwards becomes often surrounded by it, and finally lies in most cases partly within and partly without the central capsule. The chemical basis of the skeleton in the Acantharia is the curious acanthin (an organic substance allied to chitin), in the Phæodaria a silicate of carbon, and in the Nassellaria and Spumellaria silica.

17. Ground-Forms of the Radiolaria (Promorphology).—The ground-forms of the Radiolaria exhibit a greater variety than those of any other class in the organic world, greater indeed than is to be found in all the remaining groups together. For every conceivable ground-form which can be defined in the system of promorphology is actually present in the Radiolaria; their skeleton exhibits, as it were, in material existence, certain geometrical ground-forms which are found in no other organisms. The cause of this unexampled richness in different forms lies chiefly in the static relations of the Radiolaria, which swim freely in the sea, partly also in the peculiar plasticity of their protoplasm and the material of their skeletons.

18. The Principal Groups of Geometrical Ground-Forms.—The great variety of the geometrical ground-forms which are actually realised in the variously shaped bodies of the Radiolaria, renders it desirable to classify these in as small a number as possible of principal groups and a larger number of subdivisions. As extensive principal groups four at least must be distinguished; the Centrostigma or Sphærotypic, the Centraxonia or Grammotypic, the Centroplana or Zygotypic, and the Acentrica or Atypic. The natural centre of the body, about which all its parts are regularly arranged, is in the first group a point (stigma), in the second a straight line (principal axis), in the third a plane (sagittal plane), in the fourth a centre is of course wanting.

19. The Centrostigma or Sphærotypic Ground-Forms.—The first group of geometrical ground-forms, here distinguished as sphærotypic or the centrostigma, is undoubtedly the most important among the Radiolaria, inasmuch as if these be considered monophyletic, it must be the original one from which all the other ground-forms have been derived. The common character of all these sphærotypic ground-forms is that their natural centre is a point (stigma); thus there is no single principal axis (or protaxon) such as is characteristic of the two following groups. The sphærotypic ground-forms are subdivided into two important smaller groups, the spheres (Homaxonia) and the endospherical polyhedra (Polyaxonia). The spherical ground-forms, fully developed in the central capsule and calymma of Actissa and the Sphæroidea as well as in many Acantharia, present no different axes; all possible axes passing through the centre of the body are equal (Homaxonia). In the endospherical polyhedra, on the contrary, numerous axes (three at least) may be distinguished, which are precisely equal to each other and different from all the remaining axes (Polyaxonia). If the extremities of these axes, or the poles, which are all equidistant from the common centre, be united by straight lines, a polyhedral figure is produced whose angles all lie in the surface of the sphere. According as the poles of the axes are at equal, subequal, or at different distances from each other, we may divide the endospherical polyhedra into regular, subregular and irregular. (See Gener. Morphol., Bd. i. pp. 404-416.)

20. The Centraxonia or Grammotypic Ground-Forms.—The second principal group of organic ground-forms, here called grammotypic or centraxonia, is characterised by the fact that a straight line (gramma) or a single principal axis (protaxon) forms the natural centre of the body. This important and extensive group is divided into two subgroups, those with one axis (Monaxonia) and those with crossed axes (Stauraxonia); in the latter different secondary transverse or cross-axes may be distinguished, but not in the former. In the Monaxonia, therefore, every transverse section of the body perpendicular to the principal axis is a circle, in the Stauraxonia, on the contrary, a polygon. The Monaxonia are further subdivided into two groups, in one of which the two poles of the principal axis {xi}are equal and similar (Isopolar), in the other of which they are different (Allopolar); in the former the two halves of the body, which are separated by the equatorial plane (or the largest transverse plane, perpendicular to the principal axis), are equal, in the latter unequal. Among the isopolar uniaxial ground-forms (Monaxonia isopola) may be mentioned the ellipsoidal, spheroidal, lenticular, &c.; to the allopolar uniaxial forms (Monaxonia allopola) belong the conical, hemispherical, ovoid, &c. In the same way the pyramidal ground-forms with crossed axes are divisible into two groups, according as the two poles of the principal axis are equal or not. The ground-form of the former is the double pyramid, that of the latter the single pyramid. Both the double and the single pyramids may again be subdivided, each into two important lesser groups, the regular and the amphithect. In the first division the equatorial plane of the double and the basal plane of the single pyramid is a regular polygon (square, &c.), whilst in the other division it is an elongated or amphithect polygon (rhombus, &c.); the crossed axes are equal in the former, unequal in the latter. (See Gener. Morphol., Bd. i. pp. 416-494.)

21. The Centroplana or Zygotypic Ground-Forms.—The third principal group of ground-forms includes those which are bilaterally symmetrical in the ordinary sense, or zeugitic or zygotypic; the natural centre of their body is a plane. These forms are the only ones in which the distinction between right and left is possible, since their body is divided by the median plane (planum sagittale) into two symmetrical halves (right and left). In all these zeugites the position of every part is determined by three axes perpendicular to each other, and of these three dimensive axes two are allopolar, one is isopolar. The two unlike poles of the principal (or longitudinal) axis are the oral and aboral, the two unlike poles of the sagittal (or vertical) axis are the dorsal and ventral; the two similar poles of the frontal (or transverse) axis, however, are the right and left. This important group of zeugitic or bilateral forms may also be divided into two clearly distinct lesser groups, the Amphipleura and the Zygopleura. In the Amphipleura (or bilaterally radial ground-forms) the "radial two-sided" body is produced by modification of a regular pyramid (as Spatangus from Echinus), and hence is composed of several (not less than three) antimeres. In the Zygopleura (or bilaterally symmetrical ground-forms) on the other hand, the bodies consist of two antimeres (as in all the higher animals, Vertebrata, Arthropoda, &c.). (See Gener. Morphol., Bd. i. pp. 495-527.)

22. The Acentrica or Atypic Ground-Forms.—Among the acentrica or anaxonia are included all those ground-forms which are absolutely irregular, and in which neither a definite centre nor constant axes can be distinguished (e.g., most Sponges). These quite irregular ground-forms are very rare among the Radiolaria, but nevertheless there may be referred to them the amœboid central capsule of some Colloidea (Collodastrum, p. 27, Pl. 3, figs. 4, 5) among the Spumellaria, the irregular shells of many Collosphærida {xii}(Pl. 8, fig. 2), and the absolutely irregular shells of the Phorticida and Soreumida among the Larcoidea. (See Gener. Morphol., Bd. i. p. 400.)

23. The Subsidiary Groups of Geometrical Ground-Forms.—The four natural principal groups of ground-forms, which have just been defined according to the nature of the centre of their bodies, may be divided again into numerous subsidiary groups, defined by the relations of the constant axes and the two poles of each axis, as well as by the number of the axes and the differentiation of the secondary with respect to the principal axis. The most important of these subsidiary groups into which the principal ones are immediately divided are the following:—(1) The Centrostigma (or sphærotypic) are divided into spheres (Homaxonia) and endospherical polyhedra (Polyaxonia). (2) The Centraxonia (or grammotypic) into uniaxial (Monaxonia) and those with crossed axes (Stauraxonia); among the former of these may be distinguished the isopolar (phacotypic) and the allopolar (conotypic); among the latter the double and single pyramids. (3) The Centroplana (or bilaterals) are divided into amphipleura (or bilaterally radial) and zygopleura (or bilaterally symmetrical). (4) The Acentrica (or Anaxonia) or absolutely irregular ground-forms, present no special subdivisions.

24. The Spherical or Homaxon Ground-Form.—The spherical is the only absolutely regular ground-form, since only in it are all axes which pass through the centre equal; it is very often realised among the Radiolaria, especially in the Spumellaria and Acantharia, where it furnishes the common original ground-form, but it is often to be seen in the shells of many Phæodaria (in most Phæosphæria); on the other hand, it is never found among the Nassellaria. Geometrical spheres, in the strict sense of the term, are only to be found among the Spumellaria and Acantharia, namely, in the central capsule of many Collodaria (Pls. 1, 2) and all Sphæroidea (Pls. 11-30) as well as many Acanthometra and Acanthophracta (Pls. 128-138). Nevertheless, speaking generally, one includes those central capsules and skeletons which have been distinguished here as endospherical polyhedra. (On these ground-forms see Gener. Morphol., Bd. i. pp. 404-406.)

25. The Endospherical Polyhedral Ground-Form.—The endospherical polyhedron or polyaxon ground-form naturally follows the spherical or homaxon. Under it are included all polyhedra whose angles fall in the surface of a sphere; this ground-form is especially common among the Spumellaria, especially in the shells of Sphæroidea, but is also found among the Acantharia (especially in the Astrolophida and Sphærophracta), as well as among the Phæosphæria (in most genera of the Orosphærida, Sagosphærida, and Aulosphærida). Strictly speaking, all those lattice-shells which have {xiii}been incorrectly called "spherical" belong to this category, for they are none of them true spheres in the geometrical sense (like the central capsules of the Sphæroidea), but rather endospherical polyhedra, whose angles are indicated by the nodal points of the lattice shell, or the radial spines which spring from them. These endospherical polyhedra may be divided into three groups, the regular, subregular, and irregular. Of regular polyhedra, properly so-called, it may be shown geometrically that only five can exist, namely, the regular tetrahedron, cube, octahedron, dodecahedron, and icosahedron. All these are actually manifested among the Radiolaria, although but seldom. Much more common are the subregular endospherical polyhedra, e.g., spherical lattice-shells with regular hexagonal meshes of equal size; they are never exactly equal nor perfectly regular, but the divergences are so insignificant that they escape superficial observation (Pl. 20, figs. 3, 4; Pl. 26, figs. 1-3). On the contrary in the irregular endospherical polyhedra the meshes of the lattice-sphere are more or less different in size and often in form also (Pl. 28, figs. 4, 8; Pl. 30, figs. 4, 6). The five truly regular polyhedra require separate notice on account of their importance. (See Gener. Morphol., Bd. i. p. 406.)

26. The Regular Icosahedral Ground-Form.—The ground-form whose geometrical type is the regular icosahedron (bounded by twenty equilateral triangles) is rarely exemplified, but it occurs among the Phæodaria in the Circoporid genus Circogonia (Pl. 117, fig. 1), and also in certain Aulosphærida, but, apparently, only as an accidental variation (e.g., Aulosphæra icosahedra). Furthermore, this ground-form may also be assumed to occur in those Sphæroidea whose spherical lattice-shells bear twelve equidistant radial spines (e.g., many species of Acanthosphæra, Heliosphæra, and other Astrosphærida); the basal points of these spines indicate the twelve angles of the regular icosahedron. (See on this head Gener. Morphol., Bd. i. p. 411.)

27. The Regular Dodecahedral Ground-Form.—The ground-form whose geometrical type is the regular dodecahedron (or pentagonal dodecahedron), bounded by twelve equilateral and equiangular pentagons, is very rarely found perfectly developed, as in Circorrhegma dodecahedra (Pl. 117, fig. 2). This form is by no means so common among the Radiolaria as in the pollen grains of plants (e.g., Buchholzia maritima, Fumaria spicata, Polygonum amphibium, &c.). It can, however, be regarded as present in all those Sphæroidea whose spherical lattice-shells bear twenty equal and equidistant radial spines (e.g., many species of Acanthosphæra, Heliosphæra, and other Astrosphærida); the basal points of these spines mark out the twenty angles of the regular pentagonal dodecahedron. (See Gener. Morphol., Bd. i. p. 412.)

28. The Regular Octahedral Ground-Form.—The ground-form whose geometrical type is the regular octahedron (bounded by eight equilateral triangles), commonly appears among the Spumellaria in the family Cubosphærida (p. 169, Pls. 21-25). In {xiv}these Sphæroidea the typical ground-form is usually indicated by six equal radial spines, which are opposed to each other in pairs, and lie in three similar axes perpendicular to each other; these are the three axes of the tesseral crystallographic system; one of them is vertical, whilst the other two cross each other at right angles in its centre. Occasionally, too, the spherical form of the lattice-shell passes over into that of the regular octahedron (Pl. 22, figs. 8, 10). The same form recurs in Circoporus (Pl. 117, fig. 6) among the Phæodaria. In the vegetable kingdom it is exhibited by the antheridia of Chara. It is not found in the Nassellaria and Acantharia. (See Gener. Morphol., Bd. i. p. 412.)

29. The Regular Cubic Ground-Form.—The ground-form whose geometrical type is that of a die or cube, is actually presented in a very striking manner by various Radiolaria. Among the Spumellaria it occurs in certain Sphæroidea, e.g., in the Astrosphærid genera Centrocubus and Octodendron (Pl. 18, figs. 1-3); in these the central medullary shell is a complete cube, bounded by six equal squares, from the eight angles of which eight equal radial spines project. This form can also be regarded as present in those Sphæroidea whose spherical lattice-shell bears eight equal and equidistant radial spines (many Astrosphærida). Besides these the cubic ground-form is to be seen in certain Nassellaria of the family Tympanida, especially in Lithocubus (Pl. 82, fig. 12; Pl. 94, fig. 13), in many species of Acrocubus, Microcubus, &c.; the twelve bars of its lattice-skeleton correspond often exactly to the edges of the cube. (See Gener. Morphol., Bd. i. p. 413.)

30. The Regular Tetrahedral Ground-Form.—The ground-form whose geometrical type is the regular tetrahedron, bounded by four equilateral triangles, occurs less frequently in the Radiolaria than the other four regular polyhedra. Among the Spumellaria it is found in the Beloidea, and especially in those members of the Thalassosphærida and Sphærozoida whose spicules bear four equal branches, diverging at equal angles from a common centre. Precisely the same structure is seen also among the Nassellaria in some Plectoidea, as in Tetraplagia among the Plagonida, and Tetraplecta among the Plectanida. The skeleton of both these genera consists of four equal rods, which radiate at equal angles from a common centre, just as do the axes of the regular tetrahedron. The tetrahedral form of these Plectoidea is the more important and interesting since on the one hand it is related to the similar spicular form of the Beloidea, and on the other perhaps furnishes the starting point from which Cortina among the Nassellaria may be derived (Plagoniscus, Plectaniscus). (See Gener. Morphol., Bd. i. p. 415.)

31. The Isopolar-Monaxon or Phacotypic Ground-Form.—The isopolar uniaxial or phacotypic ground-form is characterised by the possession of a vertical main axis with {xv}equal poles, whilst no transverse axes are differentiated. All horizontal planes which cut the axis at right angles are circles, and increase in size from the poles towards the equator. The most important ground-forms of this group are the phacoid (the lens or oblate spheroid) and the ellipsoid (or prolate spheroid). Phacoids (or geometrical lenses with blunt margins) are very often presented by the central capsules of the Discoidea and of many Acantharia (Quadrilonchida and Hexalaspida), but the lattice-shells of many Spumellaria and Acantharia exhibit the same form, as also do a few Phæodaria (e.g., Aulophacus). True geometrical ellipsoids are seen in the central capsules of many Prunoidea among the Spumellaria, and of many Amphilonchida and Belonaspida among the Acantharia. Furthermore, the lattice shells of many species of these groups retain the same essential form, e.g., many Ellipsida, Druppulida, and Spongurida (Pls. 13-17, and 39), as well as most Belonaspida. (See Gener. Morphol., Bd. i. p. 422.)

32. Allopolar-Monaxon or Conotypic Ground-Form.—The allopolar uniaxial or conotypic ground-form is characterised by the possession of a vertical main axis whose two poles are unlike, while no transverse axes are differentiated. All horizontal planes cutting the main axis at right angles are circles, and decrease more rapidly from the largest plane towards the basal than towards the apical pole. The most important ground-forms of this group are the ovoid, the cone, and the hemisphere. They often occur (and in geometrical perfection) in the egg-shaped central capsule and podoconus of the Nassellaria, as well as in the shells of several groups of this legion, particularly in the Cyrtocalpida or Monocyrtida eradiata (Pl. 51, figs. 10-13), and in many Stichocyrtida eradiata; furthermore, they are also seen among the Phæodaria, e.g., certain Challengerida (Pl. 99, figs. 19-22). (See Gener. Morphol., Bd. i. p. 426.)

33. The Regular Dipyramidal or Quadrilonchial Ground-Form.—The ground-forms whose geometrical type is the regular double pyramid are characterised by a vertical main axis which possesses equal poles, and which is crossed at its centre by several equal transverse axes. The horizontal equatorial plane is therefore a regular polygon, and divides the body into two equal regular pyramids. The simplest and commonest form of this group is the quadratic octahedron, the ground-form of the quadratic crystallographic system; its equatorial plane is a square. This regular dipyramidal ground-form occurs among the Spumellaria in the shells of the Staurosphærida as well as of many Discoidea, in which several equidistant radial spines or arms lie in the quadratic equatorial plane of the body, and project from the margin of the lenticular disc (e.g., Sethostaurus, Pl. 31; Histiastrum, Pl. 46, &c.). It is, however, among the Acantharia that the most important part is played by this ground-form (and especially by the quadratic octahedron); it forms the basis of all those Acanthometra and Acanthophracta in which twenty radial spines are disposed according to the Müllerian Law, and in which {xvi}the four equatorial spines are of equal dimensions (Icosacantha). (See Gener. Morphol., Bd. i. p. 436-446.)

34. The Amphithect Dipyramidal or Lentelliptical Ground-Forms.—The ground-forms whose geometrical type is the lenticular or "triaxial" ellipsoid, may also be designated amphithect double pyramids; they are characterised by the possession of a vertical main axis which has similar poles, and is crossed at its middle by two transverse axes, unequal but isopolar. The horizontal equatorial plane of the body is therefore an amphithect or elongated polygon (a rhombus in the simplest case possible), and divides the whole body into two equal amphithect pyramids. The simplest and commonest form of this group is the rhombic octahedron, which is also the ground-form of the rhombic crystallographic system. It plays an important part in those Acantharia in which twenty radial spines are disposed according to the Müllerian Law, but in which the two pairs of equatorial spines are unequal (different geotomical and hydrotomical axes, see p. 719); to this category belong the Amphilonchida (Pl. 132), Belonaspida (Pl. 136), Hexalaspida (Pl. 139), and Diploconida (Pl. 140). A form essentially identical obtains also among the Spumellaria in the majority of the Larcoidea, both in their triaxial lattice-shells, and in their lentelliptical central capsules, which present geometrically accurate triaxial ellipsoids, with three unequal isopolar axes at right angles to each other. (See Gener. Morphol., Bd. i. p. 446-452.)

35. The Regular Pyramidal Ground-Forms.—The ground-forms whose geometrical type is the regular pyramid, and which are the most conspicuous in the Medusæ, Polyps, Corals, and regular Echinoderms (the Radiata of earlier authors), are almost confined among the Radiolaria to the legion Nassellaria; they occur, however, in the great majority of these, and especially in those families which may be classed together as "Cyrtoidea triradiata et multiradiata." Strictly speaking, however, almost all these Nassellaria, at all events in their origin, are bilateral or dipleuric, since the primary sagittal ring with its characteristic apophyses marks out the sagittal median plane, and further, since the three feet of the basal tripod are usually divided into an unpaired dorsal (pes caudalis) and two paired ventral or lateral (pedes pectorales, dexter et sinister). On the other hand, it is noteworthy, firstly, that among the primitive Plectoidea there are perfectly regular radial forms, without any indication of an original bilateral symmetry, and secondly, that similar forms are also very common among the Cyrtoidea, probably as secondary radial forms, developed from primitive bilateral ones. Similar cases also occur in certain Phæodaria (e.g., the Medusettida and Tuscarorida, Pls. 100, 120), but they are entirely wanting among the Acantharia and Spumellaria. The multiradial Nassellaria have arisen from the triradial by the interpolation of three, six, nine, or more interradial and adradial secondary apophyses between the three primary perradial ones. (See Gener. Morphol., Bd. i. pp. 459-874.)

36. The Amphithect Pyramidal Ground-Forms.—The ground-forms whose geometrical type is the amphithect pyramid, are distinguished from the regular pyramidal forms, just discussed, chiefly by the form of the basal plane, which is not a regular, but an amphithect or elongated polygon (in the simplest case a rhombus). Hence in this case the allopolar main axis of the body is crossed by two transverse axes which are isopolar and at right angles, but are unequal; they cannot, however be distinguished as sagittal and frontal axes as is the case in the zeugites. In the animal as well as in the vegetable kingdom, an important part is played by this ground-form, e.g., in the Ctenophora, where it is the rhombic pyramid. Among the Radiolaria it is not common, though it is clearly expressed among the Nassellaria in a number of Stephoidea (Stephanida and Tympanida), as well as in many Spyroidea (e.g., the bipedal Zygospirida). It is very accurately developed among the Phæodaria in the bivalved Phæoconchia (Pls. 121-128), where the two valves of the shell (dorsal and ventral) are generally exactly alike, their median keels corresponding to the poles of the sagittal axis. In the slit between the two valves lie the two secondary openings (right and left) of the tripylean central capsule, corresponding to the two poles of the frontal axis, and the main axis stands perpendicularly to both these, its oral pole being indicated by the astropyle, or principal aperture. (See Gener. Morphol., Bd. i. pp. 479-494.)

37. The Amphipleural Ground-Forms.—By the term amphipleural ground-forms are to be understood those usually defined as "bilaterally radial"; their geometrical type is a half amphithect pyramid. The best known examples of this form in the animal kingdom are the bilateral five-rayed Echinoderms (Spatangus, Clypeaster), in the vegetable kingdom the symmetrical five-rayed flowers (Viola, Trifolium). The three dimensive axes have the same relation as in the zygopleura, to be next discussed, and which also resemble them in being divisible only by one plane (the sagittal median plane) into two equal halves. They differ, however, the amphipleural body not being made up of two antimeres, but of at least three pairs of antimeres (or three parameres), being therefore primitively radial. Hence each of the symmetrical halves of the body contains more than one antimere. Among the Radiolaria this form does not occur in the Spumellaria, Acantharia, or Phæodaria; it is very common, however, among the Nassellaria; many Cyrtoidea multiradiata and Spyroidea multiradiata show this bilaterally radial ground-form, inasmuch as the body consists of two symmetrical halves, and is also composed of numerous (usually three, six, nine, or more) radial parameres. In the multiradiate Dicyrtida and Tricyrtida the cephalis (the first joint) is usually bilateral, whilst the thorax (the second joint) is multiradial. (See Gener. Morphol., Bd. i. pp. 495-506.)

38. The Zygopleural Ground-Forms.—As zygopleural or dipleural ground-forms, as opposed to the amphipleural, are classed those zeugites or centroplana which are known {xviii}as "bilaterally symmetrical" in the strictest sense of the term. This is the most important ground-form in the animal kingdom, inasmuch as it obtains almost exclusively among the higher animals (Vertebrata, Articulata, Mollusca, Vermes). The body consists of only two antimeres, which correspond to the two symmetrical halves of the body. Of the three dimensive axes two are allopolar, one isopolar; the oral pole of the longitudinal main axis is different from the aboral; the dorsal pole of the sagittal axis is different from the ventral; but the right pole of the frontal axis is equal to the left. The right antimere is usually precisely similar to the left (Eudipleura), more rarely it is slightly dissimilar or asymmetrical (Dysdipleura). Among the Radiolaria this ground-form is entirely wanting in the Porulosa or Holotrypasta (Spumellaria and Acantharia), but on the contrary it is very common in the Osculosa or Merotrypasta (Nassellaria and Phæodaria). In the Nassellaria it is of special importance, for the typical Cortina (the combination of the primary sagittal ring with the basal tripod) exhibits the zygopleural ground-form clearly sketched out; indeed it is usually clearly seen even in the sagittal ring itself, for its ventral segment is more strongly curved than the dorsal; its basal (or oral) pole is always different from the apical (or aboral). Of the three feet of the basal tripod the unpaired (caudal) one is directed dorsally and backwards, the two paired (pectoral) ones ventrally and forwards. The majority of the Nassellaria may be regarded as modifications of this original ground-form. Its relation to the primitively triradiate tripod presents a still unsolved problem, and the numerous relations of the zygopleural to the multiradiate ground-forms in the Nassellaria are exceedingly complicated. The zygopleural ground-form is less widely distributed among the Phæodaria, though it is very characteristically developed in the rich and varied group of Challengerida (Pl. 99). (See Gener. Morphol., Bd. i. pp. 507-527.)

39. Synopsis of the Geometrical Ground-Forms:—

Principal Groups of Ground-Forms.		Subsidiary Groups of Ground-Forms.		Geometrical Type.		Examples.
I. Centrostigma. The geometrical centre of the body is a point. Main axis wanting.		I. Homaxonia. All axes equal		1. Sphere,		Central capsule of the Sphæroidea and of many Acantharia.
		II. Polyaxonia. Endospherical polyhedra. All the angles of the body lie on the surface of a sphere. Numerous isopolar axes.		2. Endospherical polyhedron,		Lattice-spheres of the Sphæroidea, Sphærophracta, and Phæosphæria.
				3. Icosahedron,		Circogonia.
				4. Dodecahedron,		Circorrhegma.
				5. Octahedron,		Cubosphærida, Circoporus.
				6. Cube,		Centrocubus, Lithocubus, &c.
				7. Tetrahedron,		Tetraplagia, Tetraplecta, &c.
{xix} II. Centraxonia. The geometrical centre of the body is a straight line (the vertical main axis).   Constant transverse axes (perpendicular to the main axis) are wanting in the Monaxonia (which have circular transverse sections); on the contrary they are differentiated in the Stauraxonia (which have polygonal transverse sections).		III. Monaxonia. Uniaxial ground-forms or centraxonia without transverse axes. The transverse planes (perpendicular to the main axis) are circles.		8. Monaxonia isopola. (Spheroids and ellipsoids; both poles of the main axis similar.)		Central capsule and lattice-shell of of many Discoidea (lenses) and Prunoidea (ellipsoids), Belonaspida, &c.
				9. Monaxonia allopola. (Cone, ovoid and hemisphere; the two poles of the axis dissimilar.)		Central capsule and lattice-shell of many Nassellaria, especially the Cyrtoidea eradiata (Cyrtocalpida, &c.).
		IV. Stauraxonia. Pyramidal ground-forms or centraxonia with transverse axes. The transverse planes (perpendicular to the main axis) are either regular or amphithect polygons.		10. Dipyramides regulares. (Quadratic octahedron, or quadrilonchial forms and regular double pyramids.)		Acantharia with twenty radial spines, the four equatorial being equal. Multiradial Discoidea and Staurosphærida.
				11. Dipyramides amphithectæ. (Rhombic octahedron, lentellipsoid, and amphithect double pyramids.)		Acantharia with twenty radial spines, whose four equatorial spines are unequal but paired. Many Larcoidea.
				12. Pyramides regulares. (Regular pyramids.)		Many Nassellaria (triradial and multiradial). Medusettida and Tuscarorida.
				13. Pyramides amphithectæ. (Rhombic pyramids.)		Phæoconchia. Bipedal Spyroidea and Stephoidea.
III. Centroplana. The geometrical centre of the body is a plane (the sagittal plane).		V. Bilateralia (or Zeugita). Bilateral forms in the general sense, with right and left halves.		14. Amphipleura (Bilaterally radial ground-form.)		Many Cyrtoidea and Spyroidea multiradiata.
				15. Zygopleura. (Bilaterally symmetrical ground-form.)		Most Nassellaria (primitively at least), many Challengerida.
IV. Acentra. There is no geometrical centre.		VI. Anaxonia. No definite axes can be determined.		16. Irregularia. (Absolutely irregular ground-forms.)		Collodastrum, Collosphæra, Phorticida, Soreumida.

40. Mechanical Causes of the Geometrical Ground-Forms.—The great variety of ground-forms exhibited by the Radiolaria is of special interest, since in most instances their causes admit of recognition, and since they are so intimately related to each other that even in the remaining cases the assumption that they have arisen by purely mechanical causæ efficientes seems justified. In this respect the first rank is taken by statical conditions, especially the indifferent or stable equilibrium of the whole organism, which floats freely in the water. With regard to these fundamental statical relations, three principal groups of ground-forms may be distinguished, pantostatic, polystatic, and monostatic.

