Dimensions.—Diameter of the spongy shell 0.24, of its inner cavity 0.12, of the central cube 0.02; length of the spines 0.4, breadth 0.01.

Habitat.—North Pacific, Station 254, surface.

3. Octodendron pinetum, n. sp.

Radial spines eight, three-sided prismatic, three to four times as long as the diameter of the inner shell-cavity, with three spirally contorted edges. From each spine arise at equal distances (equal to the half radius of the cavity) fifteen to twenty verticils of branches which increase in size from the base of the spine. Each verticil is composed of six forked branches (two from each spine-edge); the inferior are richly branched, and form by their connection the loose network of the spongy cortical shell, the surface of which is covered with numerous bent threads. Each spine bears a resemblance to a pine tree.

Dimensions.—Diameter of the spongy shell 0.3, of its inner cavity 0.15, of the central cube 0.02; length of the spines 0.5 to 0.6, breadth 0.01.

Habitat.—Central Pacific, Station 272, surface.

4. Octodendron araucaria, n. sp.

Radial spines eight, three-sided prismatic, with three spirally contorted edges, eight to ten times as long as the diameter of the inner shell-cavity (the free distal portion twice to three times as long). From each spine arise at equal distances thirty to thirty-five verticils of branches, which decrease in size towards the distal end. Each verticil is composed of six forked branches (two from each spine-edge); the inferior are richly ramified and form by their connection the loose spongy framework; the distal branches bear on the free end elegant spathillæ (as in the following species). Surface covered with innumerable straight bristles, as long as the cavity radius, and ending with a spathilla. Diameter of the central capsule about equal to the radius of the spongy sphere, its membrane is double-edged.

Dimensions.—Diameter of the spongy shell 1.4 mm., of its inner cavity 0.14, of the central cube 0.02, of the central capsule 0.5 to 0.6; length of the spines 0.9 to 1.2, breadth 0.02.

Habitat.—South Pacific, Station 288, surface.

5. Octodendron spathillatum, n. sp. (Pl. 18, figs. 2, 4).

Radial spines eight, three-sided prismatic, with contorted edges, five to eight times as long as the diameter of the shell-cavity (the free part twice as long). From each spine arise ten to twelve verticils of lateral branches; each verticil composed of three forked branches. The proximal larger branches ramify richly, and form by their connection the loose spongy framework of the spherical shell; the distal smaller branches are simple or bifurcated, and the ramules are provided with an elegant spathillum at the end (fig. 4). Entire surface of the spongy sphere covered with innumerable bristle-shaped radial spines (half as long as the cavity radius), zig-zag, bent, with beards, and with a spathillum at the end.

Dimensions.—Diameter of the spongy sphere 1 mm., of its central cavity 0.15, of the central cube 0.02; length of the spines 1 to 1.2 mm., breadth 0.008.

Habitat.—Central Pacific, Station 271, surface.

Subgenus 2. Octodendronium, Haeckel.

Definition.—Radial spines thirty-two or more, eight primary (arising from the eight corners of the central cube) and twenty-four or more secondary (between them).

6. Octodendron verticillatum, n. sp.

Radial spines thirty-two, with three denticulate straight edges, six to eight times as long as the diameter of the shell-cavity; the distal parts half free. Eight primary spines arise from the eight corners of the central cube, twenty-four secondary from the edges of these (a verticil of every three from each primary spine). Distal free parts of all thirty-two spines equal. Each spine with eight to ten verticils of forked lateral branches, without spathillæ. Surface of the spongy sphere covered with short simple bristles.

Dimensions.—Diameter of the spongy shell 0.3, of its cavity 0.12, of the central cube 0.02; length of the spines 0.7 to 1 mm., breadth 0.02.

Habitat.—South Pacific, Station 291, surface.

7. Octodendron contortum, n. sp.

Radial spines thirty-two, disposed in a similar manner to those of the former species (eight primary and twenty-four secondary); also the spongy shell of the same shape. The difference arises in the form of the spines, the three edges of which are much broader and spirally contorted around the axis; and thus the corresponding branches of the verticil do not lie in the same meridian-plane, but alternate one with another.

Dimensions.—Diameter of the spongy shell 0.4, of its cavity 0.1, of the central cube 0.02; length of the spines 0.8 to 1.2, breadth 0.04.

