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-The Project Gutenberg eBook of The fauna of the deep sea, by Sydney John
-Hickson
-
-This eBook is for the use of anyone anywhere in the United States and
-most other parts of the world at no cost and with almost no restrictions
-whatsoever. You may copy it, give it away or re-use it under the terms
-of the Project Gutenberg License included with this eBook or online at
-www.gutenberg.org. If you are not located in the United States, you
-will have to check the laws of the country where you are located before
-using this eBook.
-
-Title: The fauna of the deep sea
-
-Author: Sydney John Hickson
-
-Release Date: January 24, 2021 [eBook #64380]
-
-Language: English
-
-Character set encoding: UTF-8
-
-Produced by: Richard Tonsing, Sharon Joiner, Bryan Ness, and the Online
- Distributed Proofreading Team at https://www.pgdp.net (This
- file was produced from images generously made available by The
- Internet Archive/American Libraries.)
-
-*** START OF THE PROJECT GUTENBERG EBOOK THE FAUNA OF THE DEEP SEA ***
-
-
- =Modern Science Series=
-
- _EDITED BY SIR JOHN LUBBOCK, BART., M. P._
-
-
-
-
- THE FAUNA OF THE DEEP SEA
-
-
-[Illustration: STOMIAS BOA. HALF NATURAL SIZE. FROM A DEPTH OF 1,900
-METRES. (AFTER FILHOL.)]
-
-
-
-
- THE FAUNA OF THE DEEP SEA
-
-
- BY
-
- SYDNEY J. HICKSON, M. A.
- (CANTAB. ET OXON.)
- D.SC. (LOND.), FELLOW OF DOWNING COLLEGE, CAMBRIDGE
-
-
- _WITH TWENTY-THREE ILLUSTRATIONS_
-
-[Illustration]
-
- NEW YORK
- D. APPLETON AND COMPANY
- 1894.
-
-
-
-
- _Authorized Edition._
-
-
-
-
- PREFACE
-
-
-The time may come when there will be no portion of the earth’s surface
-that has not been surveyed and explored by man.
-
-The work of enterprising travellers has now been carried on within a
-measurable distance of the North Pole; the highest mountain ranges are
-gradually succumbing to the geological surveyor; the heart of Africa is
-giving up to us its secrets and its treasures, and plans of all the
-desert places of the earth are being made and tabulated.
-
-The bottom of the deep sea was until quite recently one of these terræ
-incognitæ. It was regarded by most persons, when it entered into their
-minds to consider it at all, as one of those regions about which we do
-not know anything, never shall know anything, and do not want to know
-anything.
-
-But the men of science fifty years ago were not disposed to take this
-view of the matter. Pushing their inquiries as to the character of the
-sea-fauna into deeper and deeper water, they at length demanded
-information as to the existence of forms of animal life in the greatest
-depths. Unable themselves to bear the heavy expenses involved in such an
-investigation, they sought for and obtained the assistance of the
-Government, in the form of national ships, for the work, and then our
-knowledge of the depths of the great ocean may be said to have
-commenced.
-
-We know a good deal now, and in the course of time we may know a great
-deal more, about this interesting region; but it is not one which, in
-our generation at any rate, any human being will ever visit.
-
-We may be able to plant the Union Jack on the summit of Mount Everest,
-we may drag our sledges to the South Pole, and we may, some day, be able
-to travel with ease and safety in the Great Sahara; but we cannot
-conceive that it will ever be possible for us to invent a diving-bell
-that will take a party of explorers to a depth of three and a half miles
-of water. We may complete our survey of the ocean beds, we may analyse
-the bottom muds and name and classify the animals that compose their
-fauna, but there are many things that must remain merely matters of
-conjecture. We shall never know, for example, with any degree of
-certainty, how _Bathypterois_ uses its long feeler-like pectoral fins,
-nor the meaning of the fierce armature of _Lithodes ferox_; why the
-deep-sea Crustacea are so uniformly coloured red, or the intensity of
-the phosphorescent light emitted by the Alcyonaria and Echinoderms.
-These and many others are and must remain among the mysteries of the
-abyss.
-
-Our present-day knowledge of the physical conditions of the bottom of
-the deep sea and the animals that dwell there is by no means
-inconsiderable.
-
-It may be found in the reports of the scientific expeditions fitted out
-by the English, French, German, Italian, Norwegian, and American
-Governments, in numerous volumes devoted to this kind of work, and in
-memoirs and notes scattered through the English and foreign scientific
-journals.
-
-It is the object of this little book to bring together in a small
-compass some of the more important facts and considerations that may be
-found in this great mass of literature, and to present them in such a
-form that they may be of interest to those who do not possess a
-specialist’s knowledge of genera and species.
-
-When it was found that animals can and do live even at the greatest
-depths of the ocean, the interest of naturalists was concentrated on the
-solution of the following problems. Firstly, do the animals constituting
-the fauna of the abyss exhibit any striking and constant modification in
-correlation with the physical conditions of their strange habitat? And,
-secondly, from what source was the fauna of the abyss derived? Was it
-derived from the shallow shore waters, or from the surface of the sea?
-Is it of very ancient origin, or the result of, comparatively speaking,
-recent immigrations?
-
-These questions cannot be answered in a few lines. Any views that may be
-put forward regarding them require the support of a vast array of facts
-and figures; but as the limits of this little book would not permit of
-my giving these, I have endeavoured to select a few only of those which
-bear most directly upon the points at issue.
-
-To overburden my work with the names of genera or the lists of species
-would not, it seemed to me, either clear the issues or interest the
-general reader. These may be found in the ‘Challenger’ monographs, and
-other books dealing with the subject.
-
-Those who wish to pursue the subject further will find in the ‘Voyages
-of the “Blake,”’ by Alexander Agassiz, an excellent and elaborate
-discussion of deep-sea problems, and numerous illustrations of some of
-the most interesting forms of abysmal life.
-
-In Volume XXIII. of the ‘Bulletin of Comparative Zoology’ the same
-author gives a most interesting account of the deep-sea work that has
-recently been done by the ‘Albatross’ expedition.
-
-Filhol’s ‘La Vie au Fond des Mers’ is also a book that contains a great
-deal of new and interesting matter, together with some excellent
-coloured plates of deep-sea animals.
-
- SYDNEY J. HICKSON.
-
- DOWNING COLLEGE, CAMBRIDGE:
- _September, 1893_.
-
-
-
-
- CONTENTS
-
-
- CHAPTER PAGE
- I. A SHORT HISTORY OF THE INVESTIGATIONS 1
-
- II. THE PHYSICAL CONDITIONS OF THE ABYSS 17
-
- III. THE RELATIONS OF THE ABYSMAL ZONE AND THE ORIGIN OF ITS
- FAUNA 45
-
- IV. THE CHARACTERS OF THE DEEP-SEA FAUNA 59
-
- V. THE PROTOZOA, CŒLENTERA, AND ECHINODERMA OF THE DEEP SEA 86
-
- VI. THE VERMES AND MOLLUSCA OF THE DEEP SEA 109
-
- VII. THE ARTHROPODA OF THE DEEP SEA 123
-
- VIII. THE FISH OF THE DEEP SEA 148
-
- INDEX 167
-
-
-
-
- LIST OF ILLUSTRATIONS
-
-
- STOMIAS BOA. AFTER FILHOL, ‘LA VIE AU FOND DES
- MERS’ _Frontispiece_
-
- FIG. PAGE
- 1 DIAGRAM ILLUSTRATING THE PASSAGE OF AN OCEAN
- CURRENT ACROSS A BARRIER 32
-
- 2 _Sicyonis crassa._ AFTER HERTWIG, ‘“CHALLENGER”
- REPORTS’ 36
-
- 3 GLOBIGERINA OOZE. AFTER AGASSIZ, ‘VOYAGES OF THE
- “BLAKE”’ 38
-
- 4 SECTION THROUGH THE EYE OF _Serolis schythei_.
- AFTER BEDDARD, ‘“CHALLENGER” REPORTS’ 74
-
- 5 SECTION THROUGH THE EYE OF _Serolis bromleyana_.
- AFTER BEDDARD, ‘“CHALLENGER” REPORTS’ 74
-
- 6 _Opostomias micripnus._ AFTER GÜNTHER,
- ‘“CHALLENGER” REPORTS’ 78
-
- 7 HEAD OF _Pachystomias microdon_. AFTER VON
- LENDENFELD, ‘“CHALLENGER” REPORTS’ 79
-
- 8 SECTION THROUGH THE ANTERIOR SUB-ORBITAL
- PHOSPHORESCENT ORGAN OF _Pachystomias microdon_.
- AFTER VON LENDENFELD, ‘“CHALLENGER” REPORTS’ 80
-
- 9 _Challengeria Murrayi._ AFTER HAECKEL,
- ‘“CHALLENGER” REPORTS’ 90
-
- 10 _Umbellula Güntheri._ AFTER AGASSIZ, ‘VOYAGES OF
- THE “BLAKE”’ 97
-
- 11 _Rhizocrinus lofotensis._ AFTER CARPENTER,
- ‘“CHALLENGER” REPORTS’ 100
-
- 12 _Rhabdopleura normani._ AFTER LANKESTER,
- ‘CONTRIBUTIONS TO OUR KNOWLEDGE OF RHABDOPLEURA
- AND AMPHIOXUS’ 112
-
- 13 A SINGLE POLYPIDE OF _Rhabdopleura normani_. AFTER
- LANKESTER, _tom. cit._ 114
-
- 14 _Bathyteuthis abyssicola._ AFTER HOYLE,
- ‘“CHALLENGER” REPORTS’ 121
-
- 15 _Bathynomus giganteus._ AFTER FILHOL, _tom. cit._ 131
-
- 16 _Euphausia latifrons._ AFTER SARS, ‘“CHALLENGER”
- REPORTS’ 134
-
- 17 _Bentheuphausia amblyops._ AFTER SARS,
- ‘“CHALLENGER” REPORTS’ 134
-
- 18 _Polycheles baccata._ AFTER SPENCE BATE,
- ‘“CHALLENGER” REPORTS’ 136
-
- 19 _Colossendeis arcuatus._ AFTER FILHOL, _tom. cit._ 141
-
- 20 _Hypobythius calycodes._ AFTER MOSELEY,
- ‘“CHALLENGER” REPORTS’ 145
-
- 21 _Melanocetus Murrayi._ AFTER GÜNTHER,
- ‘“CHALLENGER” REPORTS’ 156
-
- 22 _Saccopharynx ampullaceus._ AFTER GÜNTHER,
- ‘“CHALLENGER” REPORTS’ 164
-
-
-
-
- THE FAUNA OF THE DEEP SEA
-
-
-
-
- CHAPTER I
- A SHORT HISTORY OF THE INVESTIGATIONS
-
-
-Our knowledge of the natural history of the deep seas may be said to
-have commenced not more than fifty years ago. There are, it is true, a
-few fragments of evidence of a fauna existing in depths of more than a
-hundred fathoms to be found in the writings of the earlier navigators,
-but the methods of deep-sea investigation were so imperfect in those
-days that naturalists were disposed to believe that in the abysses of
-the great oceans life was practically non-existent.
-
-Even Edward Forbes just before his death wrote of an abyss ‘where life
-is either extinguished or exhibits but a few sparks to mark its
-lingering presence,’ but in justice to the distinguished naturalist it
-should be remarked that he adds, ‘Its confines are yet undetermined, and
-it is in the exploration of this vast deep-sea region that the finest
-field for submarine discovery yet remains.’
-
-Forbes was only expressing the general opinion of naturalists of his
-time when he refers with evident hesitation to the existence of an azoic
-region. His own dredging excursions in depths of over one hundred
-fathoms proved the existence of many peculiar species that were
-previously unknown to science. ‘They were like,’ he says, ‘the few stray
-bodies of strange red men, which tradition reports to have been washed
-on the shores of the Old World before the discovery of the New, and
-which served to indicate the existence of unexplored realms inhabited by
-unknown races, but not to supply information about their character,
-habits, and extent.’
-
-In the absence of any systematic investigation of the bottom of the deep
-sea, previous to Forbes’s time the only information of deep-sea animals
-was due to the accidental entanglement of certain forms in sounding
-lines, or to the minute worms that were found in the mud adhering to the
-lead.
-
-As far back as 1753, Ellis described an Alcyonarian that was brought up
-by a sounding line from a depth of 236 fathoms within eleven degrees of
-the North Pole by a certain Captain Adriaanz of the ‘Britannia.’ The
-specimen was evidently an _Umbellula_, and it is stated that the arms
-(i.e. Polyps) were of a bright yellow colour and fully expanded when
-first brought on deck.
-
-In 1819 Sir John Ross published an account of his soundings in Baffin’s
-Bay, and mentions the existence of certain worms in the mud brought from
-a depth of 1,000 fathoms, and a fine Caput Medusæ (Astrophyton)
-entangled on the sounding line at a depth of 800 fathoms.
-
-In the narrative of the voyage of the ‘Erebus’ and ‘Terror,’ published
-in 1847, Sir James Ross calls attention to the existence of a deep-sea
-fauna, and makes some remarks on the subject that in the light of modern
-knowledge are of extreme interest. ‘I have no doubt,’ he says, ‘that
-from however great a depth we may be enabled to bring up the mud and
-stones of the ocean, we shall find them teeming with animal life.’ This
-firm belief in the existence of an abysmal fauna was not, as it might
-appear from the immediate context of the passage I have quoted, simply
-an unfounded speculation on his part, but was evidently the result of a
-careful and deliberate chain of reasoning, as may be seen from the
-following passage that occurs in another part of the same book:—‘It is
-well known that marine animals are more susceptible of change of
-temperature than land animals; indeed they may be isothermally arranged
-with great accuracy. It will, however, be difficult to get naturalists
-to believe that these fragile creatures could possibly exist at the
-depth of nearly 2,000 fathoms below the surface; yet as we know they can
-bear the pressure of 1,000 fathoms, why may they not of two? We also
-know that several of the same species of creatures inhabit the Arctic
-that we have fished up from great depths in the Antarctic seas. The only
-way they could get from one pole to the other must have been through the
-tropics; but the temperature of the sea in those regions is such that
-they could not exist in it, unless at a depth of nearly 2,000 fathoms.
-At that depth they might pass from the Arctic to the Antarctic Ocean
-without a variation of five degrees of temperature; whilst any land
-animal, at the most favourable season, must experience a difference of
-fifty degrees, and, if in the winter, no less than 150 degrees of
-Fahrenheit’s thermometer—a sufficient reason why there are neither
-quadrupeds, nor birds, nor land insects common to both regions.’
-
-In the year 1845, Goodsir succeeded in obtaining a good haul in Davis
-Straits, at a depth of 300 fathoms. It included Mollusca, Crustacea,
-Asterids, Spatangi, and Corallines.
-
-In 1848, Lieutenant Spratt read a paper at the meeting of the British
-Association at Swansea, on the influence of temperature upon the
-distribution of the fauna in the Ægean seas, and at the close of this
-paper we find the following passage, confirming in a remarkable way the
-work of previous investigators in the same field. He says: ‘The greatest
-depth at which I have procured animal life is 390 fathoms, but I believe
-that it exists much lower, although the general character of the Ægean
-is to limit it to 300 fathoms; but as in the deserts we have an oasis,
-so in the great depths of 300, 400, and perhaps 500 fathoms we may have
-an oasis of animal life amidst the barren fields of yellow clay
-dependent upon favourable and perhaps accidental conditions, such as the
-growth of nullipores, thus producing spots favourable for the existence
-and growth of animal life.’
-
-The next important discovery was that of the now famous Globigerina mud
-by Lieutenants Craven and Maffit, of the American Coast survey, in 1853,
-by the help of the sounding machine invented by Brooke. This was
-reported upon by Professor Bailey.
-
-Further light was thrown upon the deep-sea fauna by Dr. Wallich in 1860,
-on board H.M.S. ‘Bulldog’, by the collection of thirteen star-fish
-living at a depth of 1,260 fathoms.
-
-Previous to this Torell, during two excursions to the Northern seas, had
-proved the existence of an extensive marine fauna in 300 fathoms, and
-had brought up with the ‘Bulldog’ machine many forms of marine
-invertebrates from depths of over 1,000 fathoms; but it was not until
-1863, when Professor Lovén read a report upon Torell’s collections, that
-these interesting and valuable investigations became known to
-naturalists.
-
-Nor must mention be omitted of the remarkable investigations of Sars and
-his son, the pioneers of deep-sea zoology on the coasts of Norway, who,
-by laborious work commenced in 1849, failed altogether to find any
-region in the deep water where animal life was non-existent, and indeed
-were the first to predict an extensive abysmal fauna all over the floor
-of the great oceans. One of the many remarkable discoveries made by Sars
-was _Rhizocrinus_, a stalked Crinoid.
-
-Ever since that time the Norwegians and the Swedes have been most
-energetic in their investigations, and the publications of the results
-of the Norske Nord-havns expeditions are regarded by all naturalists as
-among the most valuable contributions to our knowledge of the deep-sea
-fauna.
-
-Notwithstanding the previous discovery of many animals that undoubtedly
-came from the abysmal depths of the ocean, there were still some persons
-who found a difficulty in believing that animal life could possibly
-exist under the enormous pressure of these great depths. It was
-considered to be more probable that these animals were caught by the
-dredge or sounding lines in their ascent or descent; and that they were
-merely the representatives of a floating fauna living a few fathoms
-below the surface.
-
-The first direct proof of the existence of an invertebrate fauna in deep
-seas was found by the expedition that was sent to repair the submarine
-cable of the Mediterranean Telegraph Company. The cable had broken in
-deep water, and a ship was sent out to examine and repair the damage.
-When the broken cable was brought on deck, it bore several forms of
-animal life that must have become attached to it and lived at the bottom
-of the sea in water extending down to a depth of 1,200 fathoms. Among
-other forms a _Caryophyllia_ was found attached to the cable at 1,100
-fathoms, an oyster (_Ostrea cochlear_), two species of Pecten, two
-gasteropods, and several worms.
