Order 2. Carpoidea.—Theca compressed in the oro-anal plane and a bilateral symmetry thus induced, affecting the food-grooves and, usually, the thecal plates and stem. Food-grooves in part epithecal and may be continued on one or two exothecal processes. No pores or folds in the stereom. Families: Anomalocystidae, Dendrocystidae. These correspond to Jaekel’s Carpoidea Heterostelea; he also includes, as Eustelea, our Comarocystidae and Malocystidae.
Order 3. Rhombifera.—Radial symmetry affects the food-grooves and, in the more advanced families, the thecal plates; probably also the nerves and ambulacral vessels, but not the gonads. The food-grooves are exothecal, i.e. are stretched out from the theca on jointed skeletal processes (brachioles). These either are close to the mouth or are removed from it upon a series of ambulacral or sub-ambulacral plates not derived immediately from thecal plates, or are separated from the oral centre by hypothecal passages passing beneath terminal plates. The stereom and stroma become arranged in folds and strands at right angles to the sutures of the thecal plates; in higher forms the stereom-folds are in part specialized as pectini-rhombs. Families: Echinosphaeridae; Comarocystidae; Macrocystellidae; Tiaracrinidae; Malocystidae; Glyptocystidae, with sub-famm. Echinoencrininae, Callocystinae, Glyptocystinae, of which examples are Cheirocrinus (fig. 10) and Cystoblastus from which Jaekel deduces the blastoids; Caryocrinidae.
Order 4. Aporita.—Pentamerous symmetry affects the food-grooves and thecal plates; probably also the nerves and ambulacral vessels, but not the gonads. Food-grooves exothecal and circumoral. The stereom shows no trace of canals, folds, rhombs or diplopores. Family: Cryptocrinidae.
Order 5. Diploporita.—Radial symmetry affects the food-grooves, and by degrees the thecal plates connected therewith, but not the interradial thecal plates; probably also the nerves and ambulacral vessels, but not the gonads. The food-grooves are epithecal, i.e. are extended over the thecal plates themselves without intermediate flooring; they are also prolonged on exothecal brachioles, which line the epithecal grooves. The stereom of the thecal plates may be thrown into folds, but the mesostroma does not so much tend to lie in strands traversing the sutures, nor are pectini-rhombs or pore-rhombs developed; diplopores are always present in the mesostereom, but often restricted to definite tracts or plates, especially in higher forms. Families: Sphaeronidae; Glyptosphaeridae, e.g. Fungocystis (fig. 9); Protocrinidae; Mesocystidae; Gomphocystidae.
The Protocrinidae lead up to Proteroblastus, in which the theca is ovoid, sometimes prolonged into a stem, the plates differentiated into (a) smooth, irregular, depressed interambulacrals, (b) transversely elongate brachioliferous adambulacrals, to which the diplopores, which lie at right angles to the main food-groove, are confined. This leads almost without a break to the Protoblastoidea.
Class II. Blastoidea.—Pelmatozoa in which five (by atrophy four) epithecal ciliated grooves, lying on a lancet-shaped plate (? always), radiate from a central peristome between five interradial deltoid plates, and are edged by alternating side-plates bearing brachioles, to which side-branches pass from the grooves. Grooves and peristome protected by small plates, which can open over the grooves. The generative organs and coelom probably did not send extensions along the rays into the brachioles; but apparently nerves from the aboral centre, after passing through the thecal plates, met in a circumoral ring, from which branches passed into the plate under each main food-groove, and thence supplied the brachioles. The thecal plates, however irregular in some species, always show defined basals and a distinct plate (“radial”) at the end of each ambulacrum; they are in all cases so far affected by pentamerous symmetry that their sutures never cross the ambulacra. All Palaeozoic.
Division A. Protoblastoidea.—Blastoidea without interambulacral groups of hydrospire-folds hanging into the thecal cavity. Families: Asteroblastidae, Blastoidocrinidae. The former might be placed with Diploporita, were it not for a greater intimacy of correlation between ambulacral and thecal structures than is found in Cystidea as here defined. They form a link between the Protocrinidae and—
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Fig. 13.—A Eublastoid, Pentremites. |
Division B. Eublastoidea.—Blastoidea in which the thecal plates have assumed a definite number and position in 3 circlets, as follows: 3 basals, 2 large and 1 small; 5 radials, often fork-shaped, forming a closed circlet; 5 deltoids, interradial in position, supported on the shoulders or processes of the radials, and often surrounding the peristome with their oral ends. The stereom of the radials and deltoids on each side of the ambulacra is thrown into folds, running across the radio-deltoid suture, and hanging down into the thecal cavity as respiratory organs (hydrospires).
These are the forms to which the name Blastoidea is usually restricted. They have been divided into Regulares and Irregulares, but it seems possible to group them according to three series or lines of descent, thus:—
Series a. Codonoblastida.—Families: Codasteridae, Pentremitidae (fig. 13).