Habitat.—Central Pacific, Station 274, surface.

8. Octodendron arboretum, n. sp.

Radial spines sixty to ninety, three-sided prismatic, with spirally contorted and denticulate edges, six to eight times as long as the diameter of the inner shell-cavity. Eight primary arise from the eight corners of the central cube, twenty-four others from their three edges (as in the two former species); the remaining thirty to sixty spines seem to arise between the former and immediately from the dense spongy framework, which is twice as thick as the diameter of the inner shell-cavity. The numerous verticils of the free distal part are of equal shape in all the spines, composed of three forked branches in the terminal, and of more ramified branches in the inferior parts. Entire surface of the spongy shell covered with simple radial bristles, without spathillæ.

Dimensions.—Diameter of the spongy shell 0.75, of its central cavity 0.15, of the central cube 0.02; length of the spines 0.9 to 1.2, breadth 0.02.

Habitat.—Tropical Pacific, Station 225, surface.

Genus 120. Spongosphæra,^[159] Ehrenberg, 1847, Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, p. 54.

Definition.—Astrosphærida with two concentric latticed medullary shells, connected by radial beams; the outer is immediately enveloped by the spongy framework, and bears numerous radial spines.

The genus Spongosphæra (in the definition here restricted) differs from Spongopila in the double medullary shell, which in the latter is simple; it exhibits to the latter the same relation that Spongodictyon among the Liosphærida bears to Spongoplegma. The outer medullary shell of Spongosphæra is immediately enveloped by the spongy wicker-work, which everywhere pierces the wall of the central capsule.

1. Spongosphæra streptacantha, Haeckel.

Spongy shell of polyhedric, irregular outline, the framework being prolonged sheath-like into the eight to twelve radial spines, which are quite irregularly distributed, very large, three-sided prismatic, with three serrated, spirally contorted edges; their length is twice to four times as great as the diameter of the spongy body; they arise with thinner bases from the outer medullary shell, which is three times as broad as the inner, both having roundish pores, twice to four times as broad as the bars; surface without radial by-spines.

Dimensions.—Diameter of the spongy shell 0.2 to 0.6, outer medullary shell 0.04 to 0.06, inner 0.012 to 0.016.

Habitat.—Cosmopolitan; common in all warmer seas, surface.

2. Spongosphæra polyacantha, J. Müller.

Spongy shell spherical, with ten to twenty large radial spines, which arise with thinner bases from the medullary shell, and are prominent on the surface at different lengths; they are three-sided prismatic (not four-sided), with three leaf-shaped straight edges. Whilst the spongy framework is much looser than in the preceding species, both medullary shells have nearly the same shape.

Dimensions.—Diameter of the spongy shell 0.2 to 0.5, outer medullary shell 0.04 to 0.06, inner 0.012 to 0.016.

Habitat.—Mediterranean (Nice); Atlantic, Station 353, surface.

Subgenus Spongosphæromma, Haeckel.

Definition.—Radial spines on the surface of the spongy shell of two different kinds; large piercing main spines and small superficial by-spines.

3. Spongosphæra helioides, Haeckel.

Spongy shell spherical, with numerous curved, bristle-shaped by-spines on the surface, as long as the radius. Main spines ten to twenty, irregularly disposed, prismatic, with three dentated, spirally contorted edges, broader toward the distal end. Outer medullary shell three times as broad as the inner, with polygonal meshes (of the same size as those in the spongy framework) and fine bars.

Dimensions.—Diameter of the spongy shell 0.2, outer medullary shell 0.02, inner 0.007.

Habitat.—Mediterranean (Messina).

4. Spongosphæra quadricuspis, n. sp.

Spongy shell spherical, with numerous curved, bristle-shaped by-spines on the surface, half as long as the radius. Main spines twenty to thirty, irregularly disposed, prismatic, with three dentated, spirally contorted edges, and at the distal end four strong pyramidal divergent teeth (three as terminations of the edges, the fourth as end of the spine axis). Outer medullary shell twice as broad as the inner, both having circular, regular pores, twice to three times as broad as the bars.

Dimensions.—Diameter of the spongy shell 0.3, outer medullary shell 0.04, inner 0.013.