-
-The discoveries that had been made indicating the existence of a
-deep-sea fauna led to the commission of H.M. ships ‘Lightning’ and
-‘Porcupine,’ and the systematic investigation that was made by the
-naturalists on these vessels brought home to the minds of naturalists
-the fact that there is not only an abysmal fauna, but that in places
-this deep-sea fauna is very rich and extensive. The ‘Lightning’ was
-despatched in the spring of 1868 and carried on its investigations in
-the neighbourhood of the Faeroe Islands, but the vessel was not suitable
-for the purpose and met with bad weather. The results, however, were of
-extreme importance; for, besides solving many important points
-concerning the distribution of ocean temperature, ‘it had been shown
-beyond question that animal life is both varied and abundant at depths
-in the ocean down to 650 fathoms at least, notwithstanding the
-extraordinary conditions to which animals are there exposed.’
-
-Among the remarkable animals dredged by the ‘Lightning’ were the curious
-Echinoderm, _Brisinga coronata_, previously discovered by Sars, and the
-Hexactinellid sponges, _Holtenia_ and _Hyalonema_, the Crinoids
-_Rhizocrinus_ and _Antedon celticus_, and the Pennatulid _Bathyptilum
-Carpenteri_, not to mention numerous Foraminifera new to science.
-
-In the spring of the following year, 1869, the Lords Commissioners of
-the Admiralty despatched the surveying vessel ‘Porcupine’ to carry on
-the work commenced by the ‘Lightning.’
-
-The first cruise was on the west coast of Ireland, the second cruise to
-the Bay of Biscay, where dredging was satisfactorily carried on to a
-depth of 2,435 fathoms, and the third in the Channel between Faeroe and
-Scotland.
-
-The dredging in 2,435 fathoms was quite successful, and the dredge
-contained several Mollusca, including new species of _Dentalium_,
-_Pecten_, _Dacrydium_, &c., numerous Crustacea and a few Annelids and
-Gephyrea, besides Echinoderma and Protozoa. A satisfactory dredging was
-also made in 1,207 fathoms.
-
-The third cruise was also successful and brought many new species to
-light, including the _Porocidaris purpurata_, and a remarkable heart
-urchin, _Pourtalesia Jeffreysi_.
-
-Concerning Pourtalesia Sir Wyville Thomson says:—
-
-‘The remarkable point is that, while we had so far as we were aware no
-living representative of this peculiar arrangement of what is called
-“disjunct” ambulacra, we have long been acquainted with a fossil
-family—the Dysasteridæ—possessing this character. Many species of the
-genera Dysaster, Collyrites, &c., are found from the lower oolite to the
-white chalk, but there the family had previously been supposed to have
-become extinct.’
-
-The discovery of two new Crinoids led to the anticipation that the
-Crinoidea, the remarkable group of Echinoderma, supposed at the time to
-be on the verge of extinction, probably form rather an important element
-in the abysmal fauna.
-
-One of the most interesting results was the discovery of three genera in
-deep water, _Calveria_, _Neolampas_ and _Pourtalesia_, almost
-immediately after they were discovered by Pourtales in deep water on the
-coasts of Florida, showing thus a wide lateral distribution and
-suggesting a vast abysmal fauna.
-
-A year before the ‘Lightning’ was despatched, Count Pourtales had
-commenced a series of investigations of the deep-sea fauna off the coast
-of Florida. The first expedition started in 1867 from Key West for the
-purpose of taking some dredgings between that port and Havana.
-Unfortunately yellow fever broke out on board soon after they started,
-and only a few dredgings were taken. However, the results obtained were
-of such importance that they encouraged Pourtales to undertake another
-expedition and enabled him to say very positively ‘that animal life
-exists at great depths, in as great a diversity and as great an
-abundance as in shallow water.’
-
-In the following years, 1868 and 1869, the expeditions were more
-successful, and many important new forms were found in water down to 500
-fathoms. Perhaps the most interesting result obtained was the discovery
-of _Bourguetticrinus_ of D’Orbigny; it may even be the species named by
-him which occurs fossil in a recent formation in Guadeloupe.
-
-By this time the interest of scientific men was thoroughly excited over
-the many problems connected with this new field of work. The prospect of
-obtaining a large number of new and extremely curious animals, the faint
-hope that living Trilobites, Cystids, and other extinct forms might be
-discovered, and lastly the desire to handle and investigate great masses
-of pure protoplasm in the form of the famous but unfortunately
-non-existent Bathybius, induced some men of wealth and leisure to spend
-their time in deep-sea dredging, and stimulated the governments of some
-civilised countries to lend their aid in the support of expeditions for
-the deep-sea survey.
-
-Mr. Marshall Hall’s yacht, the ‘Norma,’ was employed for some time in
-this work, and an extensive collection of deep-sea animals was made.
-About the same time Professor L. Agassiz was busy on board the American
-ship, the ‘Hassler,’ in continuing the work of Count Pourtales, and
-later on the Germans fitted out the ‘Gazelle,’ and the French the still
-more famous ‘Travailleur’ and ‘Talisman’ expeditions. Nor must we omit
-to mention in this connection the cruise of the Italian vessel, the
-‘Vittor Pessani,’ nor those of the British surveying vessels, the
-‘Knight Errant’ and the ‘Triton,’ and the American vessels, ‘The Blake’
-and the ‘Fish Hawk.’
-
-But of all these expeditions, by far the most complete in all the
-details of equipment, and the arrangements made for the publication of
-the results, was the expedition fitted out in 1873 by the British
-Government. The voyage of H.M.S. ‘Challenger’ is so familiar to all who
-take an interest in the progress of scientific discovery, that it is not
-necessary to do more than make a passing mention of it in this place.
-The excellent books that were written by Wyville Thomson, by Moseley,
-and by other members of the staff, have made the general reader familiar
-with the narrative of that remarkable cruise and the most striking of
-the many scientific discoveries that were made; while the numerous large
-monographs that have been published during the past fourteen years give
-opportunities to the naturalist of obtaining all the requisite
-information concerning the detailed results of the expedition.
-
-The expenditure of the large sum of money upon this expedition and the
-publication of its reports has been abundantly justified. The
-information obtained by the ‘Challenger’ will be for many years to come
-the nucleus of our knowledge of the deep-sea fauna, the centre around
-which all new facts will cluster, and the guide for further
-investigations.
-
-To say that the ‘Challenger’ accomplished all that was expected or
-required would be to over-estimate the value of this great expedition,
-but nevertheless it is difficult for us, even now, thoroughly to grasp
-the importance of the results obtained or to analyse and classify the
-numerous and very remarkable facts that were gained during her four
-years’ cruise.
-
-It is, of course, impossible, in a few lines, to give a summary of the
-more important of the Natural History results of the ‘Challenger’
-expedition. Besides proving the existence of a fauna in the sea at all
-depths and in all regions, the expedition further proved that the
-abysmal fauna, taken as a whole, does not possess characters similar to
-those of the fauna of any of the secondary or even tertiary rocks. A few
-forms, it is true, known to us up to that time only as fossils, were
-found to be still living in the great depths, but a large majority of
-the animals of these regions were found to be new and specially modified
-forms of the families and genera inhabiting shallow waters of modern
-times. No Trilobites, no Blastoids, no Cystoids, no new Ganoids, and
-scarcely any deep-sea Elasmobranchs were brought to light, but the fauna
-was found to consist mainly of Teleosteans, Crustacea, Cœlentera, and
-other creatures unlike anything known to have existed in Palæozoic
-times, specially modified in structure for their life in the great
-depths of the ocean.
-
-In 1876 the S.S. ‘Vöringin’ was chartered by the Norwegian Government
-and was dispatched to investigate the tract of ocean lying between
-Norway, the Faeroe islands, Jan Mayen, and Spitzbergen. The
-investigations extended over three years, the vessel returning to Bergen
-in the winter months.
-
-The civilian staff of the ‘Vöringin’ included Professors H. Mohn,
-Danielssen, and G. O. Sars, and the expedition was successful in
-obtaining a large number of animals from deep water by means of the
-dredge and tangles and by the trawl.
-
-The results of this expedition have been published in a series of large
-quarto volumes under the general title of the Norske Nord-havns
-Expedition.
-
-The most interesting forms brought to light by the Norwegians are the
-two genera _Fenja_ and _Aegir_, animals possessing the general form of
-sea anemones but distinguished from all Cœlenterates by the presence of
-a continuous and straight gut reaching from the mouth to the aboral
-pores which completely shuts off the cœlenteron or general body cavity
-from the stomodæum.
-
-In more recent times the work has been by no means neglected. With the
-advantage of employing many modern improvements in the dredges and
-trawls in use, the American steamer, the ‘Albatross,’ has been engaged
-in a careful investigation of the deep-sea fauna of the eastern slopes
-of the Pacific Ocean, while at the same time Her Majesty’s surveying
-vessel, the ‘Investigator,’ has been obtaining some interesting and
-valuable results from a survey of the deep waters of the Indian Ocean.
-But our knowledge of this vast and wonderful region is still in its
-infancy. We have gathered, as it were, only a few grains from a great
-unknown desert. It is true that we may not for many years, if ever,
-obtain any results that will cause the same deep interest and excitement
-to the scientific public as those obtained by the first great national
-expeditions, but there are still many important scientific problems that
-may be and will be solved by steady perseverance in this field of work,
-and if we can only obtain the same generous support from public
-institutions and from those in charge of national funds that we have
-received in the past two decades, many more important facts will
-doubtless be brought to light.
-
-
-
-
- CHAPTER II
- THE PHYSICAL CONDITIONS OF THE ABYSS
-
-
-It is not surprising that the naturalists of the early part of the
-present century could not believe in the existence of a fauna at the
-bottom of the deep seas.
-
-The extraordinary conditions of such a region—the enormous pressure, the
-absolute darkness, the probable absence of any vegetable life from want
-of direct sunlight—might very well have been considered sufficient to
-form an impassable barrier to the animals migrating from the shallow
-waters and to prevent the development of a fauna peculiarly its own.
-
-The fragmentary accounts of animals brought up by sounding lines from
-great depths might, it is true, have thrown doubts on the current views;
-but they were not of sufficient importance in themselves, nor were the
-observations made with such regard to the possibility of error, as to
-withstand the critical remarks that were made to explain them away.
-
-The absence of any evidence obtained by accurate systematic research,
-together with the consideration of the physical character of the ocean
-bed, were quite sufficient to lead scientific men of that period to
-doubt the existence of any animal life in water deeper than a few
-hundred fathoms.
-
-We now know, however, that there is a very considerable fauna at
-enormous depths in all the great oceans, and we have acquired, moreover,
-considerable information concerning some of those peculiar physical
-conditions of the abyss that fifty years ago were merely matters of
-speculation among scientific men.
-
-The relation between animals and their environment is now a question of
-such great interest and importance that it is necessary in any
-description of the fauna of a particular region to consider its physical
-conditions and the influence that it may be supposed to have had in
-producing the characteristics of the fauna.
-
-The peculiar physical conditions of the deep seas may be briefly stated
-to be these: It is absolutely dark so far as actual sunlight is
-concerned, the temperature is only a few degrees above freezing point,
-the pressure is enormous, there is little or no movement of the water,
-the bottom is composed of a uniform fine soft mud, and there is no plant
-life.
-
-All of these physical conditions we can appreciate except the enormous
-pressure. Absolute darkness we know, the temperature of the deep seas is
-not an extraordinary one, the absence of movement in the water and the
-fine soft mud are conditions that we can readily appreciate; but the
-pressure is far greater than anything we can realise.
-
-At a depth of 2,500 fathoms the pressure is, roughly speaking, two and a
-half tons per square inch—that is to say, several times greater than the
-pressure exerted by the steam upon the piston of our most powerful
-engines. Or, to put the matter in other words, the pressure per square
-inch upon the body of every animal that lives at the bottom of the
-Atlantic Ocean is about twenty-five times greater than the pressure that
-will drive a railway train.
-
-A most beautiful experiment to illustrate the enormous force of this
-pressure was made during the voyage of H.M.S. ‘Challenger.’ I give the
-description of it in the words of the late Professor Moseley.
-
-‘Mr. Buchanan hermetically sealed up at both ends a thick glass tube
-full of air, several inches in length. He wrapped this sealed tube in
-flannel, and placed it, so wrapped up, in a wide copper tube, which was
-one of those used to protect the deep-sea thermometers when sent down
-with the sounding apparatus.
-
-‘This copper tube was closed by a lid fitting loosely, and with holes in
-it, and the copper bottom of the tube similarly had holes bored through
-it. The water thus had free access to the interior of the tube when it
-was lowered into the sea, and the tube was necessarily constructed with
-that object in view, in order that in its ordinary use the water should
-freely reach the contained thermometer.
-
-‘The copper case containing the sealed glass tube was sent down to a
-depth of 2,000 fathoms and drawn up again. It was then found that the
-copper wall of the case was bulged and bent inwards opposite the place
-where the glass tube lay, just as if it had been crumpled inward by
-being violently squeezed.
-
-‘The glass tube itself, within its flannel wrapper, was found when
-withdrawn, reduced to a fine powder, like snow almost. What had happened
-was that the sealed glass tube, when sinking to gradually increasing
-depths, had held out long against the pressure, but this at last had
-become too great for the glass to sustain, and the tube had suddenly
-given way and been crushed by the violence of the action to a fine
-powder. So violent and rapid had been the collapse that the water had
-not had time to rush in by means of the holes at both ends of the copper
-cylinder and thus fill the empty space left behind by the collapse of
-the glass tube, but had instead crushed in the copper wall and brought
-equilibrium in that manner. The process is exactly the reverse of an
-explosion, and is termed by Sir Wyville Thomson an “implosion.”’
-
-It is but reasonable to suppose that the ability to sustain this
-enormous pressure can only be acquired by animals after generations of
-gradual migrations from shallow waters. Those forms that are brought up
-by the dredge from the depths of the ocean are usually killed and
-distorted by the enormous and rapid diminution of pressure in their
-journey to the surface, and it is extremely probable that shallow-water
-forms would be similarly killed and crushed out of shape were they
-suddenly plunged into very deep water. The fish that live at these
-enormous depths are in consequence of the enormous pressure liable to a
-curious form of accident. If, in chasing their prey or for any other
-reason, they rise to a considerable distance above the floor of the
-ocean, the gases of their swimming bladder become considerably expanded
-and their specific gravity very greatly reduced. Up to a certain limit
-the muscles of their bodies can counteract the tendency to float upwards
-and enable the fish to regain its proper sphere of life at the bottom;
-but beyond that limit the muscles are not strong enough to drive the
-body downwards, and the fish, becoming more and more distended as it
-goes, is gradually killed on its long and involuntary journey to the
-surface of the sea. The deep-sea fish, then, are exposed to a danger
-that no other animals in this world are subject to, namely that of
-tumbling upwards.
-
-That such accidents do occasionally occur is evidenced by the fact that
-some fish, which are now known to be true deep-sea forms, were
-discovered dead and floating on the surface of the ocean long before our
-modern investigations were commenced.
-
-Until quite recently, every one agreed that no rays of sunlight could
-possibly penetrate the sea to a greater depth than a few hundred
-fathoms.
-
-Moseley says that ‘probably all is dark below 200 fathoms excepting in
-so far as light is given out by phosphorescent animals,’ and Wyville
-Thomson speaks of the ‘utter darkness of the deep-sea bottom.’
-
-Within the last few years a few authors have maintained that it is quite
-possible that a few rays of sunlight do penetrate even to the greatest
-depths of the ocean—a view mainly based on the fact that so many
-deep-sea animals possess extremely perfect and complicated eyes and very
-brilliant colours. Verrill says: ‘It seems to me probable that more or
-less sunlight does actually penetrate to the greatest depths of the
-ocean, in the form of a soft sea-green light, perhaps at 2,000 or 3,000
-fathoms equal in intensity to our partially moonlight nights and
-possibly at the greatest depths equal only to starlight. It must be
-remembered that in the deep sea far away from land the water is far more
-transparent than near the coast.’ Packard is of a similar opinion.
-
-There seem to me to be very slight grounds for this view. The fact that,
-comparatively speaking, shallow-water fish avoid nets that are rendered
-phosphorescent by entangled jelly-fish does not justify us in assuming
-that deep-sea fish avoid regions where there are phosphorescent
-Gorgonians or Pennatulids. It is not by any means certain that fish
-avoid sunken nets on account of their phosphorescence. Most fish
-possess, as is well known, a very acute sense of smell, and it is very
-probable that they avoid such nets on account of the putrid odours of
-the dead animals that remain attached to them.
-
-Nor is there much strength in the further argument that it can hardly be
-possible that there can be an amount of phosphorescent light regularly
-evolved by the few deep-sea animals, having this power, sufficient to
-cause any general illumination, or powerful enough to have influenced,
-over the whole ocean, the evolution of complex eyes, brilliant and
-complex protective colours, and complex commensal adaptations.
-
-We have no sound information to go upon to be able to judge of the
-amount of light given off by phosphorescent animals at the bottom of the
-deep sea. The faint light they show on deck after their long journey
-from the depths in which they live to the surface may be extremely small
-compared with the light they give in their natural home under a pressure
-of 2½ tons to the square inch. The complex eyes that many deep-sea
-animals exhibit were almost certainly not evolved as such, but are
-simple modifications of eyes possessed by a shallow-water ancestry.
-
-The more recent experiments that have been made, tend to show that no
-sunlight whatever penetrates to a greater depth, to take an extreme
-limit, than 500 fathoms.
-
-Fol and Sarasin, experimenting with very sensitive bromo-gelatine
-plates, found that there was no reaction after ten minutes’ exposure at
-a depth of 400 metres on a sunny day in March.