41. Pantostatic Ground-Forms.—By pantostatic or indifferently stable ground-forms are meant those in which the centre of gravity coincides with the centre of the body, so that they are in equilibrium in any given position. Strictly speaking, the only form which possesses perfectly indifferent equilibrium is the sphere, that being the only truly homaxon and perfectly regular form. Nevertheless, in a somewhat wider sense many Polyaxonia, especially the endospherical polyhedra with very numerous sides, may be {xx}included in this category. Such indifferently stable bodies are found among the Spumellaria in many Collodaria and Sphæroidea, as well as in the Astrolophida among the Acantharia. On the contrary they are entirely wanting among the Nassellaria and Phæodaria, since their central capsule constantly presents a main axis with a differentiated basal pole, and determines the position of stable equilibrium.

42. Polystatic Ground-Forms.—Those ground forms are defined as polystatic or multistable in which the body is in equilibrium in several different positions (though not in an infinite number). The number of these positions is usually twice as many as that of the constant equal isopolar axes exhibited by the form. Hence the regular polyhedra have as many positions of equilibrium as they have angles or sides, the icosahedron twenty, dodecahedron twelve, octahedron eight, cube six, tetrahedron four. The isopolar monaxon ground-forms (lens, ellipsoid, cylinder) and the diplopyramidal ground forms (quadrilonchial and lentelliptical) have two positions of stable equilibrium, since the two poles of the vertical axis are equal and similar and the body is divided into equal halves by the equatorial plane. This is the case in many Spumellaria (especially Discoidea, Prunoidea, and Larcoidea), as well as in the great majority of Acantharia. Perhaps the same holds good also in certain Nassellaria (e.g., isopolar Tympanida) and Phæodaria (e.g., isopolar Phæosphæria), though here unistable equilibrium appears to be necessitated by the constant main axis of the central capsule and the differentiated basal pole of the main axis.

43. Monostatic Ground-Forms.—Those ground-forms are classed as monostatic or unistable in which the body is in equilibrium only in one position, since the centre of gravity of the body lies in a constant vertical axis below its centre. This fixed position is only rarely and exceptionally found among the Spumellaria (e.g., in Xiphostylus, Sphærostylus, Lithomespilus, Lithapium) and among the Acantharia (e.g., in Zygostaurus and Amphibelone). On the contrary it is quite usual among the Nassellaria and Phæodaria (with but few exceptions); for here a vertical main axis, with a differentiated basal pole, is determined even by the formation of the central capsule, and usually also by the corresponding structure of the skeleton. Among the Nassellaria this basal pole, with the porochora of the central capsule, appears always to be the lower; as also in most Phæogromia among the Phæodaria. In the peculiar bivalved Phæoconchia, on the other hand, the basal pole with the cannopyle is directed upwards; as also in the Challengerida and Tuscarorida. The Phæosphæria and Phæocystina are probably to a large extent polystatic. In general unistable equilibrium may be assumed in the following categories of ground-forms:—(1) Allopolar monaxon (conical and ovoid); (2) pyramidal (regular and amphithect); (3) Centroplana (amphipleura and zygopleura); (4) Anaxonia.

44. Principal Axes.—From the foregoing consideration of the statical conditions and their direct causal connection with the geometrical ground-forms of the Radiolaria, the great mechanical significance of the differentiation of definite axes in these unicellular free-swimming organisms will be manifest. The most important of these is the primary main axis (axis principalis, or protaxon), which in all cases has a vertical direction. It is wanting in the Centrostigma (spheres and endospherical polyhedra), and in the Anaxonia (acentra). It is isopolar in the phacotypic forms (Monaxonia isopola), and in the double pyramids (Stauraxonia isopola). It is allopolar in all monastatic ground-forms, in the conotypic forms (Monaxonia allopola), pyramids (Stauraxonia allopola), and the Centroplana (or bilateral forms).

45. Secondary or Transverse Axes.—In contrast to the vertical main axis all the other constant axes differentiated in the body may be called "secondary axes," or "transverse axes," since they cross the former at definite points. All ground-forms whose vertical axis is crossed by a fixed number of such axes at definite angles may be called "Stauraxonia." They are divided into two groups, double pyramids and single pyramids; in the former the two poles of the main axis (or the two halves of the body separated by the equatorial plane) are similar (Stauraxonia homopola), in the latter dissimilar (Stauraxonia heteropola). If all the secondary axes be equal, the stauraxon ground-form is regularly radial. If some of them be unequal they are arranged in certain relations towards two primary transverse axes, perpendicular to each other, to which all the other secondary axes are subsidiary; the ground-forms are then either amphithect or bilateral. The two primary transverse axes, which may also be designated "ideal transverse axes" (euthyni), divide the vertical main axis in its centre; one of them is the sagittal, the other the frontal. These three dimensive axes give the factors which accurately determine the ground-form and the dimensions in most Radiolaria; the vertical main axis determines the length (principal axis); one horizontal transverse axis determines the thickness (sagittal axis), and the other the breadth (frontal axis). Those ground-forms in which the transverse axes are isopolar are termed "amphithect," and those in which the one (frontal or lateral) is isopolar and the other (sagittal or dorso-ventral) is allopolar, are termed "bilateral," or better "zeugitic."

46. Primary and Secondary Ground-Forms.—The geometrical sphere must be regarded as the original ground-form of the Radiolaria; it being understood that its monophyletic derivation from a single stem-form, Actissa, is correct. The simplest forms of Actissa (Procyttarium, Pl. 1, fig. 1) are in fact geometrically perfect spheres; indeed even the individual parts which compose their unicellular bodies (nucleolus, nucleus, central capsule and calymma) are concentric spheres. But in addition the central capsules of most other Spumellaria, especially the Sphæroidea, as well as of many Acantharia {xxii}are true spheres. Furthermore the simple or concentrically composed lattice-spheres of Sphæroidea, Sphærophracta, and Phæosphæria may be regarded as spheres, although strictly speaking they are endospherical polyhedra. From the primary spherical form of the Radiolaria all other secondary forms may be derived in the following order:—1. By the development of a main axis the Monaxonia arise. 2. By the development of transverse axes the Stauraxonia arise. 3. In both groups (Monaxonia and Stauraxonia) the two poles (or upper and lower halves of the body) are at first similar (Isopola). 4. By differentiation in the two poles or halves of the body (distinction between the basal pole and the apical) the forms with different poles (Allopola) arise. 5. The transverse axes of the Stauraxonia are at first equal (regular pyramids and double pyramids). 6. By differentiation in the transverse axes (distinction between the sagittal and the frontal axis) the amphithect pyramids and double pyramids arise. 7. From the amphithect pyramids the Amphipleura arise by differentiation of both poles of the sagittal axis. 8. The zygopleural ground-form appears last, as the simplest form of the Amphipleura.

47. The Ground-Forms of the Spumellaria.—The Spumellaria, being the oldest and most primitive Radiolaria, have for the most part either indifferent or multistable equilibrium; e.g., all Colloidea and Beloidea which have a spherical central capsule, and also most Sphæroidea. Among these primitive Centrostigma true spheres and endospherical polyhedra are represented in the utmost variety, and the regular polyhedra in particular. By the development of a vertical main axis these Centrostigma have also given rise to very numerous Centraxonia, which are usually isopolar, very rarely allopolar. Sometimes they are Monaxonia (circular in transverse section), sometimes Stauraxonia (polygonal in transverse section). The vertical main axis is longer in the Prunoidea, shorter in the Discoidea than any of the other axes. The Larcoidea are distinguished by their lentelliptical or triaxial ellipsoid form; the three different but isopolar axes corresponding with those of the rhombic octahedron; but even among the Sphæroidea, Prunoidea, and Discoidea, this form is sometimes produced by the differentiation of two different transverse axes at right angles to each other. Whilst these ground-forms (Centraxonia and Centrostigma) occur in the utmost variety among the Spumellaria, the centroplanar (or true bilateral) ground-form is entirely wanting.

48. The Ground-Forms of Acantharia.—In the small family Astrolophida, which contains the most archaic forms of the legion (Actinelius, Astrolophus), the Acantharia show a direct relation to the most primitive Spumellaria (Actissa), and like these have indifferent equilibrium; their central capsule is a sphere, their calymma an endospherical polyhedron, whose angles are indicated by the distal ends of the numerous {xxiii}equal radial spines. In the great majority of Acantharia, however (all Acanthonida and Acanthophracta), twenty radial spines are present, regularly distributed, according to Müller's icosacanthan law, in five parallel circles, each containing four crossed spines (p. 717). Usually the twenty spines are equal, and the ground-form is the quadratic octahedron, or a regular double pyramid with sixteen sides. But in some groups (the Amphilonchida and Prunophracta) two opposite equatorial spines are much more strongly developed than the other eighteen, and therefore the hydrotomical axis in the equatorial plane is larger than the geotomical axis (p. 719); the isopolar stauraxonian form passes over into the allopolar, and the ground-form becomes the rhombic octahedron or the amphithect double pyramid (compare §§ 33 and 34, and p. 720). The centroplanar ground-form is entirely wanting in the Acantharia.

49. The Ground-Forms of the Nassellaria.—The Nassellaria all possess monostatic ground-forms, inasmuch as by the very structure of their monopylean central capsule a vertical main axis is necessitated, whose basal pole occupies the porochora. The same arrangement is also for the most part clearly recognisable in the corresponding structure of the skeleton, which is generally either centraxon or centroplanar. Among their manifold skeletal forms different larger groups of ground-forms may be recognised according as the vertical allopolar main axis is crossed by differentiated transverse axes or not (Stauraxonia or Monaxonia); the former are either triradial or multiradial. The triradial, with three lateral or terminal radial apophyses, constitute the greater part of the Nassellaria, and have probably been derived originally from the triradial Plectoidea (Triplagia, Triplecta); a more careful examination, however (especially with reference to the structure of the cortinar septum), reveals the fact that the ground-form is not strictly regularly pyramidal (with three equal radii), but amphipleural (with two paired ventral and one unpaired dorsal radius), and that it usually passes over into a distinctly zygopleural form. The same holds true of the multiradial Nassellaria, where for the most part three interradial or six adradial (sometimes more) apophyses are intercalated between the three primary perradial ones; sometimes here also the ground-form is a quite regular hexagonal or nonagonal pyramid, but usually it is more or less amphithect or amphipleural. Among the eradial Nassellaria, which have no radial apophyses, the ground-form is sometimes allopolar monaxon (conical, ovoid, hemispherical, &c.), sometimes amphithect pyramidal (even in the simplest Stephanida, Archicircus, &c.), or sometimes distinctly zygopleural or bilateral (many Plectellaria).

50. The Ground-Forms of the Phæodaria.—The Phæodaria agree with the Nassellaria in the possession of a primitively centraxon ground-form, and like them are monostatic, since a vertical main axis whose basal pole passes through the astropyle is present, owing to the characteristic structure of their cannopylean central capsule. In {xxiv}the great majority of Phæodaria the spheroidal central capsule also possesses a pair of parapylæ near the opposite apical pole of the main axis (Tripylea), and these determine (as the right and left secondary openings) an isopolar frontal axis. Hence, strictly speaking, in most Phæodaria the central capsule has the geometrical ground-form of the amphithect pyramid (as in the Ctenophora), with an allopolar vertical main axis, and two unequal, but isopolar, horizontal transverse axes. In many Phæodaria the skeleton also has this amphithect pyramidal ground-form, e.g., the bivalved Phæoconchia and part of the Phæogromia. On the contrary, in the rest of the Phæodaria the skeleton exhibits very various geometrical ground-forms, independent of that of the central capsule. In the Phæosphæria it forms preferably spheres or endospherical polyhedra, as also in the Castanellida and Circoporida among the Phæogromia; among the Circoporida there are also seen with remarkable distinctness the regular polyhedra (especially the dodecahedron and icosahedron). Isopolar monaxonia are found among the Aulosphærida (Aulatractus) and Orosphærida; allopolar monaxonia among the Challengerida (Lithogromia). The Medusettida and Tuscarorida show various forms of regular pyramids (allopolar Stauraxonia); and finally, the Challengerida are for the most part centroplanar or bilateral. Thus the Phæodaria present a great wealth of different geometrical ground-forms in the development of their skeleton, not in that of their central capsule.

Chapter II.—THE CENTRAL CAPSULE.

51. Components of the Central Capsule.—In all Radiolaria without exception, at some period of life or other, the central portion of the soft body is separated from the peripheral portion by an independent, anatomically recognisable membrane; this membrane with all its contents is designated the central capsule, and is the peculiar central organ of the unicellular body, which distinguishes the Radiolaria most clearly from the other Rhizopoda. In the great majority of the Radiolaria the volume of the central capsule is less than that of the surrounding peripheral soft body which we place in opposition to it as "extracapsulum." The "capsule-membrane," which separates these two constituents, arises very early in most Radiolaria, and persists throughout their whole life. In some species, however, the membrane only appears later, immediately before the formation of the spores, and hence is absent for a considerable period. Regarded as a whole, then, the capsule consists of the following parts:—(1) the capsule-membrane; (2) the enclosed endoplasm, or intracapsular protoplasm; (3) the nucleus. But in addition, many other non-essential structures may be enclosed in the central capsule, especially hyaline spheres (vacuoles), fatty spheres, pigment granules, crystals, &c.

52. The Primary Form of the Central Capsule.—The form of the central capsule is originally a geometrical sphere; and if in accordance with our monophyletic hypothesis all Radiolaria are to be derived from one common stem-form (Actissa, see p. 12), then the central capsule of this common stem-form must be regarded as perfectly spherical (Procyttarium, p. 13, Pl. 1, fig. 1). Since, further, the enclosed nucleus and the surrounding calymma of this primitive archaic form must also be spheres, and since the nucleus lies in the centre of the body, and the protoplasm is evenly distributed between it and the membrane, it follows that no axes or excentrically differentiated parts are to be distinguished in this most primitive Radiolarian. Rather in the primary central capsule all parts are concentrically and evenly arranged round its centre. This primary spherical form becomes modified in most Radiolaria into various secondary ground-forms, which are correlated partly with the structure of the capsule itself, and partly also with the development of openings in its membrane. In general the ground-form of the central capsule is polyaxon in the Porulosa (Spumellaria and Acantharia); but in the Osculosa centraxon forms are more frequently observed; in the Nassellaria the ovoid (allopolar monaxon) form is predominant, and in the Phæodaria the rhomboid or amphithect pyramid. In these latter, the astropyle indicates the basal pole of the vertical main axis, whilst the two parapylæ (right and left) mark the poles of the frontal transverse axis. In the Nassellaria the centre of the porochora corresponds with the basal pole of the main axis, whilst no transverse axes are originally present.

53. The Secondary Forms of the Central Capsule.—The original purely spherical form of the central capsule persists only in the minority of the Radiolaria, namely, the greater part of the Spumellaria and Acantharia; it passes over into various other secondary forms in the majority of the class, in the whole of the Nassellaria and Phæodaria, and in a considerable portion of the Spumellaria and Acantharia. These secondary or derived forms may be divided into two quite distinct groups, which may be designated endometamorphic and exometamorphic; in the former the cause of the divergence of the secondary form from the sphere lies in the internal structure of the central capsule; in the latter it lies in the external influence exerted by the growth of the skeleton. Obviously the former series of modifications is more significant than the latter.

54. The Endometamorphic Forms of the Central Capsule.—The secondary forms of the central capsule, which are due to internal causes connected with its growth, are as follows:—

55. The Exometamorphic Forms of the Central Capsule.—The secondary forms of the central capsule, which are brought about by external causes, chiefly dependent on the formation of the skeleton, are very various and in many cases devoid of special interest; in other instances, on the contrary, they are of great importance, because of the clear relation of cause and effect which can be traced between the development of the skeleton and of the capsule. The most important phenomena to be recorded in this connection are as follows:—

56. The Membrane of the Central Capsule.—The capsule-membrane or envelope of the central capsule is both morphologically and physiologically one of the most important parts of the Radiolarian body, for it separates its two main constituents, the capsule with its nucleus and endoplasm and the extracapsulum with the calymma and exoplasm. The capsule-membrane is invariably present at some time or other during the life of the organism, even though in a few species it may persist only for a short time. It is characterised in general by its power of resistance to chemical and physical reagents, and appears to be related to the elastic tissues or perhaps even more to the chitinous substances. Its thickness is usually less than 0.0001, though in certain groups it ranges between 0.001 and 0.002, and in many of the larger Radiolaria (such as Collida and Phæodaria) it may attain a thickness of 0.003 to 0.006 or more. In the three legions Spumellaria, Acantharia, and Nassellaria the capsule-membrane is single, while in the Phæodaria it is always double, being composed of a firm outer and a delicate inner membrane, which are in contact at only few points. Usually it is quite structureless, except for its apertures; the thicker membrane showing occasionally a fine concentric lamination. In certain large Colloidea (e.g., Thalassicolla, Pl. 1, fig. 5b) the membrane is covered on the inner surface by a network of polygonal ridges, and in some large Phæodaria with remarkable small curved rods (Pl. 114, fig. 13). In all Radiolaria the membrane is perforated by definite openings or pores, through which the intracapsular and extracapsular protoplasm are in direct communication. These openings (or "pylae") show very characteristic and constant differences in the four legions, which have given rise to the names—Peripylea, Actipylea, Monopylea, Cannopylea.

57. The Capsule-Openings of the Peripylea (or Spumellaria).—The capsule-membrane of the Peripylea is generally perforated by extremely fine and numerous pores, which are distributed at equal distances over the whole surface, and are precisely alike in all parts of the capsule. Hence the Spumellaria may be called "Holotrypasta" or "Porulosa"; they agree with the Actipylea in being devoid of an osculum or operculum; they are distinguished from the latter group mainly in that their pores are equally distributed over the whole surface of the capsule, whilst in the Actipylea the pores are disposed in definite groups or lines, separated by large imporous areas.

58. The Capsule-Openings of the Actipylea (or Acantharia).—The capsule-membrane of the Actipylea is perforated by very numerous fine pores, which are regularly distributed over the surface of the central capsule, and separated by imporous intervals. Hence the Acantharia belong to the "Holotrypasta" or "Porulosa"; they have neither osculum nor operculum, and agree in this particular with the Peripylea; but they are separated from these latter chiefly by the fact that their pores are much less numerous, and marked off into regularly arranged groups or lines by imporous intervals. In the Peripylea, on the contrary, the pores are much more numerous and are evenly distributed over the whole surface of the capsule.

59. The Capsule-Openings of the Monopylea (or Nassellaria.)—The capsule-membrane of the Monopylea always possesses a single large main-opening, an osculum, which lies at the basal pole of the main axis, and is closed by a circular perforated lid (operculum porosum). When seen from the surface this lid appears as a clearly defined porous area (porochora or area porosa), and forms the horizontal base of a peculiar cone, which stands vertically in the interior of the capsule and may be designated the "thread-cone" (podoconus). The Nassellaria may hence be termed "Merotrypasta" or "Osculosa," like the Cannopylea; the structure and significance of the circular lid (operculum), which closes the main-opening (osculum) is, however, quite different in the two legions. Whilst the lid of the Cannopylea (astropyle) is solid, traversed by radial ribs, and only perforated in its centre by a short tube (proboscis), in the Monopylea the operculum (porochora) is always perforated by numerous vertical fine pores, and is in connection with the peculiar internal "pseudopodial cone" (podoconus, Pl. 51, figs. 5, 13; Pl. 81, fig. 16; Pl. 91, fig. 5; Pl. 98, fig. 13). The pores are separated by small vertical, highly refractive rods (opercular rhabdillæ); these become intensely stained by carmine, and are either evenly distributed over the surface of the porochora or arranged in definite groups. The outer or distal end of each rod is rounded, sometimes thickened like a club or split into lobes; the inner or proximal end is usually pointed, and stands in connection with a myophane thread of the podoconus (see § 79). The primary circular form of the porochora, in which the opercular rhabdillæ are evenly distributed in a horizontal plane, undergoes various secondary modifications in many Nassellaria. The triradial structure of the skeleton, which characterises the majority of the legion, causes a splitting of the base of the central capsule into three or four lobes; this division also affects the porochora, which lies in the centre of the base, so that the rhabdillæ become arranged in three or four equal circles. If, however, the lobes of the central capsule become larger and protrude through the three or four collar pores of the cortinar septum, the central porochora may separate entirely into three or four elongated tracts, which lie on the axial side of the magnified lobes; the rhabdillæ are then arranged over the whole surface of {xxxi}these tracts, on the outer aspect of which run the longitudinal myophane fibrillæ of the podoconus (compare §§ 79 and 99).

60. The Capsule-Openings of the Cannopylea (or Phæodaria).—The capsule-membrane of the Cannopylea always possesses only a single large main-opening or osculum, which lies at the basal pole of the vertical main axis, and is closed by a circular radiated lid (operculum radiatum). This operculum appears, when seen from the surface, as a sharply defined stellate area (astropyle), from the middle of which arises a shorter or longer cylindrical tube, the proboscis. Hence the Phæodaria, like the Monopylea, belong to the "Merotrypasta" or "Osculosa"; the structure and significance of the circular operculum, which closes the main-opening (osculum), are, however, quite different in the two legions. Whilst the operculum of the Monopylea (porochora) is perforated by numerous fine vertical pores, and connected with the peculiar internal pseudopodial cone (podoconus), this structure is entirely wanting in the Cannopylea, and instead of it there is a solid operculum, with radial ribs which originate at the base of its central tubular mouth; this tube (proboscis) is cylindrical, often conical at the base, of very variable length and with a round aperture at either end. In spite of the great difference which the various families of Cannopylea exhibit in the formation of their skeleton and its appendages, the constitution of this characteristic stellate main-opening (astropyle) is always essentially the same; both the stellate operculum itself, and the proboscis which rises from its centre, show only slight differences in the various groups. In addition to this large main-opening most Phæodaria possess several small accessory openings (parapylæ); and usually two of these are present, placed symmetrically right and left of the aboral pole of the main axis and in the frontal plane (Pl. 101, figs. 2, 6, 10; Pl. 104, figs. 1, 2a). Sometimes there are more numerous accessory openings (three to six or more) regularly arranged, as in the two peculiar families, Circoporida and Tuscarorida; occasionally also there is only a single parapyle, at the aboral pole of the main axis (e.g., in Tuscaridium). The parapylæ seem to be quite absent in the families Challengerida, Medusettida, Castanellida, and perhaps also in other Phæodaria. The form and structure of the small accessory openings appear to be always the same. The {xxxii}outer capsule-membrane is elevated in the form of a short cylindrical tube or "apertural ring" (collare paraboscidis), the external margin of which bends inwards, and at the base of the ring passes over into the delicate internal capsule membrane. Upon this apertural ring is situated a longer or shorter "apertural cone" (paraboscis), which is a tubular, cylindrical or conical, prolongation of the membrane, open externally.

61. The Nucleus.—The nucleus, enclosed in the central capsule of all Radiolaria, behaves in every respect like a true cell-nucleus, and thus lies at the base of the now universal opinion, that the whole Radiolarian organism, in spite of its varied development and remarkable variations, is unicellular and remains throughout life a true individual cell. This important theory is not invalidated by the fact that the nucleus undergoes peculiar modifications in many groups, and in certain groups presents appearances seldom or never seen elsewhere.

62. Uninuclear and Multinuclear Radiolaria (Monocaryotic and Polycaryotic).—All Radiolaria present two different conditions in respect of the behaviour of the nucleus, since in their young stages they are uninuclear (monocaryotic), and in later stages multinuclear (polycaryotic). This is readily explained by the fact that each individual Radiolarian is developed from a simple unicellular swarm-spore, and that afterwards, before the formation of swarm-spores, the single nucleus divides into many small nuclei. Thus in the Radiolaria the nucleus is pre-eminently the organ of reproduction and inheritance. The division of the originally single nucleus into many small nuclei may take place, however, at very different periods, so that the Radiolaria may be divided in this respect into precocious and serotinous.

63. Serotinous and Precocious Radiolaria.—In the great majority of the Radiolaria the division of the nucleus takes place only at a late period, a short time or even immediately before the process of spore formation; it then breaks up rapidly into numerous small nuclei (always more than one hundred, sometimes many thousands), and each of these {xxxiii}either becomes itself the nucleus of a swarm-spore, or by repeated division gives rise to a group of spore-nuclei. All those Radiolaria which are uninuclear during the greater part of their existence, and in which the process of division is late, and takes place rapidly, are called "serotinous" or late-dividing forms. To this category belong all Phæodaria and Nassellaria, as well as all the solitary or monozoic Spumellaria and some Acantharia. On the other hand, the name "precocious," or early dividing, is applied to those Radiolaria in which the division of the nucleus takes place very early, and in which, therefore, the cell is multinuclear during the greater part of its existence. This is the case in all the social or polyzootic Radiolaria (Polycyttaria, Pls. 3-8), and also in the great majority of the Acantharia, both Acanthometra and Acanthophracta. In the last two groups, however, there are numerous exceptions, and these are seen in remarkably large species, characterised by the great size of the central capsule. From a phylogenetic point of view, the conclusion is allowable that the precocious forms are secondary, and have arisen by adaptive modification from the primitive serotinous stem. In the Polycyttaria (or social Spumellaria, i.e., the three families Collozoida, Sphærozoida, and Collosphærida), the cause of the adaptation lies most probably in the formation of the colony itself, for all these three families are so closely related to three corresponding families of serotinous, monozootic Radiolaria (Thalassicollida, Thalassosphærida, Ethmosphærida), that certain species of the latter are hardly to be distinguished from isolated individuals of the former. Perhaps the remarkable formation of the large central oil-globule, which particularly characterises the Polycyttaria, is the prime cause of their early nuclear division. In the Acantharia the cause is most likely to be found in the characteristic centrogenous development of their acanthin skeleton, whose radial bars first of all appear in the centre of the capsule. Hence arises directly the excentric position of the nucleus, which in the archaic stem of Acantharia (Actissa?) was probably central. In any case, but little weight is to be laid upon the precocious division of the nucleus in the Acantharia in general, inasmuch as in certain species (both Acanthometra and Acanthophracta) the more usual serotinous division persists.

64. Central and Excentric Nuclei.—The position of the nucleus in the interior of the central capsule was no doubt primitively central, and this situation in the geometrical centre of the original spherical central capsule has been accurately retained in all monozootic Spumellaria; in the polyzootic families of this legion (Polycyttaria), on the contrary, it is obscured by the precocious division of the nucleus. In the other three legions, which may be phylogenetically derived from the Spumellaria, the position of the nucleus is rarely central, but usually excentric, or at most subcentral. In the Acantharia (both Acanthometra and Acanthophracta) the central position of the nucleus is at once excluded by the constantly centrogenous development of the skeleton; the nucleus is therefore always excentric, and may lie at either side; it usually {xxxiv}divides very early into numerous separate nuclei, which are usually distributed in the peripheral portions of the central capsule. In the Nassellaria the development of the porochora, and of the podoconus which stands upon it, brings about the formation of a vertical axis, and in consequence the central capsule assumes a monaxon form (usually ovoid or conical); the nucleus then lies in the main axis, but excentrically between the apex of the podoconus and the aboral pole. In many Nassellaria, however, especially when the podoconus is so large that its apex approaches the aboral pole of the central capsule, the nucleus is pressed to one side and lies quite excentrically. The Phæodaria exhibit a different arrangement; the large spheroidal nucleus is always subcentral, so that its main axis corresponds with that of the concentric spheroidal central capsule; but since the astropyle always occupies the oral pole of the latter, and since the distance of the nucleus from this pole is always somewhat different from its distance from the other, it follows that, strictly speaking, the nucleus never lies accurately in the geometrical centre.

