Suborder III. CYRTOIDEA, Haeckel, 1862.

Cyrtida, Haeckel, 1862, Monogr. d. Radiol., pp. 272, 280.

Cyrtoidea vel Cyrtida, Haeckel, 1881, Prodromus, pp. 425-439.

Polycystina solitaria, Ehrenberg, 1847, Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, pp. 53, 54.

Monodictya nassellaria, Ehrenberg, 1875, Abhandl. d. k. preuss. Akad. d. Wiss. Berlin, pp. 156, 157.

Definition.—Nassellaria with a complete lattice-shell, exhibiting a simple or reduced cephalis, which is neither bilocular nor lobate, without sagittal constriction.

The order Cyrtoidea, described by me in 1862 as the family Cyrtida, is by far the largest of all the main groups of Radiolaria, and remarkable from the extraordinary variety of forms and the number of species. In the following system more than eleven hundred species are described, comprising about one-fourth of the number of species in the whole class of Radiolaria. This astonishing variety, however, is not effected by development of a large number of different types, but by an extraordinary variability within certain restricted boundaries, similar to what is seen among insects and birds. The number of genera, therefore, is comparatively small, and they may all be disposed into four families only, which in my Monograph (1862, p. 280) were distinguished as Monocyrtida, Dicyrtida, Tricyrtida and Stichocyrtida. If we divide these four groups in the following pages into twelve families and twenty-four subfamilies, we are guided by practical considerations only, hoping thereby to give a better survey of the difficult labyrinth of Cyrtoidean morphology.

The Cyrtoidea are characterised by this wonderful richness of specific forms not only in the present seas, but also for millions of years in the former ages of our globe. The majority of all the fossil Radiolaria which are now known belong to this group, and many species of it are so common that great rocks are formed by their union. The fact was first observed by Ehrenberg, who in his first system of Polycystina (1847, loc. cit., p. 54) enumerated forty-four genera and two hundred and eighty-two species; the Cyrtoidea, his Polycystina solitaria, form the preponderant majority of the whole class, viz., twenty-five genera and one hundred and ninety-three species.

In this first system (of 1847), as well as in the last systematic table of Ehrenberg (of 1875, loc. cit.), the Cyrtida as "Polycystina solitaria" are opposed to all other Radiolaria, as "Polycystina composita." The former bear the definition "Testæ siliceæ spatio interno ample pervio, aut passim levius transverse constricto"; the latter, however, "Testæ siliceæ spatio interno celluloso aut strictura longitudinali constricto." In reality these definitions are insufficient, and the conclusions which Ehrenberg derived from the organisation of the Polycystina solitaria and composita, were quite erroneous. So also are the definitions of the three families into which he divided the Polycystina solitaria, afterwards (in 1875) called by him "Monodictya nassellaria." These three families were the Halicalyptrina, Lithochytrina and Eucyrtidina. With these were also united the three genera of Botryodea known to Ehrenberg (Lithobotrys, Botryocampe, Botryocyrtis). We entirely separate these here from the true Cyrtida, on account of their lobate or multilocular cephalis.

Whilst Ehrenberg only knew the skeleton of the Polycystina solitaria, the first observations of living Cyrtida were published by Johannes Müller, 1858, in his fundamental treatise. He gave the first description and figures of the central capsule of this group, with the characteristic lobes developed from its basal part; and of the pseudopodia radiating on all sides (loc. cit., Taf. vi.). The forms described by him were all Mediterranean, one Dicyrtid (Lithomelissa mediterranea), two Tricyrtids (Eucyrtidium zancleum and Pterocanium charybdeum), and one Stichocyrtid (Lithocampe tropeziana).

In my monograph (1862, p. 272-341) I gave a detailed description of all known and some new Cyrtida, and characterised this family by the fundamental monaxonial form of the shell, with two different poles (an upper apical and a lower basal pole), and by the unipolar growth, beginning from the apical pole. I pointed out also the peculiar structure of the monaxonial central capsule. At that time I divided the Cyrtida into five subfamilies, in which, however, the Spyroidea (= Zygocyrtida), and the Botryodea (= Polycyrtida) were united with the true Cyrtoidea (Monocyrtida, Dicyrtida, Stichocyrtida).