-
-But although it is highly probable that not a glimmer of sunlight ever
-penetrates to the depths of the ocean, there is in some places,
-undoubtedly, a very considerable illumination due to the phosphorescence
-of the inhabitants of the deep waters.
-
-All the Alcyonarians are, according to Moseley, brilliantly
-phosphorescent when brought to the surface. Many deep-sea fish possess
-phosphorescent organs, and it is quite possible that many of the
-deep-sea Protozoa, Tunicates, Jelly-fish, and Crustacea are in their
-native haunts capable of giving out a very considerable amount of
-phosphorescent light.
-
-If we may be allowed to compare the light of abysmal animals with that
-of surface forms, we can readily imagine that some regions of the sea
-may be as brightly illuminated as a European street is at night—an
-illumination with many very bright centres and many dark shadows, but
-quite sufficient for a vertebrate eye to distinguish readily and at a
-considerable distance both form and colour.
-
-To give an example of the extent to which the illumination due to
-phosphorescent organisms may reach, I may quote a passage from the
-writings of the late Sir Wyville Thomson.
-
-‘After leaving the Cape Verde Islands the sea was a perfect blaze of
-phosphorescence. There was no moon, and although the night was perfectly
-clear and the stars shone brightly, the lustre of the heavens was fairly
-eclipsed by that of the sea. It was easy to read the smallest print,
-sitting at the after-port in my cabin, and the bows shed on either side
-rapidly widening wedges of radiance so vivid as to throw the sails and
-rigging into distinct lights and shadows.’
-
-A very similar sight may frequently be seen in the Banda seas, where on
-calm nights the whole surface of the ocean seems to be a sheet of milky
-fire. The light is not only to be seen where the crests of waves are
-breaking, or the surface disturbed by the bows of the boat, but the
-phosphorescence extends as far as the eye can reach in all directions.
-It is impossible, of course, to say with any degree of certainty whether
-phosphorescence such as this exists at the bottom of the deep sea, but
-it is quite probable that it does in some places, and hence the
-well-developed eyes and brilliant colours of some of the deep-sea
-animals.
-
-On the other hand the entire absence or rudimentary condition of the
-eyes of a very considerable proportion of deep-sea animals seems to
-prove that the phosphorescent illumination is not universally
-distributed, and that there must be some regions in which the darkness
-is so absolute that it can only be compared with the darkness of the
-great caves.
-
-It is difficult to believe that the eyes of such animals as crabs and
-prawns for example would undergo degeneration if there were a glimmer of
-light in their habitat, a light even so faint as that of a starlight
-night in shallow water. With the faintest light the eyes would be of use
-to them in seeking their prey, avoiding their enemies, and finding their
-mates, and any diminution in the keenness of this sense would probably
-be of considerable disadvantage to them and tend to their ultimate
-extinction.
-
-It might be argued that the animals of the abysses of the ocean probably
-feed chiefly upon the carcases of pelagic animals that have fallen from
-the upper regions of the sea, and that the sense of smell is probably
-the most important for them in searching for their food. That is quite
-probable; but many shallow-water animals invariably seek their food by
-their sense of smell without showing any traces of a weakness in their
-sense of sight. It may be taken as an axiom of biology that unless a
-particular sense is absolutely useless to an animal or a positive
-disadvantage to it, that sense will be retained.
-
-It may be stated then with some confidence that in the abysmal depths of
-the ocean there is no trace of sunlight. It is highly improbable, on the
-face of it, that any ray of light could penetrate through a stratum of
-water four miles in thickness, even if the water were perfectly pure and
-clear, but when we remember that the upper regions, at least, are
-crowded with pelagic organisms provided with skeletons of lime and
-silica, we may justly consider that it is impossible.
-
-The temperature of the water in the abyss is by no means constant for a
-constant depth nor does it vary with the latitude. It is true that, as a
-rule, the water is colder at greater depths than in shallower ones, and
-that the deeper the thermometer is lowered into the sea, the lower the
-mercury sinks. This is consistent with physical laws. If there is any
-difference at all in the temperature of a column of water that has had
-time to settle, the thermometer will always reach its highest point at
-the top of the column and its lowest at the bottom, for the colder
-particles being of greater specific gravity than the warmer ones will
-sink, and the warmer ones will rise.
-
-The truth of this will be clear if we imagine a locality at the bottom
-of a deep ocean with a source of great heat such as an active volcano.
-
-Such a source of heat would, it is true, raise the temperature of the
-water in its immediate vicinity, but the particles of water thus heated
-would immediately commence to rise through the superjacent layers of
-colder water, and colder particles would fall to take their places. Thus
-the effect of an active volcano at the bottom of the deep sea would not
-be apparent at any very great distance in the same plane. In fact,
-unless the bottom of the ocean was closely studded with volcanoes we
-should expect to find, as indeed we do find, that the temperature of the
-sea rises as the water shallows.
-
-If then we were to consider a great ocean as simply a huge basin of
-water, we should expect to find the water at the surface warmer than the
-water at the bottom. The temperature of the surface would vary
-constantly with the temperature of the air above it. That is to say, it
-would be warmer at the equator than in the temperate regions. The
-temperature at the bottom would be the same as the lowest temperature of
-the basin, that is, of the earth that supports it.
-
-The great oceans however cannot be regarded as simple basins of water
-such as this. The temperature of the surface water varies only
-approximately with the latitude. It is generally speaking hottest at the
-equator and coldest at the poles, but surface currents in the
-intermediate regions produce many irregularities in the surface
-temperature.
-
-Again, although we have no means of knowing what the temperature of the
-earth is at 1,000 fathoms below the surface of the ocean, it is very
-probable that in the great oceans the temperature of the deepest stratum
-of water is considerably lower than the true earth temperature. This is
-due to currents of cold water constantly flowing from the poles towards
-the equator. If these polar currents were at any time to cease, the
-temperature of the lowest strata of water would rise.
-
-Although the polar currents cannot be actually demonstrated nor their
-exact rapidity be accurately determined, the deduction from the known
-facts of physical geography that they do actually exist is perfectly
-sound and beyond dispute. A few considerations will, I think, make this
-clear.
-
-If the ocean were a simple basin somewhat deeper at the equator than at
-the poles, the cold water at the poles would gradually sink down the
-slopes of the basin towards the latitude of the equator, and the bottom
-temperature of the water would be constant all the world over.
-
-A few hills here and there would not affect the general statement that
-for a constant depth the temperature of the lowest stratum of water
-would be constant.
-
-But in some places ridges occur stretching across the oceans from
-continent to continent, and these ridges shut off the cold water at the
-bottom of the sea on the polar side from reaching the bottom of the sea
-on the equator side.
-
-If A (fig. 1) represents a ridge stretching from continent to continent
-across an ocean, and the arrow represents the direction of the current,
-then the water that flows across the ridge from the polar side to the
-equator side will be drawn from the layers of water lying above the
-level of the ridge, and consequently none of the coldest water will ever
-get across it, and from the level of the ridge to the bottom of the sea
-on the equatorial side the water will have the same temperature as the
-water at the level of the ridge on the polar side.
-
-It follows from this that in places where there are deep holes in the
-bed of the ocean surrounded on all sides by considerable elevations, the
-temperature of the water at the bottom will be the same as the
-temperature of the water on the summit of the lowest ridges that
-surround them.
-
-[Illustration: FIG. 1.—Diagram illustrating the passage of an ocean
-current across a barrier (A).]
-
-This explains why it is that we find that the bottom temperature for a
-given depth is frequently less in one place than it is in another, even
-in places of the same parallel of latitude. One or two examples may be
-taken to illustrate these points. The temperature off Rio Janeiro in
-lat. 20° S. was found by the ‘Challenger’ to be 0·6° C. at a depth of
-2,150 fathoms. In a similar latitude north of the equator at a depth of
-2,900 fathoms the temperature was found to be 2·2° C., and at a point
-near Porto Rico there is a deep hole of 4,561 fathoms with a bottom
-temperature of 2·2° C.
-
-Again it has been shown by the American expedition that the temperature
-of the water at the deepest point in the Gulf of Mexico, 2,119 fathoms,
-is the same as that of the bottom of the Straits of Yucatan, 1,127
-fathoms, namely 4·1° C. And, passing to another part of the world
-altogether, we find in the small but deep sea that lies between the
-Philippines and Borneo that, at a depth of 2,550 fathoms, the
-temperature is 10·2° C.
-
-These facts then show that, although at the bottom of the deep seas the
-water is always very cold, the degree of coldness is by no means
-constant in the same latitude for the same depth.
-
-We must now return to the polar currents. We have assumed above that
-these currents do exist, and it is probable that by this time the reader
-must have seen why they are assumed to exist.
-
-The water at the bottom of the ocean is exceedingly cold. Where does
-this coldness come from? It is obvious that in temperate and tropical
-climes it does not come from the surface. Nor is it at all probable that
-it comes from the earth upon which the water rests; for, if it were so,
-the temperature for water of a given depth would always be the same. We
-should not find the bottom temperature of 0·4° C. at 2,900 fathoms off
-Rio de la Plata and a temperature of 2·2° F. in 4,561 fathoms off Porto
-Rico.
-
-In fact the only hypothesis that can with any show of reason be put
-forward to account for the temperature of the bottom of the ocean is
-that which derives its coldness from the Polar ice.
-
-We have at present very little evidence to enable us to judge of the
-force and direction of the polar currents in the two hemispheres, but
-the researches of the ‘Challenger’ prove almost conclusively that in the
-Atlantic Ocean there is a very strong predominance of the Antarctic
-polar current. In fact it seems very probable that the Arctic polar
-current, if it exist at all, is very small and confined to the eastern
-and western shores of the North Atlantic.
-
-It is very probable, however, that these currents at the bottom of the
-ocean are extremely slow, and, as the water is never affected by tides
-or storms, the general character of the deep sea is probably one of calm
-repose. This is a matter of no little importance; for, in the
-consideration of the characters presented by the fauna of any particular
-region, it is always necessary to take into account the physical
-difficulties the animals have to contend against and the modifications
-of structure they present to combat these difficulties.
-
-Thus in a region such as that presented by the deep sea, where there are
-no rapid tides, we should not expect to find such a powerful set of body
-muscles in the free-swimming forms nor such a firm vertebral column as
-in the animals that live in more lively water.
-
-Perhaps it is of the nature of an assumption to say that there are no
-rapid currents and tides in the abysmal depths of the ocean, for we have
-no means of demonstrating or even of calculating the rate of flow of
-these waters. But it is a reasonable hypothesis and one that we may well
-use until the contrary is proved.
-
-A fact of some importance that supports this hypothesis, as regards some
-parts of the ocean at least, is presented by the sea-anemones.
-
-Many of the shallow-water Actinians are known to possess minute slits in
-the tentacles and disc, affording a free communication between the
-general body cavity or cœlenteron and the exterior.
-
-In many deep-sea forms the tentacles are considerably shorter and the
-apertures larger than they are in shallow-water forms. It is difficult
-to believe that such forms, perforated by, comparatively speaking, large
-holes, could manage to live in rapidly flowing water, for if they did so
-they would soon be smothered by the fine mud that composes the floor of
-all the deep seas. In fact anemones of the type presented by such forms
-as _Sicyonis crassa_ are only fitted for existence in sluggish or still
-water.
-
-[Illustration: FIG. 2.—_Sicyonis crassa._ M, mouth; S, ciliated groove;
-T, tentacles. Each tentacle is perforated by a single large aperture.
-(After Hertwig.)]
-
-Another character that must be taken into consideration is that
-presented by the floor of the great oceans. The floor of the ocean, if
-it were laid bare, would probably present a vast undulating plain of
-fine mud. Not a rock, not even a stone would be visible for miles.
-
-The mud varies in different parts of the globe according to the depth,
-the proximity to land, the presence of neighbouring volcanoes or the
-mouths of great rivers.
-
-The Globigerina ooze is perhaps the best known of all the different
-deep-sea deposits. It was discovered and first described by the officers
-of the American Coast Survey in 1853. It is found in great abundance in
-the Atlantic Ocean in regions shallower than 2,200 fathoms. Deeper than
-this, it gradually merges into the ‘Red mud.’ It is mainly composed of
-the shells of Foraminifera, and of these the different species of
-Globigerina are the most abundant. It is probably formed partly by the
-shells of the dead Foraminifera that actually live on the bottom of the
-ocean and partly by the shells of those that live near the surface or in
-intermediate depths and fall to the bottom when their lives are done.
-
-So abundant are the shells of these Protozoa that nearly 95 per cent. of
-the Globigerina ooze is composed of carbonate of lime. The remaining
-five per cent. is composed of sulphate and phosphate of lime, carbonate
-of ammonia, the oxides of iron and manganese, and argillaceous matters.
-The oxides of iron and manganese are probably of meteoric origin; the
-argillaceous matter may be due to the trituration of lumps of pumice
-stone and to the deposits caused by dust storms.
-
-[Illustration: FIG. 3.—Globigerina ooze. (After Agassiz.)]
-
-Globigerina ooze may be found on the floor of the ocean at depths
-ranging from 500 to 2,800 fathoms of water in equatorial and temperate
-latitudes. The reason that it is not found in Arctic seas may be that
-the cold surface waters of these regions do not bear such an abundant
-fauna of Foraminifera. This is supported by the fact that it extends ten
-degrees further north than south in the Atlantic, the warm water of the
-Gulf Stream bearing a richer fauna than the waters of a corresponding
-degree of latitude in the Southern Sea.
-
-The Pteropod ooze has only twenty-five per cent. of carbonate of lime.
-It contains numerous shells of various Pteropods, Heteropods, and
-Foraminifera, but nearly fifty per cent. of its substance is composed of
-the siliceous skeletons of Radiolaria and the frustules of diatoms.
-
-According to Murray it is found in tropical and subtropical seas at
-depths of less than 1,500 fathoms.
-
-The Radiolarian ooze is found only in the deepest waters of the Central
-and Western Pacific Ocean. In some of the typical examples, not a trace
-of carbonate of lime was to be found, but in somewhat shallower waters a
-few small fragments occurred.
-
-A Diatom ooze, mainly composed of the skeletons of diatoms, has also
-been found in deep water near the Antarctic Circle, but it has not
-apparently a very wide range.
-
-Of all the deep-sea deposits, however, the so-called ‘Red mud’ has by
-far the widest distribution. It is supposed to extend over one-third of
-the earth’s surface. It is essentially a deep-sea deposit, and one that
-is found in its typical condition at some considerable distance from
-continental land. Like the Globigerina ooze it is never found in
-enclosed seas. To the touch it is plastic and greasy when fresh, but it
-soon hardens into solid masses. When examined with the microscope it is
-seen to be composed of extremely minute fragments rarely exceeding 0·05
-mm. in diameter. It contains a large amount of free silica that is
-probably formed by the destruction of numerous siliceous skeletons, and
-a small proportion of silicate of alumina. It usually contains the
-remains of diatoms, radiolaria, and sponge spicules, and occasionally
-lumps of pumice stone, meteoric nodules, and, in colder regions, stones
-and other materials dropped by passing icebergs.
-
-In the great oceans, then, we find in the deepest places red mud, or,
-where there is an abundant Radiolarian surface fauna, Radiolarian ooze;
-in water that is not deeper than about 2,000 fathoms, we find the
-Globigerina ooze; in shallower waters and in some localities only
-Pteropod ooze.
-
-It must not be supposed that sharp limits can anywhere be drawn between
-these different kinds of deposits, for they pass gradually into one
-another and present many intermediate forms.
-
-It is probable that the sea water by virtue of the free carbonic acid it
-contains in solution is able to exert a solvent action upon the calcium
-carbonate shells of animals as they sink to the bottom, and during the
-long and very slow journey from the surface to the bottom of the deepest
-seas these shells are completely dissolved.
-
-The first to be dissolved would be the thin delicate shells of the
-Pteropods and Heteropods, for besides the fact that they present a wider
-surface to the solvent action of the water they are probably influenced
-more by tide and currents, sink more slowly and erratically, and thus
-have a longer journey to perform.
-
-Then the smaller but more solid and compact shells of the Foraminifera
-are dissolved, and lastly, in the deepest water only the siliceous
-skeletons of the radiolaria and diatoms are able to reach their last
-resting place at the bottom of the ocean.
-
-These four oozes then are characteristic of the floor of the deep
-oceans. In the proximity of land and in inland seas where deep water
-occurs, other muds are found differing from one another in accordance
-with the character of the coasts in their vicinity. It is not necessary
-to give a detailed account of them, but a few remarks on some of the
-more pronounced forms may not be without interest.
-
-The blue mud contains eighty per cent. of a mixture of quartz, mica,
-felspar, hornblende, and other minerals, mixed with a considerable
-quantity of decomposing animal and vegetable substance, the calcareous
-remains of foraminifera, mollusca, worms, echinoderms, alcyonaria and
-corals, and the siliceous skeletons of radiolaria and diatoms.
-
-The green mud is characterised by a large percentage of glauconite.
-
-The red muds characteristic of the Brazilian coast contain a large
-amount of ochreous matter brought into the sea by the great rivers.
-
-In the neighbourhood of active volcanoes there is a characteristic
-volcanic mud, and in the coral seas the deep-sea deposits contain a
-large percentage of the calcareous remains of dead corals.
-
-One more character of the deep-sea region must be referred to before we
-pass on, and that is the absence of vegetable life. It has not been
-determined yet with any degree of accuracy where we are to place the
-limit of vegetable life, but it seems probable that below a hundred
-fathoms no organisms, excepting a few parasitic fungi, are to be found
-that can be included in the vegetable kingdom. While then the researches
-of recent times have proved beyond a doubt that there is no depth of the
-ocean that can be called azoic, they have but confirmed the perfectly
-just beliefs of the older naturalists that there is a limit where
-vegetable life becomes extinct. It is not difficult to see the reason
-for this. All plants, except a few parasites and saprophytes, are
-dependent upon the influence of direct sunlight, and as it has been
-shown above that the sunlight cannot penetrate more than a few hundred
-fathoms of sea water, it is impossible for plants to live below that
-depth.