65. Homogeneous and Allogeneous Nuclei.—The nucleus of the Radiolaria not only exhibits a similar structure and composition, and suffers similar modifications to those which are found to occur in the case of other cell-nuclei, but also to some extent shows very peculiar developmental forms, which are seldom or never found in other cells. In the first place the nuclei may be divided into homogeneous and allogeneous, the former are structureless and consist of a uniform mass of nuclein, whilst the latter are composed of different substances and show various structural relations. Homogeneous nuclei, whose whole mass is uniform and exhibits no structural differentiation, are probably always to be found in the swarm-spores; in the fully developed Radiolarian body they are found only in the first legion, Spumellaria, and that both in many Monozoa (especially small Sphæroidea and Prunoidea) and in the Polyzoa (or Polycyttaria). The whole mass of these homogeneous nuclei, which are usually spherical or ellipsoidal, consists of uniform, perfectly clear and transparent nuclein, and becomes evenly stained by carmine, hæmatoxyline, &c. They may be readily distinguished by these means from the clear vacuoles or "hyaline vesicles," which are evenly distributed in the endoplasm of many Radiolaria, and may be confused with the former. Allogeneous nuclei, which are always composed of different parts and often show complicated structural relations, are found developed in the great majority of Radiolaria. The most important differentiation exhibited by these secondary forms is the separation of the nuclear mass into a firm nuclear substance (caryoplasm) and a fluid nuclear juice (caryolymph). In addition in each nucleus a nucleolus is visible, and often several or many may be seen (see §§ 67 to 70).

66. The Form of the Nucleus.—The nucleus of the Radiolaria shows greater variations in form and structure than are to be found in the majority of cell-nuclei; {xxxv}exception must, however, be made in the case of many animal ovicells, which, in their peculiar form and composition, often recall large Radiolarian nuclei. With respect to the external shape two main forms may be distinguished, as primary and secondary. The primary form of the Radiolarian nucleus is the sphere; it occurs not only in most swarm-spores, but also in most adult forms belonging to the legion Spumellaria, and in individual instances in other groups; indeed the nuclei of most Spumellaria, as also the concentric central capsules in which they lie, are true geometrical spheres. The secondary forms of the nucleus are found in the majority of adult Radiolaria, and arise from the primary spherical forms in various ways, either by the elongation or contraction of one axis, or by the formation of apophyses or processes. The most important of these secondary forms are as follows:—

67. The Nucleus of the Peripylea.—The nucleus of the Spumellaria or Peripylea shows in certain groups a very primitive arrangement, indeed the archaic structure from which the various forms of nuclei of other Radiolaria may be derived; but on the other hand, in other groups it exhibits very peculiar and remarkable differentiations. In the first place it may be noted that the monozootic or solitary Spumellaria usually possess a single serotinous nucleus, which only divides into numerous swarm-spores at a late period; {xxxvi}whilst, on the contrary, the polyzootic colonial Spumellaria (or Polycyttaria) are uninuclear only in the young state (Pl. 3, fig. 12), and speedily present numerous small homogeneous nuclei, which have arisen by precocious division of a single nucleus; these are usually spherical and 0.008 to 0.012 mm. in diameter. The serotinous nucleus of the monozootic Spumellaria, in many divisions of this large legion, and especially in the simply constituted Sphæroidea, is a homogeneous sphere of nuclein, lying in the middle of the central capsule. In many other cases it assumes the form of a spherical vesicle ("Binnen-Bläschen"), whose fluid or semi-fluid contents are enclosed by a more or less firm membrane. This vesicle often contains a single central spherical nucleolus (Pl. 1, figs. 1l, 4l), but sometimes a variable number of small excentric nucleoli (Pl. 1, figs. 1a, 2a). The nuclear membrane is often somewhat thick, presenting a double contour, and in such cases may even exhibit a fine radial striation, the expression of minute pores (Pl. 1, fig. 2a). In the colossal nuclei (as much as 1 to 2 mm. in diameter) of certain large Thalassicollida the nucleolus presents a very remarkable form, becoming stellate by the protrusion of processes, which may again branch in a dendritic fashion (as in the common Thalassicolla nucleata), or it may develop into a very long cylindrical thread, which is disposed in serpentine coils, and in Thalassophysa pelagica passes into the different cæcal processes of the stellate nucleus. In many Sphæroidea, whose skeleton is composed of numerous concentric lattice spheres, the small central spherical nucleus lies at first within the innermost of these (the medullary shell); but afterwards it grows through the meshes of the lattice-work, and the radiating club-shaped processes thus formed (Pl. 11, fig. 5) unite with each other outside the medullary shell, and form an external nuclear sphere which completely encloses the latter. In the Polysphærida (with several concentric lattice-shells) and in the Spongosphærida (with spongy lattice-spheres), this process may be several times repeated, so that eventually the central spherical nucleus attains considerable dimensions, and encloses two or more concentric lattice-shells with their radial connecting rods. The nuclear membrane is in these cases usually penetrated by radial bars, which connect the outermost of the enclosed shells with the remaining cortical shells which surround the central capsule. The same remarkable arrangement is also very common among the Discoidea. The small spherical primary nucleus is in such instances immediately surrounded by the innermost earliest developed lattice-shell, around which the concentric rings are subsequently deposited; it then grows out through the meshes, and the processes fuse outside the ring to form a homogeneous lentiform nucleus (Pl. 43, fig. 15). The same process recurs in certain Prunoidea and Larcoidea, whilst in other Spumellaria of these groups (e.g., Pylonida) the lobate processes of the nucleus remain free.

68. The Nucleus of the Actipylea.—The nucleus of the Acantharia or Actipylea shows very peculiar relations in respect of structure and division, particularly special forms of lobular budding, which belong to the characteristic peculiarities of this singular legion, and are not found among other Radiolaria. The position of the nucleus is always excentric, even in the youngest Acantharia, for the centrogeneous formation of the skeleton, the constant development of the earliest radial portions of it in the middle of the central capsule, forces the nucleus from its normal central position. The majority of the Acantharia, like most Polycyttaria, are precocious, the primary nucleus early dividing into numerous small nuclei (see note A below). Nevertheless there are many exceptions to this rule in different families, e.g., Stauracantha, Xiphacantha, Phatnacantha, and Pristacantha among the Acanthometra, and Stauraspis, Echinaspis, Dodecaspis, and Phatnaspis among the Acanthophracta. In these instances the primary nucleus remains for a long time as a simple excentric ellipsoidal or irregularly round body, even in the fully developed stage, and only at a very late period (sometimes just before the formation of the spores) divides into many small nuclei. Since this serotinous division of the nucleus takes place in different genera of very various groups, it can only be decided by further investigations how widely it is spread among the Acantharia, and upon what circumstances it is dependent (see note B). The division of the nucleus appears to be precocious in the majority of this legion, and a number of small nuclei appear to be early formed by a peculiar process of budding; in most fully developed Acantharia these are disposed in one or two layers under the surface of the central capsule, but if their numbers increase to any considerable extent, the whole space between the skeletal rods becomes filled with small nuclei; sometimes these are homogeneous, sometimes vesicular, 0.002 to 0.012 mm. in diameter; usually they are spherical and have a small nucleolus (compare Pl. 129, figs. 6-11, and note C).

69. The Nucleus of the Monopylea.—The nucleus of the mature forms of the Nassellaria or Monopylea is generally simple or lobate, homogeneous or vesicular and excentric, and appears only to divide into numerous small nuclei just before the formation of the spores. Nevertheless I have sometimes, though not often, seen in representatives of very various families of the Monopylea, the central capsule filled with many small spherical homogeneous nuclei (Pl. 53, fig. 19). Hence all the families of this legion appear to be serotinous, their simple primitive nucleus persisting for a long period. It {xxxix}is commonly placed excentrically, and most usually in the apical or aboral portion of the central capsule, either between its apex and the podoconus, or quite excentrically on the dorsal aspect. The simple nucleus of the Nassellaria usually appears to be vesicular and to possess a somewhat firm membrane, clear contents, and a rather large, dark coloured nucleolus. In many Nassellaria the nucleus is spherical or ellipsoidal (Pl. 53, fig. 11); whilst in many Stephoidea and Spyroidea, where the central capsule is constricted by the sagittal ring and divided into two symmetrical lateral lobes, the nucleus partakes of the same mode of growth and appears in the middle of the capsule as a transversely placed ellipsoid or even as a short cylinder (Pl. 90, figs. 7, 9). The most remarkable modification in the form of the nucleus is to be found in the multi-articulate Cyrtoidea. Here it is usually enclosed in the cephalis and is spherical, ellipsoidal or spheroidal, often flattened almost into a disc. If now the central capsule increase greatly in size and put forth three or four clavate lobes which hang down through the pores of the cortinar septum into the thorax (or even into the succeeding joints), the nucleus usually undergoes similar modification, and three or four finger-like apophyses are developed from its base, which project into the corresponding lobes of the central capsule (Pl. 59, figs. 4, 12, 13).

70. The Nucleus of the Cannopylea.—The nucleus presents the same remarkable structures in all species of the Phæodaria or Cannopylea which have been examined, and closely resembles the germinal vesicle of an amphibian ovum, being a large spherical or spheroidal vesicle with numerous nucleoli. Its diameter usually amounts to half or two-thirds, sometimes even three-quarters, that of the central capsule. The vertical main axis of the latter is also that of the nucleus, which usually lies somewhat nearer to the aboral pole. The nucleus is generally rather more strongly compressed in the direction of the main axis than the capsule itself. The membrane of the vesicular nucleus is thin, but firm, and encloses a clear or finely granular mass of nuclein. The number and size of the contained nucleoli are variable even in one and the same species, and stand in inverse ratio to each other, an obvious result of the gradual process of division. Commonly {xl}from twenty to fifty roundish or spherical, strongly refracting nucleoli, are present; more rarely there are several hundred very small ones. Sometimes the nucleus is penetrated by fine trabeculæ, in whose meshes lie the nucleoli (Pl. 101, fig. 2). In certain nuclei, which contained a few large nucleoli, these were of irregular form, probably the result of amœboid movements (Pl. 101, fig. 1). In the formation of spores in the Cannopylea, the nucleus apparently becomes dissolved, and its numerous nucleoli develop directly into the nuclei or mother-nuclei, which produce the nuclei of the flagellate spores. Furthermore, many Phæodaria seem to multiply by simple cell-division, since very commonly (especially in the Phæocystina and Phæoconchia) two large nuclei (right and left), may be met with in one central capsule; sometimes also a single large nucleus, in which a sagittal constriction marks the commencing division of the capsule (Pl. 101, figs. 2, 36; Pl. 104, fig. 3; Pl. 124, fig. 6, &c.).

71. The Endoplasm or Intracapsular Protoplasm.—In all Radiolaria the intracapsular protoplasm, which, for the sake of brevity, may be termed "endoplasm," constitutes originally, and especially in the earliest stages, the only important content of the central capsule, except the nucleus. In certain Spumellaria and Nassellaria, of simple structure and of small dimensions, this condition persists for a long period, and the endoplasm then appears as a homogeneous, colourless, turbid or finely granular, mucous, semi-solid mass, which cannot be distinguished from the ordinary undifferentiated protoplasm of young cells; no definite structure, and in particular, no fibrillar network, can be discovered in it even by the use of the customary reagents. In the great majority of the Radiolaria, however, this primitive homogeneous condition of the endoplasm is very transient, and it soon undergoes definite modifications, becoming differentiated into separate parts or producing new constituent contents. Such products of the internal protoplasm are in particular hyaline spheres (vacuoles and alveoles), oil-globules, pigment-bodies, crystals, &c. The most important of the differentiations which take place in the endoplasm is that into an internal, granular, medullary substance and an external, fibrillar, cortical substance; although the various legions behave somewhat differently in this respect (§§ 77-80).

72. Intracapsular Hyaline Spheres.—The central capsule of very many Radiolaria contains in its endoplasm numerous spherical bodies of varying size, which consist of watery or albuminous fluid, and have previously been regarded as nuclei, or described as products of the internal protoplasm, under various names, such as "spherical transparent {xli}vesicles" (see note A, below), "albumen spheres" (see B), "gelatinous spheres" (see C), "alveolar cells" (see D), &c. Some of these spheres are perfectly transparent, structureless and of varying refractive power, producing the impression of drops of fluid; others contain various formed constituents, such as oil-globules, fat-granules, pigment-granules, concretions, crystals, &c. From a morphological point of view they may all be divided into two categories, membraneless vacuoles and vesicular alveoles. The vacuoles are simple spherical drops of fluid or of gelatinous material, devoid of a special envelope, but immediately surrounded by the endoplasm. The alveoles, on the other hand, are true vesicles with a thin spherical envelope, enclosing a drop of fluid or jelly. This envelope is commonly very thin, homogeneous, and often scarcely discernible, so that in practice a sharp line of demarcation cannot be drawn between alveoles and vacuoles; the former are usually somewhat larger than the latter. The fact is, nevertheless, certain that the hyaline spheres, which may be isolated on rupturing the central capsule of many Radiolaria, in certain cases, particularly in large species, possess a clear, anatomically demonstrable membrane, whilst in others no such appearance is presented. It may be assumed that the vesicular alveoles are developed from the drop-like vacuoles by increase in size, and by the precipitation of a delicate envelope from the endoplasm. The character common to all these hyaline spheres, whether vacuoles or alveoles, is found in their aqueous, not adipose, constitution, and in their clear transparent appearance, which allows of no structure (the above-mentioned contained bodies excepted) being recognised. Their refractive power and consistency vary somewhat, and probably their chemical constitution still more. Sometimes they are strongly refractive and shining, and sometimes feebly refractive and pale; their consistency shows all intermediate stages between a thin fluid, which readily disappears in water, and a firm, insoluble jelly. As regards their chemical composition (which is probably very variable), the hyaline spheres may be best divided into two groups, the organic and inorganic. The inorganic hyaline spheres are simple drops of saline solution without any carbonaceous constituent; the organic, on the other hand, contain a small quantity of organic matter dissolved in the watery fluid, and may be either albuminous or gelatinous spheres. The formed contents which are commonly present are of very various natures, usually small fat-granules, more rarely larger fat-granules or pigment-granules, sometimes concretions or crystals. In many groups, especially among the large Phæodaria and Collodaria, the numerous hyaline spheres are remarkable for their equal size and even distribution throughout the endoplasm (Pl. 1, figs. 1, 4; Pl. 104, fig. 2, &c.). In some genera belonging to the Thalassicollida the alveoles are of enormous size (Pl. 1, figs. 2, 3); they then become flattened by mutual pressure into polyhedra and distend the central capsule to unusual dimensions (in Physematium and Thalassolampe 8 to 12 mm.).

73. The Intracapsular Fat-Globules.—Fat is present in the central capsule of all Radiolaria in larger or smaller quantities, and generally appears in the form of very {xliii}numerous, small, spherical granules, which are either distributed evenly in the endoplasm (as an emulsion) or enclosed in the vacuoles; the latter, in particular, is the case in most Phæodaria, perhaps generally. In this group each vacuole contains as a rule a single dark, shining fat-granule, and sometimes also an irregular bunch composed of from two to five or more granules. In addition to these small fat-granules (granula adiposa) which are always present, the central capsule of many Radiolaria contains also larger fat-globules (globuli adiposi). These appear to be generally wanting in the Phæodaria, and are on the whole rare in the Acantharia; whilst, on the contrary, they are very common in the Nassellaria and Spumellaria. The Polycyttaria or social Radiolaria are as a rule distinguished by the possession of a single large central oil-globule, which lies in the centre of the central capsule, and is on an average about one-third of it in diameter (Pl. 3, figs. 4, 5). This is absent, however, in those young capsules of the Polycyttaria in which the primary nucleus is centrally situated (Pl. 3, fig. 12). Those species of Polycyttaria whose central capsule reaches a considerable size, often enclose numerous oil-globules, and in Collophidium (species of Collozoum with an elongated cylindrical capsule, Pl. 3, figs. 1, 3) the axis of each capsule is occupied by a row of numerous oil-globules. In the monozootic Spumellaria, in which the nucleus is always centrally situated, the large oil-globules are, of course, excentric, being in apposition to the inner surface of the capsule-membrane (Pl. 1, fig. 3; Pl. 2, figs. 2, 5). In the Discoidea the oil-globules, which are often present in large numbers, form elegant concentric rings around the central nucleus, and in those species with segmented arms, there are one or more transverse rows in each segment (Pl. 43, fig. 15). In the Nassellaria the number and distribution of the oil-globules are dependent upon the form of the central capsule. When this is simple, without lobes, and ovoid or conical, they generally lie in its aboral half above the podoconus (Pl. 51, figs. 5, 13; Pl. 97, fig. 1). When, on the contrary, the basal portion of the capsule sends out three or four dependent processes (as in the majority of the Cyrtoidea), a large globule may generally be seen in the swollen distal part of each conical or ovoid lobe (Pl. 53, fig. 19; Pl. 60, figs. 4-7). In many Stephoidea and Spyroidea, whose central capsule is separated into two lateral portions by the constriction corresponding to the sagittal ring, each of these contains either a single large globule or a group of small ones (Pl. 90, figs. 7, 10). These oil-globules are usually colourless and highly refractive; rarely they are yellow or brown, sometimes rose-coloured, or an intense blood-red (e.g., in Thalassophysa sanguinolenta) or even orange (in Physematium mülleri). In many Spumellaria, and particularly in the Polycyttaria, an albuminous substratum may be recognised in them, which is sometimes disposed in layers, and after extraction of the fat presents the appearance of a laminated sphere. The physiological significance of the oil-globules is twofold; in the first place they tend to diminish the specific gravity of the organism; in the second they may be utilised as a reserve store {xliv}of nutriment. In the latter respect they are of special importance in the process of spore-formation, each flagellate spore usually containing a fat-granule.

74. The Intracapsular Pigment-Bodies.—In the majority of Radiolaria when observed alive, the central capsule is coloured, only in the minority is it colourless. The colour is never diffuse, but always due to the formation of definite pigment granules or vesicles, which are sometimes distributed evenly throughout the endoplasm, sometimes aggregated in the central or peripheral regions. Their form may be either spherical, irregularly rounded, or polyhedral. They vary much in dimensions, but in most cases are immeasurably small, and appear under a high magnifying power as fine dust; occasionally, however, their diameter may amount to from 0.001 to 0.005 or more. The chemical constitution of the intracapsular pigment is unknown in most Radiolaria, and is probably very various. In many instances the pigment-granules consist of fat, in others not. The commonest colours are yellow, red, and brown; violet and blue are rare, and green still rarer. Sometimes a definite tone of colour prevails throughout a whole group, and may then be attributed to inheritance, e.g., red is found in most Sphæroidea, and blue in the Polycyttaria (see note A). One colour is almost always constant in the members of the same species. True pigment-cells, belonging to the Radiolarian organism, do not occur within the central capsule. The peculiar yellow cells which are found in the central capsule of many Acantharia are symbiotic xanthellæ (see § 76).

75. The Intracapsular Crystals.—The crystals found in the central capsule of many Radiolaria may be divided into two groups, of very different significance; small crystals, which are very widely distributed, and large crystals, which occur in only a few genera. The small crystals may also be termed "spore-crystals," since each swarm-spore often contains such a crystal. They are rod-like or spindle-shaped, and consist of an organic substance which probably serves as a reserve of nutriment for the developing spores. Such spore-crystals have been observed in numerous Spumellaria and Acantharia belonging to various families, and are probably present throughout the two legions which make up the Porulosa. On the other hand, they have not been noticed in the Osculosa (Nassellaria and Phæodaria), the few swarm-spores belonging to these groups which have been observed not exhibiting any crystals. The large crystals, which occur in small numbers in the endoplasm, have hitherto only been observed in a few species of Spumellaria, belonging to the Polycyttaria. They were first noticed in the common Collosphæra huxleyi, and regarded as cœlestin. They are also found in the central capsule of many other Collosphærida, e.g., Buccinosphæra (Pl. 5, figs. 11, 12). Crystal-masses, crystal-sheaves, or spherical masses of radiating acicular crystals are enclosed in {xlv}the vacuoles or "albumen globules" of Thalassicola nucleata and other Thalassicollida, as well as in the central capsule of Cœlographis and some other Phæodaria (Pl. 127, figs. 4-7). All these large crystals are probably to be regarded as excretory products.

75A. The Intracapsular Concrements.—Concretions, either mineral or organic, of varying form and constitution, are to be found in the endoplasm of Radiolaria belonging to very different families. They are most abundant and multiform in Thalassicolla nucleata, being usually circular or elliptical discs, which are concentrically laminated and highly refractive, resembling starch-grains. Among them twin forms may frequently be observed, as though the concrements were in process of division (see note A). Similar amyloid concretions are to be seen in the central capsule of different Spumellaria and Nassellaria, e.g., in Cephalospyris triangulata (Pl. 96, fig. 28). Violin-shaped, highly refractive concrements have been observed in the central capsule of numerous Spumellaria, Nassellaria, and Acantharia, e.g., Thalassosphæra, Spongosphæra, Plegmosphæra, Cyrtocalpis, Peripyramis, Botryocella, &c. (see note B). The chemical constitution of these concrements is insufficiently known.

76. The Intracapsular Xanthellæ.—The xanthellæ, zooxanthellæ, or symbiotic "yellow cells" are found within the central capsule only in the Acantharia, whilst in other Radiolaria they only occur in the extracapsulum. They are most frequent in the Acanthometra, rarer in the Acanthophracta, but even in the former they are often wanting. Their number is very variable, but usually small, from ten to thirty in one capsule. They lie for the most part immediately below the capsule membrane, in the cortical layer of the endoplasm. The form of the yellow cells is either spherical or ellipsoidal, often also spheroidal or even lentiform. The diameter varies from 0.01 to 0.03 mm. They possess a distinct membrane and an excentric nucleus, and contain numerous yellow pigment-granules in the endoplasm. This yellow pigment dissolves in mineral acids to form a green fluid, and in other respects also behaves somewhat differently from the yellow pigment in the extracapsular yellow cells of the Spumellaria and Nassellaria. In both cases, however, the xanthellæ are not integral portions of the organism, but unicellular algae, living as parasites or symbiontes in the body.

77. The Endoplasm of the Peripylea.—The intracapsular protoplasm of the Spumellaria or Peripylea is usually distinguished by a more or less complete radial arrangement, which does not occur in the same form in other Radiolaria; it may be regarded as characteristic of this legion, for it probably occurs in all the species at some period of life or other, and stands in a direct causal relationship with the typical structure of the capsule-membrane in all the "Peripylea" (see note A). For as this is commonly perforated by very numerous pores distributed at equal intervals over the whole surface of the capsule, and since a communication between the intra- and extracapsular sarcode takes place through these, the radiate structure of the endoplasm may be readily explained as due to the influence of radial currents which take place continuously or intermittently in the endoplasm. This radiate structure is most obvious when the endoplasm contains no secondary products or only an insignificant amount of these, and thus appears colourless and almost homogeneous, or only finely granular. Under these circumstances, an optical section of the central capsule usually reveals a distinct radial striation; numerous narrow, straight, dark streaks alternating regularly with still narrower clear ones; the latter consist of homogeneous, the former of more or less granular protoplasm (Pl. 20, fig. 1a). Often there may be distinguished in each darker streak a single straight row of strongly refracting (fat?) granules, sometimes several such rows. Occasionally the whole endoplasm becomes divided up into a number of large "radial wedges," club-shaped, conical or pyramidal masses of granular protoplasm, separated by clear divisions of hyaline plasma (e.g., in Actissa radiata, p. 14, where in the optical section of the central capsule, between the membrane and the nucleus, twenty-five dark radial wedges of equal size were separated by thick clear partitions of hyaline protoplasm). In the majority of the Spumellaria this radial striation is partially or entirely concealed by the formation of pigment or of other products. Very often it is only visible in the cortical layer, which lies immediately below the capsule-membrane (Pl. 1, figs. 1, 3). The remarkable "centripetal cones" which characterise the Thalassicollid genus Physematium, and were formerly described as "centripetal cell-groups," are probably a special development of these cortical radial wedges; they are conical cortical bodies, regularly distributed on the inner surface of the membrane of the central capsule, and disposed with the apex turned towards the centre (see note B). More rarely than in the cortical layer, a similar radial structure is to be found in the innermost medullary layer immediately surrounding the nucleus. Here the endoplasm sometimes breaks up into fine radial threads, which are anatomically separable and hang down from the free nucleus as thin processes (see note C). In some cases it is also possible to isolate radial rods from the cortical layer of teased out central capsules.

78. The Endoplasm of the Actipylea.—The intracapsular protoplasm of the Acantharia or Actipylea is often distinguished by a partial or complete radial arrangement like that of the Peripylea, but differing in the number, size, form, and distribution of the radial portions into which the endoplasm is differentiated. For since the pores of the capsule membrane are distributed at equal distances all over the surface in the Spumellaria, whilst in the Acantharia they are arranged in definite groups, and since the number and arrangement of the pores has a direct influence upon the internal currents of the endoplasm, it follows that the radial structure in the latter legion must be very different from that in the former. In addition to this there must not be forgotten the important influence which the early centrogenous formation of the skeletal rods exercises upon the disposition and growth of the intracapsular structures. Hence the endoplasm of the Acantharia does not separate into innumerable thin, closely packed radial wedges or cortical radial rods, but into a small number of large pyramidal portions between which run the radially disposed heterogeneous portions of the contents of the capsule, viz., the radial bars of acanthin and the peculiar intracapsular "axial threads." As a direct consequence of the regular disposition of these heterogeneous radial portions, which is often characteristic of the various families of the Acantharia, a corresponding differentiation of the endoplasm is brought about; it divides into a number of conical or pyramidal portions (radial pyramids), whose bases rest upon the capsule-membrane and whose apices are directed towards the centre of {xlviii}the capsule (the central star of the skeleton). These radial pyramids are, however, but rarely visible, being usually more or less concealed by a dark pigment.

79. The Endoplasm of the Monopylea.—The intracapsular protoplasm of the Nassellaria or Monopylea is distinguished from that of any of the other three legions by the development of a quite peculiar fibrillar structure, the axial "pseudopodial cone," which may shortly be termed the "podoconus" (foot-cone). Since this is in direct correlation with the peculiar structure of the capsular opening, the large "porochora," which is situated at the basal pole of the main axis, it is quite as characteristic of the legion as the latter itself (see note A). The podoconus is primitively a vertical regular cone whose circular base occupies the horizontal porochora or "basal porous area" of the central capsule, while its vertical axis coincides with that of the latter. The apex of the cone, usually somewhat rounded off, is therefore directed towards the aboral or apical pole of the central capsule and separated from it by a larger or smaller interval. In this interval the nucleus originally lies (as in Pl. 51, fig. 13; Pl. 98, fig. 13); but it is usually displaced subsequently and lies excentrically. The cone is of very variable height; on an average its vertical height is about equal to the diameter of its horizontal base; these dimensions are, however, dependent upon the form of the central capsule; the height being greater in slender ovoid or conical capsules, and less in depressed sphæroidal or discoidal ones, than the diameter of the base. The podoconus consists of differentiated endoplasm, which becomes more deeply stained by carmine and offers greater resistance to solvents than the surrounding finely granular protoplasm. The apex, especially, becomes very intensely stained. It always exhibits a very characteristic fine but distinct striation, numerous straight radial lines diverging from the apex of the cone towards the base. The number of these striæ appears to correspond with that of the vertical rods in the porochora, and each of these latter stands apparently in direct communication with the basal end of an apical stria (§ 59). These threads are probably differentiated constant contractile threads of endoplasm, or even myophanes, comparable with the contractile cortical threads of the Cannopylea and the permanent axial threads of the Actipylea. The numerous modifications, {xlix}undergone by the form and contents of the central capsule in the different groups of Monopylea, especially those due to the formation of the skeleton, are not without influence upon the podoconus. The most important divergencies from the above described primary form are the following:—(1) The vertical axial cone becomes oblique, its axis inclining in the sagittal plane and approaching either the dorsal or the ventral wall of the capsule; the cause of this appears to be usually the excentric development of the growing nucleus or the formation of a large oil-globule. (2) The smooth mantle of the podoconus becomes divided by three longitudinal furrows into three equal prominent ridges, which correspond to three circular lobes in the porochora; the cause of this basal triradial lobular formation lies probably in the triradial development of the skeleton in many Nassellaria or in the cortinar structure of the collar septum. (3) The simple podoconus splits into three or four elongated lobes, which eventually become almost completely separated and correspond to the lobes of the central capsule, in the axial wall of which they lie as longitudinally striated bands. The behaviour of these bands justifies the hypothesis that the podoconus is a muscular differentiated portion of the endoplasm and is composed of myophane fibrillæ, whose contraction determines the opening of the central capsule.

