Further, not only the retina but also the dioptric apparatus of the vertebrate eye point to its origin from a type that combined the peculiarities of the arachnids and the crustaceans. In the former it is difficult to speak of a true lens, the function of a lens being undertaken by the cuticular surface of the cells of the corneagen (Mark's 'lentigen'), while in the latter, in addition to the corneal covering, a true lens exists in the shape of the crystalline cones. Further, these crustacean lenses are true lenses in the vertebrate sense, in that they are formed by modified hypodermal cells, and not bulgings of the cuticle, as in the arachnid. We see, in fact, that in the compound crustacean eye an extra layer of hypodermal cells has become inserted between the cornea and the retina to form a lens. So also in the vertebrate eye the lens is formed by an extra layer of the epidermal cells between the cornea and the retina. The fact that the vertebrate eye possesses a single lens, though its retina is composed of a number of ommatidia, while the crustacean eye possesses a lens to each ommatidium, may well be a consequence of the inversion of the vertebrate retina. It is most probable, as Korschelt and Heider have pointed out, that the retina of the arachnid eyes is composed of a number of ommatidia, just as in the crustacean eyes and in the inverted eyes it is probable that the image is focussed on to the pigmented tapetal layer, and thence reflected on to the percipient visual rods. In such a method of vision a single lens is a necessity, and so it must also be if, as I suppose, eyes existed with an inverted compound retina. Owing to the crustacean affinities of such eyes, a lens would be formed and the retina would be compound: owing to the arachnid affinities, the retina would be inverted and the hypodermal cells which formed the lens would be massed {116}together to form a single lens, instead of being collected in groups of four to form a series of crystalline cones. To sum up: The study of the vertebrate eyes, both median and lateral, leads to most important conclusions as to the origin of the vertebrates, for it shows clearly that whereas, as pointed out in this and subsequent chapters, their ancestors possessed distinct arachnid characteristics, yet that they cannot have been specialized arachnids, such as our present-day forms, but rather they were of a primitive arachnid type, with distinct crustacean characteristics: animals that were both crustacean and arachnid, but not yet specialized in either direction: animals, in fact, of precisely the kind which swarmed in the seas at the time when the vertebrates first made their appearance. In the opinion of the present day, the ancestral forms of the Crustacea, which were directly derived from the Annelida, may be classed as an hypothetical group the Protostraca, the nearest approach to which is a primitive Phyllopod.