-
-The absence of vegetable life is an important point in the consideration
-of the abysmal fauna, for it is in consequence necessary to bear in mind
-that the food of deep-sea animals must be derived from the surface. It
-is possible that deep-sea fish, in some cases, feed upon one another and
-upon deep-sea crustacea, that deep-sea crustacea feed upon deep-sea
-worms, that deep-sea echinoderms feed upon deep-sea foraminifera, and so
-on through all the different combinations; but the fauna would soon
-become exhausted if it had no other source of food supply. This other
-source of food supply is derived from the bodies of pelagic organisms
-that fall from the upper waters of the ocean, and is composed of
-protozoa, floating tunicates, crustacea, fish, and other animals,
-together with diatoms and fragments of sea-weed.
-
-
-
-
- CHAPTER III
- THE RELATIONS OF THE ABYSMAL ZONE AND THE ORIGIN OF ITS FAUNA
-
-
-In the study of the geographical distribution of terrestrial animals one
-of the great difficulties met with is the impossibility of defining
-exactly the limits of the regions into which we divide the surface of
-the earth. In a general way we recognise that there is an Australian
-region, an Ethiopian region, &c.; but, when we come to discuss the exact
-position of the frontier lines that separate these regions from their
-neighbours, we find all kinds of difficulties to overcome and
-inconsistencies to meet.
-
-For the sake of convenience it is useful to adopt certain arbitrary
-limits for these regions, notwithstanding these difficulties and
-inconsistencies, but we must recognise the fact that nature recognises
-no such limits, that every region overlaps its neighbours to a greater
-or less extent, and that there are many debateable grounds in the world
-where the fauna characteristic of one region is mixed with that
-characteristic of another.
-
-But this difficulty in defining the exact limits of the terrestrial
-faunistic regions is even more pronounced in the case of the regions and
-zones of the marine fauna.
-
-On the dry land we find mountain ranges, forests, deserts, and other
-barriers, that to a very considerable extent prevent the mixing of one
-fauna with another, but in the sea there are no barriers of anything
-like the same importance, but one fauna gradually merges into the
-neighbouring fauna according to the temperature, the pressure, the
-amount of light, the salinity of the water or the food supply. This then
-is one of the difficulties met with in the study of the geographical
-distribution of the marine fauna.
-
-But there is another that leads to almost greater complications. In
-considering terrestrial life it is customary to refer only to regions of
-geographical, or perhaps it would be more correct to call it—superficial
-distribution. It would be quite possible, however, to subdivide the
-geographical areas into zones of elevation above the sea-level, not very
-clearly marked off from one another, it is true, but nevertheless each
-showing a number of characteristic features. This idea is expressed, for
-example, when we speak of the Alpine fauna, the Himalayan fauna, or the
-fauna of the great Andes.
-
-In the study of the marine fauna and flora we must notice, it is the
-depth of the water, or in other words the depression of the habitats
-below the sea-level, that forms the most important consideration.
-Geographical sub-regions may be recognised and defined with a certain
-amount of accuracy, especially in the case of the fauna of the shallow
-waters, but by far the most important changes in the general characters
-of the fauna are found when we pass from one ‘zone’ of depression to
-another. Thus in describing any particular marine fauna we should
-mention first of all its zone or sub-zone of depression and then its
-geographical region and sub-region. For example, we may speak of the
-fauna of the pelagic zone of the British sub-region of the European
-region, or the fauna of the abysmal zone of the Northern sub-region of
-the Atlantic region.
-
-We can recognise three primary zones of the marine fauna which we may
-call the ‘Pelagic,’ the ‘Neritic,’ and the ‘Abysmal’ zones.
-
-The Pelagic zone includes the superficial waters of all seas extending
-from the surface to a depth which cannot at present be very accurately
-determined, but is probably the same as the limit of the influence of
-direct sunlight.
-
-The animals of this zone are frequently characterised by a general
-transparency of their tissues, a white or sea water (i.e. blue or green)
-colour, an organisation capable of prolonged swimming or floating
-movement, and by giving birth to floating eggs which hatch out
-transparent larvæ or embryos.
-
-The pelagic zone may be divided into several geographical regions and
-sub-regions, which it would be beyond the scope of this book to
-enumerate here, but there is one that calls for a few brief remarks. In
-many parts of the ocean there may be found vast areas of floating
-sea-weed, which carry with them a population of crustacea and other
-animals peculiarly their own. This ‘sargasso’ fauna presents so many
-characteristics and so many features different from that of the ordinary
-pelagic fauna, that the tracts of sea bearing this weed must be
-considered to rank as a special region of the pelagic zone, which may be
-called the Sargasso region.
-
-The zone of shallow water for which we shall adopt Professor Haeckel’s
-term—the Neritic zone—embraces all parts of the seas of less depth than
-500 fathoms, including the inland seas, the shores of great continents
-and islands, and the shallow banks in the great oceans. It does not
-include the superficial waters—which belong to the pelagic zone—but
-extends only from the actual bottom to a distance of a few fathoms above
-it. The fauna of this zone is extremely varied, consisting of animals
-that swim, crawl, or are permanently fixed to the bottom, animals of
-almost every variety of colour and marking, and of every size and shape.
-
-The exact limits of the Neritic sub-zones are not easy to define. The
-distinguished naturalist Forbes, to whom the abysmal zone was unknown,
-divided the seas from 0–50 fathoms in depth into three zones—the
-littoral zone lying between tide-marks, the laminarian zone extending
-from 0–15 fathoms, and the coralline zone from 15–50 fathoms.
-
-The first of these will stand as a sub-zone, the animals that are able
-to withstand exposure to the sun and air either in pools or upon the
-rocks and sand even for a few minutes frequently possessing features
-that distinguish them from those dwelling beyond low-water mark, just as
-those more active creatures that migrate backwards and forwards with the
-ebb and flow of every tide differ from the dwellers in the open sea.
-There is, it is true, at every low tide, a migration of part of the
-fauna of this sub-zone into the next, but still it is sufficiently well
-defined to be allowed to remain in our category.
-
-The second sub-zone is not so easy to define. The terms ‘laminarian’ and
-‘coralline’ used by Forbes are only applicable to certain geographical
-regions and must be abandoned for general use.
-
-We can only recognise one sub-zone between the littoral sub-zone and the
-abysmal zone, for notwithstanding the important varieties it exhibits in
-the nature of the bottom, whether it be rocky, sandy, or weedy, the
-amount of light, the temperature of the water, and the rapidity of the
-currents, it is not possible at present to point to any general
-characters of the fauna of its different parts to justify us in
-subdividing it.
-
-The name that may be given to this second sub-zone of the neritic zone
-is the Katantic—the sub-zone of the slopes.
-
-The last well-marked zone is the abysmal, extending from the 500–fathom
-line to the greatest depths of the ocean, one of enormous superficial
-area, one that it is most difficult to investigate, and one about which
-we know but little.
-
-In the present state of our knowledge we cannot divide it into any
-well-marked sub-zones nor even into geographical regions or sub-regions.
-It is not divided into sections by any important geographical barriers,
-and the general characters presented by its fauna are practically the
-same all the world over.
-
-Professor A. Agassiz has pointed out in his ‘Challenger’ monograph that
-the deep-sea echinoids of the Atlantic Ocean differ from those living in
-corresponding depths in the Pacific Ocean, but it is doubtful whether
-any such well-marked differences can be observed in other groups of
-animals. If, in the course of time, increased knowledge of deep-sea
-animals emphasises the difference between the abysmal fauna of the
-Pacific and that of the Atlantic, then we can divide this zone into two
-geographical regions; but at present it seems more correct to consider
-the abysmal zone as one that is indivisible either bathymetrically or
-geographically.
-
-Before passing on to the consideration of the general characters of the
-abysmal fauna, there are still one or two points that must be just
-briefly referred to.
-
-It is the function of every true naturalist to consider animals from
-every possible point of view. Not only must he regard them as members of
-a certain species belonging to a genus, a family, an order, and so on,
-presenting certain peculiarities of structure and development; not only
-must he regard them as inhabitants of a certain locality or zone of
-depth, but he must also pay attention to their habits and mode of life.
-
-Now amongst marine animals we can recognise three principal modes of
-life. Some animals simply float or drift about with the currents of the
-sea and are unable to determine for themselves, excepting, perhaps,
-within very small limits, the direction in which they travel. Such are
-the countless forms of protozoa, the jelly-fishes and medusæ, numerous
-pelagic worms and crustacea, the pyrosomas and salps, and many other
-forms well known to those who are in the habit of using the tow-net.
-This portion of the fauna has recently been called the Plankton.
-
-Then there are the animals that are capable of very considerable
-swimming movements, animals that are able to stem the tide and migrate
-at will from one part of the sea to another, such as the cetacea, most
-fishes, and perhaps also many cephalopods. This portion of the fauna has
-been called the Nekton.
-
-And lastly we have those animals that remain perfectly fixed to the
-bottom or are capable only of creeping or crawling over the rocks and
-sand, such as the sponges, hydroids, sedentary tunicates, gasteropods,
-most lamellibranchs, and many crustacea. This portion of the fauna has
-been called the Benthos.
-
-Although it will not be necessary to use these terms very frequently in
-this little book, it may be advisable for the reader to bear in mind
-that in any exhaustive treatise on the marine fauna such terms would be
-employed, and that in the chapters dealing with the fauna of the abysmal
-zone we should find accounts of the ‘bathybial plankton,’ the ‘bathybial
-nekton,’ and the ‘bathybial benthos.’
-
-Lastly we must consider quite briefly the views that have been held
-concerning the origin of the abysmal fauna.
-
-As soon as it became clear to naturalists that there is no part of the
-ocean, however deep it may be, that deserves the name ‘azoic,’ but that
-almost every part has a fauna of greater or less density, the problem of
-the origin of this fauna presented itself.
-
-Whence came the curious creatures that live mostly in total darkness and
-can sustain without injury to their delicate and complicated
-organisation the enormous pressure of the great depths? Are they the
-remnants of the fauna of shallow prehistoric seas that have reached
-their present position by the gradual sinking of the ocean basins? Or,
-are we to look upon the abysmal region as the nursery of the marine
-fauna, the place whence the population of the shallow waters was
-derived? Neither of these answers is supported by the facts with which
-we are now well acquainted. The fauna of the abysmal region does not
-show a close resemblance to that of any of the past epochs as revealed
-to us by geology, nor are we justified in assuming without much stronger
-evidence than we now possess, that the oceans have undergone any such
-great depression as this first theory presupposes.
-
-Nor can we consider for a moment that the abyss was the original source
-of the shallow-water fauna; for not only do we find but few types that
-can be considered to be, in any sense of the word, ancestral in
-character; but on the contrary most of the animals of the deep sea seem
-to be specially modified types of shallow-water forms. The most probable
-explanation of the origin of the deep-sea fauna is the one that was put
-forward by Moseley and has been since supported by almost every
-authority on the subject, namely, that the fauna of the deep sea has
-been derived from successive immigrations of the animals from the
-shallow water.
-
-This view is supported by the fact that the deep-sea fauna is much
-richer in the neighbourhood of land than it is in regions more remote
-from it. Many examples could be given to illustrate this point. The
-extraordinary richness of the deep-sea fauna on the western slopes of
-the floor of the Atlantic has been frequently commented on by the
-naturalists connected with the expeditions of the American vessels, the
-‘Blake,’ the ‘Fish Hawk,’ and the ‘Albatross.’ Moseley called attention
-several years ago to a few localities in the neighbourhood of the land
-especially rich in deep-sea forms in comparatively shallow waters, such
-as one near the island of Sombrero in the Danish West Indies, where
-within sight of the lighthouse a haul of the dredge in 450 fathoms
-brought up a rich fauna of blind crustacea, corals, echinoderms,
-sponges, &c. Another off Kermadec in 630 fathoms brought up numerous
-curious blind fishes, ascidians, cuttlefishes, crustacea, _Pentacrinus_,
-and large vitreous sponges, and there are similar localities lying
-between Aru and Ke and between the Nanusa archipelago and the Talaut
-islands. The deep water off the Norwegian, Scotch, Irish, and Portuguese
-coasts also seems to be particularly rich in various forms of animal
-life. The same is probably true of the deep sea of many other regions in
-the neighbourhood of land, and, although it cannot be taken to be a rule
-without exceptions—the abysmal fauna off the western coasts of the
-Panama region being, according to the recent researches of Alexander
-Agassiz in the ‘Albatross,’ particularly poor—yet we can assert as a
-statement of very general application that the further removed from
-continental land, the poorer is the abysmal fauna.
-
-Another argument that has been brought forward by Moseley in support of
-his view is that there is a certain relationship between the deep-sea
-fauna of any particular region and the shallow-water fauna of the
-nearest coasts. This is a point that is not easy to illustrate by
-examples, but as Moseley’s argument has not, so far as I am aware, been
-disputed by any of the naturalists who have followed him in this line of
-work, and the recent results of the ‘Albatross’ in comparing the
-deep-sea fauna of the eastern and western sides of the isthmus of Panama
-seem if anything to support it, we can take it as a point in favour of
-his view of the origin of the abysmal fauna.
-
-It is impossible to say at present at what time in the world’s history
-these migrations commenced, but, as Agassiz points out, none of the
-palæozoic forms are found in the deep sea, and this seems to indicate
-that the fauna did not commence its existence earlier than the
-cretaceous period.
-
-It is quite possible, however, that part of the fauna of the deep sea
-has been derived directly from the pelagic zone. The occurrence of
-bathybial Radiolaria, Foraminifera and Siphonophora, and among fishes
-genera and species of the pelagic families Sternoptychidæ and Scopelidæ,
-suggest that this zone may have contributed very largely to the fauna of
-the abyss.
-
-Much of course still remains to be done before we can consider any of
-these interesting problems connected with the deep-sea fauna to be
-definitely solved. All we can do at present is to speculate upon the
-direction in which the facts at our disposal seem to point, and by
-following up one clue after another hope that we may eventually arrive
-at the truth. The task may be a difficult one, but it will reward our
-efforts. If truth is hard to find when it lies at the bottom of a well,
-how much more inaccessible must it be when it lies hidden in the
-darkness of the sea’s abyss!
-
-
-
-
- CHAPTER IV
- THE CHARACTERS OF THE DEEP-SEA FAUNA
-
-
-The general characters presented by animals living in deep water may be
-considered under several headings. The most important are those that are
-directly or indirectly related to the fact that the animals live either
-in total darkness or in the faint and probably intermittent light
-emitted by phosphorescent animals; namely, the colour of the skin and
-the peculiarities of the eyes.
-
-The colours of the skin of the deep-sea animals vary to a very
-remarkable extent in the different groups. It cannot be said that there
-is any one colour at all predominant, and it is only in certain classes
-that black, white, or dull-coloured animals are more numerous than
-others. The colours are however usually very evenly distributed, and we
-find but few examples of animals with spots, stripes, or other
-pronounced markings.
-
-The majority of the fish are dark brown or black, but many other colours
-are represented. Thus _Ipnops Murrayi_, a typical deep-sea fish, is
-yellowish brown with colourless fins, and it exhibits a further
-character not uncommon in these abysmal forms, namely black buccal and
-branchial cavities. _Typhlonus nasus_, again, is said to be of a light
-brownish colour, with black fins. Many other examples could be given to
-show the prevalence in these regions of these black, dull, and pale
-uniform colours. But there are many exceptional cases. _Neoscopelus
-macrolepidotus_, for example—a form that according to Günther
-undoubtedly belongs to the bathybial region—is distinguished by its
-brilliant colours. It is bright red mixed with azure blue, the whole
-relieved by silver spots with circles of black on the abdomen.
-
-_Prorogadus nudus_ is of a pale rose colour, with the under and lateral
-sides of the head bluish black.
-
-_Rhodichthys regina_, found in 1,280 fathoms of water, is uniformly
-bright red in colour.
-
-A. Agassiz says in his reports on the dredging operations on the west
-coast of America: ‘The coloration of the deep-sea fishes is
-comparatively monotonous. The tints are all a light violet base, tending
-more or less to brownish or brownish yellow, or even to a greenish tint,
-especially among the Macruridæ. Some of the Liparidæ were of a dark
-violet, and one species was characterised by a brilliant blue band. The
-Ophidiidæ, _Nemichthys_, and the like, are usually of an ashy violet
-tint, while in _Ipnops_ and _Bathypterois_ the tints were of a decidedly
-yellowish brown.’
-
-That the deep-sea fish are usually devoid of any pronounced spots,
-stripes, and other markings, is now well recognised. It may not be
-altogether out of place, however, to refer briefly to a few exceptions.
-
-The black circles on the abdomen of _Neoscopelus macrolepidotus_ have
-already been referred to.
-
-_Halosaurus johnsonianus_, has a black spot on the tail.
-
-_Aulostoma longipes_ has three pairs of large black spots on the ventral
-side, but the specimen taken in 1,163 metres of water by the ‘Talisman’
-was probably a young one.
-
-It is very probable that in all the exceptional cases, when fish taken
-in deep-sea water have exhibited such spots and markings, they are
-examples either of fish that have quite recently adopted an abysmal
-habitat or of young specimens exhibiting ancestral inherited characters.
-
-In referring to a specimen of _Raja circularis_, taken by the ‘Triton’
-in 516 fathoms, Günther says: ‘It is notable that the spot on each side
-of the back which in littoral specimens is variegated with yellow is
-much smaller in the deep-sea specimen and uniformly black without
-yellow.’
-
-It seems to be then a very general rule among fishes that as they
-migrate into deeper water the spots and stripes, so conspicuous among
-many forms living on the surface and in shallow water, disappear, and
-the coloration of the body becomes more evenly distributed and uniform.
