Discourses: Biological & Geological Essays
Chapter 4
"For the first two or three hauls in very deep water off the coast of Portugal, the dredge came up filled with the usual 'Atlantic ooze,' tenacious and uniform throughout, and the work of hours, in sifting, gave the very smallest possible result. We were extremely anxious to get some idea of the general character of the Fauna, and particularly of the distribution of the higher groups; and after various suggestions for modification of the dredge, it was proposed to try the ordinary trawl. We had a compact trawl, with a 15-feet beam, on board, and we sent it down off Cape St. Vincent at a depth of 600 fathoms. The experiment looked hazardous, but, to our great satisfaction, the trawl came up all right and contained, with many of the larger invertebrate, several fishes.... After the first attempt we tried the trawl several times at depths of 1090, 1525, and, finally, 2125 fathoms, and always with success."
To the coral-fishers of the Mediterranean, who seek the precious red coral, which grows firmly fixed to rocks at a depth of sixty to eighty fathoms, both the dredge and the trawl would be useless. They, therefore, have recourse to a sort of frame, to which are fastened long bundles of loosely netted hempen cord, and which is lowered by a rope to the depth at which the hempen cords can sweep over the surface of the rocks and break off the coral, which is brought up entangled in the cords. A similar contrivance has arisen out of the necessities of deep-sea exploration.
In the course of the dredging of the _Porcupine_, it was frequently found that, while few objects of interest were brought up within the dredge, many living creatures came up sticking to the outside of the dredge-bag, and even to the first few fathoms of the dredge-rope. The mouth of the dredge doubtless rapidly filled with mud, and thus the things it should have brought up were shut out. To remedy this inconvenience Captain Calver devised an arrangement not unlike that employed by the coral- fishers. He fastened half a dozen swabs, such as are used for drying decks, to the dredge. A swab is something like what a birch-broom would be if its twigs were made of long, coarse, hempen yarns. These dragged along after the dredge over the surface of the mud, and entangled the creatures living there--multitudes of which, twisted up in the strands of the swabs, were brought to the surface with the dredge. A further improvement was made by attaching a long iron bar to the bottom of the dredge bag, and fastening large bunches of teased-out hemp to the end of this bar. These "tangles" bring up immense quantities of such animals as have long arms, or spines, or prominences which readily become caught in the hemp, but they are very destructive to the fragile organisms which they imprison; and, now that the trawl can be successfully worked at the greatest depths, it may be expected to supersede them; at least, wherever the ground is soft enough to permit of trawling.
It is obvious that between the dredge, the trawl, and the tangles, there is little chance for any organism, except such as are able to burrow rapidly, to remain safely at the bottom of any part of the sea which the _Challenger_ undertakes to explore. And, for the first time in the history of scientific exploration, we have a fair chance of learning what the population of the depths of the sea is like in the most widely different parts of the world.
And now arises the next question. The means of exploration being fairly adequate, what forms of life may be looked for at these vast depths?
The systematic study of the Distribution of living beings is the most modern branch of Biological Science, and came into existence long after Morphology and Physiology had attained a considerable development. This naturally does not imply that, from the time men began to observe natural phenomena, they were ignorant of the fact that the animals and plants of one part of the world are different from those in other regions; or that those of the hills are different from those of the plains in the same region; or finally that some marine creatures are found only in the shallows, while others inhabit the deeps. Nevertheless, it was only after the discovery of America that the attention of naturalists was powerfully drawn to the wonderful differences between the animal population of the central and southern parts of the new world and that of those parts of the old world which lie under the same parallels of latitude. So far back as 1667 Abraham Mylius, in his treatise "De Animalium origine et migratione, populorum," argues that, since there are innumerable species of animals in America which do not exist elsewhere, they must have been made and placed there by the Deity: Buffon no less forcibly insists upon the difference between the Faunae of the old and new world. But the first attempt to gather facts of this order into a whole, and to coordinate them into a series of generalizations, or laws of Geographical Distribution, is not a century old, and is contained in the "Specimen Zoologiae Geographicae Quadrupedum Domicilia et Migrationes sistens," published, in 1777, by the learned Brunswick Professor, Eberhard Zimmermann, who illustrates his work by what he calls a "Tabula Zoographica," which is the oldest distributional map known to me.