80. The Endoplasm of the Cannopylea.—The intracapsular protoplasm of the Phæodaria or Cannopylea is distinguished from that of the other three legions by several characteristic peculiarities, which are very important, since they stand in causal relation to the typical structure of the capsule-membrane and in particular of its {l}remarkable aperture. In the case of many and perhaps of all Phæodaria the endoplasm is differentiated into a granular medullary and a thin fibrillar cortical layer, the former of which usually encloses numerous small vacuoles, while the latter contains muscular fibrillæ. In the voluminous central capsule of large Phæodaria the whole cortical layer of the endoplasm, which lies immediately below the delicate inner capsule-membrane, sometimes appears delicately and regularly striated, and most distinctly so under the apertures, towards the centre of each of which the dark striæ are radially directed (see note A, below). These striæ are probably contractile muscular fibrillæ; or "myophanes," by whose contraction the openings are voluntarily widened. In the Tripylea this fibrillar star is much more strongly developed under the astropyle (the main opening) than under the parapylæ (or accessory openings); and probably the peculiar radial structure of the operculum of the former is due to the stronger development of these radial fibrils (being their impression). In many Phæodaria, indeed, the fine myophane fibrils are only visible under the apertures, whilst in others they form a continuous fibrillar cortical layer on the whole inner surface of the inner capsule-membrane; the fine fibrillæ run meridionally from one pole of the main axis to the other; perhaps the whole central capsule may change its form in consequence of their contractions. The medullary portion of the endoplasm, which lies below this thin cortical layer, is usually finely granular in the Phæodaria, and permeated by numerous spherical vacuoles, which are noteworthy from their equal size and regular distribution. Each clear vacuole usually contains a dark shining fat-granule, more rarely a group of such granules (see note B). Compare § 60, and Pl. 101, figs. 1-3; Pl. 104, figs. 1, 2; Pl. 111, fig. 2; Pl. 128, fig. 2, &c.

Chapter III.—THE EXTRACAPSULUM.

(§§ 81-100).

81. The Components of the Extracapsulum.—The extracapsulum or extracapsular malacoma, under which name are included all those parts of the soft body which lie outside the central capsule, consists of the following constant, and important constituents:—(1) The calymma or extracapsular jelly-veil; (2) the sarcomatrix or layer of exoplasm immediately surrounding the membrane of the central capsule; (3) the sarcodictyum or network of exoplasm, covering the surface of the calymma; (4) the pseudopodia or radial fibres of exoplasm, which may again be subdivided into intracalymmar pseudopodia, uniting the sarcomatrix and sarcodictyum, and extracalymmar pseudopodia, radiating freely into the water outside the calymma.

82. The Calymma.—The calymma or extracapsular jelly-veil of the Radiolaria is always the most voluminous portion of the extracapsulum, and in spite of its simple structureless constitution is of great morphological and physiological importance. In all Radiolaria this gelatinous mantle completely surrounds the central capsule, but is separated from its outer surface by a continuous, though thin, layer of exoplasm, the sarcomatrix. The pseudopodia radiating from the latter pierce the calymma, form the sarcodictyum at its surface, and radiate from its nodal points freely into the surrounding water. The calymma is rarely visible in living freshly captured Radiolaria, examined in sea-water, for its gelatinous substance is perfectly hyaline, colourless and pellucid, and possesses the same refractive index as sea-water; but when the object is removed from this fluid and transferred to carmine solution or some other colouring matter, the extent and figure of the calymma become apparent, for the staining fluid does not at first penetrate into the gelatinous material. When this has taken place, however (after a longer or shorter time), and the gelatinous material has become coloured, its form and size may be observed by the converse experiment; the object is transferred once more to water and the outlines of the calymma become as clear as those of the central capsule. The same is the case with dead specimens in which the sticky surface of the calymma has become covered with dust.

83. The Structure of the Calymma.—The extracapsular jelly-veil appears structureless in most Radiolaria, inasmuch as it represents a homogeneous pellucid excretion of the exoplasm and contains neither fibres nor other formed structures. In some groups, however, definite structural characters become secondarily developed. The most common and striking of these is the formation of alveoles, which takes place in the extracapsulum (see § 86). In consequence of this the calymma assumes a remarkable frothy consistency and appears to be composed of large, clear, thin-walled vesicles; this is especially the case in the Collodaria (Colloidea, Pls. 1, 3, and Beloidea, Pls. 2, 4), and in many large Phæodaria, especially among the Phæocystina (Phæodinida and Cannorrhaphida, Pl. 101, and Aulacanthida, Pls. 102-104). More rarely the calymma is not permeated by vacuoles, but there appear in it fine striæ parallel to the surface as though it were composed of thin concentric laminæ like an onion; perhaps these are the expressions of a different quantity of water in the various layers. In the calymma of many Radiolaria thin, straight, radial lines are to be seen, which are probably pseudopodia, and not to be attributed to any structural modification, or they may be slender canals which serve for the exit of the pseudopodia. On the outer surface of the calymma of different Radiolaria, and especially in the Acantharia, a peculiar network of fibres is to be found, composed of polygonal meshes, like elastic fibres, probably due to a local thickening of the jelly. These polygonal meshes are often very regularly distributed between the radial spines of the Acanthometra, and stand in a definite relation to them. The fibres which form the meshes are often rather strong, resembling elastic fibres, as above-mentioned, and either simple or composed of bundles of very fine fibrillæ (L. N. 33, p. 15, Taf. i. fig. 1, Taf. ii. fig. 4).

84. The Consistency of the Calymma.—The gelatinous material of which the calymma of the Radiolaria consists is a pellucid mass, rich in water and usually quite hyaline and structureless; its consistency is very variable. In the majority of the Radiolaria it may perhaps be about equal to that of the jelly which composes the umbrella of most Medusæ; but as in these latter it may vary between very wide extremes, constituting on the one hand a very soft jelly-mantle, offering but little {liii}resistance to mechanical influences and almost disintegrating under the eyes of the observer, and on the other hand forming a firm gelatinous shell, comparable to cartilage in hardness, elasticity, and power of mechanical resistance. In many Radiolaria of large dimensions with an alveolar calymma (especially in numerous Collodaria and Phæodaria) this may be split by means of dissecting needles and the central capsule extracted like the stone from a cherry, and then it is easy to ascertain that the firmness and elasticity of this jelly-veil are not less than those of a cherry. The different degrees of consistency in the various Radiolaria may be dependent either upon the relative amount of water which they contain, or upon qualitative or quantitative variations in the organic substance of which the jelly consists. Great importance is to be attached to the considerable consistency of the calymma, because it furnishes the indispensable groundwork for the deposition of many parts of the skeleton and particularly of the lattice-shells.

85. The Primary and Secondary Calymma.—In most Radiolaria the external form and volume of the calymma are different at different stages of growth, and this difference is mainly dependent upon the development of the skeleton. Hence it is advisable to distinguish in general the primary from the secondary calymma. The primary calymma is in the great majority of Radiolaria a perfect sphere, in the middle of which lies the concentric central capsule; on the surface of this gelatinous plate the primary spherical lattice-shell is secreted in most Spumellaria and Acanthophracta, as well as in those Phæodaria which possess a spherical shell; in the remaining Phæodaria also and in the Nassellaria, where the lattice-shell is not spherical but monaxon, it is secreted on the surface of the primary calymma. This takes place at a definite time, very important in the development of the Radiolarian, which for the sake of brevity we shall term the "lorication-period." Since the firm surface of the primary calymma furnishes the necessary foundation for the deposition of the primary lattice-shell, it is of the greatest mechanical significance in all shell-bearing Radiolaria. The secondary calymma arises only after the lorication-period by further growth of the primitive jelly-mantle and in the fully developed Radiolarian usually encloses wholly or partially the external parts of the skeleton, in consequence of which it assumes the most various forms. Very often the secondary calymma is polyhedral, being stretched between the radial spines of the skeleton, the distal ends of the latter then forming the fixed points of the gelatinous polyhedron.

86. The Extracapsular Vacuoles and Alveoles.—The calymma of the Radiolaria usually appears completely homogeneous and hyaline without any structure; sometimes it encloses numerous clear vesicles, vacuoles or alveoles, and then assumes a frothy appearance, the expression of a more or less distinct alveolar structure. {liv}The clear vesicles to which this is due are either spherical, or polyhedral from mutual pressure, and like the similar ones in the central capsule may be divided into membraneless vacuoles and vesicular alveoles. The vacuoles are simple drops of fluid, without a special envelope, and immediately surrounded by the gelatinous substance of the calymma, in which they appear as simple cavities. The alveoles on the contrary are true vesicles, with a thin envelope, which encloses a drop of fluid or a globule of jelly; in the latter case its contents are different in refracting power and amount of contained water from the substance of the surrounding calymma. A sharp boundary between the membraneless vacuoles and the vesicular alveoles cannot be drawn in the case of the extracapsular hyaline spheres any more than in the intracapsular; the envelope of the alveoles is sometimes very distinct and even anatomically separable, whilst at other times it is very thin and scarcely recognisable; it may occasionally arise and disappear within a very short time (see note A). There is no doubt that in the calymma as in the central capsule the vesicular alveoles are secondary products, which have arisen from the vacuoles by the secretion of an enveloping membrane. This membrane is either a delicate sheath of exoplasm, or a firmer and more resistant skin, distinct from the exoplasm, and probably an excretion from it (e.g., Pl. 4, figs. 2, 3). In many cases the outer surface even of the vacuoles is covered by a network of pseudopodia, which form a sarcoplegma similar to a fenestrated alveolar membrane. The colourless pellucid fluid in the vacuoles and alveoles is usually simple sea-water, more rarely it contains a small quantity of albumen ("albumen-spheres") or jelly ("gelatinous spheres"). The size of these spheres is very variable. Quite small vacuoles may be found in the calymma of many Radiolaria. Large vacuoles, on the other hand, producing the appearance of an alveolar structure, are confined to but few groups, to a part of the Spumellaria (Colloidea, Beloidea, and a few Sphæroidea), and to the Phæocystina (Phæodaria with incomplete skeleton); besides they occur only rarely in individual genera, e.g., Nassella among the skeletonless Nassellaria. Since the volume of the calymma is much increased by the development of vacuoles, and the power of mechanical resistance is at the same time much increased, the fact is explained that the vacuoles occur mainly in Radiolaria which have no skeleton or only an incomplete one (see note B). Among the monozootic Collodaria the alveolar structure is especially well developed in the following genera; Thalassicolla (Pl. 1, figs. 4, 5), Thalassophysa, Thalassoplancta, Lampoxanthium (Pl. 2, figs. 1, 2); among the Phæodaria in most genera of the Phæodinida, Cannorrhaphida and Aulacanthida (Pls. 101-104), and probably also in other voluminous Phæodaria (e.g., Phæosphæria). The alveoles or vacuoles in the calymma of these large Radiolaria lie usually in several layers, one above another, and increase in size from within outwards. The Polycyttaria or social Radiolaria (the three families Collozoida, Sphærozoida and Collosphærida) without exception have an alveolar structure, and the special form of {lv}their colonies or cœnobia is to a great extent determined by the development, number, size and arrangement of the alveoles in their calymma (compare Pls. 3-8). In these cases there is not unfrequently developed a large central alveole (see note C) whose thickened wall encloses a globe of jelly and serves as the central support of the whole colony (Pl. 5, fig. 1). Still more striking, however, is the arrangement of certain Polycyttaria, where each individual of the colony (or each central capsule with its calymma) is enclosed in a large alveole, whose firm wall often attains considerable thickness (Pl. 4, figs. 2, 3). The whole colony then appears as an aggregate of numerous cells, each of which possesses two envelopes, the inner central capsule and the outer alveolar membrane; between these lies in the Collosphærida the siliceous lattice-shell (Pl. 6, fig. 2). These pericapsular alveoles may be regarded as an outer cell-wall more correctly than the membrane of the central capsule itself, but the arrangement may also be compared to the temporary encystation of other Protista (see note D).

87. The Extracapsular Fat-Globules.—Fat is probably as widely distributed in the exoplasm as in the endoplasm of the Radiolaria; a considerable proportion of the small, dark, highly refractive granules appear to consist of fat; most likely they are for the most part direct products of metastasis. These widely-spread granules, which are sometimes coloured, and which by their passive motion produce the phenomenon of granular circulation in the exoplasm, are not the only fatty structures in the extracapsulum; larger globules sometimes occur. In certain large Collodaria (e.g., Thalassicolla melacapsa, Pl. 1, fig. 5; Thalassophysa sanguinolenta, &c.) radial series of oil-globules are found in the calymma, especially in its proximal portion; in others the central capsule is surrounded by a layer of oil-globules (situated in the sarcomatrix). In the Phæodaria a part of the phæodium appears to consist of fat-globules.

88. The Extracapsular Pigment.—The formation of colouring matters in the extracapsulum is on the whole rare in the Radiolaria, apart from the "yellow cells" (see § 91) and from the peculiar phæodium of the Phæodaria, which will be separately treated of in the next paragraph. Considerable masses of extracapsular pigment, usually black or blue, rarely brown or red, are found only in a few Radiolaria belonging to the first three legions; most often in the Spumellaria. Some large Collodaria, e.g., the common Thalassicolla nucleata and a few other species of this genus (Pl. 1, fig. 4), are characterised by a rich deposit of black or blue pigment in the sarcomatrix and in the proximal portion of the calymma. Brown pigment is deposited in the calymma of many Sphæroidea and Discoidea, as well as of some Nassellaria (Cystidium, Tridictyopus, &c.). In a part of the Acantharia red pigment granules are thickly strewn in the sarcoplegma and pass along the free pseudopodia, as for example in Actinelius purpureus and Acanthostaurus purpurascens. The composition and significance of these extracapsular pigments are not completely known.

89. The Phæodium of the Phæodaria.—The Phæodaria, which are distinguished from the other three legions of Radiolaria by the double membrane of the central capsule, and the peculiar structure of the main-opening (astropyle), differ also in other points, the most important of which is the constant presence of a voluminous mass of extracapsular pigment. This possesses a peculiar constitution and special significance, and is not to be confounded with the extracapsular pigment-granules of other Radiolaria (e.g., Thalassicolla), and hence it has been distinguished by the name "Phæodium," and the individual granules which compose it as "Phæodella" (see note A). The phæodium is always excentric in position relatively to the central capsule, of which it {lvii}surrounds the oral half in the form of a voluminous concavo-convex cap, hiding the astropyle at its basal pole so completely that the latter is rarely visible until the phæodium has been removed (Pls. 99-104; Pl. 115, fig. 8; Pl. 123, &c.). The central capsule is generally almost completely embedded in the phæodium, so that only its aboral pole (with the two parapylæ in the Tripylea) projects. In the Phæogromia, in which the lattice-shell possesses a special opening and the central capsule lies excentrically in the aboral position of its interior, the phæodium occupies the oral aspect, between the capsule and the aperture (Pls. 99, 100, 118-120, &c.). In the peculiar family Cœlographida (Pls. 126-128) a special receptacle (galea with its rhinocanna) for the phæodium is developed outside the bivalve shell, within which the central capsule lies. The proboscis, which in all Phæodaria arises from the centre of the astropyle, lies in the vertical axis of the phæodium and is entirely surrounded by it. The volume of the phæodium in the majority of the Phæodaria may be said to be about as great as that of the central capsule, although in some species it is considerably larger. Its colour is always dark, usually between green and brown, commonly olive-green or blackish-brown, rarely reddish-brown or black. The phæodellæ or pigment-granules which make up the greater part of the phæodium (see note B) are irregular in form and unequal in size and show no definite structure; usually they are spherical or ellipsoidal, and exhibit fine parallel striæ which run transversely or obliquely (Pl. 101, fig. 3, 6, 10; Pl. 103, fig. 1, &c.). Between the larger granules is usually found a thick dust-like mass of innumerable very small grains. The physiological significance of this peculiar phæodium is still unknown, but is probably considerable, judging from its large size and especially from its constant topographical relation to the astropyle; the latter consideration would lead to the supposition that it plays an important part in the nutrition and metastasis of the Phæodaria (see note C).

90. The Extracapsular Xanthellæ.—Xanthellæ or Zooxanthellæ, symbiotic "yellow cells," are very commonly found in the extracapsulum of the Radiolaria, especially in many Spumellaria and Nassellaria; whilst in the Acantharia similar yellow cells usually only occur within the central capsule, and in the Phæodaria their {lviii}presence has not been certainly demonstrated. The extracapsular Xanthellæ are found most abundantly in the Collodaria, both in the monozootic Thalassicollida and in the polyzootic Sphærozoida. They occur in smaller numbers in the Sphærellaria, and in many divisions of the latter they seem to be entirely absent. Also it sometimes happens that, though present in large numbers in some Spumellaria, they are entirely absent in others nearly related to them; indeed, this has also been observed in the case of different individuals of the same species. This fact alone is sufficient to show that the Xanthellæ are not an integral part of the Radiolarian organism (as was formerly believed) but parasites or more correctly symbiontes, which live as inhabitants of the calymma. More recent investigations have shown, that besides the yellow pigment-grains they contain starch or an amyloid substance, that is to say, vegetable reserve materials, that their thin envelope contains cellulose, and that their yellow colouring-matter resembles chlorophyll and is related to that of the Diatomaceæ ("Diatomin"). Hence they are now generally regarded as unicellular Algæ, nearly related to those which occur as symbiontes in other marine animals (Exuviella, &c.). The starch, which they develop with the formation of oxygen, may serve as nutriment to the Radiolaria, while the carbonic acid yielded by the latter is also beneficial to the Xanthellæ. The form of the Xanthellæ is usually spherical and elliptical, often also sphæroidal or discoidal. Their diameter is usually between 0.008 and 0.012 mm., rarely more or less. The differences exhibited by Xanthellæ which live in different groups of Radiolaria demand further investigation, which will perhaps lead to the establishment of several species of the genus Zooxanthella. At present Zooxanthella extracapsularis, in the calymma of Spumellaria and Nassellaria, may be clearly distinguished from Zooxanthella intracapsularis, in the central capsule of the Acantharia.

91. The Exoplasm or Extracapsular Protoplasm.—The extracapsular protoplasm, which may be shortly termed the "exoplasm" (or ectosarc), is primitively in all Radiolaria (and especially in their earliest development stages) the only important constituent of the extracapsulum, besides the calymma. Although the extracapsular and intracapsular protoplasm of the Radiolaria are everywhere in direct communication, and although the openings in the membrane of the central capsule bring about an interchange between them, still the two portions of sarcode show certain constant and characteristic differences, which are due to the physiological division of labour between the central and peripheral parts of the body and their corresponding morphological differentiation. The extracapsular, like the intracapsular, protoplasm is originally homogeneous, but may afterwards become differentiated in various ways, producing the special constituents of the extracapsulum. Such "external protoplasmic products" are vacuoles, pigment-bodies, &c. More important, however, are the topographically different sections into which the exoplasm may be divided according to its relations to the central capsule and the calymma. In this respect the following parts may be generally distinguished—(1) the Sarcomatrix, or fundamental layer of the exoplasm, which surrounds the central capsule as a continuous sheath of sarcode and separates it from the calymma; (2) the Sarcoplegma, an irregular network of the exoplasm, which spreads throughout the gelatinous material of the calymma; (3) the Sarcodictyum or network of sarcode on the outer surface of the calymma; and (4) the Pseudopodia, which project outwards from the latter and radiate into the water.

92. The Sarcomatrix.—The sarcomatrix, being "the fundamental layer of the pseudopodia" (or "matrix of the exoplasm"), constitutes the proximal innermost section of the extracapsular sarcode, and in all Radiolaria forms a thin continuous mucous layer, which covers the whole outer surface of the central capsule and separates it from the surrounding calymma (see note A, below). The sarcomatrix communicates internally {lx}through the openings of the central capsule with the endoplasm, whilst externally the pseudopodia or mucous threads arise from it, which by their union form the sarcoplegma. The sarcomatrix is only interrupted in the Spumellaria and Acantharia by those parts of the skeleton which perforate the membrane of the central capsule. In all Nassellaria and Phæodaria, as in the Collodaria, it appears as a perfectly continuous sarcode-envelope of the central capsule. Its thickness is variable; in general it is most strongly developed in the Spumellaria and Phæodaria, less so in the Nassellaria, and is thinnest in the Acantharia. The thickness seems, however, to vary even in one and the same individual, the difference depending partly upon the different stages of development and partly upon nutritional conditions. After abundant inception of nutriment the thin protoplasmic layer of the matrix is thickened and turbid, rich in granules and irregular masses, which are probably due to enclosed but only half-digested food; xanthellæ also, as well as foreign bodies taken up with the nutriment, such as frustules of Diatoms and shells of smaller Radiolaria, and of pelagic infusoria, larvæ, &c., are often, especially in large individuals, aggregated in considerable quantities in the matrix. After long fasting, on the contrary, this is poor in these enclosed bodies and in granules; it then forms a thin colourless more or less hyaline mucous coating to the central capsule. From a physiological standpoint the sarcomatrix is to be regarded as the central organ of the extracapsulum, and as of pre-eminent significance. Probably it is not only the most important organ for the nutrition of the Radiolaria (especially for digestion and assimilation in particular), but perhaps is also the central organ of perception. On the other hand the sarcomatrix belongs to those components of the Radiolarian organism which take no part in the formation of the skeleton.

93. The Sarcoplegma.—By the name sarcoplegma, as distinguished from the remaining extracapsular sarcode, is understood the intracalymmar web of exoplasm or "ectosarcode network," which ramifies within the gelatinous mass of the calymma. Internally it is in direct connection with the continuous sheath (sarcomatrix), which encloses the central capsule, whilst externally it is in contact with the superficial sarcode network (sarcodictyum) which surrounds the calymma. The configuration of this exoplasmic web, which penetrates the jelly-veil in all directions, is exceedingly variable; in most Radiolaria it is extremely irregular in form, like the protoplasmic network in the ground-substance of many kinds of connective tissue. In some groups, however, it assumes a rather regular shape which it appears to retain (e.g., in many Acantharia). It must be assumed also that in those instances where the consistency {lxi}of the calymma approaches that of cartilage, the tracks of the exoplasmic threads remain constant, but accurate observations are wanting as to how far the configuration of the sarcoplegma is constant or variable in the different groups, as well as regarding its peculiar behaviour in those Radiolaria whose calymma is characterised by the formation of vacuoles or alveoles (see § 86). Usually it envelops the larger alveoles in the form of a reticulate veil. In many Collodaria the exoplasm is aggregated at certain points of the intracalymmar web, so that large balls or amœboid bodies appear to be distributed between the alveoles, e.g., in Thalassophysa pelagica and Thalassicolla melacapsa (Pl. 1, figs. 4, 5). The sarcoplegma is metamorphosed directly into silex in the Radiolaria spongiosa, or those genera which possess a spongy cortical skeleton, and were formerly known as Spongurida; to this category belong the Spongosphærida (Pl. 18) and Spongodiscida (Pl. 47) as well as certain Nassellaria and Phæodaria. The single siliceous spicules, which are irregularly interwoven to form the spongy web, are to be regarded as the silicified threads of the intracalymmar sarcode network. From a physiological point of view the sarcoplegma is of importance both for the nutrition and motion of the Radiolaria, since it brings the sarcomatrix and the sarcodictyum, with the pseudopodia which radiate from it, into direct communication.

94. The Sarcodictyum.—The sarcodictyum may be defined as the extracalymmar network of exoplasm, and is a reticular covering which lies upon the outer surface of the gelatinous calymma. Internally, the sarcodictyum is in direct communication with the sarcoplegma, or the web of exoplasmic threads which ramifies in the gelatinous substance of the calymma; externally, on the other hand, the pseudopodia radiate freely from it; thus its relation to these is similar to that which the sarcomatrix bears to the roots of the sarcoplegma. Relations similar to those which have led to the separation of the primary from the secondary calymma, induce us to distinguish also a primary and secondary sarcodictyum. The original or primary sarcodictyum ramifies over the surface of the original or primary calymma, and like this is of pre-eminent importance in the formation of the primary lattice-shell; if we regard the surface of the primary calymma as the indispensable foundation for the deposition of this latter, then the primary sarcodictyum furnishes the material from which it is developed: silex in the Spumellaria and Nassellaria, a silicate of carbon in the Phæodaria, and acanthin in the Acantharia. It may indeed be said that the primary lattice-shell of the Radiolaria arises by a direct chemical metamorphosis of the primary sarcodictyum, by a chemical precipitation of the dissolved skeletal material (silex, silicate, or acanthin), which was stored up in the exoplasm of the sarcodictyum. Hence a deduction from the special conformation of the former to that of the latter is permissible. The particular form of the primary lattice-sphere with its regular or irregular meshes is due to the corresponding form of the primary sarcodictyum; both regular and irregular forms of this {lxii}commonly occurring. The form of the regular sarcodictyum with circular or regular polygonal, usually hexagonal, meshes is constantly maintained during the formation of the regular lattice-shells (e.g., Pl. 12, figs. 5-10; Pl. 52, figs. 8-20; Pl. 96, figs. 2-6; Pl. 113, figs. 1-6). The form of the irregular sarcodictyum, on the other hand, with irregular polygonal or roundish meshes, persists during the development of the irregular lattice-shells (e.g., Pls. 29, 70, 97, 106). All this is true also of the secondary sarcodictyum, or the exoplasmic network which ramifies over the surface of the secondary calymma. The secondary lattice-shells, which are deposited on the surface of the latter, retain the configuration of the secondary sarcodictyum, by the chemical metamorphosis of which they have originated; this is the case in many Spumellaria which develop several concentric lattice-shells (Pl. 29), in some Nassellaria (Pl. 54, fig. 5), in the Phractopeltida among the Acantharia (Pl. 133), and in the double-shelled Phæodaria, Cannosphærida, and part of the Cœlodendrida and Cœlographida (Pls. 112, 121, 128). In those Radiolaria which form no lattice-shell whatever, the conformation of the sarcodictyum is usually irregular, with meshes of irregular form and unequal size; sometimes, however, they seem to be very regular, as in many Acanthometra (Pl. 129, fig. 4).

95. The Pseudopodia.—On the whole the pseudopodia or thread-like processes of the exoplasm exhibit in the Radiolaria the same characteristic peculiarities as in all true Rhizopoda; they are usually very numerous, long and thin, flexible and sensitive filaments of sarcode, which show the peculiar phenomena of granular movement. Their physiological significance is in several respects very great, for they serve as active organs for the inception of nutriment, for locomotion, sensation, and the formation of the skeleton (see note A, below). The presence of a calymma, however, which distinguishes the Radiolaria from the other Rhizopoda, brings about certain modifications in the behaviour of the pseudopodia. If in general all the threads, which arise from the sarcomatrix or fundamental layer and radiate outwards, be called "pseudopodia," then that part of them which is included in the gelatinous substance of the calymma and forms the sarcoplegma may be termed the "collopodia" (or intracalymmar pseudopodia), and the remaining portion, which passes outwards from the sarcodictyum freely into the water, may be described as "astropodia" (or extracalymmar pseudopodia). In many Radiolaria these two portions present some differences in morphological and physiological respects, and certain distinctions are probably generally present (see note B). Apart from this universal differentiation in the different groups of the Radiolaria, specially modified forms of pseudopodia may be recognised as the axopodia and myxopodia of the Acantharia (see § 95, A), and the sarcode-flagellum of certain Spumellaria (see note C).