-
-Among the Mollusca, the deep-sea Cephalopods seem to be usually violet,
-but an _Opisthoteuthis Agassizii_ caught by the ‘Blake’ is stated to be
-of a dark chocolate colour, a _Nectoteuthis Pourtalesii_ reddish-brown,
-and a _Mastigoteuthis_ orange brown, while of the specimens brought home
-by the ‘Challenger,’ _Cirroteuthis magna_ was said to be ‘rose’ when
-captured, and the spirit specimens of _Cirroteuthis pacifica_ and
-_Bathyteuthis abyssicola_ were purplish madder and purplish brown
-respectively.
-
-The shells of the Gasteropods and Lamellibranchs living in the abyss are
-frequently so thin as to be almost transparent, and are, with very few
-exceptions, white or pale straw coloured. The colour of the only
-specimen of nudibranchiate Mollusca that has been found in the abysmal
-zone, namely, _Bathydoris abyssorum_, is described by Mr. Murray as
-follows: ‘The body of the living animal was gelatinous and transparent,
-the tentacles brown, the gills and protruding external generative organ
-orange, the foot dark purple.’
-
-Among the Crustacea various shades of red are the prevailing colours.
-‘The deep-sea types, like _Gnathophausia_, _Notostomus_, and
-_Glyphocrangon_,’ says Agassiz, ‘are of a brilliant scarlet; in some
-types, as in the Munidæ and Willemoesiæ, the coloration tends to pinkish
-or yellowish pink, while in _Nephrops_ and _Heterocarpus_ the scarlet
-passes more into greenish tints and patches.’[1] But perhaps the most
-remarkable point in the colour of the crustacea is that which
-immediately follows the paragraph I have just quoted. ‘The large eggs of
-some of the deep-sea genera are of a brilliant light blue, and in one
-genus of Macrura we found a dark metallic blue patch on the dorsal part
-of the carapace in marked contrast to the brilliant crimson of the rest
-of the body.’
-
-Footnote 1:
-
- In the recent researches of the ‘Investigator’ a few crustacea of
- rather exceptional colour were found. Whilst the great majority of
- them are described to be pink or red in colour when alive,
- _Gnathophausia bengalensis_ is deep purple lake, _Haliporus neptunus_
- lurid orange, and _Aristaeus coruscans_ bright orange.
-
-The occurrence of this blue colour in Crustaceans of the deep sea is
-very remarkable, for blue is a colour, as Moseley pointed out many years
-ago, that is rarely met with in the fauna of the abyss, and it is
-certainly very exceptional in the crustacea of that zone.
-
-Among the deep-sea Echinoderma we find a wonderful variety of
-coloration. Moseley says that many deep-sea Holothurians, for example,
-are deep purple, and Agassiz reports that in one species the colour was
-of a delicate green tinge. ‘We obtained,’ he adds, ‘a white _Cucumaria_
-and some species of _Benthodytes_ of the same colour,’ while others vary
-from transparent milky white to yellow and light yellowish brown and
-even pinkish colours. The Crinoids are described by the authorities to
-be white, purple, yellow and brownish-chestnut, and of the other groups
-of the echinoderms we read that the star-fishes are, as a rule, of
-duller colours than the crustacea, but all more or less pink or red.
-‘The Hymenasteridæ, on the contrary, vary from light bluish violet to
-deep reddish chestnut colours.’ The brittle stars are red and orange or
-dullish grey, while the urchins may be deep violet, claret coloured,
-brownish, or of a delicate pink.
-
-It is impossible to account for this extraordinary variety of colour in
-the deep-sea echinoderms. It is hardly probable that it can be
-protective or warning in function, and it is difficult to suppose that
-it is due to any peculiar excretory process. Whether it is due in any
-way to the influence of the environment, or, like the colour of autumn
-leaves, to the chemical degeneration of colours that in the
-shallow-water ancestry were functional, are problems that must be left
-for the future to decide.
-
-The colour of the deep-sea Cœlenterates has unfortunately not been
-recorded in all cases, but still the few observations that we have show
-that in this group, as in the last, almost every tint and shade are
-represented.
-
-The colouring of the deep-sea jelly-fishes is said to be usually deep
-violet or yellowish red. However ‘a species of _Stomobrachium_,’ says
-Agassiz, ‘is remarkable for its light carmine colour, a tint hitherto
-not observed among Acalephs.’
-
-Moseley records most minutely the colour of some of the deep-sea
-anemones and corals, and calls attention to the very general presence of
-madder brown in the soft parts. Agassiz says: ‘Among deep-sea Actiniæ, a
-species of a new _Cereanthus_ was of a dark brick-red, while other
-actinians allied to _Bunodes_ were of a deep violet. Actinauge-like
-forms with tentacles of a pinkish-violet tinge frequently have the
-column of a yellow shade. The Zoanthidæ were greyish-green.’ And again,
-in his narrative of the voyage of the ‘Blake,’ he records that ‘some of
-the deep-sea corals are scarlet, deep flesh-coloured, pinkish orange,
-and of other colours,’ and in referring to the Gorgonian _Iridogorgia_
-he says: ‘The species are remarkable for their elegance of form and for
-the brilliant lustre and iridescent colours of the axis, in some of a
-bright emerald green, in others like burnished gold or mother-of-pearl.’
-
-The fauna of the deep sea then, taken as a whole, is not characterised
-by the predominance of any one colour. The shades of red occur rather
-more frequently than they do in the fauna of any other zone or region,
-but whether this is in any way connected with the fact that red is the
-complementary colour to that of the phosphorescent light, in which many
-of these animals live, it is at present difficult to say; it is possible
-that, when we have further information concerning the colours of the
-animals living in the deeper parts of the Neritic zone, another
-explanation may be forthcoming.
-
-Moseley points out that there are no blue animals known to live in deep
-water, and it might be added that green is extremely rare as a colouring
-matter in abysmal animals, although the phosphorescent light given out
-by some of the echinoderms is green.
-
-Blue, as a colouring matter of marine animals, living on the surface or
-in shallow water, is not uncommonly met with, distributed in the form of
-bands or stripes, but green is extremely common in fishes, crustacea and
-cœlenterates, and it is a point of very considerable importance that in
-this respect there is a very great difference between the deep-sea and
-the shallow-sea faunas.
-
-If a considerable collection of living abysmal forms could be placed
-upon one table and a similar collection of shallow-water forms upon
-another, I believe that the first general impression upon the mind of
-one who saw them both for the first time would be the presence of green
-colours in the last-named collection, and the absence of it in the
-other.
-
-The eyes of the animals that live in deep-sea water undergo curious
-modifications. If the fauna of the abysmal region were confined to
-conditions of absolute darkness, we should expect to find either a total
-absence of eyes or mere rudiments of them only in those forms that have
-recently migrated from the shallow water. This is the case with the
-fauna of the great caves. There is probably total darkness in these
-underground lakes and streams, and there is only the remotest
-possibility of the animals living in them ever seeing, even temporarily,
-a ray of sunlight or even a glimmer of phosphorescence during the whole
-of their life-time. We find then that the cave fauna is totally blind.
-
-The conditions in the deep sea are not quite the same. In some regions
-there is probably a very considerable illumination by phosphorescent
-light, and it is quite possible that many of the characteristic deep-sea
-forms may occasionally wander into shallower regions where faint rays of
-sunlight penetrate, or even that the young stages of some species may be
-passed at or near the surface of the sea. Taking these points into
-consideration, then, it is not surprising to find that, in the deep
-seas, there are very few animals, belonging to families usually provided
-with eyes, that are quite blind.
-
-In the majority of cases we find that the eyes are either very large or
-very small. Only in a small minority of cases do we find that the eyes
-are recorded to be moderate in size. The relation between the large-eyed
-forms and the small-eyed forms is not the same in all the regions of
-deep seas. In depths of 300 to 600 fathoms the majority are large-eyed
-forms. This is as we should expect, for it is more than probable that
-many of these forms occasionally wander into shallower waters where
-there is a certain amount of sunlight.
-
-In depths of over 1,000 fathoms, the small-eyed and blind forms are in a
-majority, although many large-eyed forms are to be found.
-
-Among fishes, for example, we find the species of _Haloporphyrus_ found
-in depths of 300–600 fathoms with large eyes; and so with _Dicrolene_,
-_Cyttus abbreviatus_, and many other forms that are known to live in
-water of less depth than 700 fathoms; while on the other hand in
-_Melanocetus Murrayi_, _Ipnops Murrayi_, many deep-sea eels and other
-fish that are truly abysmal and live chiefly in depths of over 1,000
-fathoms, the eyes are either very small or absent.
-
-Some interesting examples may be found in the species of widely
-distributed genera to illustrate these points. Thus in _Neobythites
-grandis_, from 1,875 fathoms, the eye is small, only one-eleventh the
-length of the head, but in _Neobythites macrops_, _N. ocellatus_, and
-_N. gillii_ from shallower water it is much larger.
-
- N. grandis 1,785 fms. Eye 1⁄11th length of the head
- N. macrops 375 fms. Eye 2⁄9 length of the head
- N. ocellatus 350 fms. Eye ¼ length of the head
- N. gillii 111 fms. Eye 1⁄(3⅔) length of the head
-
-Similarly in the species of the widely distributed deep-sea genus
-_Macrurus_: the species _M. parallelus_, _M. japonicus_, _M. fasciatus_,
-&c., usually living in water less than 1,000 fathoms deep, have large
-and in some cases very large (_M. fasciatus_) eyes, but _Macrurus
-filicauda_, _M. fernandezianus_, _M. liocephalus_, _M. Murrayi_, _M.
-armatus_ have small eyes.
-
-Some deep-sea fish have their eyes reduced to a mere rudiment; such as
-_Ceratias uranoscopus_, _C. carunculatus_, _Melanocetus Murrayi_,
-_Typhlonus nasus_, and _Aphyonus gelatinosus_, but not even a rudiment
-of an eye is to be found in _Ipnops Murrayi_.
-
-But the fish of the greatest depths are by no means always characterised
-by small eyes. _Malacosteus_, a typical deep-sea form, has very large
-eyes, and so have _Bathylagus_, living in the enormous depth of 3,000
-fathoms, and _Bathytroctes_, in 1,090 and 2,150 fathoms.
-
-The result of recent deep-sea work, then, has been to show that as we
-proceed from shallow shore water to depths of 500 to 900 fathoms the
-eyes of the fish become larger, but in greater depths than 1,000 fathoms
-the eyes of some fish become considerably reduced, but those of others
-become still more enlarged. In the greatest depths of the ocean in fact
-it seems very probable that nearly all the fish are characterised by
-either very large eyes or very small ones.
-
-We cannot expect to learn very much at present from the study of the
-eyes of deep-sea mollusca. The Cephalopods form the only class of this
-Phylum whose genera invariably possess large and well-developed eyes,
-and there does not seem to be any very marked increase or decrease in
-the size of the eyes of the few deep-sea cuttlefish that are known to
-us.
-
-The eye of _Nautilus_ is certainly remarkably interesting, but as this
-genus is the only representative of its order, and is known at times to
-float upon the surface of the ocean, it would certainly be erroneous to
-attribute the peculiarity of the structure of its eye to its ‘temporary’
-deep-sea habits. We are still ignorant of the usual habitat of the
-remarkable genus _Spirula_, notwithstanding the fact that many of the
-tropical beaches are very largely composed of its empty shells. Whether
-it is a deep-sea dweller or not, we know nothing at present of the
-character of its eye, so that it can throw no light upon the problems we
-are now discussing.
-
-Among the deep-sea gasteropods we find the same irregularity in the
-possession of eyes that we have just described among fishes. Thus a
-species of _Pleurotoma_, dredged by the ‘Porcupine,’ in 2,090 fathoms,
-has a pair of well-developed eyes on short footstalks, but _Pleurotoma
-nivalis_, obtained by the ‘Talisman,’ is blind. Again a species of
-_Fusus_, obtained by the ‘Porcupine,’ in 1,207 fathoms, is provided with
-well-developed eyes, but _Fusus abyssorum_, obtained by the ‘Talisman,’
-is blind. Among the Lamellibranchs there are very few genera that
-possess well-marked eyes. The genus _Pecten_ is one of those that in
-shallow waters possess numerous highly complicated visual organs
-situated on the edge of the mantle. In the deep-sea species, _Pecten
-fragilis_, these eyes are wanting, but we have not sufficient evidence
-at present to enable us to assert that all the deep-sea species of this
-genus are blind.
-
-Among the Crustacea there is a very general tendency to lose the eyes at
-a depth of a few hundred fathoms of water.
-
-In _Ethusa granulata_, for example, the eyes disappear at 500 fathoms
-and the eye-stalks become firmly fixed, greater in length, and take the
-place of the rostrum which disappears. In some forms—such as
-_Thaumastocheles zaleuca_ and _Willemoesia_—the eye-stalks themselves
-have completely disappeared.
-
-In the deep-sea Isopoda some forms lose their eyes entirely, but
-_Bathynomus giganteus_ possesses a pair of enormous eyes, each provided
-with 4,000 facets.
-
-To illustrate the distribution of eyes in this group, we may take as an
-example the genus _Serolis_. All the species of this genus are provided
-with eyes except _Serolis antarctica_—a species that extends from 600 to
-1,600 fathoms.
-
-The eyes of all the deep-sea species are relatively larger than those of
-the shallow-water ones, except _Serolis gracilis_, whose eyes seem to be
-disappearing.
-
-But these large eyes of the deep-sea species of _Serolis_ are not
-capable of any greater perceptive power. In fact, the evidence of
-degeneration they show, both in minute structure and in the diminution
-of pigment, proves that they can be of very little use to these animals
-for perception (see Figs. 4 and 5).
-
-This increase in size, accompanied by degeneration of structure, is just
-what we should expect to find in the eyes of deep-sea animals, and it is
-difficult to explain why it is that we do not find more examples of it.
-
-If the animals that now live in the depths of the sea are descended from
-the shallow-water forms of bygone epochs, they must have passed through
-many different habitats with diminished light until they reached their
-present dark abode in the abyss.
-
-[Illustration: FIG. 4.—Semi-diagrammatic section through the eye of
-_Serolis schythei_, a shallow-water species (4–70 fathoms). C, lens; V,
-crystalline cone; R, rhabdom; N, nerve. (After Beddard.)]
-
-[Illustration: FIG. 5.—Diagrammatic section of the eye of _Serolis
-bromleyana_, a deep-sea species (400–1,975 fathoms), showing the
-degenerate character of the eye. The corneal facets C, and the
-crystalline cones V, are the only structures that can be recognised.
-(After Beddard.)]
-
-In every new region they came to, the forms with larger and better eyes
-would be at an advantage in the fainter light, and would be more likely
-to survive and transmit their favourable variation in this respect to
-their offspring than their less fortunate neighbours. Thus down to the
-depth of the limit of sunlight we should expect to find, as we do find
-in fishes, large-eyed species.
-
-Beyond the limit of direct sunlight the eyes would be of very little use
-to them, the pigment would disappear and the tissues become degenerate.
-This is precisely what has occurred in the genus _Serolis_.
-
-The disappearance of the sense of sight in the animals of the deep sea
-is sometimes accompanied by an enormous development of tactile organs.
-
-Thus, among fishes we find _Bathypterois_, a form that possesses
-extremely small eyes, provided with enormously long pectoral fin rays
-that most probably possess the functions of organs of touch.
-
-Among the Crustacea we find the blind form, _Galathodes Antonii_, with
-an extraordinary development in length of the antennæ, and
-_Nematocarcinus_, with enormously long antennæ and legs.
-
-The subject of the power of emitting phosphorescent light possessed by
-some deep-sea animals is much more difficult to deal with.
-
-The presence of distinct organs in many of the deep-sea fish that can
-only be reasonably interpreted as phosphorescent organs, the presence of
-well-developed and evidently functional eyes in many deep-sea animals,
-and many other considerations render it very highly probable that some,
-if not many, forms emit a phosphorescent light.
-
-The power and constancy of the light emitted, however, must for the
-present remain a matter of conjecture. We cannot judge at all of the
-amount of light given out by an animal in deep water by its appearance
-when thrown out of a dredge upon the deck. Whether the phosphorescent
-light given out by an Alcyonarian or a Crustacean is more or less at a
-temperature of 40° Fahr. and a pressure of one ton per square inch than
-it is at 60° Fahr. and the ordinary barometric pressure of the
-sea-level, is a question that has not yet been brought to an
-experimental test.
-
-Whatever the answer to this question may be, the fact remains that a
-greater percentage of animals from the deep sea exhibit some sort of
-phosphorescent light when brought on deck than animals that live in
-shallow water.
-
-The curious organs possessed by some fishes that are supposed to be
-organs for the emission of phosphorescent light have recently been
-subjected to a minute examination by von Lendenfeld.
-
-It has been known for some years now, that the slime secreted by the
-skin glands of certain sharks is highly phosphorescent. It is not
-difficult, then, to understand how it came about that certain fish
-developed complicated phosphorescent organs.
-
-From the phosphorescent slime secreted by a simple skin gland to the
-most complicated eye-like phosphorescent organ, we have a series of
-intermediate forms that are quite sufficient, even in the imperfect
-state of our knowledge at the present day, to enable us to understand
-the outlines of the evolution of these peculiar and interesting organs.
-
-We can distinguish two kinds of phosphorescent organs in the deep-sea
-fish. There are the curious eye-like or ocellar organs situated usually
-in one or more rows down the sides of the fish’s body, forming as it
-were a series of miniature bull’s-eye lanterns to illuminate the
-surrounding sea (fig. 6); and various glandular organs that may be
-situated at the extremity of the barbels or in broad patches behind the
-eyes or in other prominent places on the head and shoulders.
-
-Ocellar organs have been known for many years to occur on the sides of
-the interesting pelagic fish, _Scopelus_. Most of the species of this
-genus live in the open sea at moderate depths, coming to the surface
-only at night, but other species are found in almost every depth down to
-2,000 fathoms of water.
-
-[Illustration: FIG. 6.—_Opostomias micripnus_; 2,150 fathoms. (After
-Günther.)]