Habitat.—Central Pacific, Station 272, surface.

Genus 121. Rhizosphæra,^[160] Haeckel, 1860, Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, p. 840.

Definition.—Astrosphærida with two concentric latticed medullary shells, connected by radial beams; from the outer arise numerous radial spines, which at equal distances are connected by a latticed spherical cortical shell, surrounded by a spongy framework.

The genus Rhizosphæra exhibits the same relation to Spongosphæra that Rhizoplegma bears to Spongopila; but in the latter the latticed medullary shell is simple, in the two former double. The wall of the central capsule is pierced only by the radial spines connecting the medullary and the cortical shells.

1. Rhizosphæra trigonacantha, Haeckel.

Central cavity of the spongy cortical shell twice as broad as the diameter of the outer medullary shell. Bars of all three shells of the same breadth as the thirty to fifty (or more) radial spines, which are three-sided prismatic; their outer free distal end only as long as the diameter of the inner medullary shell.

Dimensions.—Diameter of the spongy shell 0.25, of its inner cavity 0.2, outer medullary shell 0.1, inner 0.05.

Habitat.—Cosmopolitan; Mediterranean, Atlantic, Indian, Pacific, surface from many Stations.

2. Rhizosphæra serrata, n. sp. (Pl. 18, figs. 5-7).

Central cavity of the spongy shell five times as broad as the diameter of the outer medullary shell. Bars of all three shells of the same breadth as the forty to sixty (or more) thin radial beams between them. These are three-sided prismatic, with denticulate edges, scarcely half as broad as their outer prolongations, which are half as long as the shell radius, and possess three spirally contorted serrated edges. (The figured specimen is a young one; in the older specimens the spongy framework of the cortical shell is much more developed.)

Dimensions.—Diameter of the spongy shell 0.3, of its central cavity 0.22, outer medullary shell 0.06, inner 0.02.

Habitat.—Central Pacific, Stations 270 to 274, surface.

3. Rhizosphæra leptomita, Haeckel.

Central cavity of the spongy cortical shell twice as broad as the diameter of the outer medullary shell; bars of both very thin, only one-third as broad as the bars of the inner medullary shell. Radial spines thirty to fifty (or more), curved, three-sided prismatic; inside the spongy shell as thin as their bars, outside three times as broad.

Dimensions.—Diameter of the spongy shell 0.27, of its inner cavity 0.2, outer medullary shell 0.1, inner 0.05.

Habitat.—Mediterranean (Messina); Atlantic, Stations 348 to 354, surface.

Suborder PRUNOIDEA, Haeckel, 1883 (Pls. 13-17, 39, 40).

Definition.—Spumellaria with an ellipsoidal or cylindrical central capsule, prolonged into one axis (sometimes articulate by annular transverse strictures); with an ellipsoidal or cylindrical, fenestrated siliceous shell (often articulate by annular strictures), invariably prolonged into one axis. Fundamental form monaxon, usually with the poles of the prolonged dimensive main axis equal.

The suborder Prunoidea comprises those Spumellaria in which the fenestrated spherical shell appears prolonged into one axis. The geometric fundamental form of the shell, which in the Sphæroidea was a sphere, in this case therefore becomes an ellipsoid, and whilst in the former all axes originally have the same value (Homaxonia), here one main axis is constantly larger than all other axes (Monaxonia). Usually both poles of this main axis are equal (Haplopola); but in some genera both poles become different (Diplopola).

In the Sphæroidea all planes going through the centre of the shell are circular, whereas in the Prunoidea only those planes are circular which are perpendicular to the main axis; all other planes going through the centre are elliptical; the largest of these are the meridian planes, in which is situated the main axis. Commonly all meridian planes are equal, as no transverse axes (or cross axes) are differentiated.

In my Monograph (1862) only very few forms of Prunoidea, such as Didymocyrtis and Spongurus, are described, and the greater part of them are distributed under different genera of Sphæroidea, such as Haliomma and Actinomma. In my Prodromus (1881) I separated them as the family Zygartida (p. 462). But it seems now much more convenient to restrict this term to a particular family and to give a wider extension to the whole suborder under the name Prunoidea (called after the characteristic ellipsoidal form of a plum, or Prunus, with its stone).