The astonishing number of new and interesting forms of Cyrtida which I found in the rich collection of the Challenger (beginning from 1876), and mainly in the Radiolarian ooze of the Central Pacific (Stations 263 to 274), enabled me to give in my Prodromus, in 1881, a greatly enlarged and amended system of this important group. I separated there the Spyroidea (= Zygocyrtida), and the Botryodea (= Polycyrtida) from the true Cyrtoidea by restricted definition, pointing out the essential differences in the structure of the cephalis in these groups of Cyrtellaria. The latter name, as here used, is therefore identical with the "Cyrtida" of my Monograph. In the Prodromus I divided the true Cyrtida (p. 426) into five subfamilies and thirty tribes, corresponding to the differences in the number of the shell-joints and of the radial apophyses, and in the shape of the closed or open mouth. These groups are here retained, but reduced to four families and twenty-four subfamilies, since the Tetracyrtida are better united with the Stichocyrtida (compare below).

Richard Hertwig in his work Organismus der Radiolarien (1879, pp. 74 to 86) gave the first accurate description of the finer structure of the central capsule of the Cyrtida, and pointed out their character as true Monopylea, with porochora and podoconus, and the peculiar shape of its nucleus. He also published excellent figures of some interesting new species.

O. Bütschli, 1882, in his valuable paper entitled: "Beiträge zur Kentniss der Radiolarien-Skelette, insbesondere der der Cyrtida" (Zeitschr. für wiss. Zool., vol. xxxvi. p. 485) made an attempt at a natural classification of the Cyrtida, which he derived from the Spyroidea or Zygocyrtida. As already mentioned above, we cannot accept this essay as the foundation of a true natural system, since the affinities of the Cyrtellaria (and of the Nassellaria as a whole) are far more complicated and difficult than Bütschli supposed. His views were supported by accurate observations only on the structure of the fossil Cyrtoidea of Barbados; these, however, represent the minority only of the genera, and many interesting and important forms (mainly of true "Monocyrtida") remained unknown to Bütschli. A great part, however, of his observations are very useful, and his remarks on comparative morphology are very suggestive.

The Cyrtoidea may be divided into families and subfamilies according to three different principles, viz., (1) the number of joints into which the shell is divided by transverse strictures; (2) the number of radial apophyses which arise from the shell; (3) the shape of the basal mouth, which is either open or closed by a lattice-plate. At present every attempt of classification in this large group must be more or less artificial, since the affinities of the numerous smaller and larger groups are extremely complicated, and the ontogeny, the only sure guide in this phylogenetical labyrinth, is perfectly unknown. It seems therefore the most convenient to employ for our artificial classification, first, the number of shell-joints, second, the radial structure, and third, the shape of the mouth.

A. The number of joints into which the shell is divided by transverse constrictions, serves here for the distinction of four primary groups or suborders of the Cyrtoidea, viz., (1) Monocyrtida with one joint; (2) Dicyrtida with two joints; (3) Tricyrtida with three joints; and (4) Stichocyrtida with four or more joints. In my Prodromus (1881, p. 426) I divided the latter group into Tetracyrtida (with four joints), and Stichocyrtida (with five or more joints); but these two groups may be united, since the fourth and all the succeeding joints are of rather indifferent shape and of little morphological value. The three first joints, however, are usually very different and possess a high morphological importance, so that we distinguish the first joint as cephalis, the second as thorax, and the third as abdomen. The uppermost transverse constriction, which separates the two first joints, cephalis and thorax, is the collar stricture and is usually caused by an internal fenestrated septum, the cortinar septum. The second constriction, which separates the second and third joints (thorax and abdomen) is called the lumbar constriction. The following constrictions (in the Stichocyrtida) are indifferent and of little morphological interest, and require therefore no peculiar designation.

