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 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 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 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]).
The skeletons of the Radiolaria, in addition to the general relations which have been discussed above, present numerous and important special differences in the various larger and smaller groups. These are indicated in detail in the descriptions of the legions, orders, and families in the systematic portion of this Report.
BIOGENETICAL SECTION.
A SKETCH OF OUR KNOWLEDGE OF THE DEVELOPMENT OF THE RADIOLARIA IN THE YEAR 1884.