The suborder Prunoidea comprises seven different families, of which the Ellipsida constitutes the simplest and the probable common ancestral group. In all Ellipsida the fenestrated shell is simple, and never composed of concentric or twin shells. In their primitive genus Cenellipsis, the whole shell is geometrically nothing more than a simple ellipsoid (Pl. 39, figs. 1, 2). By development of radial spines it passes into Ellipsidium, by development of spongy framework into Spongellipsis. In the greater part of this family large spines are developed on both poles of the main axis of the shell, but sometimes instead of these solid spines, two opposite hollow fenestrated tubes are developed (Pipettella, Pl. 39, fig. 6).

The second family of Prunoidea, the Druppulida, is much richer in different forms than the simple Ellipsida. In this case the ellipsoidal shell is composed of two or three (rarely more) concentric shells. Constantly one or two of these fenestrated shells are enclosed in the central capsule, and may therefore be called "medullary shells"; and one or two (rarely more) lie outside the central capsule, "cortical shells." The inner medullary shells (one or two) are either spherical or ellipsoidal; the outer cortical shells (one or two, rarely more) are always ellipsoidal. All concentric shells are connected by radial beams. In the simplest form of the subfamily, Druppula (Pl. 39, fig. 3), one medullary shell is connected with one cortical shell. By duplication of the medullary shell arises Prunulum (Pl. 39, fig. 4), and by duplication or multiplication of the cortical shell Cromyodruppa (Pl. 15, figs. 1-4) is formed. In by far the greater portion of this subfamily large spines are developed on both poles of the main axis of the cortical shell (Pls. 16, 17), but sometimes also instead of these solid spines two opposite hollow fenestrated tubes are developed (Pipetta, Pl. 39, figs. 7, 8).

A third family of Prunoidea, closely allied to the two preceding families, is the Spongurida, in which we include all Prunoidea with an ellipsoidal or cylindrical, unjointed shell, in which the lattice-work of the cortical shell is transformed into an irregular, siliceous framework. In the simplest form, Spongellipsis, the simple lattice-shell of Cenellipsis is substituted by an external spongy envelope. In other cases (Spongurus and allied genera) the whole cavity of this external spongy shell is distended with a fine spongy framework. The subfamily of Spongodruppida is distinguished by the possession of a simple or double latticed medullary shell; this lies in the midst of the central capsule, and is connected by radial beams (perforating its membrane) with the enveloping spongy cortical shell. The surface of the latter may bear either radial spines, or two opposite strong polar spines, at the poles of the main axis (Pl. 17, fig. 12).

Closely allied to the Ellipsida and Druppulida are two other families of the Prunoidea, the Artiscida and Cyphinida, which differ from the former by a circular constriction in the equatorial plane of the ellipsoidal shell; and in this way assume a characteristic twin form, like a figure of eight. In the Artiscida the shell is simple (as in the Ellipsida), whereas in the Cyphinida it is composed of two or more concentric shells (as in the Druppulida). The simplest form of the Artiscida is Artiscus (Pl. 39, fig. 9), differing from Cenellipsis in the ring-shaped, equatorial constriction. In other Artiscida polar appendages are developed on both poles of the main axis, either in the form of solid, strong spines (Stylartus), or hollow fenestrated tubes (Cannartus, Pl. 39, fig. 10).

The family Cyphinida differs from the Druppulida in the equatorial constriction of the shell, and from the Artiscida in the presence of two or more concentric shells. One or two of these concentric fenestrated shells are enclosed in the central capsule (and therefore may be called "medullary shells"); the others (one or two, rarely more) lie outside of the central capsule (therefore "cortical shells"). The internal "medullary shells" are always spherical or somewhat lenticular, compressed from both sides; the external "cortical shells" have constantly a ring-like constriction in the equatorial plane, and "twin-shells" are therefore like a figure of eight. The simplest form of this subfamily is Cyphanta, composed of a simple medullary shell and a simple cortical shell, the two being connected in the equatorial plane by radial beams. In Cyphonium (Pl. 39, fig. 12) the medullary shell is doubled, and in Cypassis (Pl. 39, fig. 13) the cortical shell likewise. On both poles of the main axis strong spines are often developed (Cyphinus, Pl. 39, fig. 14), or hollow fenestrated tubes (Cannartidium, Pl. 39, figs. 16-19).