B. The radial structure, indicated by radial apophyses arising from the shell, offers three principal differences, according to which the whole group of Cyrtoidea may be divided into three large groups or sections, viz., (1) Pilocyrtida, or Cyrtoidea triradiata, with three radial apophyses; (2) Astrocyrtida, or Cyrtoidea multiradiata, with numerous radial apophyses (four to nine or more); and (3) Corocyrtida, or Cyrtoidea eradiata, without external radial apophyses. The majority of Cyrtoidea are Pilocyrtida, with three radial apophyses, which are probably homologous to the three primary feet of the Plectoidea and of Cortina (therefore "cortinar feet"). The Astrocyrtida, or the Cyrtoidea with a variable number of radial apophyses (at least four to six) may be derived from the Pilocyrtida by interpolation of secondary or interradial apophyses between the three primary or perradial apophyses. The Corocyrtida, however, or the Cyrtoidea without external radial apophyses, may have originated by reduction and loss of the latter, either from the Pilocyrtida or from the Astrocyrtida.

C. The shape of the basal mouth in the Cyrtoidea exhibits two essential differences only, viz., (1) the terminal mouth of the shell is a simple wide opening in the Stomocyrtida, or (2) the terminal mouth is closed by a lattice-plate, in the Clistocyrtida. As these two different cases occur in all the twelve families, which we have distinguished according to the differences in the number of joints and in the radial structure, we get altogether twenty-four subfamilies which are synoptically arranged in the following table:—

Synopsis of the four sections, twelve families and twenty-four subfamilies of CYRTOIDEA.	PILOCYRTIDA. Cyrtoidea triradiata. (Three radial apophyses.)		ASTROCYRTIDA. Cyrtoidea multiradiata. (Four to nine or more apophyses.)		COROCYRTIDA. Cyrtoidea eradiata. (No radial apophyses.)
Mouth of the shell.	Aperta.	Clausa.	Aperta.	Clausa.	Aperta.	Clausa.
MONOCYRTIDA. (Cyrtoidea monothalamia).	Tripocalpida.		Phænocalpida.		Cyrtocalpida.
MONOCYRTIDA. (Cyrtoidea monothalamia).	Archipilida.	Archiperida.	Archiphormida.	Archiphænida.	Archicorida.	Archicapsida.
DICYRTIDA. (Cyrtoidea dithalamia).	Tripocyrtida.		Anthocyrtida.		Sethocyrtida.
DICYRTIDA. (Cyrtoidea dithalamia).	Sethopilida.	Sethoperida.	Sethophormida.	Sethophænida.	Sethocorida.	Sethocapsida.
TRICYRTIDA. (Cyrtoidea trithalamia).	Podocyrtida.		Phormocyrtida.		Theocyrtida.
TRICYRTIDA. (Cyrtoidea trithalamia).	Theopilida.	Theoperida.	Theophormida.	Theophænida.	Theocorida.	Theocapsida.
STICHOCYRTIDA. (Cyrtoidea polythalamia).	Podocampida.		Phormocampida.		Lithocampida.
STICHOCYRTIDA. (Cyrtoidea polythalamia).	Stichopilida.	Stichoperida.	Stichophormida.	Stichophænida.	Stichocorida.	Stichocapsida.

