They divide the area included between highwater-mark and lowwater-mark of spring tides (which is very extensive, on account of the great rise and fall of the tide on the Normandy coast about St. Malo, where their observations were made) into four zones, each characterized by its peculiar invertebrate inhabitants. Beyond the fourth region they distinguish a fifth, which is never uncovered, and is inhabited by oysters, scallops, and large starfishes and other animals. Beyond this they seem to think that animal life is absent.[3]
[Footnote 3: "Enfin plus has encore, c'est-à-dire alors loin des côtes, le fond des eaux ne paraît plus être habité, du moms dans nos mers, par aucun de ces animaux" (1. c. tom. i. p. 237). The "ces animaux" leaves the meaning of the authors doubtful.]
Audouin and Milne Edwards were the first to see the importance of the bearing of a knowledge of the manner in which marine animals are distributed in depth, on geology. They suggest that, by this means, it will be possible to judge whether a fossiliferous stratum was formed upon the shore of an ancient sea, and even to determine whether it was deposited in shallower or deeper water on that shore; the association of shells of animals which live in different zones of depth will prove that the shells have been transported into the position in which they are found; while, on the other hand, the absence of shells in a deposit will not justify the conclusion that the waters in which it was formed were devoid of animal inhabitants, inasmuch as they might have been only too deep for habitation.
The new line of investigation thus opened by the French naturalists was followed up by the Norwegian, Sars, in 1835, by Edward Forbes, in our own country, in 1840,[4] and by Oersted, in Denmark, a few years later. The genius of Forbes, combined with his extensive knowledge of botany, invertebrate zoology, and geology, enabled him to do more than any of his compeers, in bringing the importance of distribution in depth into notice; and his researches in the Aegean Sea, and still more his remarkable paper "On the Geological Relations of the existing Fauna and Flora of the British Isles," published in 1846, in the first volume of the "Memoirs of the Geological Survey of Great Britain," attracted universal attention.
[Footnote 4: In the paper in the Memoirs of the Survey cited further on, Forbes writes:—
"In an essay 'On the Association of Mollusca on the British Coasts, considered with reference to Pleistocene Geology,' printed in [the Edinburgh Academic Annual for] 1840, I described the mollusca, as distributed on our shores and seas, in four great zones or regions, usually denominated 'The Littoral zone,' 'The region of Laminariae,' 'The region of Coral-lines,' and 'The region of Corals.' An extensive series of researches, chiefly conducted by the members of the committee appointed by the British Association to investigate the marine geology of Britain by means of the dredge, have not invalidated this classification, and the researches of Professor Lovén, in the Norwegian and Lapland seas, have borne out their correctness The first two of the regions above mentioned had been previously noticed by Lamoureux, in his account of the distribution (vertically) of sea-weeds, by Audouin and Milne Edwards in their Observations on the Natural History of the coast of France, and by Sars in the preface to his Beskrivelser og Jagttayelser.">[
On the coasts of the British Islands, Forbes distinguishes four zones or regions, the Littoral (between tide marks), the Laminarian (between lowwater-mark and 15 fathoms), the Coralline (from 15 to 50 fathoms), and the Deep sea or Coral region (from 50 fathoms to beyond 100 fathoms). But, in the deeper waters of the Aegean Sea, between the shore and a depth of 300 fathoms, Forbes was able to make out no fewer than eight zones of life, in the course of which the number and variety of forms gradually diminished until, beyond 300 fathoms, life disappeared altogether. Hence it appeared as if descent in the sea had much the same effect on life, as ascent on land. Recent investigations appear to show that Forbes was right enough in his classification of the facts of distribution in depth as they are to be observed in the Aegean; and though, at the time he wrote, one or two observations were extant which might have warned him not to generalize too extensively from his Aegean experience, his own dredging work was so much more extensive and systematic than that of any other naturalist, that it is not wonderful he should have felt justified in building upon it. Nevertheless, so far as the limit of the range of life in depth goes, Forbes' conclusion has been completely negatived, and the greatest depths yet attained show not even an approach to a "zero of life":—
"During the several cruises of H.M. ships Lightning and Porcupine in the years 1868, 1869, and 1870," says Dr. Wyville Thomson, "fifty-seven hauls of the dredge were taken in the Atlantic at depths beyond 500 fathoms, and sixteen at depths beyond 1,000 fathoms, and, in all cases, life was abundant. In 1869, we took two casts in depths greater than 2,000 fathoms. In both of these life was abundant; and with the deepest cast, 2,435 fathoms, off the month of the Bay of Biscay, we took living, well-marked and characteristic examples of all the five invertebrate sub- kingdoms. And thus the question of the existence of abundant animal life at the bottom of the sea has been finally settled and for all depths, for there is no reason to suppose that the depth anywhere exceeds between three and four thousand fathoms; and if there be nothing in the conditions of a depth of 2,500 fathoms to prevent the full development of a varied Fauna, it is impossible to suppose that even an additional thousand fathoms would make any great difference."[5]
[Footnote 5: The Depths of the Sea, p. 30. Results of a similar kind, obtained by previous observers, are stated at length in the sixth chapter, pp. 267-280. The dredgings carried out by Count Pourtales, under the authority of Professor Peirce, the Superintendent of the United States Coast Survey, in the years 1867, 1868, and 1869, are particularly noteworthy, and it is probably not too much to say, in the words of Professor Agassiz, "that we owe to the coast survey the first broad and comprehensive basis for an exploration of the sea bottom on a large scale, opening a new era in zoological and geological research.">[
As Dr. Wyville Thomson's recent letter, cited above, shows, the use of the trawl, at great depths, has brought to light a still greater diversity of life. Fishes came up from a depth of 600 to more than 1,000 fathoms, all in a peculiar condition from the expansion of the air contained in their bodies. On their relief from the extreme pressure, their eyes, especially, had a singular appearance, protruding like great globes from their heads. Bivalve and univalve mollusca seem to be rare at the greatest depths; but starfishes, sea urchins and other echinoderms, zoophytes, sponges, and protozoa abound.