The section Discoidea comprises those Spumellaria in which the fenestrated shell is more or less discoidal or lenticular, flattened or compressed in the direction of one axis. The geometric fundamental form of the latticed shell, which in the Sphæroidea is a sphere, here becomes a flat disk, like a medal, or a biconvex lens, sometimes also a biconcave lens. The Discoidea can be derived from the Sphæroidea by shortening of one axis. This shortened vertical axis is the main axis of the disk; both its poles are constantly equal. Perpendicular to this axis is the equatorial plane of the disk by which it becomes divided into equal halves. In the simplest forms of Discoidea all axes of this horizontal equatorial plane (all "equatorial axes" or "cross axes") are equal; in the most of the genera and species these cross axes are different, so that rays of stronger growth ("perradii") alternate with rays of weaker growth ("interradii"). The number of these cross axes distinguishable is commonly two to four, rarely more. In the direction of these are developed either radial marginal spines or spongy arms.
The order Discoidea was founded in my Monograph (1862, p. 476) as the family "Discida" (Radiolaria with flat discoidal or biconvex lenticular shell), comprising the "Calodictya and Lithocyclidina" of Ehrenberg and a great part of his "Haliommatina." As three different subfamilies of that family I separated the Coccodiscida (with five genera), the Trematodiscida (with seven genera), and the Discospirida (with two genera; Monogr. d. Radiol., p. 485). A fourth group of Discoidea was constituted by the Spongodiscida (with eight genera, including the Spongocyclida), which at that time I united with the Spongurida, because of their spongy structure (loc. cit., p. 452).
As the number of fossil Discoidea found in the Tertiary rocks of Barbados and of the Mediterranean shores (Sicily and Greece) is comparatively very large, we find even in the first system of Polycystina of Ehrenberg (1847), not less than twelve genera distinguished, viz., six Calodictya, two Haliommatina, and four Lithocyclidina (Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, 1847, p. 53). The whole number of Radiolarian genera distinguished in that first system was forty-four. The diagnoses of them given by Ehrenberg were as usual very insufficient. The characters of the three families given by him were the following:—Calodictya—"Testarum intus spongiosarum et nucleo destitutarum orbes; Haliommatina—Testæ subglobosæ nucleus radiatus; Lithocyclidina—Testarum disci in media parte nucleati margine celluloso." In the latest work of Ehrenberg (1875, p. 157) the same system was repeated, but some new genera added; and thirty-eight different species, appertaining to the Discoidea, were figured in the same work (Abhandl. d. k. Akad. d. Wiss. Berlin, 1875, Tafs. xx.-xxx.).
Richard Hertwig, 1879, in his excellent work, Der Organismus der Radiolarien (pp. 57-68), gave a detailed description of the skeleton of some Discoidea, and arrived at the conclusion that this whole family had a spirally constructed skeleton, and should therefore be derived from the Lithelida. But this conclusion is certainly erroneous, and in my opinion the whole explanation of that spiral structure, and of its signification in the development of Discoidea, is the weakest part of that otherwise very important work.
In my Prodromus (1881, p. 456) I gave a provisional system of the Discida or Discoidea from the immense quantity of new material collected by the Challenger, and could distinguish not less than eighty-four genera. This number is from subsequent research only augmented by seven, so that in the following pages ninety-one genera with five hundred and one species are described. In the Prodromus I had disposed them in four different families, which number is now increased to six. These six families can be again disposed in two main groups or sections, the Phacodiscaria and the Cyclodiscaria, each section with three families.
The Phacodiscaria are characterised by the possession of a typical "phacoid shell," and contain the three families Cenodiscida, Phacodiscida, and Coccodiscida. On the other hand, the Cyclodiscaria are distinguished by the absence of such a "phacoid shell," and contain the three families Porodiscida, Pylodiscida, and Spongodiscida. Both sections exhibit an analogous development.