95A. The Myxopodia and Axopodia.—The two forms of pseudopodia which we distinguish as myxopodia and axopodia differ markedly from each other both morphologically and physiologically. The myxopodia, or ordinary free pseudopodia, which are found in large numbers in all Radiolaria, and constitute their most important peripheral organs, are simple homogeneous exoplasmic threads, which arise from the sarcodictyum or extracalymmar sarcode network, and radiate freely into the water; here they may branch and combine by anastomosis to form a changeable network, but they never contain an axial thread. The axopodia, on the other hand, are differentiated pseudopodia, which consist of a firm radial thread, and a soft covering of exoplasm; they penetrate the whole calymma in a radial direction and project freely from its surface, and generally (if not always) they are produced inwards to the middle of the central capsule, perforating its membrane; their proximal end is lost in a dark central heap of granules. Such axopodia are at present known with certainty only in the Acantharia, where they are widely, and perhaps universally, distributed. Their development in this legion probably stands in direct causal relation to the peculiar structure of the central capsule and the centrogenous formation of the skeleton. Since the radial skeletal rods of the Acanthometra possess originally a thin coating of protoplasm, it may be said that the centrogenous axopodia of this group became differentiated in two ways, the firm axial threads of one section remaining very thin and covered by protoplasm, whilst those of the other section became metamorphosed into radial bars of acanthin. This hypothesis acquires more probability from the regular distribution and arrangement of the axopodia in the Acantharia; they usually stand at fixed intervals {lxiv}between the radial bars, singly or in groups; sometimes their number seems to be not greater than that of the bars, whilst in other cases a circlet or group of axopodia corresponds to each radial bar. Perhaps their fine axial thread consists of acanthin. At all events the axopodia are constant organs (probably sensory, like the "palpocils") and not retractile like the movable myxopodia.

96. The Myophriscs of the Acanthometra.—The Acanthometra are characterised by a very peculiar differentiation of the exoplasm, namely, by the formation of myophriscs or contractile threads from the sarcodictyum. In most (and perhaps in all) Acantharia of this order each radial bar is surrounded by a circlet of such contractile threads, which was first described as a "ciliary corona" (see note A, below). The number of contractile threads in each circlet usually amounts to from ten to twenty, rarely being more than thirty and less than eight; it often appears to be constant in the individual species (see note B). In the living state the myophriscs are long, thin filaments, the pointed distal end of which is inserted into the radial bar, whilst the thicker proximal end is attached to the surface of the calymma, which is elevated round the base of each rod into the form of a gelatinous cone or skeletal sheath (see note C). Probably the myophriscs lie on the outer surface of the apical portion of this gelatinous cone, and are hence to be regarded as exoplasmic threads differentiated from the sarcodictyum. Sometimes, however (as in Acanthochiasma), they fuse into a contractile membrane and form the envelope of a cone, whose interior is occupied by a gelatinous papilla of the calymma. On mechanical irritation the myophriscs contract rapidly and suddenly, like muscle-fibrillæ, becoming at the same time thicker, and hence are very different from pseudopodia. Their distal point of insertion being fixed to the firm acanthin rod, they raise by their contraction the skeletal sheath, to which their bases are attached or in the surface of which they lie. The result of their contraction is therefore a distention and increase in volume of the calymma, with which is no doubt connected an inception of water into the gelatinous mass, and hence a diminution in its specific gravity. Probably the Acanthometra contract their myophriscs voluntarily when they wish to rise in the water; when these relax the calymma collapses owing to its elasticity, water is then expelled and the specific gravity increases. From a physiological point of view, then, the myophriscs are to be regarded as a hydrostatic apparatus, morphologically as myophanes or muscular fibrillæ, such as also occur in the intracapsular protoplasm (see §§ 77-80). On more violent irritation and after the death of the Acanthometra the myophriscs separate from the radial bars and remain attached to the distal ends of the conical gelatinous sheaths as free "ciliary coronas." At the same time, {lxv}they melt into short, thick, hyaline rods, the so-called "gelatinous cilia." The myophriscs are found only in the order Acanthometra, and are wanting in the Acanthophracta, as well as in the other three legions of Radiolaria.

97. The Exoplasm of the Peripylea.—The extracapsular protoplasm of the Spumellaria or Peripylea is in communication with the intracapsular sarcode by the innumerable fine pores of the capsule-membrane, and like these pores is evenly distributed over the whole surface. The sarcomatrix which immediately surrounds the central capsule is moderately strong, and sends out innumerable long, thin pseudopodia, which probably correspond to the pores of the membrane. Their number is markedly greater in the Spumellaria than in the other three legions. The ramifications and communications which the radiating fibres of the sarcomatrix undergo within the calymma, apparently present the most manifold variations, so that the sarcoplegma or intracalymmar network thus formed has very diverse forms. On the surface of the calymma the exoplasmic threads constitute a variously disposed sarcodictyum, a regular or irregular exoplasmic network, by the silicification of which a primary lattice-shell arises in the majority of the Spumellaria. The free ends of the pseudopodia, which arise from this extracalymmar network and radiate out into the water, appear in most Spumellaria to be relatively short, but exceedingly numerous. Specially modified pseudopodia and axial threads in particular do not seem to occur in this legion. Perhaps, however, among the latter may be reckoned the remarkable pseudopodia which combine to form the sarcode flagellum in many Discoidea (and perhaps in other Spumellaria). This axoflagellum is a particularly strong thread of sarcode, arising from a definite point in the central capsule; it is cylindrical or slenderly conical in form, much longer, stronger, and more contractile than the ordinary pseudopodia; it contracts in a serpentine fashion on mechanical irritation and seems to originate by the fusion of a bundle of pseudopodia (compare § 95, C).

98. The Exoplasm of the Actipylea.—The extracapsular protoplasm of the Acantharia or Actipylea differs in several important respects from that of other {lxvi}Radiolaria, and appears to undergo more significant differentiations than that of the three other legions. Since the pores in the wall of the central capsule are not distributed evenly and at equal intervals over its whole surface (as in the Peripylea), but rather exhibit a regular disposition in groups at unequal intervals, the number of projecting pseudopodia is much less and the law of their arrangement different from that which obtains in the Peripylea (§ 58). In many and probably in all Acantharia they are divided into two groups, those which arise from the centre of the capsule and possess firm axial threads, and those which have not these characters (compare § 95, A). The axopodia, or stiff pseudopodia with axial threads, arise from the centre of the capsule, are present in much smaller numbers than the soft and flexible myxopodia, and are regularly disposed between the radial bars of acanthin, usually so that they are as far removed from them as possible, i.e., in the centre between each three or four bars; these latter may indeed be regarded as strongly developed axial threads, which have become changed into acanthin (§ 95, A). The soft myxopodia, or pseudopodia without axial threads, are much more numerous than the others, and arise from the sarcodictyum or exoplasmic network which ramifies over the surface of the calymma. Their number and arrangement seem, however, in many (if not in all) Acantharia to be regular and not to possess the extraordinary variability seen in the other three legions. In many Acanthometra the sarcodictyum exhibits a symmetrical conformation, with regular or subregular, polygonal (mostly hexagonal) meshes, and generally the stronger threads of the sarcodictyum secrete a firm, homogeneous or fibrillar, striated substance, which forms a network of ridges on the surface of the calymma. In the Acanthophracta the place of this is taken by the acanthin network of the primary lattice-shell. The axopodia of the Acanthometra are usually about as long as the radial spines between which they stand; their stiff axial thread is surrounded by a soft sheath of protoplasm, communicating with the thin sarcomatrix which surrounds the central capsule. Numerous branches pass into the calymma from the exoplasmic sheath of the axial threads, and form by their interweaving a loose sarcoplegma. The most peculiar differentiated products of the exoplasm of the Acantharia, however, are the myophane fibrillæ of the Acanthometra, which have already been described under the name of myophriscs (§ 96).

99. The Exoplasm of the Monopylea.—The extracapsular protoplasm of the Nassellaria or Monopylea arises only from the porochora, or the intracapsular podoconus, the oral base of which is formed by this porous area. The pseudopodia or protoplasmic threads which pass through the pores of the latter, united into a bundle, are not very numerous (in most Nassellaria probably between thirty and ninety), and unite just outside it to form a thick discoid sarcomatrix; this covers the porochora completely below, and spreads out in the form of a thin envelope of exoplasm over the whole {lxvii}surface of the central capsule; at the apical portion of the latter the sarcomatrix is often so thin that it can only be recognised by the aid of reagents; it separates the membrane of the central capsule from the surrounding calymma. The pseudopodia, which penetrate the latter and by loose anastomoses from a wide-meshed sarcoplegma within it, are usually not very numerous. The greater part of them radiate in a bunch downwards from the basal disc of the sarcomatrix, and a smaller number arise from the thinner envelope which covers the remainder of the central capsule (Pl. 51, fig. 13; Pl. 65, fig. 1; Pl. 81, fig. 16). On the outer surface of the calymma the collopodia, which have passed through it, unite to form the sarcodictyum, and through the silicification of this the primary lattice-shell arises in the great majority of the Nassellaria. From the surface of the sarcodictyum arise the astropodia, or free pseudopodia which radiate outwards into the water. Their number in most Monopylea is relatively small, but their length appears to be very great.

100. The Exoplasm of the Cannopylea.—The extracapsular protoplasm of the Phæodaria or Cannopylea is much better developed as regards volume than in the other three legions, and is connected with the intracapsular sarcode by only a few apertures in the capsule-membrane. In most Phæodaria three of these are present, the astropyle or main-opening at the oral pole of the main axis, and the two lateral parapylæ or accessory openings on either side of the aboral pole (§ 60). In several families the latter appear to be wanting, whilst in others their number is increased; these families have not yet, however, been observed during life. The protoplasm projects both from the oral main-opening and from the two aboral accessory openings in the form of a thick cylindrical rod; the tube into which each opening is produced in many Phæodaria (longer in the case of the astropyle, shorter in the parapylæ) being regarded as an excretion from this protoplasmic cylinder. The sarcode threads within the tube appear like a bundle of fibrils, either quite hyaline or finely striated. After issuing from the mouth of the aperture they pass over into a thick sarcomatrix, which surrounds the central capsule entirely and separates it from the enclosing calymma. In the neighbourhood of the basal astropyle the sarcomatrix is usually swollen into a thick lenticular disc, which is in direct contact with the peculiar phæodium of this legion (§ 89). The pseudopodia, which radiate from the sarcomatrix, and form by anastomosis a wide-meshed sarcoplegma within the calymma, are usually not very numerous in the Phæodaria, but are very strong. Sometimes two stronger bundles of collopodia may be distinguished at the two poles of the main axis, an oral bundle (in the direction of the proboscis of the astropyle) and an aboral bundle (at the opposite pole between the parapylæ). The collopodia of the sarcoplegma unite at the surface of the calymma into a regular or irregular sarcodictyum, which, in most Phæodaria produces by the secretion of a peculiar silicate the primary lattice-shell. {lxviii}The free astropodia, which pass outwards from the sarcodictyum into the water, are in most Phæodaria very numerous (Pl. 101, fig. 10). Since, however, only a few species of this great legion have been observed in a living state, their pseudopodia require further accurate examination.

Chapter IV.—THE SKELETON.

(§§ 101-140).

101. The Significance of the Skeleton.—The skeleton of the Radiolaria is developed in such exceedingly manifold and various shapes, and exhibits at the same time such wonderful regularity and delicacy in its adjustments, that in both these respects the present group of Protista excels all other classes of the organic world. For, in spite of the fact that the Radiolarian organism always remains merely a single cell, it shows the potentiality of the highest complexity to which the process of skeleton formation can be brought by a single cell. All that has been brought to pass in this direction by single tissue-cells of animals and plants does not attain the extremely high stage of development of the Radiolaria. Only very few Rhizopoda of this very rich and varied class fail to exhibit the power of forming this firm supporting and protecting organ—indeed, only ten of the seven hundred and thirty-nine genera which are enrolled in the list of the Challenger collection, namely, six genera of Spumellaria (five Thalassicollida, Actissa, Thalassolampe, Thalassopila, Thalassicolla, Thalassophysa, Pl. 1, and one genus of Collozoida, Collozoum, Pl. 3), and in addition two genera of Nassellaria (the Nassellida, Cystidium and Nassella, Pl. 91, fig. 1), and two genera of Phæodaria (the Phæodinida, Phæocolla and Phæodina, Pl. 101, figs. 1, 2). These skeletonless forms of Radiolaria are, however, of extreme interest, since they include the original stem-forms of the whole class as well as of its four legions. All Radiolaria which form skeletons have originated from soft and skeletonless stem-forms by adaptation, and that polyphyletically, for the skeletal types of the four legions have been developed independently of each other (§ 108).

102. The Chemical Peculiarities of the Skeleton.—The chemical composition of the skeleton shows very marked variations in the different legions of the Radiolaria. The two legions Spumellaria and Nassellaria (united formerly as "Polycystina") form their skeleton of pure silica (see note A, below); the legion Phæodaria of a silicate of carbon (see note B), and the Acantharia of a peculiar organic substance—acanthin (see note C). This explains the well-known fact that the deposits of fossil Radiolaria (or Polycystine marls) are composed exclusively of the skeletons of Spumellaria and Nassellaria, those of the Acantharia and Phæodaria being entirely absent (in the case of the last group, however, exception must be made in favour of the Dictyochida, or those Phæodaria {lxix}whose skeleton is made up of isolated scattered tangential siliceous fragments). The enormous deposits of Radiolarian skeletons in the deep sea of today, which constitute the Radiolarian ooze, consist, like the fossil Polycystine marls, almost exclusively of the shells of Spumellaria and Nassellaria, though here the acanthin skeletons of the Acantharia may be present in very small numbers, and the silicate skeletons of the Phæodaria, which offer more resistance to the solvent action of sea-water, somewhat more abundantly. Calcareous skeletons do not occur in the Radiolaria (see note D).

103. The Physical Properties of the Skeleton.—The skeletons of all Radiolaria are characterised pre-eminently by a high degree of firmness, which fits them to serve as protective and supporting apparatus. This is obvious in the case of the pure siliceous shells of the Polycystina; but the acanthin framework of the Acantharia also possesses a degree of stiffness but little inferior, whilst the silicate skeletons of the Phæodaria seem on the whole to be not so firm. The hollow skeletal tubes of the last-named, which are filled with gelatinous material, are very brittle on account of the delicacy of their walls. Their elasticity also is very small, whilst that of the acanthin spines is considerable. The thin long needles of many Acantharia are very elastic, as are also the bristle-like siliceous spicules of many Spumellaria. The refractive power of the skeleton in the various legions is very different, depending upon the chemical constitution. The siliceous skeleton of the Polycystina (Spumellaria and Nassellaria) and the silicate skeleton of the Phæodaria have the same refractive index as glycerine, and hence become invisible when mounted in that fluid; they then become visible only on addition of {lxxi}water, and are clearer in proportion to the quantity of water which is added. The refractive index of acanthin is, however, very different from that of glycerine, so that the skeletons of Acantharia are readily visible when mounted in this fluid. In water, the skeletons of all Radiolaria appear about equally refractive, as also in Canada balsam. The substance of the skeleton appears almost entirely hyaline, colourless, and transparent. Very rarely it is faintly coloured (in some Acantharia). A cloudy opaque constitution is seen in some Phæodaria (especially in the "porcellanous shells" of Tuscarorida and Circoporida, Pls. 100, 114-117); when dried, these appear by reflected light milky-white or yellowish-white; the cause of this opacity lies partly in the peculiar "cement-like structure" of these porcellanous shells, partly in their fine porosity, and the minute air-bubbles contained in their thick walls.

104. The Elementary Structure of the Skeleton.—The general constitution of the skeleton—or more accurately expressed, of the morphological elements of which the skeleton consists—is of such a nature that it may be termed structureless. Both the organic acanthin skeletons of the Acantharia and the silicate skeletons of the Phæodaria, as well as the inorganic siliceous skeletons of the Spumellaria and Nassellaria, appear under the microscope perfectly homogeneous, transparent, colourless, and crystalline. Only very rarely do they show traces of a concentric striation, which arises from the deposition of the skeletal substance in layers; as, for example, the thick spines of some Phæodaria (Pls. 105-107, &c.). Some of the Phæodaria, however, form an exception to this rule, inasmuch as their partially tubular skeletal elements possess a remarkable porcellanous structure. In the tubular or Cannoid skeleton, which occurs in most Cannopylea, the lumen of the thin-walled flinty tube is filled with jelly, and frequently a thin siliceous thread runs in its axis, and is connected with the wall by transverse threads (§§ 127, 139). The elementary structure of the opaque porcellanous shells, which distinguish the two families Circoporida (Pls. 114-117) and Tuscarorida (Pl. 100), is quite peculiar. Numerous fine siliceous spicules lie scattered irregularly in a finely granular or porous matrix.

105. Complete and Incomplete Lattice-Shells.—In the great majority of Radiolaria (in all four legions) the skeleton has the form of a delicate lattice-shell or a receptacle in which the central capsule is enclosed. In a small minority, however, this is not the case. The skeleton then consists only of isolated rigid pieces (radial or tangential spicules), or of a simple ring (sagittal ring of the Stephoidea), or of a basal tripod with or without a loose tissue of trabeculæ, &c. (Plectoidea); the central capsule is then not surrounded by a special latticed receptacle, but only rests upon the skeletal trabeculæ. According to these different arrangements, two principal groups or sublegions may be distinguished in each legion, of which one set (Cataphracta) are characterised by a complete {lxxii}lattice-shell, whilst the others (Aphracta) are without it. The Radiolaria aphracta, then, or Radiolaria without a complete skeleton, are the Collodaria (p. 9), the Acanthometra (p. 725), the Plectellaria (p. 895), and the Phæocystina (p. 1543). On the other hand, the Radiolaria cataphracta, or Radiolaria with a complete skeleton, are the Sphærellaria (p. 49), the Acanthophracta (p. 791), the Cyrtellaria (p. 1015), and the Phæocoscina (p. 1590).

106. The Ectolithia and Entolithia (Extracapsular and Intracapsular Skeletons).—The relation of the skeleton to the central capsule in the Radiolaria is very various in many respects; in the first instance two great groups, Ectolithia and Entolithia (see note A), may be distinguished topographically by mere external observation; in the former the skeleton lies entirely outside the central capsule; in the latter, partially at all events, within it. The Ectolithia, with a completely extracapsular skeleton, include all Nassellaria and Phæodaria, as well as a great part of the Spumellaria (all Collodaria and the most archaic forms of Sphærellaria); the Entolithia, on the other hand, in which the skeleton lies partly within, partly without the central capsule, include all Acantharia and the majority of the Spumellaria (most Sphærellaria, see note B).

107. Perigenous and Centrogenous Skeletons.—Much more important than the topographical relation of the skeleton to the central capsule, according to which the Ectolithia and Entolithia are separated from each other (§ 106), is the original development of the skeleton within or without the central capsule, which gives rise to the distinction between perigenous and centrogenous skeletons. Centrogenous skeletons are found only in the Acantharia, which are further distinguished from all other Radiolaria by their skeleton being formed of acanthin; in all Acantharia the formation of the skeleton begins in the middle of the central capsule, from which twenty (the number is inconstant only in the {lxxiii}small group Actinelida) radial spines are centrifugally developed. The three other legions, on the contrary, possess on the whole a perigenous skeleton, which originally develops outside the central capsule and never in its middle. In the Nassellaria and Phæodaria the skeleton retains this extracapsular position, as also in the Beloidea and part of the Sphærellaria among the Spumellaria; in the great majority of the latter, however, the primary perigenous skeleton is subsequently enveloped by the growing central capsule, so that it lies partially within it (§ 109).

108. Polyphyletic Origin of the Skeleton.—The skeleton of the Radiolaria has undoubtedly originated polyphyletically, for it is impossible to derive its manifold varieties from a single ground-form, or to regard them as modifications of one type. It is much more probable that the different skeletonless Radiolaria have entered upon different ways of skeleton formation quite independently of each other. At the outset it is quite clear that the skeletons of the four legions have originated independently of each other. Further, it is certain that within the legion of the Spumellaria the Beloid skeletons of the Collodaria are not connected with the Sphæroid skeletons of the Sphærellaria and the forms derived from them (see § 109). In the same way the skeletons of the Phæodaria are polyphyletic; probably in this legion the Beloid, Sphæroid, Cyrtoid, and Conchoid skeletons have been developed quite independently (see § 112). In the Nassellaria, on the other hand, it is possible that all the skeletal forms are to be derived monophyletically from a single simple primitive form (either the sagittal ring or basal tripod?) (see § 111). Still more probable is it that the Acantharia have arisen monophyletically, for all the forms of their acanthin skeleton may be derived without violence from Actinelius (see § 110).

109. The Skeleton of the Spumellaria.—The skeletons of the Spumellaria or Peripylea consist of silica, and are very different and of independent origin in the two orders of this legion. The first order, Collodaria, have either no skeleton whatever (Colloidea, p. 10, Pls. 1, 3), or their skeleton is Beloid, a loose extracapsular envelope of spicules, consisting of numerous unconnected portions; the separate parts are usually disposed tangentially, either as simple or compound siliceous spicules (Beloidea, p. 28, Pls. 2, 4). The second order of Spumellaria, on the other hand (Sphærellaria, p. 49), develops a siliceous lattice-shell which consists of a single piece, and is remarkable for the extraordinary variety of its forms (pp. 50-715, Pls. 5-50). To this order belong not less than three hundred genera and seventeen hundred species of the Challenger Radiolaria (that is, about two-fifths of all the genera and species). In spite of this extreme richness in different forms this large group must be regarded as monophyletic, since all its forms may be quite naturally derived from a common stem-form, a simple lattice-sphere (Cenosphæra, p. 61, Pl. 2). The twenty-eight families of Sphærellaria may be distributed in four suborders, among which the Sphæroidea constitute the {lxxiv}stem-forms, since they retain the original spherical shape (Pls. 5-8, 11-30). In the other three suborders a vertical main axis is developed, which in Prunoidea is longer, in Discoidea shorter than the other axes of the shell. Hence the shell of the Prunoidea (p. 284, Pls. 13, bis, 17, 39, 40) is ellipsoidal or cylindrical, that of the Discoidea, on the other hand, lenticular or discoidal (p. 402, Pls. 31-38, 41-48). Finally, the shell of the fourth suborder, Larcoidea, is lentelliptical; it has the ground-form of a triaxial ellipsoid, and is characterised by the possession of three unequal dimensive axes, or three isopolar axes of different lengths perpendicular to each other (p. 599, Pls. 9, 10, 49, 50).

110. The Skeleton of the Acantharia.—The skeletons of the Acantharia or Actipylea are distinguished from those of all other Radiolaria by two very important peculiarities; in the first place, they consist not of silica but of a peculiar organic substance, Acanthin, and secondly, their development is centrogenous, numerous radial spines or acanthin spicules being formed which are united in the middle of the central capsule. Hence the Acantharia are the only Radiolaria in which the skeleton originates from the first in the middle of the central capsule. The number of radial spines is primitively indefinite, variable, and often considerable (more than a hundred), but in the great majority it is limited to twenty. In accordance with this the legion may be divided into two orders, the more archaic small group Adelacantha, with an indefinite number of spines, and the more recent group, Icosacantha, which has been developed from them and possesses twenty regularly disposed spines; of the three hundred and seventy-two species of Acantharia which have been hitherto described, about five per cent. belong to the former, about ninety-five per cent. to the latter division (see note A, below). The numerous genera of Icosacantha may then be again divided into two suborders, of which the Acanthonida (p. 740, Pls. 130-132) produce no complete lattice-shell, and thus agree with the Actinelida, with which they may be united as Acanthometra in the broader sense (or Acantharia without a lattice-shell). The Acanthophracta, on the other hand (p. 791, Pls. 133-140), produce a complete lattice-shell, usually by means of two opposite or four crossed transverse processes, which arise from each radial spine and unite with each other (see note B, below). In most Acanthophracta the lattice-shell remains single; only in the Phractopeltida does it consist of two concentric lattice-spheres (p. 847, Pl. 133, figs. 1-6). Furthermore, the whole order Acanthophracta may be subdivided into two suborders according to the different ground-form of the lattice-shell; this remains spherical in the Sphærophracta (the three families Sphærocapsida, Dorataspida, Phractopeltida, Pls. 133-138). On the other hand, it assumes another form in the Prunophracta; it becomes ellipsoidal in the Belonaspida (Pl. 136, figs. 6-9), discoidal or lentiform in the Hexalaspida (Pl. 139); and finally takes the shape of a double cone in the Diploconida (Pl. 140).

111. The Skeleton of the Nassellaria.—The skeletons of the Nassellaria or Monopylea consist of silica, and are never composed of separate portions, but constitute always a single continuous piece. The ground-form is originally monaxon, corresponding to that of the central capsule, with a constant difference between the two poles of the vertical main axis. The ground-form is never spherical or polyaxon as in the lattice-shells of the Spumellaria, and the skeleton never consists of hollow tubes, as in the Phæodaria. The legion Nassellaria may be divided into two orders; in the Plectellaria (three suborders Nassoidea, Plectoidea, Stephoidea) the skeleton does not form a complete lattice-shell; in the Cyrtellaria, on the other hand, which are derived from these, the siliceous skeleton forms a complete lattice-shell enclosing the central capsule. The number of forms thus developed is astonishingly great, so that among the Nassellaria no less than two hundred and seventy-four genera and sixteen hundred and eighty-seven species may be distinguished, almost as many as in the Sphærellaria. In spite of this great variety of forms the legion Monopylea is probably monophyletic; at least all the different skeletal forms may be derived from three elements which are combined in the most manifold fashion; (1) the sagittal ring, a simple siliceous ring, which lies vertically in the sagittal plane of the body, encircles the central capsule and comes into contact with it at the basal pole of the main axis (§ 124); (2) the basal or oral tripod, composed of three diverging radial spines, which meet in the middle of the basal pole of the central capsule (or in the centre of the porochora) (§ 125); (3) the cephalis, or lattice-head, a simple ovoid or subspherical lattice-shell, which encloses the central capsule and stands in connection with it at the basal pole of its main axis. Any one of these three important structural elements of the Nassellarian skeleton may possibly be the starting-point {lxxvi}for all the remaining forms of the Monopylea; the great difficulty in their phylogenetic derivation lies in the facts that, on the one hand, any one of the three elements may alone constitute the skeleton, and on the other hand, in the great majority of the legion, two or three are united together (compare §§ 182-185).

112. The Skeleton of the Phæodaria.—The skeleton of the Phæodaria or Cannopylea is always extracapsular, usually consists of a silicate of carbon (more rarely of pure silica), and in the majority of the legion is composed of hollow cylindrical tubes, whose siliceous wall is very thin, and whose lumen is filled with gelatinous material (§ 127). The manifold and remarkable skeletal forms occurring in this legion are not monophyletic, since they cannot be derived from a common stem-form; they are, on the contrary, polyphyletic, various skeletonless Phæodaria (Phæodinida) have independently acquired skeletons of different form and composition. The legion Phæodaria can be subdivided into four orders, the skeletons of which present the following important distinctions:—(1) The Phæocystina possess only incomplete Beloid skeletons (§ 115), composed of many separate pieces, sometimes tangentially (Cannorrhaphida, Pl. 101), sometimes radially arranged (Aulacanthida, Pls. 102-105). (2) The Phæosphæria form Sphæroid skeletons (§ 116), usually only a simple lattice-shell without special aperture (Pls. 106-111); two concentric shells united by radial bars occur only in the Cannosphærida (Pl. 112). (3) The Phæogromia are distinguished by the formation of a simple Cyrtoid skeleton (§ 123) resembling that of the Monocyrtida; the monothalamus lattice-shell is usually ovoid or helmet-shaped, more rarely polyhedral or almost spherical; a vertical main axis can always be distinguished, at the basal pole of which is an aperture usually armed with teeth or spines (Pls. 99, 100, 113-120). (4) The Phæoconchia are distinguished from all other Radiolaria by the possession of a bivalved shell like that of the Conchifera; the two valves of this Conchoid skeleton must be distinguished as dorsal and ventral, as in the Brachiopoda (Pls. 121-128). The fifteen families of Phæodaria which are arranged in the four orders just mentioned, present such great differences among themselves, that the skeleton must be regarded as probably polyphyletic even within the limits of each order.