In regard to matters of fact, Zimmermann's chief aim is to show that among terrestrial mammals, some occur all over the world, while others are restricted to particular areas of greater or smaller extent; and that the abundance of species follows temperature, being greatest in warm and least in cold climates. But marine animals, he thinks, obey no such law. The Arctic and Atlantic seas, he says, are as full of fishes and other animals as those of the tropics. It is, therefore, clear that cold does not affect the dwellers in the sea as it does land animals, and that this must be the case follows from the fact that sea water, "propter varias quas continet bituminis spiritusque particulas," freezes with much more difficulty than fresh water. On the other hand, the heat of the Equatorial sun penetrates but a short distance below the surface of the ocean. Moreover, according to Zimmermann, the incessant disturbance of the mass of the sea by winds and tides, so mixes up the warm and the cold that life is evenly diffused and abundant throughout the ocean.
In 1810, Risso, in his work on the Ichthyology of Nice, laid the foundation of what has since been termed "bathymetrical" distribution, or distribution in depth, by showing that regions of the sea bottom of different depths could be distinguished by the fishes which inhabit them. There was the _littoral region_ between tide marks with its sand-eels, pipe fishes, and blennies: the _seaweed region_, extending from low- water-mark to a depth of 450 feet, with its wrasses, rays, and flat fish; and the _deep-sea region_, from 450 feet to 1500 feet or more, with its file-fish, sharks, gurnards, cod, and sword-fish.
More than twenty years later, M.M. Audouin and Milne Edwards carried out the principle of distinguishing the Faunae of different zones of depth much more minutely, in their "Recherches pour servir à l'Histoire Naturelle du Littoral de la France," published in 1832.
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.
It is obvious that the _Challenger_ has the privilege of opening a new chapter in the history of the living world. She cannot send down her dredges and her trawls into these virgin depths of the great ocean without bringing up a discovery. Even though the thing itself may be neither "rich nor rare," the fact that it came from that depth, in that particular latitude and longitude, will be a new fact in distribution, and, as such, have a certain importance.
But it may be confidently assumed that the things brought up will very frequently be zoological novelties; or, better still, zoological antiquities, which, in the tranquil and little-changed depths of the ocean, have escaped the causes of destruction at work in the shallows, and represent the predominant population of a past age.
It has been seen that Audouin and Milne Edwards foresaw the general influence of the study of distribution in depth upon the interpretation of geological phenomena. Forbes connected the two orders of inquiry still more closely; and in the thoughtful essay "On the connection between the distribution of the existing Fauna and Flora of the British Isles, and the geological changes which have affected their area, especially during the epoch of the Northern drift," to which reference has already been made, he put forth a most pregnant suggestion.
In certain parts of the sea bottom in the immediate vicinity of the British Islands, as in the Clyde district, among the Hebrides, in the Moray Firth, and in the German Ocean, there are depressed areas, forming a kind of submarine valleys, the centres of which are from 80 to 100 fathoms, or more, deep. These depressions are inhabited by assemblages of marine animals, which differ from those found over the adjacent and shallower region, and resemble those which are met with much farther north, on the Norwegian coast. Forbes called these Scandinavian detachments "Northern outliers."
How did these isolated patches of a northern population get into these deep places? To explain the mystery, Forbes called to mind the fact that, in the epoch which immediately preceded the present, the climate was much colder (whence the name of "glacial epoch" applied to it); and that the shells which are found fossil, or sub-fossil, in deposits of that age are precisely such as are now to be met with only in the Scandinavian, or still more Arctic, regions. Undoubtedly, during the glacial epoch, the general population of our seas had, universally, the northern aspect which is now presented only by the "northern outliers"; just as the vegetation of the land, down to the sea-level, had the northern character which is, at present, exhibited only by the plants which live on the tops of our mountains. But, as the glacial epoch passed away, and the present climatal conditions were developed, the northern plants were able to maintain themselves only on the bleak heights, on which southern forms could not compete with them. And, in like manner, Forbes suggested that, after the glacial epoch, the northern animals then inhabiting the sea became restricted to the deeps in which they could hold their own against invaders from the south, better fitted than they to flourish in the warmer waters of the shallows. Thus depth in the sea corresponded in its effect upon distribution to height on the land.
The same idea is applied to the explanation of a similar anomaly in the Fauna of the Aegean:--
"In the deepest of the regions of depth of the Aegean, the representation of a Northern Fauna is maintained, partly by identical and partly by representative forms.... The presence of the latter is essentially due to the law (of representation of parallels of latitude by zones of depth), whilst that of the former species depended on their transmission from their parent seas during a former epoch, and subsequent isolation. That epoch was doubtless the newer Pliocene or Glacial Era, when the _Mya truncata_ and other northern forms now extinct in the Mediterranean, and found fossil in the Sicilian tertiaries, ranged into that sea. The changes which there destroyed the _shallow water_ glacial forms, did not affect those living in the depths, and which still survive."[6]
[Footnote 6: _Memoirs of the Geological Survey of Great Britain_, Vol. i. p. 390.]