-
-In _Opostomias micripnus_, a dark black fish living at a depth of over
-2,000 fathoms, there are two rows of ocellar organs running down the
-sides of the body from the head to the tail. In the living animal they
-are said to shine with a reddish lustre. In addition to these, the
-conspicuous organs, there are groups of fifty, a hundred, or even more
-very much smaller organs situated on the sides and back of the fish,
-each of which is lenticular in shape and consists of a number of short
-polygonal tubes containing a granular substance with rounded bases
-resting on the subjacent tissue. The whole organ is covered by a simple
-continuation of the cuticle of the body-wall. The granular substance
-contained in the tubes is most probably the seat of luminosity.
-
-[Illustration: FIG. 7.—Head of _Pachystomias microdon_ (after Von
-Lendenfeld). A, anterior sub-orbital phosphorescent organ; B, posterior
-sub-orbital phosphorescent organ.]
-
-As a type of the glandular organs we may take one of the sub-orbital
-organs found on the head of _Pachystomias microdon_.
-
-In this fish there are two very conspicuous white organs immediately
-below the eye. The anterior one, which lies below and in front of the
-eye, is oval, with its upper margin slightly concave. In section it is
-seen to be surrounded by a thin layer of black pigment, and to consist
-of a reticular glandular substance in which is embedded a hammer-shaped
-lens-like body. Between these two structures there is interposed a thick
-layer of light reflecting spicules.
-
-The exact part that is played by the different components of these
-curious phosphorescent organs is not yet known, but sufficient has been
-said to indicate to the reader the degree of complexity that these
-organs may reach in the fish of the great depths of the ocean.
-
-[Illustration: FIG. 8.—Section of the anterior sub-orbital
-phosphorescent organ of _Pachystomias microdon_ (after Von Lendenfeld).
-L, lens; O, phosphorescent gland; P, pigment sheath.]
-
-But the power of emitting phosphorescent light is by no means confined
-to the group of fishes. Some of the Macrurous Decapoda among the
-Crustacea are known to be phosphorescent. In the case of _Heterocarpus
-Alphonsi_, for example, the naturalists of the ‘Investigator’ found that
-‘clouds of a pale blue highly luminous substance, which not only
-illuminated the observers’ hands and surrounding objects in the vessel
-in which the creature was confined, but also finally communicated a
-luminosity to the water itself, were poured out apparently from the
-bases of the antennæ.’
-
-‘The _Willemoesia_, too, was luminous at two circumscribed points
-somewhere near the orifices of the genital glands.’
-
-Again, all the Alcyonarians dredged by the ‘Challenger’ in deep water
-were found to be brilliantly phosphorescent when brought to the surface,
-the light consisting, according to Moseley, of red, yellow, and green
-rays only.
-
-Among the Echinoderms we have not many recorded instances of a
-phosphorescent light being emitted, but it is quite possible that many,
-if not all of them, may possess this power. The curious deep-sea form
-_Brisinga_, that was first discovered by Ch. Asbjörnsen, is known to be
-so brilliantly phosphorescent that it has been called a veritable
-_gloria maris_, and writing of the curious brittle-star _Ophiacantha
-spinulosa_ (dredged by the ‘Porcupine’ in 584 fathoms of water),
-Professor Wyville Thomson remarks that the light was of a ‘brilliant
-green, coruscating from the centre of the disc, now along one arm, now
-along another, and sometimes vividly illuminating the whole outline of
-the star-fish.’
-
-According to Filhol many of the abysmal Annelid worms are in the habit
-of emitting a vivid phosphorescent light, and capable thereby of
-illuminating the medium in which they live.
-
-We have now considered all those characters exhibited by deep-sea
-animals that may be associated with the absence of direct sunlight. To
-run through them again briefly we may say: that the deep-sea species,
-belonging to classes of animals that usually possess eyes, show some
-modification in the size of their eyes, in that they are either very
-large, very small, or altogether wanting. That deep-sea animals are
-nearly always uniformly coloured. Very frequently they are black or grey
-or white, less frequently bright red, purple, or blue. But whatever the
-colour may be, spots, stripes, bands, and other markings of the body are
-very rarely seen. That deep-sea animals are brilliantly phosphorescent,
-the light being emitted either by special organs locally situated on the
-head, body, or appendages, or by the general surface of the body.
-
-But there are some other characters that cannot be thus associated with
-the absence of sunlight.
-
-In the first place bathybial fish, mollusca, crustacea, and other
-animals usually possess a remarkably small amount of lime in their bones
-and shells.
-
-In fishes we are told that the bones have a fibrous, fissured, and
-cavernous texture, are light, with scarcely any calcareous matter, so
-that the point of a fine needle will readily penetrate them without
-breaking. In some the primordial cartilage is persistent in a degree
-rarely met with in surface fishes, and the membrane bones remain more or
-less membranous or are reduced in extent, like the operculum, which is
-frequently too small to cover the gills.
-
-This cannot be due in all cases to a deficiency of carbonate of lime in
-the sea water, for we find these characters well marked in some of the
-fish, such as _Melanocetus Murrayi_, _Chiasmodus niger_, and _Osmodus
-Lowii_, that are found on the Globigerina mud.
-
-Then again, the shells of the deep-sea Lamellibranchs, Gasteropods,
-Brachiopods, and Crustacea are very frequently remarkably thin and
-transparent, a character that is probably more generally due to a
-weakness in absorptive or secretive activity than to a deficiency in the
-supply of lime.
-
-There are one or two characters of the deep-sea fish that it is not easy
-to account for, and it is necessary only to mention their occurrence
-without attempting to offer any explanation of them.
-
-One of the most common of these is the very dark pigment occurring in
-certain parts of the epithelium of the mouth and respiratory passages
-and the endothelium of the peritoneum. For example, in _Bathysaurus
-mollis_, living at a depth of 2,000 fathoms, the mouth and buccal
-cavities are black. The same thing occurs in _Ipnops Murrayi_, and
-indeed in all the strictly deep-sea forms.
-
-Another important character of very frequent occurrence is the reduction
-in size, length, and number of the gill laminæ.
-
-Among invertebrates we may mention as a fact of some interest, dependent
-perhaps on the soft character of the bottom, the preponderance of
-stalked forms over those of more sessile habits.
-
-Thus among the Alcyonaria the characteristic forms of the deep water are
-the Pennatulids, and more particularly the genus _Umbellula_ with its
-long graceful stem and terminal tuft of polyps. Among the Echinoderma we
-find many forms of stalked Crinoids. Among the Tunicates several curious
-genera characterised by their long peduncles.
-
-Taking the fauna as a whole, Moseley regarded it as similar in some
-respects to the flora of the high mountains. Some forms are dwarfed in
-size, such as the species of Radiolaria, Cerianthus, some of the
-Cephalopods, &c., while others are very much larger than their
-shallow-water allies, such as the Pycnogonids, nearly all the Crustacea,
-Alcyonarians (as regards the size of the polypes), Siphonophora, and
-many others.
-
-
-
-
- CHAPTER V
- THE PROTOZOA, CŒLENTERA, AND ECHINODERMA OF THE DEEP SEA
-
-
-The most important, but perhaps somewhat disappointing, result of the
-deep-sea researches of recent years has been to prove that the abysmal
-fauna does not possess many very extraordinary forms.
-
-It seemed probable, before the dispatch of the ‘Challenger’ expedition,
-that when the dredge and the trawl should be successfully employed in
-depths of over 2,000 fathoms, a new and remarkable fauna would be
-brought to light. Some naturalists thought it even possible that, not
-only would many genera be found alive that are known to us only by their
-fossilised skeletons in the secondary and tertiary rocks, but that there
-might be many other new creatures whose anatomy would throw much light
-on the theories of the evolution of the animal series.
-
-But none of the great expeditions that have sailed since the year 1874
-have yet succeeded in showing that the hopes and wishes of these
-naturalists were really justified. Although thousands of species of
-animals have been described in the volumes that have been devoted to
-deep-sea work, the number of the sub-kingdoms and classes remains the
-same, and indeed the number of new families and genera has not been
-increased in any very unprecedented manner.
-
-We have found no animals in the depths of the sea of such interest and
-importance as Ornithorhynchus, Amphioxus, Balanoglossus, Peripatus,
-Millepora, or Volvox among the living, or Hipparion, Archæopteryx,
-Ammonites, Slimonia, and the Trilobites among extinct animals.
-
-The abysmal fauna is not in fact remarkable for possessing a large
-number of primitive or archaic forms. It is mainly composed of a number
-of species belonging to the families and genera of our shallow-water
-fauna that have, from time to time, migrated into greater depths and
-become modified in their structure in accordance with the extraordinary
-conditions of their new habitat.
-
-There is very good reason to believe that this migration has been going
-on from time immemorial, and consequently we find a few forms typical of
-the bygone times, left to struggle for existence with the more recent
-immigrants from shallow waters. But after all the proportion of ancient
-forms to modern ones in the fauna of the abyss is not larger than it is
-in the fauna of fresh-water lakes and streams or even of the dry land.
-Nor is there any reason why it should be. The land and the fresh water
-have been peopled by migrations from the shallow water of the sea from
-generation to generation in precisely the same way, and they each can
-show a certain number of archaic forms.
-
-We must now consider briefly some of the most interesting deep-sea
-representatives of the various classes of the animal kingdom, referring
-as we pass on to the extent to which these classes contribute to the
-fauna of the abyss.
-
-We find a great difficulty in determining with any degree of certainty
-the actual depths at which the supposed abysmal forms of Protozoa
-actually live. All the Radiolaria and Foraminifera—the only Protozoa
-that are largely represented in the fauna of the open seas—are
-planktonic in habit; that is to say, they float or drift about in the
-water without ever becoming attached to the sea bottom; and when the
-contents of a dredge, that has been hauled up from a great depth, are
-examined, it is impossible to say at what points in its long journey
-from the bottom the Protozoa it contains were caught. Even if dredges
-and nets are used which can be closed by a messenger at any particular
-depth, the problem cannot be very easily settled; for even if the
-protozoa shells that are captured are found to contain a certain amount
-of protoplasm, it must be proved that that protoplasm is actually alive
-when brought on deck before we know for certain that the species
-actually live on the bottom. When the pelagic Foraminifera and
-Radiolaria die and sink to the bottom, their protoplasm probably
-disintegrates very slowly, and it is quite probable that the floor of
-the ocean is littered with the shells of truly pelagic protozoa, each
-containing a greater or smaller amount of undecomposed protoplasm.
-
-However, there is little doubt that there are some truly abysmal
-Protozoa. Among the Radiolaria, for example, it seems extremely probable
-that the majority of the Phæodaria and many Spumellaria live only in
-very deep water. ‘A character common to these abyssal forms,’ says
-Haeckel, ‘and not found in those from the surface or slight depths, is
-found in their small size and massive heavy skeletons, in which respects
-they strikingly resemble the fossil Radiolaria of Barbadoes and Nicobar
-islands.’ The Phæodaria are very widely distributed over the floor of
-the ocean, and occur in some districts in such numbers that the
-‘Challenger’ was able to bring home some hundreds of thousands of
-specimens. They are distinguished from other Radiolaria by the thick
-outer and thin inner capsule, by the typical main opening or atropyle
-placed on the oral pole of the main axis with a radiate operculum
-provided with a tubular proboscis, and lastly by the presence of the
-phæodium, a voluminous pigment body which lies invariably on the oral
-half of the calymma and is composed of numerous singular pigment
-granules of green, olive, brown, or black colour.
-
-[Illustration: FIG. 9.—_Challengeria Murrayi_, one of the Phæodaria
-(2,250 fathoms). A, phæodium; B, central capsule; C, strands of
-protoplasm in the calymma. After Haeckel.]
-
-There are many genera belonging to the Foraminifera that are very
-probably inhabitants of abysmal depths, but they do not seem to possess
-any special characters, unless it be a greater thickness and density of
-their shells, to distinguish them from their shallow-water allies.
-
-Passing now to the group of the sponges or Porifera, we find that the
-calcareous sponges are not represented at all in the abysmal zone. Two
-species are found at a depth of 450 fathoms, but none are truly
-bathybial in habit. The same remark applies to the horny sponges.
-These forms chiefly belong to the littoral or very shallow-water
-fauna, and never descend to greater depths than 400 fathoms. Of the
-other groups of Porifera—the Monaxonia, the Tetractinellidæ, and the
-Hexactinellidæ—several genera are known to extend down to some of the
-greatest depths at which trawling operations have been successfully
-carried on. It is difficult to point to any characters in these
-sponges that can be attributed in any way to the conditions of
-deep-sea life, but nevertheless we do find in deep water some of the
-most remarkable and beautiful forms of sponge skeleton that can be
-found anywhere.
-
-Amongst the Cœlentera we find in the deep water a remarkable sub-family
-of Medusæ, which has been named by Haeckel the Pectyllidæ. It is
-distinguished from the other jelly-fish by the curious sucking cups
-situated at the ends of the tentacles. It seems probable that they are
-used for purposes of locomotion, the animal walking over the muddy
-bottom as on a series of stilts.
-
-Like most of the deep-sea Hydroids, the Pectyllidæ are usually devoid of
-sense organs, but a single specimen of _Periphylla mirabilis_, captured
-by the naturalists of the ‘Challenger,’ possessed well-marked eyes.
-
-There is also a peculiar family of the Siphonophora, called the
-Auronectæ, consisting of a few specimens that have been hitherto found
-only in very deep water. Like the well-known Portuguese man-of-war
-_Physalia_ of the surface waters, the Auronectæ possess a large swimming
-bladder or pneumatophore, but they have in addition another peculiar
-bladder-like cavity, called the aurophore, communicating with it, which
-may be an organ for secreting gas.
-
-A very interesting genus allied to Velella was also found in depths of
-over 2,000 fathoms by the ‘Challenger’ expedition. It is supposed to be
-a survival of the ancestral form of the Disconectæ, or, at any rate, to
-be a link connecting the Siphonophora with the Medusæ. The very well
-marked octoradial arrangement of the parts of _Discalia_, as this genus
-has been termed, is certainly a point of great interest and importance.
-
-There is no large family of the sea anemones that is peculiar to deep
-water, but several genera that occur only in the abyss exhibit some
-curious modifications. The manner in which the tentacular pores have
-become enlarged, and the tentacles themselves diminished in size and
-flexibility, has already been referred to in a previous chapter (p. 36).
-
-The family of sea anemones that has been named the Corallimorphidæ,
-characterised by the stiffness and slight contractility of the body, the
-knobbed nature of the tentacles, and their distribution in several
-series, was, until quite recently, considered to be a true abysmal
-family. The two species, _Corallimorphus rigidus_ and _C. profundus_,
-are known to occur only in very deep waters, and present some curious
-modifications of structure in relation to their habit; but it seems
-probable that to this family should be added the remarkable littoral
-form _Thelaceros rhizophoræ_ found on the coast of Celebes attached to
-the roots of the mangrove trees in the swamps.
-
-The fact that all the principal groups of the Actiniaria, except,
-perhaps, the group that includes those forms with only eight
-mesenteries, the Edwardsiæ, have representative genera or species in the
-great depths of the ocean, points to the conclusion that the sea
-anemones have migrated from the shallow waters in comparatively recent
-times, and that the migrations have been successive, each period of
-their history sending some specimens to survive or to become extinct in
-the struggle for life in the deep sea.
-
-Of the Madreporarian corals, several genera are now known to inhabit
-very deep water, but they do not present many very remarkable points of
-divergence from the shallow-water forms.
-
-It is true that as we pass from the shallow waters, of those parts of
-the world where the great colonial madrepores build up the greater part
-of the vast coral reefs, into the deeper water beyond them, the solitary
-forms become relatively more abundant, but no new groups characterised
-by any special deep-sea attributes make their appearance. We must
-remember, not only that a great many solitary corals occur in shallow
-water in different parts of the world, but that some colonial forms,
-such as _Lophohelia prolifera_ for example, are found only in very deep
-water.
-
-Until quite recently it was usually stated in works dealing with the
-structures of coral reefs that the so-called reef-building corals, that
-is to say the large madrepores, astræids, and others, are confined to
-water not deeper than thirty fathoms. This limit must now be somewhat
-extended, in consequence of the discovery by Captain Moore of an
-abundance of growing coral at a depth of forty-four fathoms in the China
-seas; but, nevertheless, it is perfectly true that the corals do not
-grow in such profusion in very deep water as to form anything that can
-be compared with the reefs of the shores. It is quite possible that the
-advantages afforded by the light, warmth, and abundance of food of the
-shallow water may account for the luxuriance and vigour of the reef
-corals, and that where the food is scarce, and the water cold and dark
-as it is below fifty fathoms, the power of continuous gemmation is lost,
-and the rapidity of the growth and reproduction of the individual polyps
-is considerably diminished.
-
-The fact remains, however, that, as with the sea anemones, so with the
-madrepores, nearly all the great divisions have a few isolated
-representatives in the abyss, and that no great family occurring in
-large numbers has yet been discovered peculiar to this zone.
-
-The Alcyonaria, on the other hand, do present us with at least one
-example of a true deep-sea family. This great class of Anthozoa,
-distinguished from the Zoantharia by the presence of not more than eight
-tentacles and mesenteries and by the pinnate character of the former,
-falls into four principal divisions. The Stolonifera, the Alcyonidæ, the
-Gorgonidæ, and the Pennatulidæ. The first three of these divisions
-principally inhabit the shallow water. Each of them sends a few
-representatives into the great depths, but by far the greater number of
-the genera and species are to be found between tide-marks or in depths
-of less than fifty fathoms.
-
-The Pennatulids, on the other hand, are rarely found in very shallow
-water, and nearly half the known genera live in deep water. At least two
-families may be said to be characteristically abysmal. These are the
-Umbellulidæ and the Protoptilidæ.
