
Discussion
Of the modern struthiolariids, Tylospira would seem to be the closest to Pelicaria in the characters of the animal, radula and operculum, as well as in the scaphelloid apex and the probable loss of the free-swimming larval stage. Both are also close—though rather less so—to Struthiolaria, and in the more fundamental characters of the animal have shown great conservatism to change. Pelicaria is of rather late (Upper Miocene) New Zealand origin, and its resemblances to the Australian Tylospira therefore involve the same difficulties as Marwick (1952) has pointed to, in discussing the Pelicaria-Singletonaria relationship. Tylospira must have been restricted in its range by the loss of the free-swimming veliger stage, and—rather than to postulate a post-Miocene crossing of the Tasman—it seems much more reasonable on geographical grounds to suppose that Tylospira took its origin from Struthiolaria in New Zealand at a much earlier date, possibly in the Oligocene. As has been already pointed out (Morton, 1950), it is not necessary to insist that the scaphelloid apex was developed only once in the Struthiolariidae. Its occurrence is a normal corollary of the incubation of the eggs in a brood pouch, with the consequent need of a larger volume of yolk; and it is very possible that parallel evolution of incubation has taken place in the two most advanced genera of the family—namely, Pelicaria and Tylospira. In the same manner, the evolution of the radula in these genera, with its elongated central cusps and loss of denticles on the marginals, might be

regarded as a minor “orthogenetic” trend, that was followed more than once among “advanced” genera of the Struthiolariidae. We may on the whole, then, in morphological features and in life history, regard Tylospira as most closely related to Pelicaria, while remembering that none of these facts would exclude the hypothesis of parallel evolution, if the geographical evidence should seem to require it.
The Australian Singletonaria (Marwick, 1952) is probably another advanced genus having a scaphelloid apex, and showing restraint of the callus of the inner lip and simplification of the outer lip, in general similar to the condition of Pelicaria. In both cases also the brephic whorls are convex and spirally sculptured. Because of the geographical difficulties of a Pelicaria-Singletonaria connection, Marwick (1952) turned to consider the relationships of the two Australian struthiolariids, concluding that Tylospira, also possessing convex and spirally sculptured brephic whorls, might represent a stock of which Singletonaria is the neotenic descendant. Alternatively, the modern Tylospira could have been derived by the pressing back of the earlier convex, spirally sculptured stage into the initial part of the life history, while the overgrowth of the later whorls by a callus layer arose as a later gerontic feature.
