The typical importance of the phæodium for all members of this legion is proved by the following five facts:—(1) its constant presence in all Phæodaria; (2) its constant excentric position in the oral half of the calymma; (3) its constant relation to the astropyle, the operculum and the proboscis of which is always surrounded by it; (4) its constant considerable size, its volume being usually greater than that of the central capsule, the aboral half of which is covered by it; (5) its constant colour and morphological as well as chemical composition. These five facts together demonstrate by their absolute constancy the high morphological and physiological importance of the phæodium for this peculiar subclass of Radiolaria, although its true nature is difficult to make out, and its main function is not yet sufficiently known.
The first remarks that have been made on the phæodium were published in 1862 in my Monograph, where I noticed the peculiar dark brown extracapsular pigment body and its excentric position covering only one-half of the central capsule, in Aulacantha (p. 263, Taf. ii. figs. 1, 2), in Thalassoplancta (p. 262, Taf. iii. fig. 10), and in Cœlodendrum (p. 361, Taf. xxxii. fig. 1). Its general presence and peculiar composition were first recognised by Dr. John Murray, who had, during the Challenger Expedition, the first opportunity of examining many big living Phæodaria brought up from great depths. He gives in his first Report (1876, loc. cit., p. 536), the following important notice:—"The sarcode of all these deep-sea Rhizopods has many large black-brown pigment-cells. Small bioplasts are scattered through the sarcode. These collect into capsular-like clumps when the animal is at rest, and are quickly coloured by carmine." In 1877 Dr. John Murray sent me the wonderful collection of Phæodaria brought home by the Challenger, and I had now the best opportunity for examining the phæodium in hundreds of well-preserved specimens. Supported by these extensive observations, I gave, in 1879, in a preliminary paper,[[273]] a fuller description of the phæodium, and of the phæodella (or the peculiar dark pigment-granules composing it). At the same time R. Hertwig published his observations on the big living Tripylea examined by him at Messina, and pointed out particularly that the constant presence, composition, and arrangement of this excentric extracapsular pigment-body was most characteristic, and sufficient in itself to distinguish this group from all the other Radiolaria (1879, loc. cit., p. 99).
The most striking peculiarity of the phæodium, and the most important on account of its absolute constancy, is its excentric position, covering only the oral hemisphere of the central capsule, and wholly or partly wanting on the aboral hemisphere. This constant topographical relation to the capsule never fails, and may be always observed at the first glance, when the body is seen from the side (the main axis of the capsule being perpendicular to the axis of the eye of the observer), (compare Pl. [101], figs. 6, 10; Pl. [102], fig. 1; Pl. [103], fig. 1, &c.). The phæodium envelops, therefore, the oral half of the capsule completely, and especially the astropyle in its centre; hence, the radiate operculum and the proboscis arising from the latter cannot be seen usually before the former is removed. The general form of the entire phæodium, in consequence of this excentric position, is concavo-convex (or crescentic in longitudinal section), its concave face embracing the oral or anterior face of the capsule, and its convex face being turned to the surface of the calymma.
The topographical relation of the phæodium to the surrounding shell is also very characteristic in the suborder Phæogromia, or in those Phæodaria which possess a peculiar shell-mouth placed on the oral pole of its main axis. Here the capsule is always placed in the aboral half of the shell-cavity, the phæodium in its oral half, separating the astropyle from the mouth of the shell, both of which lie in the main axis; as in the Challengerida (Pl. [99]), Tuscarorida (Pl. [100]), Castanellida (Pl. [113]), Circoporida (Pl. [115]), and Medusettida (Pl. [118]-[120]). In this suborder (the Phæogromia), the volume of the phæodium may be, on an average, about as great as that of the central capsule, whilst in the majority of other Phæodaria it is much greater than the latter.
A similar topographical relation between the phæodium and the enclosing shell, as in the Phæogromia, also exists in the suborder Phæoconchia, or in those Phæodaria, which are distinguished from all the others by the possession of a bivalved shell (Pl. [121]-[128]). The two valves of this curious shell, which resembles that of the Brachiopoda, are dorsal and ventral, and the tripylean central capsule is always so placed between them that its two lateral parapylæ (right and left) lie in the frontal plane of the shell, where a large frontal fissure opens between the opposed margins of the two hemispherical or cap-shaped valves. The phæodium is also placed here on the oral half of the capsule and surrounds its astropyle; but it exhibits some differences in the three families of Phæoconchia.
