Turning to fossil Asteroidea, we find the earlier ophiurids scarcely distinguishable from the asterids, while in the alternation of the ambulacrals, which undoubtedly correspond to the flooring-plates of Edrioaster, both groups approach the Pelmatozoan type.
It is spheroidal, with the mouth and anus at opposite poles; there are five ambulacra, and the ambulacral plates are large, simple and alternating, each being pierced by two podial pores which lie in a small oval depression; the ambulacrals next the mouth form a closed ring of ten plates; the interambulacrals lie in single columns between the ambulacra, and are separated from the mouth-area by the proximal ambulacrals just mentioned, and sometimes by the second set of ambulacrals also; the ambulacra end in the five oculars or terminals, which meet in a ring around the anal area and have no podial pores, but one of them serves as a madreporite; within this ring is a star-shaped area filled with minute irregular plates, none of which can safely be selected as the homologues of the so-called basals or genitals of later forms; within the ring of ambulacrals around the mouth are five somewhat pointed plates, which Jaekel regards as teeth, but which can scarcely be homologous with the interradially placed teeth of later echinoids, since they are radial in position; small spines are present, especially around the podial pores.
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.
- 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.
- 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.
- 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.
Tegmen flexible, with distinct ambulacrals and numerous small interambulacrals; mouth and food-grooves remain supra-tegminal and open.
The radial water-vessels lie in grooves on the ventral side of flooring-plates (usually called "ambulacrals"); they and their podia are limited to the oral surface of the body and their extremities are separated from the FIG.
A separation of the latter as Palasterida, because of their alternating ambulacrals, from the recent Euasterida with opposite ambulacrals, is now discarded and an attempt made to arrange the Palasterida in divisions originally established for Euasterida.
P. Sladen, who divided the Euasterida into two orders: (I) Phanerozonia, with marginals large and highly developed, the supero-marginals and inferomarginals contiguous, with papulae confined to the dorsal surface, with ambulacrals well spaced and usually broad, adambulacrals prominent in the mouth-skeleton, with pedicellariae sessile; (2) Cryptozonia, with marginals inconspicuous and somewhat atrophied in the adult, the supero-marginals separated from the inferomarginals by intercalated plates, with papulae distributed over the whole body, with ambulacrals crowded and narrow, either ambulacrals or adambulacrals prominent in the mouth-skeleton, with pedicellariae stalked or sessile.
- Ambulacrals not yet forming complete vertebrae; plates of disk not yet specialized into mouth, radial or genital shields.
- Ambulacrals alternating and unfused, groove uncovered by ventral arm-plates.
- Ambulacrals opposite and, except in Ophi urinidae, fused; ventral arm-plates developed in some.
The Silurian genera Eucladia and Euthemon have the rays greatly reduced and merged in the disk, so that the ambulacrals are unseen.
- A vertebral arm-ossicle (fused ambulacrals) of a Zygophiuran, Ophiolepis.
- Ambulacrals simple, each with one or two pores, which sometimes pass between rather than through the plates, in 2 columns; interambulacrals, unior multi-tuberculate, in numerous (say io or more) columns, none passing on to peristome; mouth central with jaws, no external gills or sphaeridia; position of anus doubtful, acyclic, i.e.
- Ambulacrals generally compound, with two pores obliquely juxtaposed, in 2 columns as in all subsequent orders; interambulacrals usually with large radioles surrounded by smaller ones, as in Cidaroida, in 2 columns as in all subsequent orders, only one plate resorbed; mouth central, with jaws and external gills, sphaeridia present; anus aboral endocyclic. J.
- Ambulacrals sometimes compound, with one or two pores to a plate, some dorsal podia begin to assume respiratory function; interambulacrals multi-tuberculate, none resorbed; mouth central, with jaws weak or wanting, with external gills and sphaeridia; anus exocyclic. Families: Pygasteridae, Discoidiidae, Galeritidae, Conoclypeidae; Jurassic to Recent.
- Ambulacrals simple, with two pores juxtaposed, dorsal podia respiratory; interambulacrals bearing numerous small spines, none resorbed; mouth central or shifted forwards, with no jaws or external gills, sphaeridia numerous; anus exocyclic. As the mouth moves forward and the anus downward, the posterior interambulacrals between them are enlarged and strengthened so as to form a sternum.
- Ambulacrals simple or compound, with two pores juxtaposed, dorsal podia respiratory; interambulacrals multi-tuberculate, none resorbed; mouth central with flattened unequal jaws, reduced external gills, and few sphaeridia; anus exocyclic. Families: Fibulariidae, Laganidae, Scutellidae, Clypeastridae; Cretaceous to Recent.