Series b. Troostoblastida.—Families: Troostocrinidae, Eleutherocrinidae.
Series c. Granatoblastida.—Families: Nucleocrinidae, Orbitremitidae, Pentephyllidae, Zygocrinidae.
Class III. Crinoidea.—Pelmatozoa in which epithecal extensions of the food-grooves, ambulacrals, superficial oral nervous system, blood-vascular and water-vascular systems, coelom and genital system are continued exothecally upon jointed outgrowths of the abactinal thecal plates (brachia), carrying with them extensions of the abactinal nerve-system. The number of these processes is primitively and normally five, but may become less by atrophy. The brachia rise from a corresponding number of thecal plates, “radials (RR).” Below these is always a circlet, or traces of a circlet, of plates alternating with the radials, i.e. interradial, and called “basals (BB).” Through all modifications, which are numerous and vastly divergent, these elements persist. A circlet of radially situate infrabasals (IBB) may also be present. Below BB or IBB there follows a stem, which, however, may be atrophied or totally lost (see fig. 1).
The classification here adopted is that of F. A. Bather (1899), which departs from that of Wachsmuth and Springer mainly in the separation of forms with infrabasals or traces thereof from those in which basals only are present. These two series also differ from each other in the relations of the abactinal nerve-system. O. Jaekel (1894) has divided the crinoids into the orders Cladocrinoidea and Pentacrinoidea, the former being the Camerata of Wachsmuth and Springer (Monocyclica Camerata, Adunata and Dicyclica Camerata of the present classification), and the latter comprising all the rest, in which the arms are either free or only loosely incorporated in the dorsal cup. In minor points there is fair agreement between the American, German and British authors. The families are extinct, except when the contrary is stated.
Sub-class I. Monocyclica.—Crinoidea in which the base consists of BB only, the aboral prolongations of the chambered organ being interradial; new columnals are introduced at the extreme proximal end of the stem.
Order 1. Monocyclica Inadunata.—Monocyclica in which the dorsal cup is confined to the patina and occasional intercalated anals; such ambulacrals or interambulacrals as enter the tegmen remain supra-tegminal and not rigidly united. Families: Hybocrinidae, Stephanocrinidae, Heterocrinidae, Calceocrinidae, Pisocrinidae, Zophocrinidae, Haplocrinidae, Allagecrinidae, Symbathocrinidae, Belemnocrinidae, Plicatocrinidae, Hyocrinidae (recent), Saccocomidae.
Order 2. Adunata.—Monocyclica with dorsal cup primitively confined to the patina and an occasional single anal; tegmen solid; portions of the proximal brachials and their ambulacrals tend to be rigidly incorporated in the theca. Arms fork once to thrice, and bear pinnules on each or on every other brachial. BB fused to 3, 2 or 1. (Eucladocrinus and Acrocrinidae offer peculiar exceptions to this diagnosis.) Families: Platycrinidae, Hexacrinidae, Acrocrinidae.
Order 3. Monocyclica Camerata.—Monocyclica in which the first, and often the succeeding, orders of brachials are incorporated by interbrachials in the dorsal cup, while the corresponding ambulacrals are either incorporated in, or pressed below, the tegmen by interambulacrals; all thecal plates united by suture, somewhat loose in the earliest forms, but speedily becoming close, and producing a rigid theca; mouth and tegminal food-grooves closed; arms pinnulate.
Sub-order i. Melocrinoidea.—RR in contact all round; first brachial usually quadrangular. Families: Glyptocrinidae, Melocrinidae, Patelliocrinidae, Clonocrinidae, Eucalyptocrinidae, Dolatocrinidae.
Sub-order ii. Batocrinoidea.—RR separated by a heptagonal anal; first brachial usually quadrangular. Families: Tanaocrinidae, Xenocrinidae, Carpocrinidae, Barrandeocrinidae, Coelocrinidae, Batocrinidae, Periechocrinidae.
Sub-order iii. Actinocrinoidea.—RR separated by a hexagonal anal; first brachial usually hexagonal. Families: Actinocrinidae, Amphoracrinidae.
Sub-class II. Dicyclica.—Crinoidea in which the base consists of BB and IBB, the latter being liable to atrophy or fusion with the proximale, but the aboral prolongations of the chambered organ are always radial; new columnals may or may not be introduced at the proximal end of the stem.
Order 1. Dicyclica Inadunata.—Dicyclica in which the dorsal cup primitively is confined to the patina and occasional intercalated anals, and no other plates ever occur between RR (Grade: Distincta); Br may be incorporated in the cup, with or without iBr, but never rigidly, and their corresponding ambulacrals remain supra-tegminal (Grade: Articulata); new columnals are introduced at the extreme proximal end of the stem.