The Cenodiscida (Pl. [48], figs. 1-3) open the series of the Discoidea as their simplest forms; a discoidal or lenticular simple lattice-shell encloses a central capsule of the same form, and is separated from it by the calymma or jelly-veil. The common ancestral form of this family is Cenodiscus, without radial marginal spines; it can be derived from Cenosphæra in the most simple way, by flattening in one axis. If on the equatorial margin of the lens a peculiar solid girdle be developed, we obtain Zonodiscus; in all other genera of the Cenodiscida radial spines are developed on the margin. As the simple lenticular cortical shell of Cenodiscus, in which the central capsule is enclosed, is most characteristic not only of this family, but also of the two following families, we call it the phacoid shell (that is, a lenticular extracapsular or cortical lattice-shell).
The Phacodiscida (Pls. [31]-[35]), the second family, have the same extracapsular "phacoid shell" as the Cenodiscida, but differ from these by the possession of one or two intracapsular concentric medullary shells, which are connected with the former by radial beams, perforating the lenticular central capsule. The radial beams are commonly numerous, and arranged in two opposite bunches around the shortened main axis. But often also besides these occur other longer radial beams, situated in the equatorial plane; the number of these is commonly four, and they form a regular rectangular cross, lying opposite in pairs in two equatorial diameters, perpendicular one to another. In the simplest genera of this family (the Sethodiscida) the equatorial margin of the phacoid shell is simple or surrounded by a solid smooth girdle; in all other genera are developed on the margin solid radial spines lying in the equatorial plane, either regularly disposed in a somewhat constant number (two to eight, Heliosestrida), or irregularly disposed, in a larger and more variable number (ten to twenty or more, Heliodiscida).
The Coccodiscida (Pls. [36]-[38]) form a third family of the Discoidea, directly associated with the Phacodiscida; both have the same characteristic extracapsular "phacoid shell," which is connected by radial beams with a simple or double, intracapsular medullary shell. But whilst in the foregoing family the equatorial margin of the phacoid shell is simple or only armed with radial spines, in the Coccodiscida it is surrounded by peculiar concentric chambered girdles, or rings, which resemble those of the following family, the Porodiscida. Each of these "chambered girdles" is composed of a circular ring in the equatorial plane, a variable number of radial beams dividing it into incomplete chambers, and two porous cover-plates or "sieve-plates," covering the upper and lower face of the disk. These sieve-plates may be regarded as incomplete lenticular cortical shells, which are only developed in the peripheral part of the disk, whilst their central part is represented by the only complete cortical shell, the "phacoid shell." The number of these concentric chamber-girdles amounts to from one to ten or more. The margin of the disk is either simple (Lithocyclida) or armed with radial spines (Stylocyclida), or provided with two to five chambered radial arms (Astracturida); the structure of the arms is the same as that of the girdles.
The Porodiscida (Pls. [41]-[47]), the largest family of all Discoidea, begins the series of Cyclodiscaria, or those Discoidea in which there is no "phacoid shell," but a small simple central chamber surrounded by a number of small latticed chambers of nearly the same size and form. In the Porodiscida these chambers are arranged in complete circular concentric rings or spiral convolutions; in the small family of Pylodiscida the central chamber is surrounded by three radial arm-chambers separated by three open spaces; in the third family of Cyclodiscaria, the Spongodiscida, all the chambers are arranged more or less irregularly, and the whole disk becomes spongy; also the surface of the disk is spongy, whilst in both former families it is covered by two regular even porous plates or "sieve-plates." Probably all Cyclodiscaria can be derived phylogenetically from Archidiscus, a very small and simple lenticular disk, which is composed of a small spherical latticed central chamber and of a single concentric chambered ring or girdle; the margin of this ring is connected with the central chamber by a variable number of radial beams. This Archidiscus can be derived either from Saturnalis (Pl. [13], fig. 16.) by the development of lattice-work between the equatorial ring and both polar faces of the concentric central chamber, or from Sethodiscus (Pl. [33], figs. 1-3) by flattening of the lenticular shell, so that the enclosed inner medullary shell (the central chamber) meets the outer phacoid shell at both poles.