Synopsis of the four sections, twelve families and twenty-four subfamilies of CYRTOIDEA.	PILOCYRTIDA. Cyrtoidea triradiata. (Three radial apophyses.)
Mouth of the shell.	Aperta.	Clausa.
MONOCYRTIDA. (Cyrtoidea monothalamia).	Tripocalpida.
MONOCYRTIDA. (Cyrtoidea monothalamia).	Archipilida.	Archiperida.
DICYRTIDA. (Cyrtoidea dithalamia).	Tripocyrtida.
DICYRTIDA. (Cyrtoidea dithalamia).	Sethopilida.	Sethoperida.
TRICYRTIDA. (Cyrtoidea trithalamia).	Podocyrtida.
TRICYRTIDA. (Cyrtoidea trithalamia).	Theopilida.	Theoperida.
STICHOCYRTIDA. (Cyrtoidea polythalamia).	Podocampida.
STICHOCYRTIDA. (Cyrtoidea polythalamia).	Stichopilida.	Stichoperida.
	ASTROCYRTIDA. Cyrtoidea multiradiata. (Four to nine or more apophyses.)
Mouth of the shell.	Aperta.	Clausa.
MONOCYRTIDA. (Cyrtoidea monothalamia).	Phænocalpida.
MONOCYRTIDA. (Cyrtoidea monothalamia).	Archiphormida.	Archiphænida.
DICYRTIDA. (Cyrtoidea dithalamia).	Anthocyrtida.
DICYRTIDA. (Cyrtoidea dithalamia).	Sethophormida.	Sethophænida.
TRICYRTIDA. (Cyrtoidea trithalamia).	Phormocyrtida.
TRICYRTIDA. (Cyrtoidea trithalamia).	Theophormida.	Theophænida.
STICHOCYRTIDA. (Cyrtoidea polythalamia).	Phormocampida.
STICHOCYRTIDA. (Cyrtoidea polythalamia).	Stichophormida.	Stichophænida.
	COROCYRTIDA. Cyrtoidea eradiata. (No radial apophyses.)
Mouth of the shell.	Aperta.	Clausa.
MONOCYRTIDA. (Cyrtoidea monothalamia).	Cyrtocalpida.
MONOCYRTIDA. (Cyrtoidea monothalamia).	Archicorida.	Archicapsida.
DICYRTIDA. (Cyrtoidea dithalamia).	Sethocyrtida.
DICYRTIDA. (Cyrtoidea dithalamia).	Sethocorida.	Sethocapsida.
TRICYRTIDA. (Cyrtoidea trithalamia).	Theocyrtida.
TRICYRTIDA. (Cyrtoidea trithalamia).	Theocorida.	Theocapsida.
STICHOCYRTIDA. (Cyrtoidea polythalamia).	Lithocampida.
STICHOCYRTIDA. (Cyrtoidea polythalamia).	Stichocorida.	Stichocapsida.

The cephalis, or the first shell-joint of the Cyrtoidea, is in the majority homologous with the cephalis of the Spyroidea, from which it differs in the reduction of the sagittal ring and the absence of the corresponding sagittal constriction; its cavity is therefore simple, not bilocular. Its homology with the original cephalis of the Spyroidea cannot be doubted, when its base exhibits the typical basal pores of the Semantida. But in many cases these are wanting, and in a great number of Cyrtoidea (mainly of Monocyrtida) there is more or less evidence that the original cephalis is lost, and that the real first joint is the thorax, the original second joint. At present it is quite impossible to distinguish between the former and the latter shells, and therefore in the following descriptions the first joint is always named cephalis and the second thorax. In future, when the affinities of the Cyrtoidea become better known, it will be necessary to distinguish the "Archicephalis," or true cephalis of all Spyroidea and of the majority of Cyrtoidea, from the "Pseudocephalis" or the false cephalis of the minority (e.g., of many Monocyrtida aperta, Archipilida, Archiphormida, Archicorida, &c.).

The thorax, or the second shell-joint of the Cyrtoidea, is in the majority homologous with the thorax of the Phormospyrida and Androspyrida, and therefore developed by apophyses, which arise from the base of the cephalis and become united by transverse branches forming a lattice-plate. Its size is generally in inverse proportion to that of the cephalis. The more the cephalis becomes reduced, the more the thorax is developed. Its form is very variable, usually three-sided pyramidal or prismatic in the triradiate, polyhedral in the multiradiate, and conical or cylindrical in the eradiate Cyrtoidea. Its terminal mouth is either a simple wide opening, or closed by a lattice-plate. In the majority of Cyrtoidea the thorax is separated from the cephalis not only by the external collar constriction, but also by the internal cortinar septum, a horizontal lattice-plate which exhibits the typical basal pores of the Semantida (usually two smaller jugular and two larger cardinal pores). But this septum is often reduced or perfectly lost, and then the external collar constriction alone indicates the separation of the cephalis and the thorax.

