# Popular Science Monthly/Volume 27/September 1885/The Fauna of the Sea-Shore

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* THE FAUNA OF THE SEA-SHORE.
By Professor H. N. MOSELEY, F. R. S.
THE marine fauna of the globe may be divided into the littoral, the deep-sea, and the pelagic faunas. Of the three regions inhabited by these faunas, the littoral is the one in which the conditions are most favorable for the development of new forms through the working of the principle of natural selection. As Professor Lovèn writes, "The littoral region comprises the favored zones of the sea where light and shade, a genial temperature, currents changeable in power and direction, a rich vegetation spread over extensive areas, abundance of food, of prey to allure, of enemies to withstand or evade, represent an infinitude of agents competent to call into play the tendencies to vary which are embodied in each species, and always ready by modifying its parts to respond to the influences of external conditions." It is consequently in this littoral zone, where the water is more than elsewhere favorable ​for respiration, and where constant variation of conditions is produced by the tides, that all the main groups of the animal kingdom first came into existence; and here also, probably, where the first attached and branching plants were developed, thus establishing a supply of food for the colonization of the region by animals.
The animals inhabiting the littoral zone are most variously modified, to enable them to withstand the peculiar physical conditions which they encounter there. Hence the origin of all hard shells and skeletons of marine invertebrata, various adaptations for boring in sand, the adoption of the stationary fixed condition, and similar arrangements. Almost all the shore forms of animals, however inert in the adult condition, pass through, in embryological development, free swimming larval stages which are closely alike in form for very widely different groups of animals. Thus the oyster and. most other mollusca of all varieties and shapes when adult, develop from a free-swimming pelagic trochosphere larva, and so do many annelids. Such larva? can not be of subsequent origin to the adults of which they are phases. If such were the case, they would not have become so closely alike in structure. In reality they represent the common ancestors from which all the forms in which they occur were derived, and, as all these larvæ are pelagic in habits and structure, it follows that the inhabitants of the shores were derived from pelagic ancestors. The earliest plants were also probably free-swimming.
In the case of the cirripedia there can be no doubt, from the history of their development, that they were originally pelagic, and have become specially modified for coast-life; and in the case of the echinoderms the only possible explanation of the remarkable similarity of the larval forms of the various groups of widely differing adults is that these pelagic larvae represent a common ancestor of the group. The madreporarian corals all spring from a pelagic larva. The colonial forms probably owe their origin and that of their skeletons to the advantage gained by them in the formation of reefs, and the increase in facilities of respiration consequent on the production of surf. In the deep sea they are very scarce.
The vertebrata are sprung from a very simple free-swimming ancestor, as shown by the ciliated gastrula stage of Amphioxus. The ascidians afford another evident instance of the extreme modification of pelagic forms for littoral existence.
The peculiar mode of respiration of vertebrata by means of gill-slits occurs in no other animal group except in Balanoglossus, which will probably shortly be included among vertebrata. Possibly gill-slits as a respiratory apparatus first arose in a littoral form, such as Balanoglossus, and hence their presence at the anterior end of the body, that nearest to the surface in an animal buried in sand. The connection of Balanoglossus with the echinoderms through Tornaria is very remarkable. Possibly Amphioxus once had a Tornaria stage, ​and has lost it just as one species of Balanoglossus has lost it, as Mr. Bateson has lately discovered.
The littoral zone has given off colonists to the other three faunal regions. The entire terrestrial fauna has sprung from colonists contributed by the littoral zone. Every terrestrial vertebrate bears in its early stages the gill-slits of its aquatic ancestor. All organs of aerial respiration are mere modifications of apparatus previously connected with aquatic respiration, excepting, perhaps, in the case of Tracheata, tracheæ being most likely modifications of skin-glands, as appears probable from their condition in Peripatus. The oldest known air-breathing animals are insects and scorpions, which have lately been found in Silurian strata. Professor Ray Lankester believes the lungs of scorpions to be homogeneous with the gill-plates of Limulus. Birds were possibly originally developed in connection with the sea-shore, and were fish-eaters like the tooth-bearing Hesperornis.
The fauna of the coast has not only given rise to the terrestrial and fresh-water fauna; it has from time to time given additions to the pelagic fauna in return for having thence derived its own starting-points. It has also received some of these pelagic forms back again, to assume a fresh littoral existence.
The deep-sea fauna has probably been formed almost entirely from the littoral, not in the remotest antiquity, but only after food derived from the débris of the littoral and terrestrial faunas and floras became abundant.
It is because all terrestrial and deep-sea animal forms have passed through a littoral phase of existence, and that the littoral animals retain far better than those of any other faunal region the recapitulative larval phases by means of which alone the true histories of their origins can be recovered, that marine zoölogical laboratories on the coast have made so many brilliant discoveries in zoölogy during late years.