In the cases where the hypoblast is not invaginated in the form of a sack the formation of the mesenteron is somewhat complicated, and is described in the sequel.
From the trochosphere stage the larva passes into what has been called by Lankester the veliger stage ([fig. 103]), which is especially characteristic of Gasteropod and Pteropod Mollusca.
The shell-gland (with a few exceptions to be spoken of subsequently) of the previous stage flattens out, forming a disc-like area, on the surface of which a delicate shell becomes developed, while the epiblast of the edges of the disc becomes thickened. The disc-like area is the mantle. The edge of the area and with it the shell now rapidly extend, especially in a dorsal direction. Up to this time the embryo has been symmetrical, but in most Gasteropods the shell and mantle extend very much more towards the left than towards the right side, and a commencement of the permanent spiral shell is thus produced.
Fig. 103. Larvæ of Cephalophorous Mollusca in the veliger stage.
(From Gegenbaur.)
A. and B. Earlier and later stage of Gasteropod. C. Pteropod (Cymbulia).
v. velum; c. shell; p. foot; op. operculum; t. tentacle.
The edge of the mantle forms a projecting lip separating the dorsal visceral sack from the head and foot. An invagination appears, usually on the right in Gasteropods, and eventually extends to the dorsal side ([fig. 103] B). It gives rise to the pallial or branchial cavity, and receives also the openings of the digestive, generative and urinary organs. In most Pteropods it is also formed to the right, and usually eventually extends afterwards towards the ventral surface ([fig. 103] C). In the pallial cavity the gills are formed, in those groups in which they are present, as solid processes frequently ciliated. They are coated by epiblast and contain a core of mesoblast. They soon become hollow and contractile.
The velum in the more typical forms loses its simple circular form, and becomes a projecting bilobed organ, which serves the larva after it is hatched as the organ of locomotion ([fig. 103] B and C). The extent of the development of the velum varies greatly. In the Heteropods especially it becomes very large, and in Atlanta it becomes six-lobed, each lateral half presenting three subdivisions. It is usually armed on its projecting edge with several rows of long cilia, and below this with short cilia which bring food to the mouth. It persists in many forms for a very long period. Within the area of the velum there appear the tentacles and eyes ([fig. 103] B). The latter are usually formed at the base of the tentacles.
The foot grows in most forms to a very considerable size. On its hinder and dorsal surface is formed the operculum as a chitinos plate which originates in a depression lined by thickened epiblast, much in the same way as the shell ([fig. 103] B and C, op). In the typical larval forms it is only possible to distinguish the anterior flattened surface of the foot for locomotion and the posterior opercular region, but special modifications of the foot are found in the Pteropods and Heteropods, which are described with those groups. The foot very often becomes richly ciliated, and otic vesicles are early developed in it ([fig. 101], ot).