The abdomen, or the third shell-joint of the Cyrtoidea, absent in the Monocyrtida and Dicyrtida (as also in all Spyroidea), occurs constantly in all Tricyrtida and Stichocyrtida. It is a simple large chamber in the Tricyrtida, but forms an annulated body, composed of a variable number of successive joints, in the Stichocyrtida. The constrictions between these joints, and also the lumbar constriction, between abdomen and thorax, are usually provided with a lattice-girdle, projecting into the cavity of the shell, like a diaphragm. Usually this horizontal girdle bears only a single circle of pores, rarely two or more. In many Cyrtoidea it is replaced by a solid horizontal ring of silex, and often it is wanting. It originates by the insertion of the following shell-joint, which takes place not on the terminal mouth of the preceding joint, but somewhat above it.

The annular joints of the Stichocyrtida succeeding the third joint, and very variable in number, may be regarded either as a series of new postabdominal chambers, succeeding the true abdomen, or as secondary joints of the annulated abdomen itself. The latter view may be sustained by the fact that these joints are usually of an indifferent shape, and do not possess the characteristic features which we find in the first three joints, the abdomen, the thorax and the cephalis.

The lattice-work of the shell exhibits in the Cyrtoidea an extraordinary variety, similar to that of the Sphæroidea; it serves in the first place for the distinction of species. The three first joints of the shell are often distinguished by the different character of the lattice-work. The cephalis has usually very small and simple pores. The lattice-work of the thorax is often characterised by radial structures. The pores of the abdomen are usually very numerous and regular. The numerous joints in the annulated abdomen of the Stichocyrtida commonly exhibit little variety.

The closure of the mouth, effected by a convex or horizontal terminal lattice-plate, has a different signification in the Monocyrtida and in the jointed Cyrtoidea. In the Monocyrtida clausa this closing plate is the original cortinar plate or the basal plate of the cephalis. In the jointed Cyrtoidea, however, the lattice-plate which closes the terminal mouth of the thorax or of the abdomen (of the last annular joint in the Stichocyrtida), is produced by central union of the convergent edges, which grow centripetally from the margin of the mouth of the last joint towards its centre.

The radial apophyses arising from the shell of the Cyrtoidea may probably be always derived from that tripodal structure which is found in all Plectoidea, in Cortina and Cortiniscus among the Stephoidea, and in the majority of Spyroidea. Therefore the prototype of this radial structure would be Plagoniscus and Cortina, with four radial spines united in a common point, the cortinar centrum; an ascending apical horn and three descending basal feet. The odd posterior or caudal foot is usually similar in shape to the two paired anterior or pectoral feet, but may be distinguished from these latter by its relation to the apical horn. Very frequently an internal vertical free columella arises in the cephalis, or instead of it an ascending rib in the dorsal wall of the cephalis, which connects the base of the apical horn with the origin of the caudal foot. This is probably the remaining part of the sagittal ring. More rarely also a part of the ventral rod of the latter is preserved, or on the anterior pole of the basal rod of the cephalis an ascending procolumella arises which is inserted on the frontal face of the cephalis, and sometimes prolonged into a nasal horn (the rod, C, of Bütschli). These two odd horns, the posterior apical horn and the anterior nasal horn, are usually different and divergent. In some genera a variable number of accessory radial horns is developed on the convex face of the cephalis. In many hornless genera the free apical horn is lost, but not unfrequently the columella is preserved which connects the caudal foot with that point of the cephalis, in which formerly the apical horn was inserted.

