The evidence of the rocks, therefore, proves that the pair of median eyes which were originally the principal eyes (Hauptaugen), had already, in the dominant arthropod group been supplanted by a pair of lateral eyes, and had, in consequence, become small and insignificant, at the time when vertebrates first appeared. This dwindling process thus initiated in the arthropod itself has steadily continued ever since through the whole development of the vertebrates, with the result that, in the highest vertebrates, these median or pineal eyes have become converted into the pineal gland with its 'brain-sand.'
In the earliest vertebrate these median eyes may have been functional; they certainly were more conspicuous than in later forms. Alone among living vertebrates the right median eye of Ammocoetes is so perfect and the skin covering it so transparent that I have always felt doubtful whether it may not be of use to the animal, especially when one takes into consideration the undeveloped state of the lateral eyes in this animal, hidden as they are under the skin. Thus the one living vertebrate which is comparable with these extinct fishes is the one in which one of the pineal eyes is most well defined, most nearly functional.
Before passing to the consideration of the structure of the median eyes of Ammocoetes, it is advisable to see whether these median eyes in other animals, such as arachnids and crustaceans, belong to any particular type of eyes, for then assuredly the median eyes of Ammocoetes ought to belong to the same type if they are derived from them.
In the specialized crustacean, as in the specialized vertebrate, the median eyes have disappeared, at all events in the adult, but still exist in the primitive forms, such as Branchipus, which resemble the trilobites in some respects. On the other hand, the median eyes have persisted, and are well developed in the arachnids, both scorpions and spiders possessing a well-developed pair. The characteristics of the median eyes must then be especially sought for in the arachnid group.
Both scorpions and spiders possess many eyes, of which two are { 76}always separate and median in position, while the others form lateral groups; all these eyes possess a simple retina and a simple corneal lens. Grenacher was the first to point out that in the spiders two very distinct types of eye are found. In the one the retina is upright; in the other the retina is inverted, and the eye possesses a tapetal layer. The distribution of these two types is most suggestive, for the inverted retina is always found in the lateral eyes, never in the two median eyes; these always possess a simple upright retina.