The equatorial constriction of the ellipsoidal shell, which characterises the Artiscida and Cyphinida, is repeated or multiplied in the two following families, in the Panartida and Zygartida; in the former we find three ring-like strictures, in the latter five or more (lying in parallel transverse planes); therefore the fenestrated shell is composed in the one instance of four chambers, in the other of six or more; all the chambers form a single series and have a common main axis. All constrictions lie in planes parallel to the equatorial plane of the original ellipsoid; in the centre of the latter constantly lies a double "medullary shell," composed of two concentric, either spherical or lenticular, compressed shells. In all Panartida we call the two inner chambers (on both sides of the equatorial constriction) "proximal chambers," the two outer chambers (on the poles of the main axis) "distal chambers." The four-chambered cortical shell of the Panartida is either simple (in Panartus, Pl. 40, figs. 1-4) or double, with an external mantle (as in Peripanartus, Pl. 40, figs. 5-7). The simplest form of the subfamily is Panartus (loc. cit.). In this case also on both poles of the main axis may be developed solid spines, or hollow fenestrated tubes (Panarium, Pl. 40, fig. 9).

The seventh and last family of the Prunoidea, the Zygartida is most nearly allied to the Panartida, and appears as a further developmental step from that family. Whilst in the Panartida the cortical shell is constantly four-chambered, with three parallel ring-like constrictions, in the Zygartida it is always prolonged and composed of six or more chambers, separated by five or more ring-shaped constrictions, in the middle of which is the equatorial stricture. In the centre of the latter (as also in the Panartida) always lies the double medullary shell, composed of two concentric, spherical, or lenticular shells. The number of the chambers of the cortical shells is commonly six or eight (with five to seven ring strictures), but it often mounts to ten and sometimes to twenty (with nineteen strictures), as in some species of Zygartus (Pl. 40, fig. 13). All the chambers lie in one series, one behind another, with a common main axis. The cortical shell is usually simple (in Ommatocampe, Pl. 40, fig. 10), sometimes double (in Desmocampe, Pl. 40, fig. 12), rarely triple (in Zygocampe, Pl. 40, fig. 13). In all three cases hollow fenestrated tubes may be developed on the poles of the main axis.

The morphological references and the phylogenetic affinities of all Prunoidea are so complex, that they seem to represent a quite natural group; all forms of it may be derived from the common ancestral form Cenellipsis. But a far more difficult question is the manner in which its pedigree may be constructed. The oldest family is probably the simplest, namely, Ellipsida. From this the Druppulida may be derived by production of medullary shells, the Artiscida by equatorial constriction. The Cyphinida can be produced either from the Druppulida by equatorial constriction or from the Artiscida by development of medullary shells. The Panartida appear as further developmental steps of the Cyphinida, by duplication of the chamber number; and the Zygartida as further productions of the Panartida, by increasing the number of the chambers.

The seven subfamilies of the Prunoidea can be arranged in two sections according to the presence or absence of medullary shells. The Ellipsida, Spongellipsida, and Artiscida possess a simple cortical shell, without a medullary shell; they represent the section Cenoprunida. All other families possess medullary shells, and so represent the section Coccoprunida.

Another character, which can be employed in the arrangement of the seven subfamilies in some larger groups, is the presence or absence of ring-like constrictions, by which the cortical shell is divided into chambers. I. The Monoprunida comprise all forms without any constriction, of which the Ellipsida are without a medullary shell, the Druppulida with one or two medullary shells, and the Spongurida with a spongy cortical shell. II. The Dyoprunida contain all forms with a cortical twin shell, or with two chambers separated by one equatorial constriction, of which the Artiscida are without a medullary shell and the Cyphinida have one or two medullary shells. III. The Polyprunida comprise all forms with several (three or more) constrictions, which separate four or more chambers, of which the Panartida have three constrictions and four chambers, and the Zygartida five or more constrictions and six or more chambers.