The conception that the inhabitants of local depressions of the sea bottom might be a remnant of the ancient population of the area, which had held their own in these deep fastnesses against an invading Fauna, as Britons and Gaels have held out in Wales and in Scotland against encroaching Teutons, thus broached by Forbes, received a wider application than Forbes had dreamed of when the sounding machine first brought up specimens of the mud of the deep sea. As I have pointed out elsewhere,[7] it at once became obvious that the calcareous sticky mud of the Atlantic was made up, in the main, of shells of _Globigerina_ and other _Foraminifera_, identical with those of which the true chalk is composed, and the identity extended even to the presence of those singular bodies, the Coccoliths and Coccospheres, the true nature of which is not yet made out. Here then were organisms, as old as the cretaceous epoch, still alive, and doing their work of rock-making at the bottom of existing seas. What if _Globigerina_ and the Coccoliths should not be the only survivors of a world passed away, which are hidden beneath three miles of salt water? The letter which Dr. Wyville Thomson wrote to Dr. Carpenter in May, 1868, out of which all these expeditions have grown, shows that this query had become a practical problem in Dr. Thomson's mind at that time; and the desirableness of solving the problem is put in the foreground of his reasons for urging the Government to undertake the work of exploration:--
[Footnote 7: See above, "On a Piece of Chalk," p. 13.]
"Two years ago, M. Sars, Swedish Government Inspector of Fisheries, had an opportunity, in his official capacity, of dredging off the Loffoten Islands at a depth of 300 fathoms. I visited Norway shortly after his return, and had an opportunity of studying with his father, Professor Sars, some of his results. Animal forms were _abundant_; many of them were new to science; and among them was one of surpassing interest, the small crinoid, of which you have a specimen, and which we at once recognised as a degraded type of the _Apiocrinidoe_, an order hitherto regarded as extinct, which attained its maximum in the Pear Encrinites of the Jurassic period, and whose latest representative hitherto known was the _Bourguettocrinus_ of the chalk. Some years previously, Mr. Absjornsen, dredging in 200 fathoms in the Hardangerfjord, procured several examples of a Starfish (_Brisinga_), which seems to find its nearest ally in the fossil genus _Protaster_. These observations place it beyond a doubt that animal life is abundant in the ocean at depths varying from 200 to 300 fathoms, that the forms at these great depths differ greatly from those met with in ordinary dredgings, and that, at all events in some cases, these animals are closely allied to, and would seem to be directly descended from, the Fauna of the early tertiaries.
"I think the latter result might almost have been anticipated; and, probably, further investigation will largely add to this class of data, and will give us an opportunity of testing our determinations of the zoological position of some fossil types by an examination of the soft parts of their recent representatives. The main cause of the destruction, the migration, and the extreme modification of animal types, appear to be change of climate, chiefly depending upon oscillations of the earth's crust. These oscillations do not appear to have ranged, in the Northern portion of the Northern Hemisphere, much beyond 1,000 feet since the commencement of the Tertiary Epoch. The temperature of deep waters seems to be constant for all latitudes at 39°; so that an immense area of the North Atlantic must have had its conditions unaffected by tertiary or post-tertiary oscillations."[8]
[Footnote 8: The Depths of the Sea, pp. 51-52.]
As we shall see, the assumption that the temperature of the deep sea is everywhere 39° F. (4° Cent.) is an error, which Dr. Wyville Thomson adopted from eminent physical writers; but the general justice of the reasoning is not affected by this circumstance, and Dr. Thomson's expectation has been, to some extent, already verified.
Thus besides _Globigerina_, there are eighteen species of deep-sea _Foraminifera_ identical with species found in the chalk. Imbedded in the chalky mud of the deep sea, in many localities, are innumerable cup- shaped sponges, provided with six-rayed silicious spicula, so disposed that the wall of the cup is formed of a lacework of flinty thread. Not less abundant, in some parts of the chalk formation, are the fossils known as _Ventriculites_, well described by Dr. Thomson as "elegant vases or cups, with branching root-like bases, or groups of regularly or irregularly spreading tubes delicately fretted on the surface with an impressed network like the finest lace"; and he adds, "When we compare such recent forms as _Aphrocallistes, Iphiteon, Holtenia_, and _Askonema_, with certain series of the chalk _Ventriculites_, there cannot be the slightest doubt that they belong to the same family--in some cases to very nearly allied genera."[9]
[Footnote 9: _The Depths of the Sea_, p. 484.]
Professor Duncan finds "several corals from the coast of Portugal more nearly allied to chalk forms than to any others."