113. Types of Skeletal Formation.—No less than twelve different principal forms may be distinguished as morphological types of the formation of the skeleton in the Radiolaria; some of these are peculiar to a single legion or even to a smaller group; but sometimes the same form occurs in several legions. Some types occur only in an isolated manner, independently of the others, but most exist in various combinations with other types. Of the twelve described below the Conchoid and Cannoid occur only in the Phæodaria; the Plectoid and Circoid only in the Nassellaria; the Astroid only in the Acantharia; the remaining seven types are found in several legions in the same form and hence are polyphyletic.

114. The Astroid Skeleton.—Under the name "Astroid" we place the peculiar star-shaped skeletons of the Acantharia in opposition to those of all other Radiolaria, for they are separated from them not only fundamentally by reason of the chemical nature of their substance (Acanthin, § 102), but also by their centrogenous origin, and the resulting stellate form (Pls. 129-140). The Acantharia are the only Radiolaria in which the skeleton arises within the central capsule by the formation of numerous rays or radial spines of acanthin which project on all sides from the centre. Originally these are united at this point, their conical or pyramidal points meeting and being supported one upon another. In the great majority of Acantharia this loose apposition is constant, so that when the soft parts are destroyed the skeleton falls to pieces. Only in a few forms in this legion are the central ends of the spines fused so that the whole skeleton forms a connected star (Astrolithium). The small group Chiastolida (or Acanthochiasmida) is characterised by the fact that the two rays which are opposite to one another in each axis unite and form a diametral bar. The skeleton is almost always composed of twenty radial spines, which are regularly disposed (Icosacantha), only in the small primitive group Actinelida is the number variable (Adelacantha, § 110).

115. The Beloid Skeleton.—As Beloid or spicular skeletons are grouped together all those which consist of several disconnected portions; these always lie outside the central capsule, either within the calymma or on its surface. Such extracapsular Beloid skeletons are entirely wanting in the Acantharia and Nassellaria; they occur only in the Beloidea among the Spumellaria, and in the Phæocystina among the Phæodaria; the individual Beloid portions of the former are solid, those of the latter hollow. In both groups the simplest forms of the separate portions are simple unbranched needles (Thalassosphæra, Thalassoplancta, Physematium, Belonozoum, among the Spumellaria; Cannobelos and Cannorrhaphis among the Phæodaria); usually these spicules are disposed tangentially over the surface of the calymma. Among the Beloidea branched spicules occur more commonly than these simple ones; they are either stellate (with many rays united in a centre) or twin-like, with a tangential bar, from each pole of which two or three (seldom more) radial branches project (Pls. 2, 4). Among the Phæodaria the subfamily Dictyochida is characterised by the annular shape of its Beloid portions, either simple rings, or hat-shaped or pyramidal bodies with a latticed cap over the ring (Pl. 101, figs. 3-14; Pl. 114, figs. 7-13). The family Aulacanthida among the Phæodaria, alone possesses hollow radial tubes, which penetrate the whole calymma, and project distally over its surface, whilst their proximal ends rest upon the surface of the central capsule. Although in these cases the enclosed proximal end is always simple, the free distal end develops the most various processes in adaptation to its prehensile functions (Pls. 102-105).

116. The Sphæroid Skeletons or Lattice-Spheres.—The "lattice-spheres" or sphæroid skeletons are the simplest and most primitive forms of lattice shells, and are widely distributed in the three legions Spumellaria, Acantharia, and Phæodaria, whilst they are entirely wanting in the Nassellaria. The round lattice-shell is either a true sphere in the geometrical sense, or an endospherical polyhedron, i.e., a polyhedron, all whose angles lie in the surface of a sphere (§ 25). In general, primary and secondary lattice-spheres may be distinguished, of which the former are secreted on the outer surface of the primary, the latter on that of the secondary calymma (§ 85). Furthermore, simple and compound lattice-spheres may be distinguished, the latter of which consist of two or more concentric lattice-spheres firmly united by radial bars; in such cases the innermost lattice-sphere is always to be regarded as the oldest or primary, all the succeeding ones as secondary, and the outermost as the youngest (§ 129). The simple lattice-spheres are usually to be regarded as primary; they may, however, occasionally be secondary, in which case the primary shell, originally enclosed, has been lost by degeneration (as, for example, in the case of the Aulosphærida and some Sphærellaria).

117. The Lattice-Spheres of the Spumellaria.—The lattice-spheres or Sphæroid skeletons of the Spumellaria exhibit in spite of their simple type of structure, an extraordinary variety in the formation of the lattice-work and radial apophyses, so that in the systematic portion of this work no less than one hundred and seven genera and six hundred and fifty species are distinguished; these are united in one suborder, the Sphæroidea (pp. 50-284, Pls. 5-8, 11-30). It may be divided into two main divisions, the Monosphærida with a single primary lattice-sphere (Pls. 12-14, 21, 26, 27), and Pliosphærida (or Sphæroidea concentrica) whose skeleton consists of two or more concentric lattice-spheres united by radial bars. The latter are subdivided into Dyosphærida with two concentric lattice-spheres (Pls. 16, 19, 20, 22, 28); Triosphærida, with three spheres (Pls. 17, 24, 29); Tetrasphærida, with four (Pls. 23, 30); Polysphærida, with five or more (Pls. 15, 23); and Spongosphærida, with spongy lattice-spheres (Pls. 18, 25). A special group is made up of the simple lattice-spheres of the social Collosphærida (or Sphæroidea polyzoa) (Pls. 5-8); these are usually more or less irregular, and characterised by the development of peculiar tubular processes; the latter are generally wanting in the Sphæroidea monozoa, whose lattice-shell is very regularly formed. This distinction is interesting and important, inasmuch as the regular lattice-spheres are explained by the independent development of the free-swimming Monozoa, whilst the irregular spheres are due to the mutual dependence of the social Polyzoa.

118. The Lattice-Spheres of the Acantharia.—The lattice-shells or Sphæroid skeletons of the Acantharia are immediately distinguishable from those of all other Radiolaria by their centrogenous development and the central union of the radial spines by which they are supported; the only exception is furnished by the remarkable genus Cenocapsa {lxxix}(Pl. 133, fig. 11), in which the radial spines are absent, not primitively, however, but in consequence of degeneration; for the twenty cross-shaped perspinal pores, originally due to the twenty radial spines, are still present. In the most nearly allied genera, Porocapsa (Pl. 133, fig. 7) and Cannocapsa (Pl. 133, fig. 8), the proximal part of the twenty radial spines is still present, while their distal portion has degenerated; hence in this case they do not stand in direct communication with the spherical shell. On the other hand, this primitive connection persists in the genera Astrocapsa (Pl. 133, figs. 9, 10), and Sphærocapsa (Pl. 135, figs. 6-10). The five genera just mentioned form the peculiar family Sphærocapsida (pp. 795-802); the spherical shell is in these cases composed of very numerous small plates disposed like a pavement, each plate or aglet being perforated by a pore canal; in addition to which there are twenty larger (perspinal) pores (or twenty cross-shaped groups each of four aspinal pores) at those important points where primitively the twenty radial spines penetrate the calymma. This peculiar porous "pavement shell" has probably been developed (independently of the twenty radial spines) upon the calymma of the Acanthonida (Acanthonia, p. 749) by the action of the sarcodictyum; it has, therefore, quite a different morphological significance from the spherical lattice-shell of the Dorataspida, which is composed of tangential apophyses of the twenty Acanthonid spines (pp. 802-847, Pls. 134-138). Each radial spine here forms either two opposite or four crossed transverse processes, and since their branches spread over the surface of the spherical calymma and are united suturally at their extremities, the peculiar lattice-sphere of the Dorataspida arises. This extensive family is again divided into two subfamilies:—the Diporaspida (Pls. 137, 138) possess always only two opposite apophyses, and form by the union of their branches two opposite primary apertures or aspinal meshes. The Tessaraspida, on the other hand (Pls. 135, 138), have always four crossed transverse processes, and form by their union four primary aspinal meshes. From the Diporaspida are probably to be derived the Phractopeltida (p. 847, Pl. 133, figs. 1-6), the only Acantharia which possess a double lattice-sphere; their double concentric spherical shell may be compared with that of the Dyosphærida.

119. The Lattice-Spheres of the Phæodaria.—The lattice-spheres or Sphæroid skeletons of the Phæodaria, which are generally developed quite regularly, though occasionally in a modified form, fall in the order Phæosphæria into two groups of very different structure, each of which includes two families. The first group (Phæosphæria inarticulata) contains the families Orosphærida (Pls. 106, 107) and Sagosphærida (Pl. 108); the lattice-work of the former consists of irregular polygonal meshes and very coarse, partially hollow trabeculæ; in the latter, on the other hand, it consists of triangular meshes and very slender filiform trabeculæ; in both families the whole sphæroid skeleton forms a single unsegmented piece as in most Sphæroidea. In the second group of {lxxx}Phæosphæria (Phæosphæria articulata), on the other hand, the lattice-sphere is segmented in quite a peculiar manner, and composed of hollow cylindrical tangential tubes, which are separated by astral septa at the nodal points of the network; this remarkable structure characterises the two families, Aulosphærida (Pls. 109-111) and Cannosphærida (Pl. 112); the segmented lattice-sphere of the former is simple and hollow; while that of the latter is connected by centripetal radial tubes with a simple concentric inner shell, which is sometimes solid, sometimes latticed, and provided with a main-opening corresponding to the astropyle of the enclosed central capsule. Since in the Aulosphærida also, hollow centripetal radial tubes project from the segmented lattice-sphere, it is possible that they have been derived from the Cannosphærida by the loss of the primitive internal shell. A special peculiarity of many Phæosphæria (Oroscena, Sagoscena, Auloscena, &c.) consists in the fact that the whole surface of the lattice-sphere is regularly covered with pyramidal or tent-shaped prominences (Pl. 106, fig. 4; Pl. 108, fig. 1; Pl. 110, fig. 1). A simple lattice-sphere quite similar to that of most Monosphærida also constitutes the skeleton of the Castanellida (Pl. 113), but since it possesses a special main-opening, it must be referred promorphologically to the Cyrtoid shells of the Phæogromia.

120. The Prunoid Skeleton or Lattice-Ellipsoid.—The "lattice-ellipsoids" or Prunoid skeletons have arisen from the lattice-spheres or Sphæroid skeletons by more energetic growth and elongation of one axis; this is the main axis of the body and is probably always vertical; its two poles are commonly equal. The Prunoid skeleton is either a true ellipsoid in the geometrical sense or an "endellipsoidal polyhedron" (i.e., a polyhedron, all the angles of which lie in an ellipsoidal surface). By further elongation of the main axis, the ellipsoidal form passes over into the cylindrical, the polar surfaces of the cylinder being usually rounded, rarely truncated. The rich order Prunoidea (pp. 284-402) contains numerous modifications of this form of shell which arise on the one hand by the formation of transverse constrictions, on the other by the apposition of concentric secondary shells. In respect of the latter, simple and compound Prunoid shells can be distinguished as in the case of the Sphæroid shells. In the compound Prunoid shells either all the concentric lattice-shells may be ellipsoidal or the inner may be spherical. More important differences are found in the transverse annular constrictions, which give the Prunoid skeleton a segmented appearance; in this respect, three principal forms may be distinguished (p. 288):—(A) Monoprunida, with unsegmented shell, having no transverse constriction (Pls. 15-17); (B) Dyoprunida, having a shell with two segments and one (equatorial) transverse constriction (Pl. 39); (C) Polyprunida, with three or more parallel transverse constrictions, by means of which the shell is divided into four or more segments (Pl. 40). In the same manner as the Prunoidea have arisen from the Sphæroidea among the Spumellaria by greater {lxxxi}development of the vertical main axis, the ellipsoidal Belonaspida have arisen from the spherical Dorataspida among the Acantharia (p. 859; Pl. 136, figs. 6-9; Pl. 139, figs. 8, 9). The main axis of the ellipsoid in this case is always occupied by the opposite equatorial spines of the hydrotomical axis (pp. 719, 860). In the legion Phæodaria a similar prolongation of the main axis rarely occurs; it is found, however, in Aulatractus (Pl. 111, figs. 6, 7), the lattice-shell of this Aulosphærid being sometimes truly fusiform, sometimes rather ellipsoidal or even double-conical.

121. The Discoid Skeletons or Lattice-Discs.—The "lattice-discs" or Discoid skeletons are characteristic of the Spumellarian group Discoidea, and have arisen from the lattice-spheres of the Sphæroidea by a less development of one axis, which is the main axis of the body, and is probably usually vertical; its two poles are always equal. The Discoid lattice-shell is either a biconvex lens (with a thin margin), or a plane disc (a shortened cylinder with thick margin), or some form intermediate between the two. All Discoid shells show a horizontal median plane or equatorial plane, by which they are divided into two equal halves, an upper and lower; the margin of the lens itself is originally the equator. The main axis, the shortest of all the axes of the shell, stands vertically in the centre of the equatorial plane. Among the Phæodaria Discoid shells rarely occur (Aulophacus), as also among the Acantharia (Hexalaspida).

122. The Larcoid Skeleton or Lentelliptical Lattice-Shell.—The lentelliptical lattice-shells, which may be shortly designated "Larcoid," are especially characteristic of the Larcoidea, a large order of Spumellaria (pp. 599-715; Pls. 9, 10, 49, 50). In addition they recur among the Acantharia, in the small family Hexalaspida (p. 872, Pl. 139), and the family Diploconida (p. 881, Pl. 140), which is derived from it. These lentelliptical lattice-shells are all characterised by the clear differentiation of three unequal, but isopolar dimensive axes, i.e., the three geometrical axes, perpendicular to one another, which determine the form of the shell, are of unequal length; the two poles of each are, however, equal. The geometrical ground-form is, therefore, a triaxial ellipsoid (§ 34). In the rich order Larcoidea the lentelliptical lattice-shell shows many variations in its development.

123. The Cyrtoid Skeleton.—Cyrtoid skeletons are those lattice-shells which possess a vertical main axis with two different poles (Monaxonia allopola); the upper pole is usually termed the apical, the lower the basal. Such Cyrtoid shells are characteristic of the great majority of the Nassellaria or Monopylea (and especially of the Cyrtellaria); they are also found in a large division of the Phæodaria (the Phæogromia), and in some Spumellaria. In general the manifold Cyrtoid shells may be divided into two large groups, those with one and those with several chambers. The monothalamous Cyrtoid shells are usually ovoid, conical, cap- or helmet-shaped; their {lxxxii}internal cavity is simple, without constrictions or septa. Among the Nassellaria they occur in the Monocyrtida (Pls. 51-54, 98), where they have received the name "Cephalis." A form of shell, essentially the same, is found amongst the Phæodaria in the order Phæogromia, more especially in the Challengerida (Pl. 99), Medusettida (Pls. 118-120), and Tuscarorida (Pl. 100), many of these latter closely resembling many Monocyrtida. Such monothalamous Cyrtoid shells occur much more rarely among the Spumellaria (e.g., among the Prunoidea in Lithapium, Lithomespilus, Druppatractus, Pls. 13, 14, &c.). Polythalamous Cyrtoid shells (Pls. 55-80) occur exclusively in the Nassellaria, and exhibit in this legion an astonishing variety of structure; they are distinguished from the monothalamous forms by the development of internal septa, or of annular incomplete diaphragms, which usually correspond to the external constrictions; their interior is thus divided into two or more communicating compartments. Among the polythalamous Cyrtoid shells may be distinguished three principal groups, the Stichocyrtid, Zygocyrtid, and Polycyrtid. Zygocyrtid shells are characteristic of the Spyroidea (Pls. 84-90), and are distinguished by a bilobate cephalis (cephalis bilocularis); the median sagittal ring, or a corresponding constriction, divides the shell into right and left compartments. Polycyrtid shells (Pl. 96) are peculiar to the Botryodea, and characterised by a multilobate cephalis (cephalis multilocularis). Stichocyrtid shells are those in which the primary cephalis remains simple, and new joints are successively added to its basal pole; such shells occur in the majority of the Cyrtoidea. Secondary chambers are sometimes added in the other two groups (Botryodea and Spyroidea). When, as often happens in these polythalamous Cyrtoid shells, two or three distinct joints follow each other, the first is called the "cephalis," the second the "thorax," and the third the "abdomen" (Tricyrtida Pls. 64-75).

124. The Circoid Skeleton.—This is a very important and remarkable type of skeletal formation, which occurs exclusively in the legion Nassellaria, where it plays a very prominent part; its characteristic element is the "sagittal ring," a simple, vertical, siliceous ring, which surrounds the central capsule in its sagittal plane, and is specially differentiated in its basal portion. This "primary sagittal ring" whose vertical allopolar main axis coincides with that of the Monopylean central capsule embraced by it, is characteristic of all members of the order Stephoidea (p. 931, Pls. 81-83, 92-94); here it forms by itself the skeleton of the Stephanida (Pl. 81); in the Semantida (Pl. 92) it is combined with a horizontal basal ring, in the Coronida (Pls. 82, 93) with a vertical frontal ring and in the Tympanida (Pls. 83, 94) with two horizontal rings, an upper mitral and a lower basal. In the great majority of these Stephoidea there often develop in definite places characteristic processes or apophyses, whose branches combine to form a loose tissue or an incomplete lattice-shell. This becomes complete in the Cyrtellaria, the majority of which retain more or less {lxxxiii}distinct traces of the sagittal ring. Hence the skeletons of all Nassellaria may be derived monophyletically (Hypothesis A, p. 893) from a simple sagittal ring (Archicircus and Lithocircus, Pl. 81). This theory, however, encounters the great difficulty that in many Stephoidea (Cortina, Cortiniscus, &c.) it is combined in a remarkable manner with the basal tripod of the Plectoidea, whilst in these latter it is entirely wanting (compare p. 894).

125. The Plectoid Skeleton.—Those forms are distinguished as Plectoid in which three, four, or more radial siliceous spines proceed from a common point, which lies excentrically outside the central capsule and at the basal pole of its vertical allopolar main axis. This peculiar type of skeletal formation only occurs in the legion Nassellaria, and is specially characteristic of the order Plectoidea (p. 898, Pl. 91). But since the essential elements of this remarkable skeleton also occur in many other Nassellaria, sometimes combined with the Circoid, sometimes with the Cyrtoid skeleton, it perhaps has a fundamental significance in this legion; at all events it is possible to derive monophyletically all the other forms of this legion from it (Hypothesis B, p. 893). The simplest form of the Plectoid skeleton is a tripod, the three feet of which either lie in a horizontal plane (Triplagia, Pl. 91, fig. 2), or correspond to the three edges of a low pyramid (Plagiacantha). A fourth ray is sometimes added, which stands vertically upon the summit of the pyramid (Plagoniscus, Plagiocarpa, Pl. 91, figs. 4, 5). In other Plectoidea three secondary rays are intercalated between the three primary (Hexaplagida, &c.); seldom the number is greatly increased (Polyplagida, &c.). The rays are rarely simple, but usually branched; in the Plagonida (Pl. 91, figs. 2-6) the branches remain free; in the Plectanida (Pl. 91, figs. 7-13) they are united to form a loose wicker-work. From such a web a perfect Cyrtoid shell may arise. Several forms of Plagonida may also be readily confounded with the isolated triradiate or quadriradiate spicula of many Beloid skeletons (Sphærozoum, Lampoxanthium, &c.).

126. The Spongoid Skeleton.—From the simple lattice-skeleton which the majority of Radiolaria possess, some of them develop a spongy shell; the trabeculæ of the lattice-work, situated in one plane in the former, are developed in the latter in different planes and cross irregularly in all directions; thus arises a kind of wicker-work of more or less spongy structure, usually with very thin trabeculæ and irregular meshes. Such Spongoid shells are most common among the Spumellaria, especially in the Sphæroidea (Spongosphærida, Pl. 18) and Discoidea (Spongodiscida, Pls. 41-47), more rarely in the Prunoidea and Larcoidea. Lattice-work of similar spongy structure occurs very seldom among the Nassellaria, e.g., in some Plectoidea (Pl. 91) and Cyrtoidea (Spongocyrtis, Spongopyramis, Spongomelissa, &c., Pl. 56, fig. 10; Pl. 64, figs. 5-10, &c.). Among the Phæodaria spongy skeletons are very rare; they {lxxxiv}are to be seen in some Phæosphæria (Oroplegma, Pl. 107, fig. 1; Sagoplegma, Pl. 108, fig. 2; Auloplegma, Pl. 111, fig. 8). No Spongoid skeletons are known among the Acantharia.

127. The Cannoid Skeleton.—Cannoid or tubular skeletons are those which are composed of hollow tubes; they occur exclusively in the Phæodaria or Cannopylea. Tubular processes, nevertheless, occur in some other Radiolaria, as, for example, among the Spumellaria in a portion of the Collosphærida (Siphonosphæra, Caminosphæra, Pls. 6, 7), and of the Prunoidea (Pipetta, Cannartus, &c., Pl. 39, figs. 6-10, &c.), also among the Nassellaria in Theosyringium (Pl. 68, figs. 4-6), Cannobotrys (Pl. 96, figs. 3, 4, 8-11, 20-22), &c. In all these cases, however, the tubes are direct processes of the cavity of the shell, the trabeculæ of the lattice-work being solid. Only in the Cannopylea are the lattice-bars themselves, the radial spines and appendicular organs, generally tubular (hence the designation "Pansolenia"). The lumen of the thin-walled siliceous tubes is filled with jelly, and hence the specific gravity of the relatively large skeleton is considerably diminished. This peculiarity is not found in all Cannopylea; it is wanting in all Sagosphærida and Concharida, as well as in a part of the Orosphærida and Castanellida; in the latter there are found intermediate stages between hollow and solid skeletal rods. Very often a fine siliceous thread runs in the axis of the tubes, which is connected with its wall by lateral branches (Pl. 110, figs. 4, 6; Pl. 115, figs. 6, 7). More seldom the tubes are divided by horizontal septa into a series of chambers (Medusettida, Pls. 118-120). The two families Aulosphærida (Pls. 109-111) and Cannosphærida (Pl. 112) are distinguished from all other Phæodaria by the fact that their tubes are separated by astral septa in the nodal points of the lattice-shell (§§ 112, 134).

128. The Conchoid Skeleton.—By the name "Conchoid skeletons" are distinguished the bivalved lattice-shells which occur exclusively in the legion Phæodaria; they are quite characteristic of the Phæoconchia or Phæodaria bivalvia, which embrace three families:—Concharida (Pls. 123-125), Cœlodendrida (Pls. 121, 122), and Cœlographida (Pls. 126-128). The two valves of the lattice-shell of the Concharida are simple, hemispherical, or boat-shaped, whilst in the Cœlodendrida and Cœlographida tubes grow out from them, which branch and usually give rise by anastomosis to a second external bivalved shell. In all Phæoconchia the two valves are so disposed about the central capsule that an open slit remains between them, into which open the apertures of the central capsule; and since all these Phæodaria conchoidea are Tripylea, with three typical openings in the central capsule, and since the two lateral accessory openings lie at either side of the aboral pole, and the unpaired main-opening at the oral pole of the main axis, it follows that the two valves are to be regarded as dorsal and ventral as in the Brachiopoda (not right and left as in the Lamellibranchiata). The dorsal and ventral {lxxxv}valves are usually equal, but in a portion of the Concharida they present constant differences. In this family the two valves are attached to each other by their free edges, just as in the bivalved Mollusca and Diatoms; and these edges may either be smooth (Conchasmida, Pl. 123, figs. 1-6), or dentate (Conchopsida, Pls. 124, 125); the valvular connection of the latter is sometimes strengthened by a special ligament which unites the two valves at the aboral pole (Pl. 123, figs. 8, 9). The form of the valve is sometimes hemispherical, sometimes boat-shaped, with a sagittal keel.

129. Medullary and Conical Shells.—In all Radiolaria whose skeleton consists of a double shell or of two concentric lattice-shells united by radial bars, an inner medullary shell (testa medullaris) and an outer cortical shell (testa corticalis) may be distinguished (see note A, below). The medullary shell is usually to be regarded as a primary, the cortical as a secondary structure. Such double shells occur among the Spumellaria in the Dyosphærida (Pls. 19, 20), as well as in many Prunoidea (Pls. 39, 40), Discoidea (Pls. 33, 34), and Larcoidea (Pls. 9, 10); among the Acantharia only in the family Phractopeltida (Pl. 133); among the Nassellaria only in very few Cyrtoidea (e.g., Periarachnium, Pl. 55, fig. 11), and finally among the Phæodaria in the Cannosphærida (Pl. 112) as well as in part of the Cœlodendrida (Pl. 121) and Cœlographida (Pls. 127, 128). In most cases (if not always?) the cortical shell arises by the growth of radial spines from the surface of the medullary shell; these become united at equal distances from the centre by transverse apophyses, the surface of the secondary calymma furnishing the basis for their secretion (§ 85). Nevertheless, it seems that in many Sphærellaria the formation of the whole cortical shell proceeds simultaneously (at a definite dictyotic period) like that of the primary medullary shell (see note B). Whilst in the Phæodaria, Acantharia, and Nassellaria, at most two concentric shells are formed, in many Spumellaria their number increases continuously with additional growth; in many Sphærellaria it rises to four, eight, or even more, as well as in many Discoidea (if the concentric, peripherally disposed rings of chambers be regarded as incomplete flattened shells). In these cases either only the innermost primary lattice-shell is to be styled "medullary shell," or at most the two innermost (inner and outer medullary shells), all the others being cortical.

130. Dictyosis or Lattice Formation of the Skeleton.—In the great majority of Radiolaria the dictyosis or formation of lattice-work, and especially the formation of a variously-shaped "lattice-shell," plays such an important part that the whole class has long been popularly known in Germany by the name "lattice animalcules" ("Gitterthierchen" or "Gitterlinge") (Protista dictyota). The old name Polycystina also (1838), although referring only to the Spumellaria and Nassellaria, is derived from the lattice-work of the siliceous skeleton. The extremely various forms in which this is manifested furnish the means of distinguishing species. The specific conformation of the skeletal lattice-work is usually caused by the special disposition of the sarcodictyum (§ 94), whose exoplasmatic threads become silicified or (in the Acantharia) converted into bars of acanthin. In many cases, however, the form of the lattice is mainly dependent upon the situation and form of the radial spines or of special processes from them. With respect to their origin, two varieties of lattice may be distinguished—simultaneous and successive. Simultaneous dictyosis occurs especially in the simple lattice-shells of the Sphærellaria and Phæodaria, where, at a given moment ("dictyotic moment") the whole lattice of the shell is excreted on the surface of the calymma. Successive dictyosis, on the other hand, is found more particularly in the lattice-shells of the Acantharia (and in the concentric cortical shells of many Sphærellaria), which develop from the separate lattice-plates formed by the apophyses of the radial spines, and hence not at the same moment. The lattice-shells of the Cyrtellaria, which gradually grow out from a sagittal ring or a basal tripod, arise by successive dictyosis.

131. Dictyosis of the Spumellaria.—Siliceous lattice-structures are wanting in the first section of the Spumellaria, the Collodaria, but in the second section, Sphærellaria, they are developed in extraordinary variety of details. In spite of this extreme richness in different forms, the lattice-shells of the Spumellaria may all be derived from one and the same primitive ground-form, a simple lattice-sphere with regular hexagonal meshes (Phormosphæra, p. 61, Pl. 12, figs. 9-11; Heliosphæra, &c.). {lxxxvii}The siliceous bars which bound these regular and subregular meshes are at first exceedingly then and filiform; afterwards they become thicker or spread out laterally, whence the meshes often become round with a hexagonal frame (Pl. 12, fig. 5; Pl. 28, fig. 1). If the latter vanish, a lattice-shell with simple circular meshes is formed. Very commonly the regular form of the meshes or pores becomes more or less irregular, polygonal, or roundish. Hence, in general, four different principal forms of dictyosis may be distinguished among the Spumellaria; viz. (1) regular or subregular hexagonal meshes; (2) regular or subregular circular meshes; (3) irregular polygonal meshes; (4) irregular roundish meshes. The three latter forms are to be regarded as secondary, derived from the primary first form. In those Spumellaria which possess several concentric lattice-shells enclosed one within another, either these have all the same form of dictyosis, or the lattice-work of the innermost primary shell is different from that of the outer secondary shells (Pls. 19, 20); sometimes these latter also differ more or less among themselves (§ 129).