-
-The Pennatulidæ are regarded by naturalists as the most complicated or
-highly organised group of the Alcyonaria. Three different forms of
-polype build up the colony or sea-pen as it is called. There is a single
-very much modified and enormously large polype, without tentacles,
-forming the axis, a large number of ordinary Alcyonarian polypes
-(autozooids) arranged in the form of leaves, or simply scattered
-irregularly on the surface of the central polype, and a number of very
-small undeveloped polypes (Siphonozoids) without tentacles, whose
-function seems to be to pump water into the canals of the colony, and
-thus to keep up the circulation of water.
-
-[Illustration: FIG. 10.—Umbellula Güntheri. Nat. size. After Agassiz.]
-
-The deep-sea genus _Umbellula_ possesses a very long and delicate axial
-polype, and the Autozooids and Siphonozooids form a little cluster only
-at its extreme summit. The small number of these polypes and the very
-limited area over which they extend are the two most characteristic
-features of the genus. It would take me too far into the anatomy of the
-group if I were to add any further details; but I cannot pass on without
-noting that the whole structure of Umbellula shows that it is far more
-primitive and simple than the shallow-water genera. And, generally
-speaking, this holds good for all the deep-sea Pennatulids. In fact, we
-have here one of the rare examples of a series of genera, that can be
-regarded as a slightly modified ancestry of the shallow-water genera,
-that has been brought to light by the exploration of the abysmal depths
-of the ocean.
-
-We have seen, then, that of the Cœlentera, the only order that has a
-large proportion of its genera living in deep water, is the only one
-whose members all possess a stalk by which they fix themselves into the
-mud or sand at the bottom of the sea.
-
-It is not uninteresting to note, then, in passing on to the Echinoderma,
-that the stalked Crinoids, the only Echinoderms that can permanently fix
-themselves to the bottom, are nearly all found in deep water.
-
-Several years before the ‘Lightning’ was despatched on her memorable
-pioneering voyage, Vaughan Thomson had proved that the common feather
-star of the shallow waters of the British coasts passes through a stage
-in its development which resembles the fossil genera of the order in
-being provided with a stalk for attachment.
-
-But it was left for the naturalists of the ‘Porcupine,’ the
-‘Challenger,’ the ‘Talisman,’ and other vessels employed in deep-sea
-researches to prove that adult stalked Crinoids are still living in
-nearly all parts of the world at the great depths of the sea.
-
-The genera of stalked Crinoids now living are the remains of a family
-that at one time had many representatives in all parts of the world.
-Nearly all the marine deposits of bygone epochs, including even those of
-such remote periods as the Cambrian and Sub-Silurian, contain the
-fossilised skeletons of these Crinoids. In some strata they are
-represented by only a few genera, but in others they are found in such
-enormous numbers that the seabeds of those early times must have been
-literally carpeted with them.
-
-At the present day the few genera that survive have been driven from the
-shore waters, and are chiefly found at depths of more than 200 fathoms,
-a few only extending into 140 and even 70 fathoms.
-
-[Illustration: FIG. 11.—_Rhizocrinus lofotensis_, one of the deep-sea
-stalked Crinoids. (After Carpenter.)]
-
-There are six genera known, and of these, two, _Hyocrinus_ and
-_Bathycrinus_, are not found in less than 1,000 fathoms of water.
-
-There can be no doubt that these modern stalked Crinoids are closely
-related to many of those that flourished in bygone periods of the
-history of the earth. As Carpenter has pointed out, the family
-Pentacrinidæ are remarkable for their long geological history. The genus
-_Pentacrinus_ itself first appeared in the Trias and persisted through
-the Secondary and Tertiary times to the present day.
-
-The general character of the fossil Pentacrinidæ is essentially the same
-as that of their recent representatives, except that they often had much
-longer stems which reached to a length of as much as 50 or even 70 feet;
-while the number of arms was frequently limited to ten, which is not the
-case in any recent species but _Pentacrinus naresianus_.
-
-But the deep-sea Echinoids, or sea-urchins, also present some features
-of particular interest. Professor Agassiz in his report says, ‘One of
-the very first results clearly indicated by the deep-sea dredgings of
-Count Pourtales and the subsequent investigations of the “Porcupine”
-expedition was the antique character of the new genera discovered in
-deep water, and especially their resemblance to the cretaceous genera;
-and the study of the “Challenger” Echinoids has brought this out still
-more clearly.’
-
-No fewer than twenty-four genera extend into the abysmal regions; of
-these no less than sixteen, nearly all belonging to a new group of
-Spatangoids, do not live at all in shallow water.
-
-The most interesting forms among these are the Pourtalesiæ, a group that
-has existed since the Chalk. These are heart-shaped urchins with a very
-peculiar test. ‘They all have large coronal plates, recalling the
-Echini, with a disconnected apical system characteristic of many
-cainozoic spatangoids; they have a sunken anal system, some of them a
-most remarkable anal beak and a very striking pouch in which the mouth
-is placed.’ They are found only in very deep water, and have no allies
-among the modern littoral fauna.
-
-The genera _Calveria_ and _Phormosoma_ are two of the most abundant
-Echinoids found in deep water, and they are both representatives of
-forms that were very abundant in cretaceous times. They are remarkable
-for the extreme flexibility of their shells. In shallow-water
-sea-urchins the shells are composed of a great number of little plates
-that fit so closely to one another that no movement is possible between
-them. When the animal dies all the soft tissues decay and the shell
-remains, to be tossed about by the waves until crunched or dashed to
-pieces. In _Phormosoma_, however, the tiny plates of which the shell is
-composed are freely movable on one another, and when the animal is alive
-very considerable contractions and expansions can take place.
-
-None of the modern shallow-water Echinoids present this peculiarity, and
-it is a very interesting and surprising fact that in this respect the
-fossils of the chalk should resemble so closely the living urchins of
-the abyss.
-
-But before leaving the Echinoids reference must be made to two more
-points that have been made by the illustrious American naturalist.
-Agassiz points out that all those genera that have the greatest
-bathymetrical range, extending from the littoral to the abysmal region,
-are at the same time genera which date back to the Cretaceous period,
-while those having a somewhat more limited range go back to the
-tertiaries, and those that extend only slightly beyond the littoral area
-go back only to the later tertiaries.
-
-This interesting generalisation brings home to our minds the enormous
-length of time that it must have taken these animals to migrate from the
-shallow to the deep sea. In the struggles for existence between marine
-animals it must always have been the last resort of those unable to
-compete with the younger generations in shallow water to migrate into
-the deeps.
-
-The scarcity of food, the darkness, and the pressure of these regions
-can never be so favourable for the support of animals as the conditions
-of the shores. We can well imagine that a species would take every
-opportunity that is afforded to return from such inhospitable habitats,
-and that only when, as it were, every door is closed, when no island,
-continent, or cape can afford it a free scope for life in shallow water,
-does it become a true deep-sea species.
-
-Steps taken towards the darkness in one period may be retrieved in the
-next. The competing species may itself have become extinct or have moved
-to another locality. Organs may have become modified or a new source of
-food supply tapped which enable them to return once more to shallower
-waters. No wonder that the steps in the progress, or rather retreat, to
-the abyss have been the work of a time that can be counted only by
-geological periods; and no wonder then at the remark made by many
-deep-sea naturalists that the abysmal fauna becomes poorer the farther
-it is from shallow water.
-
-The group of the Asteroidea, or star-fishes, contributes largely to the
-fauna of the abyss.
-
-During the voyage of the ‘Challenger’ no fewer than 109 different
-species were found in depths of over 500 fathoms, and in some localities
-a very large number of star-fish were taken in one haul of the dredge.
-
-Nevertheless, there are not many abysmal genera that differ to any
-remarkable degree from the littoral ones; and indeed it may be said that
-the recent work on deep-sea Asteroids does not throw much new light
-either on the phylogeny of the group or on their palæontological
-history.
-
-The genus _Brisinga_, at one time supposed to be a connecting link
-between the star-fishes and the brittle stars (Ophiurids), has recently
-been shown to be closely related to the families Heliasteridea,
-Echinasteridea, and others typical of the class Asteroidea; and, as
-Sladen has pointed out, the peculiarities of structure that it exhibits
-are probably due to its extreme isolation and the influence of its
-abysmal habitats.
-
-But no work on the deep-sea fauna would be complete without some
-reference to Brisinga. Discovered by Asbjörnsen in 1853, in 200 fathoms
-of water in the Hardanger fjord, and described in a splendid memoir by
-the elder Sars, it excited great interest among naturalists. The great
-brilliancy of the phosphorescent light that it gave out on being brought
-on deck, the remarkable tendency that it had to cast off some of its
-numerous long, thin, ophiurid-like arms, and some of the general
-features of its internal anatomy were points that were considered at the
-time to be sufficient to justify the establishment of a separate
-sub-order for the family Brisingidæ.
-
-The more recent discovery, however, of genera allied to Brisinga has
-bridged over the gap separating it from other star-fish, and it is now
-considered simply as the type of a family of the order.
-
-The numerous species of the genus that have been found since
-Asbjörnsen’s original discovery are all inhabitants of deep water, some
-of them going down to the enormous depth of 2,000 fathoms; indeed there
-are very few genera in the animal kingdom, containing so many species as
-the genus Brisinga, that have such a uniform deep-sea habitat.
-
-The last group of Echinoderms that we have to consider is the
-Holothurians. It contains one order—the Elasipoda—that may be considered
-to be truly bathybial, as there is only one species belonging to it,
-_Elpidia glacialis_, that extends into water as shallow as fifty
-fathoms.
-
-The Elasipoda are remarkable for their strongly-developed bilateral
-symmetry. Adult Echinoderms as a rule possess a well-marked radial
-symmetry, as we see exemplified in the feather-star, star-fish, and
-sea-urchin, but this radial symmetry is only adopted when they undergo
-their metamorphosis from the free swimming and bilaterally symmetrical
-larval stage. They are not born radially symmetrical, but become so as
-they grow up. Moreover, we must bear in mind that the radial symmetry of
-the adult only obscures, it does not obliterate, the bilateral symmetry
-of the larva.
-
-In the Holothurian, however, we can always discover a clear bilateral
-symmetry even in the adult. That is to say, we can recognise an anterior
-and a posterior end, a right and a left side of the body. It is an
-organisation which emphasises, as it were, the anterior and posterior
-ends, the right and left sides and the dorsal and ventral surfaces that
-characterise this interesting deep-sea order, the Elasipoda.
-
-Here, then, we have an example of a character common to all the larvæ of
-the sub-kingdom and exceptionally well marked in the adults of a family
-confined to deep-sea habitats.
-
-Now we know that there is a tendency for some of the peculiar characters
-of the ancestors of animals to be recapitulated in the course of their
-development from the egg, and accordingly most naturalists are agreed
-that all the Echinoderms have descended from some form of bilaterally
-symmetrical ancestor. Are we, then, to believe that the Elasipoda
-brought from the depths of the sea are more closely related to these
-ancestral forms than the shallow-water families?
-
-The state of our knowledge at the present day hardly allows us to answer
-this question very definitely. However nearly they are related to such
-ancestral Echinoderms in general form, they are probably profoundly
-modified by a deep-sea life. Nevertheless, in the simple shape of the
-calcareous corpuscles of the skin, the simple form of the calcareous
-ring, the communication of the water-vascular system with the exterior
-by one or several pores, and in some other anatomical characters, they
-give evidence of their primitive characters.
-
-
-
-
- CHAPTER VI
- THE VERMES AND MOLLUSCA OF THE DEEP SEA
-
-
-It has not been my intention in this volume to confine my attention to
-the truly abysmal forms, but rather to consider all those animals living
-in deep water that show any characters strikingly different from their
-relatives living in shallow water.
-
-The term deep water is, after all, only a relative one.
-
-To one accustomed only to shore collecting, ten fathoms is deep water,
-while on the other hand, to such naturalists as those on board the
-‘Challenger,’ who are accustomed to dredge in all seas, nothing under
-1,000 fathoms is considered deep water.
-
-We must bear in mind, however, that at a depth of only 200 fathoms, the
-conditions of life are very different to those of the shore waters. We
-find a very great diminution in the amount of light, for instance, that
-can penetrate through sea water teeming with floating organisms of all
-kinds to reach the fauna attached to the bottom at such a depth. The
-diminution in the amount of light must mean a diminution in the rapidity
-of growth of chlorophyll-bearing plants, and consequently a diminution
-in the food supplies of animals drawn from that source.
-
-We might expect then to find, even in such shallow water as this, some
-forms of particular interest. It is true that the greater part of the
-fauna is made up of ordinary shallow-water forms that have migrated
-quite recently, and perhaps only temporarily, into the depths, but we
-expect to find, and actually do find, the outposts of a new fauna.
-
-These remarks lead me to the consideration of one or two very remarkable
-animals that have recently been brought to light.
-
-In that strange assembly of animals which, for want of a better word,
-the authorities call the Vermes, there are three groups whose relations
-to one another and to the other groups of Vermes have been and still
-remain a puzzle to naturalists.
-
-These three groups are the Gephyrea, the Polyzoa, and the Brachiopoda.
-In external form they are as different from one another as possible.
-
-The Gephyrea are solitary worm-like forms burrowing in the sand or
-perforating rocks; the Polyzoa are minute creatures that frequently
-build up by budding large colonies which assume in some cases dendritic
-forms like corals, and the Brachiopoda are protected by thick bivalve
-shells simulating in a striking manner the shells of the
-Lamellibranchiate mollusca.
-
-But external form is not the only character that can be relied upon for
-purposes of classification. The general and minute anatomy, together
-with the story of the development of these animals, teach us that they
-are in some way closely related.
-
-It is not within the scope of this book to enter into the discussion of
-what these relations are; suffice it to say that the controversy has
-within recent years to a great extent turned upon the position in our
-classification of three interesting genera. These are Phoronis,
-Rhabdopleura and Cephalodiscus.
-
-Phoronis occurs only in shallow water, Rhabdopleura has been found in
-water from 40 to 200 fathoms deep off the Shetlands and on the Norwegian
-coasts, while Cephalodiscus was discovered by the ‘Challenger’ at a
-depth of 245 fathoms off Magellan Straits.
-
-Rhabdopleura forms colonies consisting of branched tubes growing upon
-the tests of Ascidians, on sea-weeds, corals, or other objects fixed to
-the sea bottom. In the open, free extremity of each of the branches may
-be found the polypide attached to a filament or stalk which connects it
-with the other polypides of the colony (fig. 12).
-
-[Illustration: FIG. 12.—A portion of a colony of _Rhabdopleura normani_.
-(After Lankester.)]
-
-Each polypide is provided with a single pair of large pinnate arms,
-resembling the arms of a Brachiopod, and a broad muscular epistome by
-means of which it is able to creep up or down the tube.
-
-The affinities of this interesting creature are by no means sufficiently
-well understood. It is one of those forms that, without being, strictly
-speaking, a connecting link between large and well-known groups of
-animals, indicates to us some of the lines of evolution that these
-groups may have passed through; and, in so far as it does this, it has
-its value and importance.
-
-Cephalodiscus, though related to Rhabdopleura in the presence of a
-structure corresponding to the arms, and a broad epistome, seems to be
-more closely connected with such a form as Balanoglossus in the presence
-of a single pair of gill-slits, a small rudimentary notochord and the
-position of the central nervous system.[2]
-
-Footnote 2:
-
- A rudimentary notochord projecting forward from the buccal cavity into
- the epistome has quite recently been discovered in Rhabdopleura.
-
-[Illustration: FIG. 13.—A single polypide of _Rhabdopleura normani_. M,
-mouth; B, epistome; S, polypide stalk. (After Lankester.)]
-
-Whatever position these genera may ultimately occupy in our systems of
-classification, there can be little doubt that much valuable information
-will be obtained by a further study of their structure and
-development—information that will probably shed much light on the
-relationships to one another of the many groups of Vermes. Their
-occurrence in water of moderate depths only indicates perhaps that they
-are gradually being crowded out from the more favourable localities of
-shallow water, and are tending towards extinction on the one hand, or a
-deep-sea habitat on the other.
-
-The Brachiopoda need not detain us long. Some species are capable of
-existing at a great variety of depths without any observable
-modification of shape or characters. Thus _Terebratulina caput
-serpentis_ has the extraordinary bathymetrical distribution of 0–1,180
-fathoms, and _Terebratula vitrea_ 5–1,456 fathoms. _Atretia_ is the only
-genus peculiar to deep water. It is a noteworthy fact in connection with
-this order that the two genera, _Lingula_ and _Glottidia_, which compose
-the sub-order Ecardines, are both confined to shallow water. Now the
-Ecardines are anatomically, at any rate, the most primitive of the
-Brachiopoda, and Lingula has the most ancient geological history of any
-living genus of the animal kingdom, shells almost identical with those
-of the living species being found abundantly in the Cambrian strata. Why
-it is that Lingula has been able to maintain itself almost unchanged
-through all the countless generations that have elapsed since Cambrian
-times, and can now flourish amid the desperate struggle for existence in
-the shallow waters of the tropics, while its companions, the corals,
-mollusks, arthropods, &c., have changed or passed away, is one of those
-problems in natural history that seem to us impossible of solution. The
-time may come when we shall be able to appreciate better than we do now
-the complicated relations between animals and their environment, and
-then perhaps the peculiar fitness of Lingula will be made manifest; but
-at present we can but mention the fact as a fact, and leave the solution
-of the problem to the future.
-
-The order Gephyrea is probably another very ancient group of animals,
-although in the absence of any hard calcareous, siliceous, or horny
-skeleton the geological record can give us no confirmation of their
-antiquity. As with the Brachiopods so with the Gephyrea, some of the
-species have a very wide bathymetrical distribution. _Sipunculus nudus_,
-for example, the commonest and best known of all the Gephyrea, extends
-from quite shallow water to a depth of over 1,500 fathoms. As Selenka
-has pointed out, it is those Gephyreans that live in holes in stones, or
-in shells such as _Phascolion_ and _Phascolosoma_, that are more
-frequently found at the greater depths; but, apart from this, there are
-no characters that exclusively belong to the abysmal Gephyrea or are
-more frequently found in them than in the shallow-water forms. Nor are
-there any genera, at present brought to light, that are confined to
-those regions of the sea.