The Central Capsule of the Prunoidea is originally ellipsoidal (monaxial), and preserves this form in the greater part of the genera. In some groups, where the axis of the ellipsoid is much prolonged, it passes over to the cylindrical form (with hemispherical vaultings on both poles), as in Spongurus and Spongocore, in many Panartida and Zygartida. Very often the ellipsoidal or cylindrical capsule gets annular transverse constrictions, corresponding to those of the enveloping cortical shell (one single, equatorial stricture in the Artiscida and Cyphinida, three strictures in the Panartida, five or more in the Zygartida). In the Cenoprunida (Ellipsida and Artiscida, also in Spongellipsis) the central capsule lies freely in the cavity of the cortical shell, separated from its inner surface by the jelly-envelope; in all other groups it contains a part of the skeleton, the medullary shell and the beams which connect it with the enveloping cortical shell.

Synopsis of the Families of Prunoidea.

Family XI. Ellipsida, Haeckel, 1882 (Pls. 13, 14, 39).

Definition.—Prunoidea with simple ellipsoidal shell, without equatorial stricture (without enclosed medullary shell); network a simple lattice lamella, not spongy. Central capsule ellipsoidal or cylindrical, without annular equatorial constriction.

The family Ellipsida comprises the simplest forms of Prunoidea, and probably represents the ancestral forms of this whole suborder. The fenestrated shell, which encloses the ellipsoidal central capsule, is a perfectly simple "cortical shell" of the same form, without enclosed "medullary shell." Its form is commonly a regular monaxial ellipsoid; sometimes a little modified by unequal growth of the two poles of the main axis. Two opposite large spines are often developed at these poles, or it may be that instead of these, two hollow fenestrated tubes are present.

The ellipsoidal fenestrated shell exhibits in the regular Ellipsida all the characters of a geometric ellipsoid; one main axis surpasses in length all other possible axes. All sections going through this main axis are "meridian sections," with elliptical periphery; all sections perpendicular to the main axis are "transverse sections," with circular periphery. The largest of these is the equatorial section, which divides the main axis into halves. The diameter of this equatorial plane is the "minor axis" of the ellipsoid.

The proportion of the two axes of the ellipsoidal shell, of the major vertical or main axis and the minor horizontal or equatorial axis, is commonly between 6 : 5 and 3 : 2. In the former case it approaches the spherical shell, from which it is derived; in the latter case it becomes almost fusiform or cylindrical. The network of silex, constituting the shell, is constantly a simple latticed lamella, never composed of concentric shells (as in Druppulida) or spongy (as in Spongurida). The network is often very regular and elegant, in other cases irregular.

The simplest genus among the Ellipsida, and probably the common ancestral form of the whole subfamily, is the genus Cenellipsis, possessing a simple ellipsoidal shell without any appendages. It is derived from Cenosphæra (the simplest spherical shell) by the prolongation of one axis. Cenellipsis passes over into Ellipsidium by the production of radial spines on the surface (corresponding to Heliosphæra). Axellipsis is a peculiar genus differing from Cenellipsis in an axial rod, which corresponds to the minor or equatorial axis. In all other genera of the subfamily both poles of the main axis are distinguished by peculiar polar prolongations, either hollow fenestrated tubes (as in Pipettella) or strong solid spines. Both polar spines are of equal size and similar form in Ellipsoxiphus, unequal in Ellipsostylus. From the latter is derived Lithapium, by reduction and loss of one spine (so that only one remains); Lithomespilus, by production of a bunch of several spines at one pole. In the three latter genera both poles of the main axis are unequal, in all others equal.

The central capsule of the Ellipsida is in all cases ellipsoidal, and occupies the largest part of the shell, being separated from its inner surface by a thinner or thicker jelly-mantle.

Synopsis of the Genera of Ellipsida.

Genus 122. Cenellipsis,^[161] n. gen.

Definition.—Ellipsida with simple ellipsoidal shell, without radial spines and without polar tubes.

The genus Cenellipsis is the simplest and most primitive form, not only among the Ellipsida, but also among the Prunoidea, and it may therefore be regarded as the common ancestral form of the whole family. It corresponds to Cenosphæra among the Sphæroidea, to Cenodiscus among the Discoidea, to Cenolarcus among the Larcoidea. Probably it is derived from Cenosphæra by prolongation of one axis.

Subgenus 1. Cenellipsium, Haeckel.

Definition.—Network of the shell regular, with meshes of equal size and similar form.