132. Dictyosis of the Acantharia.—The lattice-structures of the Acantharia differ essentially from those of other Radiolaria in several particulars. Firstly, they consist not of silica but of acanthin (§ 102); secondly, they are always secondary formations, usually developed from transverse processes of the primary centrogenous radial spines; thirdly, their formation is not simultaneous (at the same time over the same shell), but successive (proceeding from the individual radial spines tangentially towards the middle of the intervals); fourthly, the configuration of the network is due to the relative position of the spines and the mode of union of their transverse apophyses. Since they are at right angles to the spines, and since the branches of the apophyses are at right angles to them, the original ground-form of their dictyosis is a lattice-work with quadrangular meshes; these are often quite regular and square (Pl. 130, figs. 5, 6; Pl. 136, figs. 2, 9, &c.); more commonly they are rectangular or irregularly quadrangular (Pl. 131, fig. 10; Pl. 133, figs. 2, 3, &c.). In the majority of the Acantharia the quadrangular form of the meshes passes over into an irregularly polygonal or roundish one (Pls. 137, 138). Very often the primary meshes of the lattice-shells, which immediately surround the radial spines, are larger and more regular ("aspinal pores"), whilst the numerous secondary meshes between them are smaller and irregular ("coronal pores"; Pl. 135, figs. 1-4, &c.).

133. Dictyosis of the Nassellaria.—The siliceous lattice-structures of the Nassellaria are formed on the whole like those of the Spumellaria, with which they were formerly united under the name "Polycystina." In this group also there may be distinguished as two main forms the regular and irregular. In the Nassellaria the regular lattice-structures generally exhibit hexagonal or circular meshes, whilst the irregular are either polygonal or roundish; the irregular forms are, however, much more abundant than the {lxxxviii}regular, and a further distinction from the Spumellaria consists in the fact that the primary skeletal elements, from which the lattice is secondarily developed, exercise a predominant influence upon their form. These primary elements in the majority of the Nassellaria are to be seen in two morphologically most important structures:—first, the primary sagittal ring, which embraces the central capsule in the median plane (§ 124); and secondly, the basal tripod (§ 125), whose three diverging rays proceed from the base of the central capsule, whilst commonly a fourth vertical ray supports the dorsal side of latter (compare Pls. 81-91, p. 892). In the majority of the Nassellaria these two primary elements appear in combination, whilst in others only one of them is recognisable. In addition there occur numerous monaxon lattice-shells in which neither of these elements can be recognised, but a simple ovoid lattice-shell (cephalis) alone forms the whole skeleton or its primary part (Pl. 51, fig. 13; Pl. 98, fig. 13). The great difficulty in the morphological interpretation and phylogenetic derivation of the Nassellarian skeleton lies in the fact that each of these three elements—the primary sagittal ring, the basal tripod, and the latticed cephalis—may form the whole skeleton by itself or be combined with one or both of the others (p. 893). Even nearly related or at all events very similar forms may differ very greatly in this respect. With regard to the manifold forms of their dictyosis it follows that it is partly dependent upon one of the two first elements, partly independent. In the Plectellaria (or those Nassellaria which do not possess a complete lattice-shell) the lattice-work is usually irregular and arises by union of the ramifications, which proceed either from the primary sagittal ring (Pls. 81, 82, 92-94) or from the basal tripod (Pl. 91). In the Cyrtellaria (or Nassellaria with a complete lattice-shell, Pls. 51-80), on the other hand, the lattice-work is sometimes regular, sometimes irregular, being often very different in the different joints of a segmented shell (Pl. 72); a great part of it arises independently of the two chief morphological elements, and develops according to laws similar to those which regulate the dictyosis of the Spumellaria.

134. Dictyosis of the Phæodaria.—The lattice-structures of the Phæodaria, which consist of a silicate of carbon (§ 102), are on the whole not developed in such variety as those of the other Radiolaria, but exhibit several essentially different types of structure, not reducible to a common primitive type of lattice-work. In one portion of this legion there occurs an ordinary simple lattice-work (as in Spumellaria and Nassellaria), with solid trabeculæ; of these the Castanellida (Pl. 113) and Concharida (Pls. 123-125) have usually regular or subregular, circular meshes, sometimes hexagonally framed; the Orosphærida (Pls. 106, 107) large irregular polygonal meshes with thick trabeculæ, the Sagosphærida (Pl. 108) large triangular meshes with thin filiform trabeculæ. The Challengerida (Pl. 99) are characterised by a very delicate regular lattice-work, with minute hexagonal pores, like a Diatomaceous frustule. The Medusettida (Pls. 118-120) {lxxxix}show a peculiar alveolar structure, numerous small compartments being enclosed between two parallel plates. In the Circoporida (Pls. 114-117) and Tuscarorida (Pl. 100) the opaque porcellanous shell has a peculiar cement structure (§ 104), and the lattice-structure is confined for the most part to characteristic rings of pores at the base of the hollow tubes, which arise from the shell. The most peculiar lattice-work, however, appears in the segmented shell of the Aulosphærida (Pls. 109-111) and Cannosphærida (Pl. 112). In the former the large meshes of the lattice-work are usually subregular and triangular, in the latter polygonal; the trabeculæ are hollow cylinders, filled with jelly, and containing usually a central axial thread. In each nodal point of the lattice, in which three or more tangential tubes meet, these are separated by stellate or astral septa.

135. Radial Spines of the Skeleton.—The skeleton in the great majority of Radiolaria is armed with radial spines, which are of great importance in the development of their general form and of their vital functions. From a morphological point of view the number, arrangement, and disposition of the spines is usually the determining factor as regards the general form of the skeleton. Physiologically they discharge distinct functions, as organs of protection and support; they act also, like the tentacles of the lower animals, as prehensile organs, since their points, lateral branches, barbed hooks, &c. serve to hold fast nutritive materials. In general main-spines and accessory spines may be distinguished in most Radiolaria; the former are of pre-eminent importance in determining the figure of the skeleton; the latter are merely appendicular organs. The main-spines present such characteristic and important differences in the various legions of Radiolaria that they must be considered separately.

136. Radial Spines of the Spumellaria.—The radial spines, which exhibit most manifold variations in the large order Sphærellaria, present characteristic differences in its four suborders. In the Sphæroidea their number and disposition serve for the separation into families (p. 59); the Cubosphærida (Pls. 21-25) always possess six radial main-spines, which stand opposite to each other in pairs and lie in three diameters of the shell, which are at right angles to each other and correspond to the axes of the regular crystallographic system. The Staurosphærida (Pl. 15) have four spines, which form a regular cross and stand opposite to each other in pairs, in two axes at right angles. The Stylosphærida (Pls. 13-17) show only two main-spines, which are opposed to each other in the vertical main axis of the body. Finally, the Astrosphærida (Pls. 18-20, 26-30) are characterised by a larger and variable number of radial spines (eight, twelve, twenty or more), sometimes regularly, sometimes irregularly arranged. Among the other Sphærellaria the Prunoidea (Pls. 13-17, 39, 40) are most allied to the Stylosphærida with two opposite main-spines; the Discoidea (Pls. 31-47), on the other hand, to the Staurosphærida with four crossed spines; there exist, however, Discoidea with two opposite, three marginal, or numerous radial main-spines; it is {xc}characteristic of this suborder that they all usually lie in the horizontal median plane of the lenticular shell, arising from its equatorial margin. The Larcoidea (Pls. 9, 10, 49, 50) show a great variety in the number and arrangement of their radial main-spines, which in the different families of this suborder stand in direct causal relation to the various forms of growth of the shell; usually the primary main-spines lie either in the three different dimensive axes, at right angles to each other, whose differentiation is characteristic of the lentelliptical Larcoid shell (§§ 34, 122) or in definite diagonal axes, which cut the former obliquely. The radial spines of the Spumellaria are never united in the centre of the body, but arise separately from the surface of the primary central lattice-shell (medullary shell), more rarely from one of the secondary (cortical) shells, which enclose it. Their form is originally three-edged (sometimes pyramidal, sometimes prismatic); the cause of this is to be found in their origin from the nodal points of the lattice-shell, whose meshes are primitively hexagonal; hence three trabeculæ unite in each nodal point, and are produced into three edges of the spine. Very commonly, however, the spines are round (conical or cylindrical), more rarely polygonal. The three edges are often delicately toothed, not unfrequently spirally twisted around the axis of the spine (Pl. 21, figs. 1, 12).

137. Radial Spines of the Acantharia.—The radial spines of this legion have a much greater significance than in the other three classes of Radiolaria, since here alone they are the primary determining factors in the skeletal structure, and grow outwards from the middle of the central capsule. This centrogenous origin of the radial spines is as characteristic of the Acantharia as their chemical constitution, which is not siliceous but acanthinic (§ 102). Furthermore, their form is in most cases so peculiar that even an isolated Acantharian spine can be generally distinguished from one belonging to either of the other three legions. In the great majority of the Acantharia (all Acanthonida and Acanthophracta) twenty radial spines are constantly present, which, disposed according to a definite geometrical law, make up the skeleton (compare § 110 above and p. 717). The twenty spines are generally simply apposed to each other in the centre (either by the surfaces or the edges of their pyramidal base); more rarely they are completely united and form a single star-like piece of acanthin (Astrolithium). Very rarely (Acanthochiasma) each two opposite spines are united so that ten diametric bars cross in the middle of the central capsule. Whilst in the great majority of Acantharia these twenty radial spines are present, the small group Actinelida is characterised by the possession of an inconstant, often very large number, sometimes over one hundred. Among these Actinelida are probably to be found the stem-forms of the whole legion. The variously modified spines of the Acantharia may be grouped in three main categories: (1) round (cylindrical or conical); (2) four-edged (prismatic or pyramidal); (3) two-edged (leaf- or sword-shaped). The latter very commonly bear two {xci}opposite transverse processes, the former four crossed ones. By ramification and union of these apophyses arise the lattice-shells of the Acanthophracta (excepting the Sphærocapsida).

138. Radial Spines of the Nassellaria.—The radial spines in this legion show as great a variety in their form as in the Spumellaria, and, as in them, are solid, siliceous bars, usually three-edged (prismatic or pyramidal), or round (cylindrical or conical); more seldom they are polygonal in section. The great majority of the Nassellaria are, however, distinguished by a triradial structure, three primary radial bars diverging from the base of the central capsule (usually from the centre of the porochora); there is usually in addition a fourth apical spine, which passes upwards vertically or obliquely on the dorsal aspect of the central capsule. These three or four typical radial spines of the Nassellaria may be derived with great probability from the basal tripod of the Plectoidea (Plagoniscus, Plectaniscus, &c., Pl. 91); and since this tripod is very characteristically combined in Cortina and Cortiniscus with the primary sagittal ring of the Stephoidea, the three typical rays may be generally designated "cortinar feet," in contradistinction to the other radial processes of the Nassellarian skeleton. One of the three descending basal feet ("pes caudalis," Pls. 91-95, C) is always unpaired, and lies in the vertical median plane (or sagittal plane), just as does the vertically directed apical spine, which originally forms the dorsal bar of the sagittal ring, and is produced upwards into the "apical horn," (marked a on the plates). The other two basal feet are paired, and diverge right and left, forwards and downwards ("pedes pectorales," p.p.). Six-rayed Nassellaria, in which three secondary (interradial) feet are intercalated between the three primary (perradial) cortinar feet, are less common than the three-rayed forms. In some groups the number rises still higher, nine, twelve, or even more secondary feet being intercalated between the three primary. Besides, accessory radial spines may be developed on different parts of the shell, which have sometimes a definite relationship to the typical radial spines, sometimes not. Their form and ramification are very various (Pls. 51-98).

139. Radial Spines of the Phæodaria.—The radial spines of the Phæodaria are very clearly distinguished from those of other Radiolaria by the fact that they are usually hollow tubes, rarely solid bars. As a rule, the tubes are cylindrical, often slightly fusiform or conical, their siliceous wall is very thin, and their lumen filled with jelly; a fine thread of silica usually runs in the axis, and in several families is connected by fine transverse threads with the wall of the tube (Pl. 110, figs. 4, 6; Pl. 115, figs. 6, 7). The peculiar family Medusettida is characterised by a very remarkable segmentation of the hollow spines (Pls. 118-120). Each tube is divided by a series of septa into chambers, which communicate by a central or excentric opening in each septum, an arrangement resembling the siphuncle of the chambered Cephalopod shells. The number and {xcii}arrangement of the radial tubes in most Phæodaria is indefinite and very variable; only in a few families is the number constant in each species and genus, and the disposition regular. The Medusettida (Pls. 118-120) resemble the Nassellaria, inasmuch as equal radial feet diverge from the base of their shell, sometimes three in number (Cortinetta, Pl. 117, fig. 9), sometimes four (Medusetta, Pl. 120, figs. 1-4), sometimes six (Gazelletta); Gorgonetta is specially distinguished by the possession of six ascending and six descending spines regularly alternating (Pl. 119). The Tuscarorida (Pl. 100) usually have three or four equidistant feet. The Circoporida (Pls. 115-117), on the other hand, rather approach the Sphæroidea, their spherical or regular polyhedral shell having a definite number of tubular radial spines, which arise at regular intervals from their angles; Circoporus has six, Circospathis nine, Circogonia twelve, and Circorrhegma twenty radial tubes. Very rarely the tubes of the Phæodaria are angular, usually they are round, more or less cylindrical, though they are often bifurcated or even ramified, and exhibit a great wealth of the most delicate appendages; siliceous hairs, bristles, spines, barbed or anchor-like hooks, spathillæ, brushes, circlets, &c. (compare Pls. 99-128).

140. Main-Spines and Accessory Spines.—As accessory spines (Paracanthæ) we oppose to the main-spines (Protacanthæ), just described, all those processes which have no determining influence upon the formation of the skeleton as a whole, but are to be regarded as secondary constituents of the skeleton, or appendicular organs of inferior significance. They are developed in the utmost variety, sometimes as hairs or bristles, sometimes as thorns or clubs, either straight or curved (often zigzag), smooth or barbed; sometimes standing vertically upon the shell, or directed towards the centre, sometimes obliquely, or rising at a definite angle. In those Spumellaria whose lattice-shell consists of several concentric spheres, the accessory spines generally arise from the outermost, the main-spines, on the contrary, from the innermost. In the Nassellaria, multifarious forms of accessory spines are especially developed in the order Plectellaria. In the Phæodaria they are often furnished with delicate appendages, e.g., anchor-hooks, spathillæ, coronets, &c. Among the Acantharia the accessory spines which arise from the surface of the shell in the Acanthophracta are very characteristic. They are not radially disposed (like the similar superficial spines of the Spumellaria), but parallel to the radial main-spines from whose transverse processes they arise. Since in all these Acanthophracta the twenty radial main-spines are opposite to each other in pairs, all the accessory spines (often several hundred) are parallel to ten different regularly disposed axes of the lattice-shell (Pls. 134-138).

BIOGENETICAL SECTION.

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

Chapter V.—ONTOGENY OR INDIVIDUAL DEVELOPMENT.

(§§ 141-152.)

141. Individual Developmental Stages.—The germinal history of the Radiolaria presents great obstacles to direct observation, and hence is very incompletely known. The fragmentary observations, however (having been made on Radiolaria of very various groups and supplemented by comparative anatomical considerations), allow us to draw a general picture of the essential developmental processes in this great class. It may probably be assumed that in all Radiolaria, after maturation, the central capsule discharges the function of a sporangium, and its contents are broken up into numerous flagellate swarm-spores (zoospores). After these flagellate swarm-spores (resembling Astasia) have emerged from the ruptured central capsule, they probably pass over into a Heliozoan-stage (Actinophrys) and then after the formation of a jelly-veil into the condition of Sphærastrum. Afterwards, when a membrane is formed between the outer jelly-veil and the inner nucleated cell-body, an Actissa-stage arises, which exhibits in its simplest form the differentiation of the spherical unicellular body into the central capsule and calymma. Actissa thus represents both ontogenetically and phylogenetically the primitive condition of the Radiolarian organism, and may thus be regarded as the point of departure of all other forms.

142. The Astasia-Stage.—The formation of flagellate zoospores in the mature central capsule is probably to be regarded as the common form of individual development in all Radiolaria; since the whole contents are utilised in the formation of these swarm-spores, and since the extracapsulum takes no share in the process and perishes after they are evacuated, the central capsule may be regarded as a sporangium (see note A, below). The zoospores of the Radiolaria generally arise in the following way:—the nucleus of the unicellular organism, sometimes early, sometimes late (and in several different ways, §§ 63-70) breaks up into numerous small nuclei, and each of these surrounds itself with a small portion of the endoplasm. Very often, perhaps generally, this endoplasm contains one or several fat-granules and sometimes also a small oblong crystal; from the protoplasm {xciv}of the small roundish or ovoid cells protrudes one or more vibratile flagella. The fully developed spores, which commence their vibrations even within the central capsule, emerge when it ruptures, and swim about freely in the surrounding water by means of the flagellum. At this stage of its existence the young Radiolarian represents essentially the simplest form of the Flagellata, such as Astasia or Euglena; the unicellular body is for the most part ovoid or subcylindrical, sometimes fusiform or reniform, usually from 0.004 to 0.008 mm. in diameter (Pl. 1, fig. 1c; Pl. 129, fig. 11). In the anterior part of the flagellate cell, immediately behind the base of the flagellum, lies a homogeneous, spherical nucleus, whilst in the posterior part are usually several small fat-granules and often also a small oblong crystal (hence the name "crystal-spore," "Krystall-Schwärmer"). The number of vibrating flagella, which are extremely long and fine, seems to be variable, usually one, sometimes two, occasionally perhaps three, or even four or more (see note B).

143. The Actinophrys-Stage.—The fate of the flagellate zoospores which emerge from the mature central capsule of the Radiolaria has not hitherto been decided by actual observation; all attempts to rear the swarming zoospores have been in vain, for they have soon died. From what we know, however, of the comparative morphology of the Protista, the hypothesis is fully justified, that between the Astasia-stage of the flagellate swarm-spores, and the well-known Actissa-stage of the simplest Radiolaria, there lies an intermediate developmental stage, which may be regarded as being essentially the simplest Heliozoan form, Actinophrys or Heterophrys. The swarm-spore is very probably converted directly in to a simple floating Heliozoon by its elongated or ovoid body {xcv}becoming spherical and by fine pseudopodia protruding all round instead of a single flagellum; the nucleus at the same time assuming a central position.

144. The Sphærastrum-Stage.—The Actinophrys-stage of the young Radiolaria, which proceeds immediately from the flagellate zoospore, is probably connected with the Actissa-stage by an intermediate form, which may be regarded as a simple skeletonless Heliozoon with a jelly-veil; a well-known example of such a form is Sphærastrum (in the solitary, not the social condition) and Heterophrys. This important intermediate form has arisen from the simple Actinophrys-stage by the excretion of an external structureless jelly-veil, such as is formed in many other Protista (e.g., in the encystation of many Infusoria). The young Radiolarian in this second Heliozoon-stage becomes a simple cell with pseudopodia radiating on all sides; its body consists of three concentric spheres, the central nucleus, the protoplasmic body proper, and the surrounding calymma or jelly-veil. When a firm membrane is developed between the last two spheres this Sphærastrum-stage passes over into the Actissa.

145. The Actissa-Stage.—The first Spumellarian genus, Actissa, is not only the simplest form actually observed among the Radiolaria, and the true prototype of the whole class, but also the simplest form under which the Radiolarian organisation can be conceived. It is therefore extremely probably that Actissa not only forms the common stem-form of the whole class in a phylogenetic sense, but is also its common ontogenetic or germinal form. Probably in all Radiolaria the Sphærastrum-stage develops immediately into the typical Actissa-stage, by the formation of a firm membrane between the protoplasmic body of the spherical Heliozoan cell and its jelly-veil. Thus arises the characteristic central capsule, which is wanting in the nearly related Heliozoa. It is further probable that all Radiolaria in their early stage will so far conform to the state of things in Actissa as to have the capsule-membrane of the spherical skeletonless cell perforated everywhere by fine pores. This structure is retained in all Spumellaria, whilst in the other three legions those structural relations of the capsule which are characteristic of each develop from the Actissa-stage.

146. The Ontogeny of the Spumellaria.—In the simplest case the individual development in the Spumellaria ceases with the Actissa-stage. In all other genera of this legion diverging forms proceed from this, of which the different growth of the three dimensive {xcvi}axes on the one hand (§§ 44, 45), and the differentiation of the various parts of the unicellular organism with the formation of the skeleton on the other, are of pre-eminent significance. Even in the varying growth of the central capsule in the different dimensions of space in the skeletonless Colloidea, four different modes may be distinguished, which further, in the corresponding development of the skeleton, furnish the basis for the origin of the four orders of Sphærellaria. The most primitive and simplest form of growth, equal extension in all directions, is found in the spherical central capsule and the concentric spherical skeletons (Procyttarium, Sphæroidea). When the growth of the central capsule proceeds more rapidly in the direction of the vertical main axis than in any other direction, the ellipsoidal or cylindrical central capsule (Actiprunum) arises, and the vertically elongated skeleton of the Prunoidea, which is derived from it. When, on the contrary, the growth of the central capsule and lattice-shell is less in the direction of the vertical main axis than in any other direction, the lenticular or discoid central capsule (Actidiscus) arises, and the corresponding lenticular shell of the Discoidea. Finally, even quite early in many Spumellaria, the growth of the central capsule and of the corresponding lattice-shell in the three dimensive axes is different, and hence arise the lentelliptical forms whose geometrical type is the triaxial ellipsoid or the rhombic octahedron (Actilarcus, Larcoidea). Thus the origin of the four orders of Sphærellaria is simply explained by a varying growth in the different dimensive axes. The primary (innermost) lattice-shell is in this legion always simultaneously developed (suddenly excreted at the moment of lorication from the sarcodictyum). The secondary lattice-shells, on the other hand, which surround the former concentrically, and are united with it by radial bars, arise successively from within outwards.

147. The Ontogeny of the Acantharia.—The individual development of the Acantharia in the simplest case (Actinelius) stops at a point which differs from the Actissa-stage only in the change of radial axial threads into acanthin spines. In the small group Actinelida, their number remains variable and usually indeterminate (Adelacantha), whilst in the great majority of the legion (Acanthonida and Acanthophracta) the number is constantly twenty, and those spines are regularly arranged according to the Müllerian law in five parallel circles, each containing four crossed spines (Icosacantha). The simplest form among these latter is Acanthometron, which may be regarded both ontogenetically and phylogenetically as the common starting-point of all the Icosacantha. Within this extensive group variations in the length of the dimensive axes appear, similar to those observed in the Spumellaria. In the Astrolonchida and Sphærophracta the central capsule remains spherical, extending equally in all directions; and correspondingly the lattice-shell, which is excreted on the surface of the spherical calymma, remains spherical. In the Belonaspida (just as in the Prunoidea) {xcvii}this form passes over into an ellipsoid by prolongation of one axis; on the contrary, in the Hexalaspida (as in the Discoidea) the discoidal or lenticular form arises by shortening of an axis. Finally, in the Diploconida, and in some Hexalaspida in which the growth is different in all three dimensive axes (as in the Larcoidea), both the central capsule and the shell assume the lentelliptical form. The lattice-shell of the Acanthophracta is usually successive in its development, since from each of the twenty radial spines two or four tangential apophyses proceed, whose branches subsequently unite and combine to form the lattice-shell. Only in the peculiar Sphærocapsida can the pavement-like shell arise simultaneously or in a moment of lorication.

148. The Ontogeny of the Nassellaria.—The individual development of the Nassellaria in the simplest instance remains stationary at the skeletonless Nasselid stage (Cystidium, Nassella), which can be immediately derived from the foregoing Actissa-stage by the disappearance of the pores in the upper (apical) hemisphere of the central capsule, whilst in the lower (basal) portion they are modified to form a porochora; the podoconus is developed within the endoplasm upon this latter. Usually the spherical form of the central capsule passes over into an ovoid or ellipsoidal one, the vertical axis which passes through the centre of the porochora being elongated. From the skeletonless Nassellida the other Nassellaria may be derived both ontogenetically and phylogenetically by the excretion of an extracapsular siliceous skeleton. Unfortunately, the earliest stages in the formation of this skeleton are unknown, and hence no answer can at present be given to the important question, in what order the three primary skeletal elements of the Nassellaria (the basal tripod, sagittal ring, and latticed cephalis) appear (compare §§ 111 and 182). If, for example, in Cortina and Tripospyris the basal tripod were to appear first in the ontogeny, and the sagittal ring were developed from this, then the Plectoidea would be rightly considered to be the oldest forms in the phylogeny of the skeleton-forming Nassellaria; and in the contrary case the Stephoidea would be so regarded. The relations of growth in the three dimensive axes are very variable in the Nassellaria; the three most important factors in this respect (partly separately and partly in combination) are; (1) the development of the basal tripod to a triradial stauraxon form (the ground-form being a three-sided pyramid); (2) the development of the sagittal ring in the median plane of the body (the vertical axis having the poles different); (3) the development of the latticed cephalis outside the central capsule (the poles of the vertical axis being again different). Since the development both of the skeleton and of the malacoma is characterised in most Nassellaria by the stronger growth of the vertical axis and the differentiation of the two poles, the allopolar monaxon ground-form acquires a predominant significance in this legion (§ 32); the starting point of most of the further modifications is the basal pole of the vertical main axis. Next to this the sagittal axis is usually the most important determining factor (its dorsal and ventral poles being {xcviii}usually different), more rarely the frontal axis (with equal right and left poles). In the zygothalamous Spyroidea (as in the Stephoidea) the formation of the shell proceeds from the sagittal ring, whilst in the polythalamous Cyrtoidea the latticed cephalis is always the starting point, from which a series of joints (thorax, abdomen, and in the Stichocyrtida, the numerous post-abdominal joints) successively arise (unipolar growth).

149. The Ontogeny of the Phæodaria.—The individual development of the Phæodaria in the simplest case stops with the skeletonless condition of the Phæodinida (Phæodina, Phæocolla), which can be immediately derived from the foregoing Actissa-stage by the disappearance of the pores in the greater part of the central capsule, the characteristic astropyle being developed at the basal pole (§ 60). Since this particular form and structure of the spheroidal central capsule remains the same in all Phæodaria, whilst the formation of their skeleton follows very different directions, it follows that further common paths of development are excluded both ontogenetically and phylogenetically. What will be laid down in this respect as regards the phylogeny of the different groups of Phæodaria (§§ 194-199) holds true also of their ontogeny. The relations of growth in the three dimensive axes are hence very different in the skeletons of the various groups of Phæodaria. This difference is best marked in the Phæoconchia, whose bivalved lattice-shells have as their ground-form the rhomboid pyramid of Ctenophora. In most Phæogromia the monaxon lattice-shell may develop simultaneously by sudden excretion at a particular moment of lorication; this is also the case with the polyaxon lattice-shells of the Phæosphæria. In their future growth the development of basal or radial apophyses is of special importance. In the majority of the Phæodaria these apophyses are tubes of silicate filled with jelly (often provided with an axial siliceous thread); thus their development is distinguished by complications which are absent in the case of the other three legions.

