Besides these three general and never failing marks of the Phæodaria, the majority of this legion (but by no means all) possess the three following peculiarities; (1) two parapylæ or accessory openings of the central capsule, placed laterally (at the right and left) on the aboral pole of the main axis (wanting in the Challengerida, Medusettida, Castanellida, and perhaps in some other families); (2) a characteristic skeleton which is always extracapsular, wanting only in the Phæodinida, incomplete in the Cannorrhaphida and Aulacanthida, but perfectly developed and of very various shapes in the twelve other families; usually this silicated skeleton is composed of hollow tubules, which are filled up by jelly (Pansolenia); but in some families it is composed of ordinary solid network, not different from that of the other Radiolaria, e.g., especially in the Castanellida and Sagosphærida; (3) an extraordinary size of the body, as well of the central capsule and its nucleus, as of the extracapsular skeleton; the majority of Phæodaria have a diameter of 1 to 2 mm., and are therefore from ten to twenty times as large as the majority of the other Radiolaria; in some gigantic forms the diameter of the unicellular body reaches 20 or even 30 mm.
The Central Capsule of the Phæodaria is the most important part of their body and preserves in all families of this legion the same essential structure, and nearly the same form, in striking contrast to the extraordinary variety and complication of the skeleton. As already mentioned, it differs from the central capsule of all the other Radiolaria in two most important peculiarities; firstly, the double membrane of the spheroidal capsule, and secondly, the singular structure of its constant main-opening, the astropyle. A third constant character is the considerable size of the enclosed nucleus, the diameter of which usually equals about half of that of the capsule itself.
The diameter of the capsule is, in the majority of Phæodaria, 0.1 to 0.2 mm., often also 0.3 to 0.4, rarely more than 0.5, or less than 0.05 mm.
The position of the central capsule is somewhat different in the four orders into which we have divided the Phæodaria. It lies in the centre of the spherical, concentric calymma in all the Phæocystina, or in those Phæodaria which possess no complete lattice-shell—Phæodinida, Cannorrhaphida, and Aulacanthida (Pls. [101]-[105]). The Phæosphæria (comprising the Orosphærida, Sagosphærida, Aulosphærida, and Cannosphærida, Pls. [106]-[112]) all possess a very big, usually spherical lattice-shell, and here the central capsule is much smaller than the latter and lies in its centre. Another position is constantly occupied by the central capsule in all Phæogromia (Pls. [99], [100], [113]-[120]; the families Challengerida, Medusettida, Castanellida, Circoporida, Tuscarorida), which have a peculiar mouth on the lattice-shell, placed on the oral pole of its main axis; the central capsule lies in the opposite aboral half of the shell cavity. The Phæoconchia finally (Pls. [121]-[128], the families Concharida, Cœlodendrida, and Cœlographida) all possess a bivalved shell, and the capsule is here enclosed between the two valves of the shell.
The form of the central capsule is constantly spheroidal, slightly depressed in the direction of the main axis, and therefore comparable to the spheroidal form of our globe. The depression is generally very slight, so that the proportion of the minor vertical main axis to the major horizontal or equatorial axis is about 4 : 5 or 5 : 6, often even 8 : 9 or less; but sometimes the proportion becomes 3 : 4, or even 2 : 3, so that the capsule becomes nearly lenticular; very rarely its form becomes almost perfectly spherical (Pl. [101], fig. 1). The main axis stands always vertically, and is distinctly marked by the peculiar shape of the astropyle, placed on its oral pole. Usually this pole, in the living and freely floating Phæodaria seems to be the lower pole, directed downwards (as also in the Nassellaria); but in some families it seems to be inversely directed upwards, as in the Challengerida and Tuscarorida, (Pls. [99], [100]).
The two membranes of the central capsule possess in all Phæodaria a very different shape, and were in all preparations which I could accurately examine (some hundreds belonging to all families) separated by a clear, rather wide interval, filled up either by a colourless fluid or by a structureless jelly (Pl. [103], fig. 1; Pl. [123], figs. 8, 9, &c.). They are in direct connection only at the openings. In the living Phæodaria, however, their distance is very small, or they are in immediate contact without any interval (Pl. [101], fig. 10; Pl. [102], fig. 1). According to the observations of Hertwig, the two membranes are always in close contact, and without interspace, in the living Phæodaria; and the space between them is an artificial product due to the influence of the preserving fluid or of certain chemical agents. In every case it is very easy to separate both membranes completely, except at the openings, where they are in direct connection. We distinguish both membranes shortly as ectocapsa and endocapsa.
The ectocapsa, or the outer membrane of the central capsule, is rather firm and durable, double-contoured, elastic and difficult to destroy. Its physical and chemical qualities seem to approach those of chitin. It becomes, however, stained red by carmine, and yellow by nitric acid. Usually it appears structureless and refracts the light strongly. In a few cases, however, it exhibits, when examined by strong lenses, a fine punctation; and in some Aulacanthida (especially in some big forms of Aulographis and Aulospathis) the entire ectocapsa was densely covered with peculiar curved, or S-shaped dark corpuscles (Pl. [114], fig. 13). They were all of the same length, about 0.01, and seemed to lie on its inner face.
The endocapsa, or the inner membrane of the central capsule, is much thinner than the outer, with which it is in immediate connection only at the openings. It encloses the entire contents of the capsule, and becomes very distinct, as soon as the latter are dissolved by chemical agents, or stained by carmine. In the majority of well-preserved preparations it is irregularly plicated, and resembles a thin, but firm, crumpled paper. Isolated pieces of the endocapsa are completely structureless, but exhibit also a considerable resistance, in spite of their minute thickness.
The openings of the central capsule exhibit in the Phæodaria a greater variety than their discoverer, R. Hertwig, supposed. The majority of the legion, certainly, possess the three openings described by him, and are therefore true Tripylea. Some families, however, have only one opening, the astropyle, which is generally present (Challengerida, Medusettida, Castanellida, and Phæocolla among the Phæodinida, Pl. [101], fig. 1). In some other families there is a variable number of accessory openings or parapylæ, one, three, or more, e.g., especially in the Circoporida and Tuscarorida. The former may be called Astropylea, the latter Sporopylea. Thus only a single opening to the central capsule is constant in all Phæodaria without exception, and that is the astropyle, or the large main-opening with its peculiar structure.
The astropyle, or the single constant main-opening of the central capsule, is distinguished by a very remarkable structure, and is sufficient of itself to separate the Phæodaria from all the other Radiolaria, and from the other Rhizopoda in general. It is always placed on the oral pole of the main axis, forming here a peculiar cap-shaped or flatly conical elevation, the centre of which is prolonged into a short, cylindrical tubule; we call the latter shortly a proboscis, and the former an operculum. To understand better the different forms which these important parts assume in the different families, compare Pl. [101], figs. 1-6; Pl. [102], fig. 1; Pl. [103], fig. 1; Pl. [104], figs. 1-3; Pl. [111], fig. 2; Pl. [123], figs. 1-9; Pl. [127], figs. 4-6; Pl. [128], fig. 2, &c. Compare also the first very accurate figures which R. Hertwig has given in 1879 (loc. cit.) in his Taf. x.