The three primary radial beams, corresponding to the three basal feet of Plectaniscus and Cortina, exhibit in the Cyrtoidea the greatest variety in form and size, and chiefly in their relation to the shell, the latter serving mainly for the distinction of genera. Originally these three cortinar beams arise from the basal plate of the cephalis, the odd caudal foot appearing as a prolongation of the basal rod of that plate, and the paired pectoral feet as prolongations of its coracal rods (between the jugular and cardinal pores). The lattice-work of the thorax is developed usually between the three cortinar feet, more rarely inside or outside of them. Therefore the three beams appear commonly as three divergent ribs in the wall of the thorax, and continue over its basal mouth as three free terminal feet. With the increasing length of the shell and the number of its joints the three radial ribs are also prolonged, and their free distal ends may be prominent at very different points, either as three lateral wings or as three terminal feet. These are either solid spines or lattice-plates, sometimes more or less ramified.

The three radial apophyses are prevalent in the majority of the Cyrtoidea, which we call "Pilocyrtida" (or Cyrtoidea triradiata). Their number increases in the Astrocyrtida (or Cyrtoidea multiradiata). The most frequent cases of multiplication are here caused by the development of six or nine radial apophyses; these may be enclosed ribs, or lateral wings, or terminal feet. In the sexradial Cyrtoidea there are three secondary or interradial apophyses interpolated between the three primary or perradial; in the nine-radial Cyrtoidea, however, there are six adradial apophyses interpolated.

A third and last great group is formed by the Corocyrtida or Cyrtoidea eradiata. These exhibit no radial apophyses, neither enclosed ribs, nor free lateral wings, nor terminal feet. But in a great number of them internal traces of an original triradiate structure are visible, mainly in the cortinar septum between cephalis and thorax; this often exhibits three or four, and sometimes six cortinar or collar pores, of the same typical shape as in the triradial Spyroidea. Sometimes even an internal columella with three radial branches is preserved, as in Axocorys. It is therefore very probable that a great part of these Cyrtoidea eradiata (if not all) may be derived from triradiate or multiradiate ancestral forms, by reduction and loss of the radial apophyses. In another part of this group, mainly in the Monocyrtida eradiata (Cyrtocalpida) it is possible, or even probable, that their eradiate shell has originated independently from Nassellida, and that they have no true relation to radial Cyrtoidea.

The Central Capsule of the Cyrtoidea, first observed by J. Müller (1858), and more fully described in my Monograph (1862), was very accurately examined by Richard Hertwig (1879). His observations were confirmed by numerous new forms, which I was able to examine in well-preserved preparations of the Challenger. The central capsule, according to these, exhibits the same typical shape, which is characteristic of all Monopylea (with porochora and podoconus), and may be derived with the latter from the common ancestral forms, Cystidium and Nassella (= the skeletonless Nassellida). In the majority, however, of Cyrtoidea, the capsule develops on its basal face a number of depending lobes, as were also found in some Spyroidea (and probably also Botryodea). In this respect we may distinguish two main forms of the capsule in the Cyrtellaria, viz., the primary simple, not lobate form, and the secondary lobate form. The central capsule is originally always enclosed in the cephalis, and has there a simple, subspherical, ellipsoidal or ovate form. As soon as their growth increases, and the enclosing cephalis becomes too narrow, it sends out prolongations in the form of basal lobes, which depend from its base, and proceed through the pores of the basal lattice of the cephalis, or the cortinar pores. In the great majority of Cyrtoidea in which the capsule was observed, either three or four such lobes were seen (already described by J. Müller). Of course this number depends upon the number of cortinar pores, which is either three or four; therefore in the Cyrtoidea with three pores in the cortinar plate, we find three lobes of the central capsule (an odd posterior and two paired anterior); in the Cyrtoidea, however, with four pores in the cortinar plate (the majority) we find four lobes of the central capsule (two smaller anterior jugular and two larger posterior cardinal lobes). Usually each lobe is ovate or pear-shaped and encloses a large oil-globule, and often also an apophysis of the cell-nucleus.