Acanthodesmida, R. Hertwig, 1879, Organismus der Radiol., p. 68.
Spyrida vel Spyroidea, Haeckel, 1881, Prodromus, p. 440.
Definition.—Nassellaria with a complete lattice-shell, exhibiting constantly a bilocular cephalis with a sagittal constriction.
The suborder Spyroidea, and the two following closely allied suborders, Botryodea and Cyrtoidea, represent together that large group of Radiolaria which I first described in my Monograph (1862, pp. 272, 280) as the family Cyrtida, but afterwards as a separate order (or sublegion) under the name Cyrtellaria (1883, Jena. Sitzungsber., Feb. 16, p. 18). This group comprises all those Monopylea or Nassellaria which possess a complete lattice-shell, whilst the preceding Plectellaria never develop a perfect fenestrated shell enveloping the central capsule. The Spyroidea differ from the other Cyrtellaria (Botryodea and Cyrtoidea) in the bilocular shape of the cephalis, which is bisected by the sagittal ring and a corresponding longitudinal constriction into two symmetrical halves.
The Spyroidea appear in the first system of Polycystina of Ehrenberg (1847, loc. cit., pp. 53, 54) as the fourth of his seven families, under the name Spyridina, with the following definition:—"Testæ nucleo destitutæ (associatæ et coalitæ); cellulæ binæ clathratæ, nucis forma amplæ, strictura longitudinali levius discretæ." Ehrenberg united them with his "Polycystina composita or Spumellaria" and separated them from the closely allied "Polycystina solitaria or Nassellaria." He distinguished among them five genera, two of which have no external appendages (Dictyospyris and Pleurospyris), whilst the other three possess spiny or lamellar appendages (Ceratospyris, Cladospyris, and Petalospyris). These five genera and the accompanying definitions were also repeated in the same terms in the last system of Ehrenberg (1875, loc. cit., p. 157). In my Monograph (1862, pp. 280, 291) the Spyroidea are enumerated as a subfamily of the Cyrtida, under the name Zygocyrtida, with the following definition:—"Lattice-shell divided by an annular sagittal longitudinal constriction into two neighbouring equal joints or chambers." I adopted at that time the above five genera of Ehrenberg, only eight species of which were known.
Richard Hertwig gave in 1879 the first accurate description of the central capsule of the Spyroidea, and demonstrated that they possess the same characteristic structure of the capsule, with a porochora and a podoconus, that is common to all Monopylea. He separated the Zygocyrtida from the other Cyrtida, and united them with the Stephoidea (Lithocircus) in the family Acanthodesmida. In my Prodromus (1881, p. 449) I adopted that division and put the "Spyrida or Spyroidea" as a separate family between the Cyrtoidea and Stephoidea. I there gave the following definition:—"Monopylaria gemina, testa silicea clathrata gemina, cameris binis juxta compositis, annulo verticali sagittali contiguis, capsula centrali a testa inclusa. Promorpha dipleura vel bilaterali, dextro et sinistro antimero symmetricis, plano sagittali annuloque separatis. Polum superiorem axis verticalis vel principalis plerumque spina apicalis (vel occipitalis) occupat, polum inferiorem ostium clathratum (poris tribus aut quatuor aut pluribus) et spina caudalis (posterior). Ad dextram et sinistram duæ spinæ laterales distant. Ab hac forma tripoda (Monocyrtida triradiata acuta cum annulo mediano) diversæ Spyridum formæ derivandæ sunt." I there divided the Spyrida into seven subfamilies and forty-eight genera. Bütschli in 1882 published accurate researches on the Zygocyrtida in his Beiträge zur Kenntniss der Radiolarien-Skelete, insbesondere der der Cyrtida (Zeitschr. f. wiss. Zool., vol. xxxvi. p. 501, Taf. xxxii.). He gave a very exact anatomical description of the skeleton of several fossil Spyroidea of Barbados, and pointed out their close affinity to the Acanthodesmida on the one hand, and to the Cyrtida on the other. Regarding their phylogenetical relation, he supposed that the Zygocyrtida have descended from the Acanthodesmida (our Stephoidea) and are the ancestors of all other Cyrtida. The best part of this valuable paper of Bütschli is his exact description of the sagittal ring and its apophyses, producing the basal plate; and the general conception thus derived of the essential parts of the skeleton in all Monopylea, bearing a sagittal ring. But his general conclusions were partly erroneous, since the fossil Spyroidea and Cyrtoidea of Barbados, upon which they were founded, contain only the minority of genera of the large group; and many important and typical forms remained unknown to him.