150. Growth.—The growth of the body in the Radiolaria, as in all other organisms, is the fundamental function of individual development (see note A). All structural relations which this richest class of the Protista exhibits may be referred to different forms of growth, either of the unicellular malacoma or of the skeleton which it produces. In general the special development of the skeleton is dependent upon that of the central capsule, and of the sarcodictyum on the surface of the calymma; in the further growth, however, the conditions are reversed, and the condition of the skeleton already formed directly determines the further development of the central capsule and of the calymma with its sarcodictyum. The four legions of Radiolaria show, speaking generally, certain characteristic differences in growth, which are due in great part to the different structure and ground-form of their central capsule. In the two legions of the Porulosa (Spumellaria and Acantharia), in which the central capsule is originally spherical and {xcix}the ground-form of the skeleton either polyaxon or isopolar monaxon, two fundamental and variously combined directions of growth are recognisable; firstly, the concentric growth (equal increase of volume in all directions), and secondly, multipolar or diametral growth (hypertrophy of certain parts in the direction of definite pairs of radii). A different state of things obtains, however, for the most part, in the two legions of the Osculosa (Nassellaria and Phæodaria), in which the central capsule possesses a vertical main axis with different poles, and the structure of the skeleton is determined by this allopolar monaxon ground-form. The two fundamental directions of growth here combined in the most various ways are, firstly, unipolar growth (starting from the basal pole of the vertical main axis), and secondly, radial or pyramidal growth (characterised by the different development of separate parts in the direction of definite radii). Whilst the growth of the malacoma is dependent on intussusception (as in most organic structures capable of imbibing), the growth of the skeleton in all Radiolaria takes place by apposition (see note B).

151. Regeneration.—Whilst the general course of individual development (perhaps without any exception in the Radiolaria), begins with the formation of zoospores in the central capsule, there yet occurs in some groups a different form of ontogeny, introduced by simple division of the unicellular organism, and coming under the term "regeneration" in its wider sense. This spontaneous division occurs quite commonly in the Polycyttaria (or social Spumellaria), and produces their colonies (compare the chapter on Reproduction, § 213). On the contrary, it has not been observed in the solitary Spumellaria, nor in the Acantharia and Nassellaria; possibly, however, the peculiar Acantharian family, Litholophida, has arisen by the division of Acanthonida (compare p. 734). Among the Phæodaria division is commonly observed in the order Phæocystina (which have an incomplete Beloid skeleton or none), and also in the Phæoconchia. In all these cases the increase by division is nothing else than an ordinary case of cell-division, in which bisection of the nucleus precedes that of the central capsule. The regeneration by {c}which each of the two daughter-cells develops to a complete mother-cell depends upon simple growth. Another form of regeneration, different from this, has been observed in Thalassicolla. If the central capsule be extracted artificially from the large concentric calymma, the enucleated central capsule produces a new extracapsulum, with sarcomatrix, pseudopodia, and calymma. This experiment may be repeated several times with the same result. (Compare A. Schneider, 1867, L. N. 20.)

152. The Formation of Colonies.—The individual development of colonies takes place in all three families of the Polycyttaria (Collozoida, Sphærozoida, Collosphærida) in the same simple way, by the repeated division of a single monozootic Spumellarian. Since these divisions only affect the central capsule and not the extracapsulum, the sister-cells, which arise by repeated division of the mother, remain enclosed in a common rapidly growing calymma. Probably in all Polycyttaria the commencement of the formation of colonies immediately follows the Actissa-stage of the monozootic mother-cell (or takes place in the Thalassicolla-stage, which arises from the former by the development of alveoles in the calymma). The simple central nucleus separates (by direct nuclear division) into two halves, and the central capsule follows this process of bisection, becoming constricted in the middle between the two daughter nuclei (Pl. 3, fig. 12). In the further growth of the colony the process of division proceeds in the older, now multinucleate, central capsules, in which an oil-globule has taken the place of the original nucleus; then the division of the oil-globules precedes that of the central capsule (Pl. 5, fig. 1). Another mode of growth of the colonies is the multiplication of the central capsules by gemmulation, or the formation of the so-called "extracapsular bodies" (Gemmulæ, § 214). The characteristic skeletal structure of the different species appears at a later stage. Whether ripe central capsules can emerge from the social bond of a cœnobium, and, having become isolated, establish the formation of a new colony, is very doubtful. The various forms which the cœnobium assumes in the different species of Polycyttaria, are due partly to simple growth, partly to the development of large vacuoles in the calymma.

Chapter VI.—PHYLOGENY OR GENEALOGICAL DEVELOPMENT.

(§§ 153-200.)

153. Sources of Phylogenetic Knowledge.—For the purpose of constructing a hypothetical genealogical tree of the Radiolaria, as of all other organisms, three sources of information are open to us, viz., palæontology, comparative ontogeny, and comparative anatomy. In the present case, however, these three sources are of very different value; the first two are at present only very inadequately known and have only been partially investigated, hence they can only be utilised to a very slight extent. The comparative anatomy of the Radiolaria, on the other hand, is so completely known, and affords such certain glimpses into the morphological relations of the related groups, that by its aid we are in a position at all events to lay down the general features of their phylogeny with some probability, and to lay the foundation of a natural system.

154. Natural and Artificial Systems.—Although in the classification of the Radiolaria, as in the case of all other organisms, the natural system must be regarded as the goal of systematic classification, our phylogenetic knowledge of the Radiolaria is too fragmentary and inadequate to admit of the systematic arrangement here adopted being regarded as a thoroughly consistent natural system, that is, as representing the true genealogical tree of the class. Owing, however, to the extraordinary variety of form of the Radiolaria, and the complicated relationships of the larger and smaller groups, a synoptical grouping of the different categories and the erection of a complete, even if to some extent artificial, system, becomes a logical necessity. Under these circumstances, and regard being had to both these conditions, the following systematic treatment of the Radiolaria will appear as a compromise between the natural and artificial systems, like all other zoological and botanical classificatory attempts. On the one hand, the attempt is made to arrange the larger and smaller groups as nearly as possible according to their phylogenetic relationships, whilst, on the other hand, the practice of circumscribing each by a definition as clear and logical as possible has been carried out. Since these two efforts naturally often come into contact, the insufficiency of many parts of the arrangement is obvious, hence its hypothetical and provisional character is emphatically stated.

155. Systematic Categories.—The categories or different orders of divisions have in the Radiolaria, as in all other organisms, no absolute significance, but only a relative value. In itself it is quite unimportant whether the whole group be regarded, as at first, as a family (Ehrenberg, 1847), or as an order (J. Müller, 1858), or as a class (Haeckel, {cii}1881). These different views are regulated, on the one hand, by the known extent of the group and by the amount of our acquaintance with it, and on the other, by comparison with related groups and by reference to their conventional disposition. When, therefore, the whole class, Radiolaria, is here divided into two subclasses, four legions, eight orders, eighty-five families, &c., these artificial categories are drawn up only in the conviction that by this means the easiest survey and most thorough insight into the system as a whole may be attained; this latter will indeed approach as far as possible the ideal of a natural system, but must on numerous practical grounds always remain more or less artificial. Since it is to be expected that with the progress of our systematic knowledge the rank of the various categories will rise, it is possible that in the future the arrangement of the group may be somewhat as follows:—Phylum, RADIOLARIA; Four Classes, Spumellaria, Nassellaria, Acantharia, Phæodaria; Eight Legions (Nos. I.-VIII. in the following Table); Twenty Orders (Nos. 1-20 in the Table), &c.

Four Legions.		Eight Sublegions.		Twenty Orders.		Typical Families.
I. Legion (or Subclass) Spumellaria (Peripylea) [Porulosa peripylea.]		I. Collodaria (Spumellaria palliata)		1. Colloidea,		1a. Thalassicollida.
						1b. Collozoida.
				2. Beloidea,		2a. Thalassosphærida.
						2b. Sphærozoida.
		II. Sphærellaria (Spumellaria loricata)		3. Sphæroidea,		3a. Ethmosphærida.
						3b. Collosphærida.
				4. Prunoidea,		4a. Ellipsida.
						4b. Zygartida.
				5. Discoidea,		5a. Phacodiscida.
						5b. Porodiscida.
				6. Larcoidea,		6a. Larnacida.
						6b. Pylonida.
II. Legion (or Subclass) Acantharia (Cannopylea). [Osculosa cannopylea.]		III. Acanthometra (Acantharia palliata)		7. Actinelida,		7a. Astrolophida.
						7b. Litholophida.
						7c. Chiastolida.
				8. Acanthonida,		8a. Astrolonchida.
						8b. Quadrilonchida.
						8c. Amphilonchida.
		IV. Acanthophracta (Acantharia loricata)		9. Sphærophracta,		9a. Sphærocapsida.
						9b. Dorataspida.
						9c. Phractopeltida.
				10. Prunophracta,		10a. Belonaspida.
						10b. Hexalaspida.
						10c. Diploconida.
{ciii} III. Legion (or Subclass) Nassellaria (Monopylea) [Osculosa monopylea.]		V. Plectellaria (Nassellaria palliata)		11. Nassoidea,		11. Nassellida.
				12. Plectoidea,		12a. Plagonida.
						12b. Plectanida.
				13. Stephoidea,		13a. Stephanida.
						13b. Tympanida.
		VI. Cyrtellaria (Nassellaria loricata)		14. Spyroidea,		14a. Zygospyrida.
						14b. Androspyrida.
				15. Botryodea,		15a. Cannobotryida.
						15b. Lithobotryida.
						15c. Pylobotryida.
				16. Cyrtoidea,		16a. Monocyrtida.
						16b. Dicyrtida.
						16c. Tricyrtida.
						16d. Stichocyrtida.
IV. Legion (or Subclass) Phæodaria (Actipylea) [Porulosa actipylea.]		VII. Phæocystina (Phæodaria palliata)		17. Phæocystina,		17a. Phæodinida.
						17b. Cannorrhaphida.
						17c. Aulacanthida.
				18. Phæosphæria,		18a. Orosphærida.
						18b. Aulosphærida.
						18c. Cannosphærida.
		VIII. Phæocoscina (Phæodaria loricata)		19. Phæogromia,		19a. Challengerida.
						19b. Castanellida.
						19c. Circoporida.
				20. Phæoconchia,		20a. Concharida.
						20b. Cœlodendrida.
						20c. Cœlographida.

156. Formation of Species.—The totality of similar forms, which we unite in one species, and which in the earlier dogmatic systems was regarded as a category of absolute value, possesses only a relative value like all other systematic categories (§ 155). According to the individual views of the systematist and the general survey which he has attained of the smaller and larger systematic groups, the conception of a species adopted in his practical work will be wider or narrower. In the present systematic arrangement a medium extent has been adopted. It is shown that in the Radiolaria, as in all other extensive groups of organisms, the constancy of the species is very variable in the different groups. Many families of Radiolaria are very rich in "bad species," i.e., very variable forms, in which the process of the formation of species is seen in progress; such, for example, are—among the Spumellaria, the Sphærozoida, Stylosphærida, Phacodiscida and Pylonida; among the Acantharia, the Amphilonchida and Phractopeltida; among the Nassellaria, the Stephoidea and Botryodea; and among the Phæodaria, the Aulacanthida, Sagosphærida, Castanellida and Concharida. On the {civ}other hand, in some families numerous "good species" may be distinguished, since the intermediate connecting forms are no longer present and the forms have become relatively constant. As instances of such families may be mentioned, among the Spumellaria, the Astrosphærida, Cyphinida, Porodiscida and Tholonida; among the Acantharia the Quadrilonchida and Dorataspida; among the Nassellaria, the Spyroidea and Cyrtoidea; among the Phæodaria, the Challengerida, Medusettida, Circoporida and Cœlographida. The more carefully the different groups are studied, the more numerous the individuals of each species under comparison, the greater becomes the number of "bad" species among the Radiolaria, and the smaller the number of good ones. Originally, no doubt, all "species bonæ" were "malæ." There may be observed in the manifold skeletal forms of the Radiolaria, on the one hand, the utmost accuracy of configuration, and on the other, the greatest variability, and hence a careful comparative study of them leads to a firm conviction of the gradual "Transformation of Species," and of the truth of the "Theory of Descent."

157. Palæontological Development.—The palæontology of the Radiolaria already offers very considerable material for study; but in consequence of its incompleteness this is of little value for the study of the phylogeny of the class. By far the larger portion of the fossil Radiolaria belong to the Tertiary period; only quite recently have numerous well-preserved fossil Radiolaria been described from the Mesozoic period, and especially from the Jura. Of Palæozoic Radiolaria (from the coal measures) only slight traces are known. Moreover, the fossil Radiolaria hitherto known have been found only in very circumscribed and widely separated localities. The majority of all the species belong to the small island of Barbados. Although our palæontological acquaintance with the Radiolaria must necessarily be incomplete for this reason, it is still more so since at least thirty out of the eighty-five families (that is more than a third) could not possibly leave any fossil remains, either because they possess no skeleton, or because of its chemical composition.

158. Origin of the Four Legions.—The agreement of all Radiolaria in those constant and essential characters of the unicellular body, which distinguish them from all other Protista (especially the differentiation of the malacoma into a central capsule and extracapsulum), justifies the conclusion that all members of this class have been developed from a common undifferentiated stem-form. Only the simplest form of the Spumellaria, a skeletonless spherical cell with concentric spherical nucleus and calymma, can be regarded as such. The simplest form of the Thalassicollida which is now extant (Actissa, Procyttarium, p. 12), corresponds so exactly to the morphological idea of that hypothetical stem-form that it may unhesitatingly be regarded in a natural system as the common point of origin of the whole class. On the other hand, Actissa is so closely related to the simple Heliozoa (Actinophrys, Actinosphærium, Heterophrys, Sphærastrum, &c.) that its origin from this group of Rhizopoda is exceedingly probable. The three legions Acantharia, Nassellaria, and Phæodaria are to be regarded as three main diverging branches of the genealogical tree, which have been developed in different directions and are only connected by their simplest stem-forms (Actinelius, Nassella, Phæodina) with the stem-form of the Spumellaria, the primordial Actissa.

159. Phylogeny of the Spumellaria.—The legion Spumellaria or Peripylea is to be regarded as the common stem-group of the Radiolaria, and its simplest form, Actissa, as the primitive genus or radical form of the whole class; for it possesses in the simplest and most undifferentiated form all those characters by which the Radiolaria are distinguished from other Protista; all the other genera of the class may be derived from it by successive modifications. Considered as a legion the whole group Spumellaria is undoubtedly monophyletic, for all its members possess those essential characters by which it is distinctively marked off from the other three legions, more especially a simple capsule-membrane, which is everywhere evenly perforated by innumerable small pores; the nucleus lies originally in the centre of the spherical central capsule. Furthermore, all Spumellaria lack those positive characters which distinguish the three remaining legions—the centrogenous acanthin skeleton of the Acantharia, the basal porochora and the monaxon podoconus of the Nassellaria, the astropyle and phæodium of the Phæodaria.

160. Origin of the Spumellaria.—The genus Actissa (p. 12, Pl. 1, fig. 1) presents the Radiolarian type in its simplest and most primitive form—a spherical central capsule, which encloses in its middle a spherical nucleus, and which is surrounded by a spherical calymma. The whole unicellular body consists, therefore, of three concentric spheres, {cvi}and possesses neither skeleton nor alveoles, nor other differentiated parts. The innumerable fine pseudopodia, which issue from the central capsule through the evenly distributed pores in its membrane, radiate in all directions through the calymma and pass out over its surface. Actissa can, therefore, be directly derived phylogenetically from the simplest skeletonless Heliozoa (Actinophrys, Heterophrys, Actinosphærium, Sphærastrum). The only essential difference between the two consists in the development of the central capsule, which in Actissa separates as a distinct membrane the endoplasm from the exoplasm. This differentiation which we regard is the most important distinguishing character of the Radiolaria, has been transmitted by inheritance, along with the formation of flagellate spores in the central capsule, from Actissa, the primitive parent to all the other Radiolaria.

161. Hypothetical Genealogical Tree of the Spumellaria:—

Larcoidea

Discoidea

Streblonida

Phacodiscaria

Tholonida

Coccodiscida

Prunoidea

Soreumida

Zygartida

Zonarida

Lithelida

Sphæroidea

Cyclodiscaria

Spongodiscida

Stylosphærida

Pylodiscida

Phorticida

Panartida

Artiscida

Phacodiscida

Spongodruppida

Staurosphærida

Spongel- lipsida

Pylonida

Cyphinida

Porodiscida

Astrosphærida

Larnacida

Cubosphærida

Cenodiscida

Druppulida

Collosphærida

Archidiscida

Larnacilla (Trizonium)

Spongurida

Ellipsida (Cenellipsis)

Larcarida (Cenolarcus)

Liosphærida (Cenosphæra)

Cenodiscida (Cenodiscus)

[Actiprunum?]

[Actilarcus?]

[Procyttarium]

[Actidiscus?]

Cenosphæra (Common stem-form of all Sphærellaria?)

Polycyttaria

Collosphærida

Collozoida

Sphærozoida

Beloidea

Thalassosphærida

Ethmosphærida

Colloidea

Thalassicollida

Actissa

Larcoidea

Streblonida

Tholonida

Prunoidea

Soreumida

Zygartida

Zonarida

Lithelida

Phorticida

Panartida

Artiscida

Spongodruppida

Spongel- lipsida

Pylonida

Cyphinida

Larnacida

Druppulida

Larnacilla (Trizonium)

Spongurida

Ellipsida (Cenellipsis)

Larcarida (Cenolarcus)

[Actiprunum?]

[Actilarcus?]

≈

≈

Discoidea

Phacodiscaria

Coccodiscida

Sphæroidea

Cyclodiscaria

Spongodiscida

Stylosphærida

Pylodiscida

Phacodiscida

Staurosphærida

Porodiscida

Astrosphærida

Cubosphærida

Cenodiscida

Collosphærida

Archidiscida

≈

Liosphærida (Cenosphæra)

Cenodiscida (Cenodiscus)

Actiprunum Actilarcus

[Procyttarium]

[Actidiscus?]

Cenosphæra (Common stem-form of all Sphærellaria?)

Polycyttaria

Collosphærida

Collozoida

Sphærozoida

Beloidea

Thalassosphærida

Ethmosphærida

Colloidea

Thalassicollida

Actissa

162. Collodaria and Sphærellaria.—Whilst in all Spumellaria the malacoma agrees in possessing the characteristic features of the legion, and thus justifies its derivation monophyletically from the common stem-form Actissa, the different forms of skeleton, on the other hand, cannot all be referred to the same fundamental form. More especially the spherical lattice-shell, from which all the numerous skeletal forms of the Sphærellaria may be derived, cannot have arisen from the incomplete Beloid skeleton which characterises the Beloidea among the Collodaria. It is probable rather that the formation of the skeleton has taken place independently in those two groups of Spumellaria. From the skeletonless Colloidea, as the common stem-group of the Spumellaria, two different main groups have diverged, on the one hand the Beloidea, whose skeleton consists of separate spicules scattered in the extracapsulum, and on the other hand, the Sphærellaria, which have formed a simple lattice-sphere around the central capsule; from this the manifold forms of the remaining Spumellaria may be derived.

163. Descent of the Sphærellaria.—The extensive order Sphærellaria, which includes all Spumellaria with a complete lattice-shell, develops an extraordinary variety of skeletal structures; these may, nevertheless, all be derived without violence from a common stem-form, or simple spherical lattice-shell, Cenosphæra. The main stem of the order, the extensive suborder Sphæroidea (Pls. 5-30), is derived immediately from Cenosphæra (p. 61, Pl. 12); three diverging branches of it being represented by the other three suborders, the Prunoidea (Pls. 16, 17, 39, 40) being developed by elongation, and the Discoidea (Pls. 31-48) by shortening of the vertical main axis, whilst the Larcoidea (Pls. 9, 10, 49, 50) have originated by the modification of the spherical lattice-shell into a lentelliptical or triaxial ellipsoidal one. Although the monophyletic derivation of all Sphærellaria from Cenosphæra is exceedingly probable, the possibility of a polyphyletic origin for the group is by no means excluded. For even in the skeletonless primitive genus of all the Spumellaria, Actissa (as well as in the social Collozoum), there are found, in addition to the usual spherical types, other species (or subgenera, p. 12) whose central capsule is not spherical but a modification of the sphere; in Actiprunum ellipsoidal; in Actidiscus lenticular; in Actilarcus lentelliptical; if such modified forms of Actissa were to develop their lattice-shells independently, then their form would correspond to that of the central capsule; and such simple ellipsoidal, discoidal, and lentelliptical lattice-shells might have been the primitive forms of the Prunoidea, Discoidea and Larcoidea.

164. Genealogical Tree of the Sphæroidea.—Cenosphæra, the simplest form of the spherical lattice-shell, may be unhesitatingly regarded as the common stem-form of all the Sphæroidea (pp. 50-284, Pls. 5-30). Cenosphæra (p. 61, Pl. 12) arose directly from Actissa simply by the silicification of the spherical exoplasmatic network of the sarcodictyum around the central capsule, on the surface of the concentric calymma. From this simple siliceous extracapsular lattice-sphere all other forms of Sphæroidea have arisen, in the main by the manifold combination of two simple processes, first by the formation of radial spines on the surface of the lattice-sphere, and second, the addition of concentric spherical lattice-shells. Both processes may be utilised as the foundation for a systematic treatment of the Sphæroidea (compare pp. 52-58).

165. Genealogical Tree of the Prunoidea.—The suborder Prunoidea is very closely related to the Sphæroidea, and is distinguished from it by the elongation of one axis; from the simple lattice-sphere (Cenosphæra) is developed a latticed ellipsoid (Cenellipsis, Pl. 39, fig. 1). The development of this vertical isopolar main axis is foreshadowed even among the Sphæroidea, in that family in which two opposite radial spines grow out of the primary lattice-sphere at the two poles of the vertical main axis (Stylosphærida, Pls. 13, 14). These latter pass over without any sharp boundary into those forms of Prunoidea whose ellipsoidal lattice-shell bears two opposite main-spines (Stylatractida, Pls. 15, 16). Other very intimate relationships between the Sphæroidea and Prunoidea are indicated in certain of the latter by the fact that of the two concentric lattice-shells the inner (medullary) shell is spherical, the outer (cortical) shell ellipsoidal (Pl. 39, figs. 3, 7, 8, 14, 19); often three concentric lattice-shells are present, of which the two inner are spherical intracapsular medullary shells, whilst the outer is an extracapsular cortical shell, ellipsoidal or cylindrical in form (Pl. 39, figs. 4, 12, 17, 18). Owing to the manifold nature of these phylogenetical relations and the variety of their combinations, the derivation of the individual Prunoidea from the Sphæroidea is rendered very difficult; in addition to which it is possible that the simplest Prunoidea (Cenellipsis, Ellipsidium) have been directly developed from the skeletonless Actiprunum (a form of Actissa with ellipsoidal central capsule, p. 14) by the excretion of a simple ellipsoidal lattice-shell on the surface of their calymma.

166. Genealogical Tree of the Discoidea.—The suborder Discoidea is closely related to the Sphæroidea, but separated from it by shortening of one axis; from a simple lattice-sphere (Cenosphæra) a latticed lens or flattened spheroid is developed, whose circular equatorial plane is larger than any other section (Cenodiscus, Pl. 48, fig. 1). The formation of this horizontal equatorial plane is perhaps indicated in that family of Sphæroidea in which four crossed radial spines, lying in one plane, are developed (Staurosphærida, Pls. 15, 31, 42). The morphological and phylogenetical relations of the Discoidea to the Sphæroidea are precisely the converse of those of the Prunoidea; in the latter the vertical axis appears longer, in the former shorter than any {cx}other axis of the body. The Discoidea are probably polyphyletic, having originated from several different groups of Sphæroidea; at least two essentially different main groups may be distinguished among them; of these the one is characterised by the formation of a large extracapsular lenticular cortical shell (Phacodiscaria), whilst in the other this typical "Phacoid shell" or lattice-lens is wanting (Cyclodiscaria, compare pp. 403-409).

167. Genealogical Tree of the Larcoidea.—The suborder Larcoidea presents in the structure, composition, and development of its variously formed lattice-shells much more complicated relations than the other Sphærellaria; it is essentially distinguished from them by the characteristic ground-form of its lattice-shells, which is a "lentellipsis" or a triaxial ellipsoid (also the ground-form of the rhombic crystallographic system, the rhombic octahedron). Hence all parts of the body are regularly disposed with respect to three different dimensive axes; all three axes, perpendicular one to another, are isopolar but of different lengths; the longest is the vertical main axis, the mean the horizontal frontal axis, the shortest the horizontal sagittal axis. In the great majority of the Larcoidea the lentelliptical ground-form is indicated in the central capsule, even when it is not at once obvious in the skeleton. Since such lentelliptical central capsules are developed even in Actissa (Actilarcus, p. 16), it is possible that the simplest Larcoidea may have arisen directly from these by deposition of a simple lentelliptical lattice-shell in the sarcodictyum, on the surface of the calymma (Cenolarcus, Pl. 50, fig. 7). It is more probable, however, that these simplest forms (Cenolarcus, Larcarium) have been developed from the simplest Sphæroidea (Cenosphæra), by the spherical body growing unequally in the three dimensions of space. It appears especially likely {cxi}from a study of the concentrically disposed lattice-shells of some Larcoidea (Coccolarcus, Larcidium, Pl. 50, fig. 8), in which the inner medullary shell is spherical, the outer cortical shell more or less elliptical. In the great majority of Larcoidea the latter arises in quite a peculiar manner, three broad lattice-zones, which are developed in three planes at right angles to each other, growing out from a small spherical or lentelliptical medullary shell, Trizonium, Larnacilla (compare pp. 600, 615, 628, &c.).

168. Descent of the Polycyttaria.—The polyzootic or colonial Radiolaria, which we unite in the group Polycyttaria (sometimes known as "Sphærozoea"), belong without doubt to the legion Spumellaria, for they possess all the peculiarities by which these Peripylea are distinguished from the other legions of the Radiolaria. Only the morphological position of the Polycyttaria in that legion, and their phylogenetic relation to the monozootic or solitary Spumellaria, can be variously interpreted. The three families which we distinguish among the Polycyttaria are so closely related to three different families of the Monocyttaria, that they may be directly derived from them by the formation of colonies. According to this triphyletic hypothesis the social skeletonless Collozoida (Pl. 3) would be descended from the solitary Thalassicollida (Pl. 1), the polyzootic Sphærozoida with a Beloid skeleton (Pl. 4) from the monozootic {cxii}Thalassosphærida (Pl. 2), and the colonial Collosphærida with a Sphæroid skeleton (Pls. 5-8) from the solitary Ethmosphærida (Pl. 12, &c.). Many species of monozootic and polyzootic forms in all three groups are so alike that they can only be distinguished by the fact that the one series are colonial, the others solitary. On the other hand, there are some reasons which would justify a monophyletic hypothesis for the Polycyttaria, e.g., the precocious nuclear division; in this case it would be most natural to hold that the Sphærozoida and Collosphærida have arisen as two diverging branches from the Collozoida, whilst the latter are nothing else than colonial Thalassicollida.

169. Phylogeny of the Acantharia.—The legion Acantharia or Actipylea is distinguished by its peculiar acanthin skeleton, which develops centrogenously, as well as by the disposition in groups of the pores in its central capsule, and its excentric usually precocious nucleus; it is thus so different from all other Radiolaria as undoubtedly to furnish, phylogenetically considered, an independent stem (§ 7). This stem is only connected at the root by Actinelius with the primitive form of the Spumellaria, Actissa. The stem is monophyletic, since all the forms belonging to it may be derived without violence from Actinelius as a common primitive form.

170. Origin of the Acantharia.—The genus Actinelius (p. 730, Pl. 129, fig. 1), which may naturally be regarded as the common primitive form of all Acantharia, possesses a spherical central capsule, which in consequence of the early division of the nucleus (§ 63), encloses numerous small nuclei; from its centre arise many simple radial spines of equal size, which penetrate the central capsule. A large number of radial pseudopodia issue between the spines from the sarcomatrix which surrounds the capsule. Actinelius may have been directly derived from Actissa, the common stem-form of all Radiolaria, by the division of the pseudopodia into two groups, myxopodia, which remained soft, and axopodia, which became firm (§ 95A). As the latter became changed into strong acanthin rods, and touched each other in the centre, they forced the nucleus from its originally central position and brought about its early division. Actinelius is also of all Radiolaria the form which, next to Actissa, most nearly approaches the Heliozoa. If the stiff axial threads of Actinosphærium be conceived of as partially converted into acanthin spines, and its nucleated medullary substance as separated from the alveolar cortical layer by a membrane (central capsule), then Actinelius would be produced.

171. Hypothetical Genealogical Tree of the Acantharia:—