Reviewing the whole group of the Pelmatozoa, we see that in the Cambrian they begin with the extremely primitive Thecoidea and Carpoidea, together with some obscure forms which, combining a stem with pentamerous symmetry in the calyx, are supposed to be the forerunners of the Crinoidea. In the Lower Silurian or Ordovician the two groups of the Cystoidea make their appearance, possibly independently developed from either Carpoidea or primitive Crinoidea, which in this period are present in unmistakable form. In the Upper Silurian the Blastoidea appear, distinguishable from the most regular Cystoidea only by their hydrospires. It seems practically certain that they were developed from Cystoidea, and we follow Jaekel in believing that they arose from Dichoporita. The Carpoidea do not extend beyond the Ordovician, and by the end of the Carboniferous period Cystoidea and Blastoidea die out, leaving only the Crinoidea, which at that period were at their maximum development. From the Carboniferous to the present day the Crinoidea have continually decreased, leaving in recent seas, as sole representatives of the Pelmatozoa, only the few forms described at the beginning of this chapter.
CHAPTER XXI
ECHINODERMATA (CONTINUED): DEVELOPMENT AND PHYLOGENY
In Chapter XVI. it was stated that whilst a more or less perfectly developed radial symmetry was one of the characteristic features of the phylum Echinodermata when in the adult condition, yet in the immature or larval condition the members of the group have a strongly marked bilateral symmetry. In this feature larval Echinodermata resemble the other Phyla of the animal kingdom which have a well-developed coelom, such as Annelida, Mollusca, Vertebrata, etc. Since, then, the peculiar radial symmetry is gradually acquired during the growth of the Echinoderm, we may possibly discover by a close scrutiny of the life-history what is the nature and meaning of this departure from the ordinary type of structure among coelomate animals.
There are two kinds of development met with amongst Echinodermata, which may be roughly characterised as the "embryonic" and the "larval" type respectively, although neither description is exact. In developmental histories of the first type so much reserve material is laid up in the egg in the form of food-yolk that the young animal whilst in the bilateral stage requires little or no food. In some cases, however, as in Amphiura squamata, the mother pours out a nourishing exudation; but whether this is so or not, the parent in nearly every case carries the young about with her until they have reached the adult condition. In some Asteroidea, as for instance in the Antarctic species Asterias spirabilis (Fig. 280), the young become fixed to the everted lips of the mother; in Amphiura squamata, and some other Ophiuroidea the eggs remain in the genital bursae, which serve as nurseries; in some Spatangoidea, as for instance in Hemiaster philippi (Figs. 250, 281), the eggs are carried in some of the deeply grooved petaloid ambulacra; whilst in Holothuroidea they may develop in the body-cavity (Phyllophorus urna), or they may adhere to the back of the mother (Cucumaria crocea, Fig. 259, p. [573]), or they may be protected in special brood-pouches either on the ventral side of the parent (Cucumaria laevigata) or on the dorsal surface (Psolus ephippifer, Fig. 261).
The majority of these cases of embryonic development have been recorded from Arctic or Antarctic waters; it appears as if conditions there were not favourable to the larval type of development. In Pelmatozoa the development of Antedon rosacea alone is known, and that is of the embryonic type.
Fig. 280.—Oral view of Asterias spirabilis, slightly enlarged, showing embryos attached to the everted lips. emb, Embryos. (After Perrier.)
So far, however, as their mode of propagation is known, it may confidently be affirmed that the development of the majority of the species of Eleutherozoa is of the second or larval type. In this type there is little food-yolk in the egg, and the young animal or larva is forced from a very early period of development to seek its own living, and hence it is usually a considerable time (from a fortnight to two months) before the adult form is attained. When the embryos of different groups of Eleutherozoa are compared, there is no obvious agreement in structure between them; but the larvae of the four classes of Eleutherozoa exhibit with differences in detail a most remarkable fundamental similarity in type, and we are accordingly justified in regarding the larval development as primitive, and the embryonic type as derived from it and differently modified in each case.