The numerous new genera discovered in the rich collection of the Challenger, and mainly in the wonderful Radiolarian ooze of the Central Pacific (Stations 263 to 274) throw a new light upon the complicated affinities of the whole group, and manifest a far greater richness in developmental variations and admirable morphological productions than Bütschli could suppose. The following description, however, must remain very incomplete, since the exact and thorough study of all individual forms, in the same manner as Bütschli had done in the case of a small number of species, would require several years. I can therefore only give the basis of further researches.
In the majority of Spyroidea the three essential elements of the Nassellarian skeleton are easily recognisable, viz.:—A, the vertical sagittal ring of the Stephoidea; B, the basal tripod of the Plectoidea, and C, the latticed cephalis of the Cyrtellaria. These three elements are constantly so combined that the primary sagittal ring bisects the cavity of the lattice-shell and produces an external, longitudinal, more or less distinct constriction, separating the lateral inflated halves of the bilocular cephalis; while the three divergent feet of the basal tripod descend from the cortinar or basal plate of the cephalis. The number of the feet is often augmented; they are rarely wanting (probably reduced). The sagittal ring is constantly present, though its relation to the shell-wall exhibits many modifications. The bilocular form of the fenestrated cephalis, with its sagittal constriction, is characteristic of all true Spyroidea, and separates them from the Botryodea (with multilobate cephalis) on the one hand, and from the Cyrtoidea (with simple cephalis) on the other. There are, however, some intermediate forms between these three suborders of Cyrtellaria, which show a direct transition to one another. Almost constantly the length of the three dimensive axes in the cephalis of the Spyroidea is so different that the lateral or frontal axis is the longest, the sagittal or dorso-ventral axis the shortest, and the principal or longitudinal axis intermediate between them. As in all dipleuric or bilaterally symmetrical forms, the two poles of the frontal axis are equal, whilst the two poles of the sagittal and of the principal axis are more or less different. In only a few forms of Spyroidea these differences are difficult to make out; usually the dorsal side is distinctly different from the ventral, and the apical side from the basal. A perfect knowledge of the dipleuric shell requires therefore accurate observation from all six sides.
The suborder of Spyroidea is here divided into four different families. In the first and original family, the Zygospyrida, the shell is represented by the bilocular cephalis only. The second family, Tholospyrida, is distinguished by a fenestrated cupola or galea, a hemispherical or conical dome, arising from the upper or coryphal face of the cephalis. In the third family, Phormospyrida, a thorax is developed, or a fenestrated basal chamber, arising from the lower or basal face of the cephalis. The fourth family, Androspyrida, combines the characters of the second and the third family, a galea arising from the upper and a thorax from the lower face of the cephalis.
The bilocular cephalis, as the most important and constant part of the skeleton in all Spyroidea, requires a further general consideration. Its lattice-work, very variable in the numerous species, is usually more or less different on the six sides of the nut-shaped shell. Its most important part is the horizontal basal plate or lower face, and the sagittal ring arising from it in the vertical median plane. This part of the shell corresponds exactly to the skeleton of the Semantida, and exhibits the same modifications as in that family; it is therefore probable that the majority of the Spyroidea (if not the whole group) have been derived directly from the Semantida by further development of network from the fenestrated ring of Clathrocircus. Whilst in this latter Semantid the two lateral sides of the central capsule remain naked, in the Spyroidea they become enclosed and covered by lattice-work, which arises on the right and left from the ring. In some forms, however, as in the Ceratospyris acuminata of Hertwig, and in my Perispyrida, the frontal ring also of the Coronida and the mitral ring of the Tympanida appear as the initial parts of a complete latticed cephalis.