Concharida, Haeckel, 1879, Sitzungsb. med.-nat. Gesellsch. Jena, Dec. 12, p. 6.
Definition.—Phæodaria with a bivalved lattice-shell, which is spherical or lenticular, and composed of two equal or unequal boat-shaped valves, a dorsal and a ventral. The valves bear neither an apical latticed cupola or galea, nor hollow radial tubes. The central capsule is placed in the aboral half of the shell-cavity, and so enclosed between both valves, that its three openings lie in the open frontal fissure between them (the astropyle on the oral pole of the main axis, the two parapylæ on both sides of its aboral pole, at right and left).
The family Concharida and the two following closely allied families, the Cœlodendrida and Cœlographida, compose together the most remarkable and interesting suborder of Phæoconchia (or "Phæodaria bivalva"), differing from all the other Radiolaria in the possession of a bivalved lattice-shell, composed of two separate valves, like the shell of a Brachiopod. The central capsule is so enclosed between the two fenestrated valves that its three openings lie in the horizontal open (frontal) fissure between them, the astropyle or main-opening on the oral pole of the main axis; the two secondary openings or parapylæ on the two sides of its aboral pole, at right and left. The plane in which the three openings lie is therefore the frontal plane, dividing the entire body into a dorsal and a ventral half. The two valves, accordingly, must be considered as dorsal and ventral valves (as in the Brachiopoda), and the symmetrical halves of each valve as right and left. These halves may be always easily distinguished, since the oral pole of each valve is constantly different from the aboral pole. The voluminous phæodium always lies in the oral half, and the central capsule in the aboral half of the shell-cavity, whilst the calymma encloses the whole shell.
The Concharida differ from the other two families of bivalved Phæodaria in the absence of the apical galeas, and the branched hollow tubes arising from them. Each of these two cupolas, which are at the opposite poles of the sagittal axis (one cupola on the apex of each valve), is in the Cœlographida connected by a simple or double frenulum with a peculiar rhinocanna, or an open nasal tube directed towards the mouth; whilst the cupolas of the Cœlodendrida possess neither a rhinocanna nor a frenulum. The three families of Phæoconchia may therefore represent a phylogenetical series, the common root of which are the Concharida. From these are developed the Cœlodendrida by development of an apical cupola or galea on each valve, and of hollow radial tubes arising from it; whilst the Cœlographida are developed from the latter by production of a rhinocanna on the base of each cupola, and of one or two frenula connecting the former with the latter.
All the Concharida described in the following pages (seven genera and thirty species), are perfectly new to science, and not a single form of this interesting family was known before the explorations of the Challenger. Some species (mainly of the genera Conchidium and Conchopsis) are by no means rare, and are found in great numbers at some stations of the tropical seas (in the Pacific as well as in the Atlantic). All described species are closely allied, agree in the majority of characters, and are easy to distinguish from all the other Radiolaria. Some few forms of Concharida, however, form a direct passage to the Cœlodendrida.
Regarding the probable origin of the Concharida (and therefore also of all other Phæoconchia derived from the latter), two different hypotheses are possible. They have either been derived directly from the skeletonless Phæodina, by development of a bivalved lattice-shell; or they may be derived from Phæodaria with a simple spherical lattice-shell (Castanellida), by the halving of this latter, or its splitting into two hemispherical valves; the former hypothesis is more probable than the latter.
The two valves of the lattice-shell (Pls. [123]-[126]) must in the Concharida (as in all other Phæoconchia) be distinguished as dorsal and ventral, and may therefore be compared with the two valves of the Brachiopoda, not with those of the Lamellibranchia. This important morphological distinction is expressed by the constant position of the central capsule within the shell-cavity. The capsule always exhibits the character of the "Tripylea" and has three tubular openings, placed in the frontal or lateral plane of the unicellular body. In the same plane lies the open frontal fissure between the two valves, and the three openings are so disposed in it that the large anterior main-opening (or the astropyle) is placed on the oral pole of the main axis, whilst the two accessory small lateral openings or parapylæ are placed on both sides of the aboral pole, at the right and left. Therefore in a dorsal or ventral view all three openings are visible (Pl. [123], figs. 1, 8a); in the usual lateral view, however, from the right or left side, only two openings are visible, the astropyle on the anterior, and one parapyle (right or left) near the posterior pole of the main axis (Pl. [123], figs. 8, 9; Pl. [124], figs. 6, 10). The posterior view (from the aboral pole) shows the two parapylæ, at right and left; in the anterior view (from the oral pole) the astropyle may be visible, but usually it is completely hidden in the dark voluminous phæodium. This latter envelops sometimes nearly the whole capsule as an opaque conglomeration of green or brown phæodella (Pl. [123], figs. 8, 9); but usually the phæodium fills up the anterior (oral) half of the shell-cavity, whilst the capsule occupies the posterior (aboral) half (Pl. [124], figs. 6, 10).
The dorsal shell-valve is in almost all Phæodaria smaller or somewhat different in shape from the ventral valve, and this difference is often very striking (Pl. [124], figs. 3-16); but in a few species both valves are so similar, that I could not discover any certain difference. This equality of the two valves occurs mainly in those Concharida which pass over into the Cœlodendrida; in these latter as well as in the Cœlographida, both valves are usually equal in size and form. Whilst the main axis (or the longitudinal axis of the body) in the two latter families of Phæoconchia seems to be normally vertical (in the living and freely floating body), in the living Concharida it is probably horizontal, so that the larger and heavier ventral valve lies below the smaller and lighter dorsal valve.
The geometrical fundamental form of the body is therefore in the Concharida dipleural or bilaterally symmetrical, and we distinguish in it the same three dimensive axes, as in all other dipleural forms. On the anterior or oral pole of the main axis (or longitudinal axis) lies the mouth of the shell, and behind it the phæodium; on the opposite posterior or aboral pole lies the hinge of the shell (comparable to the shell-hinge of the Brachiopoda) and in front of it the central capsule. The sagittal (or dorso-ventral) axis, determining the height of the shell, has on its dorsal (or upper) pole the apex or highest point of the dorsal valve, on its ventral (or lower) pole the apex or lowest point of the ventral valve. The two poles of the frontal (lateral or transverse) axis are equal and are determined by the two parapylæ of the capsule, and the corresponding points of the shell-fissure between both valves. Usually the main-axis is the longest, the frontal axis the shortest, and between both the sagittal axis.
In regard to the three dimensive planes which are determined by these three axes, perpendicular to one another, they are rarely of nearly equal size (as in some subspherical species), usually the sagittal plane (separating the right and left halves of the body) is the largest; the cinctural or equatorial plane (separating oral and aboral halves) is the smallest, and the frontal or lateral plane (separating dorsal and ventral halves) is intermediate in size. The relation of the three perimeters of these three planes corresponds to that proportion; the sagittal perimeter (in which the keel of the compressed valves lies) is the largest; the cinctural or equatorial perimeter (separating the anterior phæodium and the posterior central capsule) is the smallest, and the frontal or lateral perimeter (in which the fissure between the valves lies) is intermediate in size.