The Concharida, the bivalved shell of which is simple, and without tubular apophyses (Pl. [123]-[125]), possess a relatively small central capsule, which usually fills up only the third or fourth part of the shell-cavity. This is the aboral or posterior part, on the apex of which both valves are united by a ligament in some Concharida (Pl. [123], figs. 8, 9). The oral or anterior part of the shell-cavity (usually two-thirds or more) is filled up by the phæodium, and this is usually bifid, being divided by a frontal constriction into two wings or lobes; the dorsal wing is hidden in the upper valve of the shell, the ventral wing in the lower valve; both wings are usually united only by a small central bridge, and this bridge of the phæodium is pierced in its centre by the proboscis of the astropyle (Pl. [124], figs. 6, 10; Pl. [123], figs. 8, 9).
The Cœlodendrida have a different shape (Pl. [121]). Their bivalved shell is relatively small and tiny, and bears on the two poles of the sagittal axis two conical apophyses or galeæ, from each of which three or four very large, dichotomously branched tubes arise. The central capsule fills up the cavity of the bivalved shell almost entirely, and the voluminous dark phæodium envelops both to such an extent that the shell and the enclosed capsule are often hidden in it completely. Therefore I arrived in my first description of Cœlodendrum (1862, loc. cit.) at the erroneous conclusion that the capsule lies outside, not inside the shell. The first accurate figure and description of its structure was given in 1879, by Hertwig (loc. cit., p. 99, Taf. x. fig. 3). The central capsule (v) is here separated from the bivalved shell (m) only by a very small distance, and the oral part of both is hidden in the phæodium. I find, however, in the majority of the numerous preparations of the Challenger collection, the volume of the phæodium much greater, and it often envelops the entire shell.
The Cœlographida, finally, have a phæodium of the most remarkable shape, since in their bivalved shell a peculiar reserve store or magazine of phæodella, which we call the "phæocapsa" is developed for it (Pl. [126]-[128], g.t.m.). The bivalved shell has in these most perfect Phæodaria a structure similar to that in the Cœlodendrida; but they differ from the latter in the stronger development, and greater differentiation of the two apical galeæ, and the large hollow tubes arising from them. These two helmet-shaped cupolæ, the galeæ (g), which arise from the two valves on the poles of the sagittal axis, are in the Cœlographida usually larger than the valves themselves, and are not closed, as in the Cœlodendrida, but open by a tubular apophysis at their base, the nasal tube or rhinocanna (t). The apex of the galea is connected with the open mouth of the rhinocanna by a single or double frenulum (b). The two nasal tubes or rhinocannæ (a dorsal and a ventral) lie in the sagittal plane of the body and run from the base of each galea along the anterior convexity of the valve to its oral margin. Here is placed the proboscis of the astropyle, between the two opposed mouths of the rhinocannæ (Pl. [128], fig. 2). The phæodium is usually hidden entirely in the two phæocapsæ, which are composed of the two galeæ (g), and the two rhinocannæ arising from them (Pl. [127], figs. 4-9). A part of the phæodella is usually thrown out by the mouth of the latter (m).
The characteristic colour of the phæodium exhibits numerous different tints between green, brown, and black. It seems to be in the majority blackish-brown or greenish-brown, very often olive, more rarely almost quite green or red-brown. Usually the colour is so dark, intense and opaque, that the parts enclosed by the phæodium, mainly the oral hemisphere of the central capsule and the astropyle, are completely hidden in it. The chemical composition of the phæodium demands further accurate researches; unfortunately I have not been able to make out its true nature, since numerous different experiments furnished no certain general results.
The phæodella, or the pigment-corpuscles, which compose the phæodium, aggregated in hundreds, and in the bigger species in thousands, are usually spherical, sometimes somewhat ellipsoidal, at other times spheroidal or lenticular; but usually numerous smaller, irregular, roundish particles are intermingled between the larger and more regular corpuscles, and often the main mass forms a very fine black powder. The size of the phæodella is very variable, not only in the different species, but also in one and the same individual. The larger phæodella have a diameter of 0.01 to 0.02, the smaller of 0.004 to 0.008 mm.; but there also occur very big forms of 0.04 to 0.05 mm., or even more, and very small ones of 0.001 mm. or less. Usually the phæodium appears as an aggregate of numerous larger and smaller phæodella, which are very different in size as well as in the intensity and tint of their colour, and are irregularly crowded in a black, powder-like substance.