-
-The group of the Annelida is not very well represented in the deep-sea
-fauna. The genera _Serpula_ and _Terebella_ have been found very widely
-distributed over the earth, at all depths from the shore to the abyss,
-but there do not seem to be many genera that are confined to deep water.
-In some cases, where there is a scarcity of lime in the water, the thin
-protecting tubes of the sedentary forms are strengthened by the adhesion
-of foreign particles, such as sponge spicules and arenaceous
-foraminifera, but in others, the tubes are formed of successive layers
-of a transparent quill-like substance (_Nothria Willemoesii_) which is
-frequently armed with spiny projections.
-
-Most of the errant Polychætes found at great depths are said to be most
-brilliantly coloured, and some of these, such as _Eunice amphiheliæ_,
-have the power of emitting a bright phosphorescent light; but there seem
-to be no very definite and constant characters separating these forms
-from the Polychætes of shallow waters.
-
-As is the case with many other orders of animals, the species of
-Annelida living in deep water are either blind or possess eyes of a
-remarkably large size. _Genityllis oculata_ may be taken as an example
-of a deep-sea annelid with large eyes. This annelid, belonging to the
-family Phyllodocidæ, was found at a depth of 500 fathoms in the Celebes
-sea. It possesses two enormous eyes which cover almost the whole of the
-head, and there can be no doubt, from the investigations of Dr. Gunn on
-their minute anatomy, that they are perfectly functional.
-
-Before leaving the Annelida a brief notice must be made of the very
-extraordinary form _Syllis ramosa_, found parasitic on a hexactinellid
-sponge at depths of about 100 fathoms. It is chiefly remarkable for the
-very complicated manner it has of producing buds which do not
-immediately become detached from the parent, but form a compound network
-which ramifies through the interstices of the sponge like the colony of
-a Hydromedusan.
-
-Passing now to the sub-kingdom Mollusca, we shall find that all the
-classes are represented in the abysmal fauna.
-
-The Lamellibranchiata, or bivalves, occur in almost all depths of the
-ocean, _Callocardia pacifica_ and _Callocardia atlantica_ having been
-found at the enormous depth of 2,900 fathoms. Some species, such as
-_Venus mesodesma_, have a very wide bathymetrical distribution, but
-others are only known to occur in deep water.
-
-Concerning the characters of the deep-sea Lamellibranchiates, Mr. Smith,
-in his report on the Lamellibranchia of the ‘Challenger’ expedition,
-says ‘very deep-water benthal species certainly have a tendency to be
-without colour, and of thin structure, no doubt resulting from the
-absence of light the difficulty of secreting lime, the scarcity of food
-and other unfavourable conditions of existence.’ But notwithstanding
-this, the same author continues: ‘The species are apparently few in
-number in comparison with those of shallow water; and new and peculiar
-generic forms which we naturally expected would have been discovered are
-of even still rarer occurrence.’
-
-As in the group of the Lamellibranchia, so in the Gasteropoda, no very
-remarkable new genera or species have been found in deep sea. Some
-shallow-water genera, such as _Fusus_ for example, have representative
-species in the abyss; but, with the exception of a want of brilliant
-coloration and marking and a thinness of the shell, the deep-sea forms
-do not exhibit any characteristic features. One of the most brightly
-coloured shells found at great depths is that of _Scalaria mirifica_,
-which is tinted rose and white, but this seems to be quite an
-exceptional character among the deep-sea Gasteropods. Several new genera
-were found in the deep water, but their general characters do not call
-for any special remark.
-
-Among the Cephalopoda there seems to be little doubt that the genera
-_Cirroteuthis_, _Bathyteuthis_, and _Mastigoteuthis_ are entirely
-abysmal, and the same applies probably to one or two species of octopus;
-but as Hoyle remarks, ‘apart from the single fact that _Bathyteuthis_
-and _Mastigoteuthis_ both have slender filiform tentacles with minute
-suckers, no structural features have been discovered which will serve to
-diagnose a deep-sea form from a shallow-water one.’
-
-The exact habitat of the interesting genus _Spirula_ is unfortunately
-still unknown. In some parts of the tropics the shores are covered with
-spirula shells, and yet the animals that secrete them are still to be
-reckoned amongst the greatest rarities of our museums. The numerous
-dredgings of the ‘Challenger’ only brought to light one specimen of this
-animal, and that from a depth of 360 fathoms, and the ‘Blake’ caught one
-at a depth of 950 fathoms, so that there can be little doubt that
-_Spirula_ lives in deep water.
-
-[Illustration: FIG. 14.—_Bathyteuthis abyssicola._ (After Hoyle.)]
-
-It seems to be very probable that some day, when the right place and
-depth are discovered, _Spirula_ may be discovered in great abundance,
-but we have at present no means of judging whether this will be in very
-deep water or not.
-
-Almost precisely similar remarks apply to the distribution of the pearly
-Nautilus. The shells of this Cephalopod are sometimes found in great
-number on the shores of some of the islands of the Southern Pacific
-Ocean and the Malay Archipelago, but the living animal is but rarely
-captured. It has been asserted by some travellers that the pearly
-Nautilus floats on the surface of the ocean and possesses the power of
-suddenly diving to great depths on being disturbed; but it must be
-remembered that Rumphius originally caught his Amboyna specimens of
-Nautilus in traps set at a depth of 200 fathoms baited with sea-urchins,
-and that the ‘Challenger’ captured a single living specimen off Matuku
-island in 300 fathoms.
-
-The probability, then, is that both _Nautilus_ and _Spirula_ should be
-included in the deep-sea fauna, but we are still in want of a great deal
-more information concerning their life and habits before this point can
-be definitely determined.
-
-
-
-
- CHAPTER VII
- THE ARTHROPODA OF THE DEEP SEA
-
-
-The deep-sea fauna seems to be particularly rich in marine Arthropoda,
-many curious and interesting forms being brought up with almost every
-haul of the dredge. The Arthropoda, too, being very highly organised
-animals, afford interesting and instructive examples of the effect of
-abysmal life in the modification of the sense organs and the production
-of varieties specially modified for the conditions of the struggle for
-existence in their strange habitat.
-
-Concerning the groups of Ostracoda and Copepoda it may be said that the
-evidence is not yet conclusive that they include any truly deep-sea
-species. The largest known Ostracod, measuring somewhat more than an
-inch in length and probably allied to the genus _Crossophorus_, has
-quite recently been captured by Professor Agassiz at depths of less than
-200 fathoms, but he could obtain no evidence that it descended into much
-deeper water than this.
-
-Mr. Brady, in writing the report of the ‘Challenger’ Ostracoda, came to
-the conclusion that they do exist in very limited numbers in the most
-profound depths of the sea; but it is nevertheless quite possible that
-all the Ostracods brought on deck by the trawl or dredge were really
-captured either on the way down or on the way up, and are, strictly
-speaking, pelagic in habit.
-
-Similar caution must be taken in dealing with the Copepoda, an order of
-Crustacea that is essentially pelagic in habit. The only species that
-has been regarded as undoubtedly abysmal is _Pontostratiotes
-abyssicola_, a form whose carapace and antennæ are armed with
-exceedingly long and strongly toothed spines, and was found in the mud
-brought up by the trawl from a depth of 2,200 fathoms.
-
-_Calamus princeps_, the largest species of its genus of a deep reddish
-brown colour, may also belong to the fauna of the deep sea, but we have
-less evidence concerning the habitat of _Hemicalamus aculeatus_,
-_Phyllopus bidentatus_, and some of the Euchætæ.
-
-The Amphipoda seem to be but poorly represented in the fauna of the
-abyss; in fact it may be considered to be still an open question whether
-any Amphipods habitually live in very deep water.
-
-In the reports on the ‘Challenger’ Amphipoda, the Rev. T. R. Stebbing
-states that thirty-one specimens are known to come from great depths,
-but it would be more correct to say that these specimens were found in
-the dredges and trawls that had been lowered into the great depths. It
-should be noticed, however, that some of these specimens do show
-characters that suggest, at any rate, that they come from deep water.
-Thus the genus _Lanceola_, for example, is characterised by the
-smallness of the eyes and a soft membranous integument, while _Cystisoma
-spinosum_, found in a dredge that had been at work at a depth of over a
-thousand fathoms, has very large eyes.
-
-In his report on the Crustacea of the ‘Norske Nord-havns’ expedition,
-Professor Sars gives a full description of many species of Amphipoda
-brought by the dredge from depths of over 1,000 fathoms, and nearly all
-of these were found to be quite blind.
-
-The form that seems to be most peculiar to the great depths of the
-Northern Ocean is _Harpinia abyssi_. It was found at no less than
-fifteen different stations at depths ranging from 350 to 2,215 fathoms,
-and is characterised by its large size and the total absence of eyes.
-
-Another point that should be considered in coming to any conclusion on
-the supposed habitat of such forms, is the similarity or dissimilarity
-of widely distributed species.
-
-I have had occasion to point out in a previous chapter the general
-similarity of the abysmal fauna all over the world, a very striking
-phenomenon, commented on by almost every naturalist who has had a wide
-experience of this kind of investigation.
-
-Among the Amphipoda we have a very striking example of this. The species
-_Orchomene musculosus_ was taken by the ‘Challenger’ off the southern
-part of Japan at a depth of 2,425 fathoms, the bottom being red clay and
-the temperature 35·5° Fahr. The species _Orchomene abyssorum_ was taken
-off the east coast of Buenos Ayres at a depth of 1,900 fathoms, the
-bottom being blue mud and the temperature 33·1° Fahr. To the description
-of this last-named species Mr. Stebbing adds, ‘had this species been
-taken within reasonable distance of _O. musculosus_, the resemblance is
-so great that one might have been tempted to disregard the points of
-difference as due to some other cause than difference of species.’
-
-Such a striking similarity between two species living so far apart from
-one another may, when we take into consideration the depth, the
-character of the bottom, and the temperature from which they are
-supposed to have been dredged, be taken to support very strongly the
-view that these species are really abysmal in habit.
-
-Among the Isopoda we have several very characteristic forms—no fewer
-than nine distinct genera peculiar to the abysmal zone have been
-described by Beddard—and of these two, _Bathynomus_ and _Anuropus_, are
-to be regarded as types of sub-families. They seem to be very unevenly
-distributed over the floor of the ocean, some regions, such as the whole
-of the Central and Southern Atlantic and the Central and Western
-Pacific, produce none; whilst the waters of the east coast of New
-Zealand, the Crozets, and others, produce a great many varieties. Many
-of the deep-sea Isopoda exhibit characters that are usually associated
-with the bathybial life. Thus, according to Beddard, thirty-four of the
-deep-sea species are totally blind, and eighteen have well-developed
-eyes. In four species there are eyes which are evidently degenerating.
-If we compare, for instance, the structure of the eye of _Serolis
-schythei_, a species found in shallow water ranging from 4 to 70
-fathoms, with the eyes of _Serolis bromleyana_, a species living in deep
-water ranging from 400 to 1,975 fathoms, we cannot fail to see that the
-latter are undergoing a process of degeneration; the retinulæ and
-pigment being absent, and nothing left of the complicated structure of
-the Isopod eye but the remnants of the crystalline cones and corneal
-facets (see figs. 4 and 5, p. 74).
-
-Taking the genus _Serolis_ alone, it has been said ‘that in all the
-shallow-water forms the eye is relatively small but very conspicuous
-from the abundant deposition of pigment; in all the deep-sea forms, with
-the exception of _S. gracilis_, where the eye seems to be disappearing,
-it is relatively larger but not so conspicuous, owing to the fact that
-little or no pigment is present.’
-
-In many groups of animals it has been shown that some of the deep-sea
-species are relatively much larger in size than the shallow-water
-species, and that others, more rarely, are much smaller, the abysmal
-fauna reminding us in this respect of the characters of the alpine
-flora.
-
-The Isopoda show many examples of this largeness in size, thus
-_Bathynomus giganteus_, dredged by Professor Agassiz off the Tortugas at
-a depth of over 900 fathoms, reaches the enormous size, for an Isopod,
-of 9 inches (fig. 15). _Stenetrium haswelli_, again, is larger than any
-of the shallow-water species of the genus, and the same remark applies
-to the deep-sea species of the genus _Ichnosoma_, while _Iolanthe
-acanthonotus_, from a depth of nearly 2,000 fathoms, is considerably
-larger than most of the shallow-water Asellidæ.
-
-There is another very common character of deep-sea Crustacea that is
-also well exemplified in the group of the Isopods, and that is the
-extraordinary length and number of the spines covering the body.
-
-I have already referred to this character in the supposed deep-sea
-Copepod _Pontostratiotes abyssicola_, and I shall have again to refer to
-it in treating of the Decapoda and other groups of the Crustacea.
-
-Besides its enormous size _Bathynomus_ possesses some other characters
-that may be correlated with its deep-sea environment. The respiratory
-organs are quite different from those of other Isopods; instead of being
-borne by the abdominal appendages, they are in the form of branched
-outgrowths from the body-wall containing numerous blood-lacunæ, and the
-appendages simply act as opercula to cover and protect them. The eyes of
-the Bathynomus too are remarkably well developed, each one bearing 4,000
-facets, and they are directed not dorsally as in the Cymothoadæ, but
-ventrally. The cause of these curious modifications of structure in
-Bathynomus is by no means clear, but it is quite probable that they are
-connected with the conditions of pressure and light in the deep sea. It
-is a remarkable fact that the other deep-sea Isopods do not exhibit
-precisely these modifications, and it might be supposed that the same
-causes would produce the same or similar effects on the structure of
-animals belonging to the same order. That is perfectly true, but we
-cannot yet determine how long ago any one species has taken to a
-deep-sea life, or what length of time, in other words, these conditions
-have been at work in modifying the structure of the organism. A recent
-immigrant into the abyss will naturally exhibit closer affinities with
-its shallow-water allies than those that have dwelt in the region since
-secondary or tertiary times. If we take this into consideration we
-should expect to find considerable differences occurring between
-deep-sea species of the same order, which is precisely what we do find.
-
-[Illustration: FIG. 15.—_Bathynomus giganteus._ From a depth of 1,740
-metres. (From Filhol.)]
-
-Concerning the Cirripedia, that curious group of profoundly modified
-Crustacea that includes the barnacles and acorn shells, Dr. Hoek writes
-in the ‘Challenger’ monograph:—
-
-‘Though unquestionably by far the greater part of the known Cirripedia
-are shallow-water species, and though some of the species are capable of
-living at a considerable variety of depths, as, for instance,
-_Scalpellum stroemii_, yet it must be granted that the number of true
-deep-sea species of Cirripedia is very considerable.’ Only two genera,
-however, occur in depths of over 1,000 fathoms, and these—_Scalpellum_
-and _Verruca_—occur also as fossils in secondary and tertiary deposits.
-The oldest of all fossil cirripedes, however, namely, _Pollicipes_,
-never occurs, at the present day, in deep water, but is purely littoral
-or neritic in habit. But what is perhaps more interesting still is the
-fact, that, when we come to compare the living and the fossil species,
-we find that in the one genus (_Scalpellum_) the deep-sea forms have
-preserved the more archaic characters, and in the other (_Pollicipes_)
-the shallow-water forms.
-
-Here then we are presented with a veritable puzzle for which we can at
-present frame no manner of answer. Pollicipes on the one hand—like
-Lingula among the brachiopods—has been able to maintain itself almost
-unchanged amid the tremendous struggle for life of the shallow water of
-the tropics ever since the Lower Oolite epoch; while Scalpellum, on the
-other hand, has either become profoundly modified, or been driven into
-the abysmal depths of the ocean.
-
-The group of the Thoracostraca, or stalk-eyed Crustacea, including
-lobsters, crabs, hermit crabs, prawns, and shrimps, is well represented
-in the deep sea. Most of them are characterised by being quite blind (in
-many cases even the eye-stalks are obliterated), by being protected with
-a dense covering of spines, by the thinness of their shells, and by
-their bright red or carmine colour.
-
-The order Stomatopoda is almost entirely confined to the shallow waters
-of the tropical or temperate shores. Not a single species is known to
-inhabit the deep sea, and only a very few specimens have been captured
-in more than a few fathoms of water.
-
-The Schizopoda, however, present us with many curious abysmal forms.
-Most of the genera of this order belong to the pelagic plankton, and
-many of them are known to possess the power of emitting a very strong
-phosphorescent light. Several genera, however, such as _Gnathophausia_,
-_Chlaraspis_, _Eucopia_, _Bentheuphausia_, &c., never seem to leave the
-great depths of the ocean, and nearly all of these genera are
-distinguished by being quite blind or possessing very much reduced or
-rudimentary eyes.
-
-[Illustration: FIG. 16.—_Euphausia latifrons_, from the surface of the
-sea. (After Sars.)]
-
-[Illustration: FIG. 17.—_Bentheuphausia amblyops_, from 1,000 fathoms.
-(After Sars.)]
-
-If we compare, for example, the pelagic _Euphausia latifrons_ (fig. 16)
-with the nearly allied but abysmal _Bentheuphausia amblyops_ (fig. 17),
-the difference in this respect between a Schizopod living in the
-sunlight and one living in the darkness of the deep-sea is very
-apparent.
-
-The pelagic Schizopoda are usually quite pale and transparent; the
-deep-sea forms on the other hand are frequently if not invariably of a
-bright red colour, as is the case with many other deep-sea Crustacea to
-which reference will be made later on.
-
-Passing on to the group of the Decapoda, we find that the most
-interesting of all the abysmal cray-fish is the family of the Eryonidæ;
-indeed, in some respects the discovery of these curious forms may be
-reckoned among the most valuable results of the ‘Challenger’ Expedition.
-They are characterised by the dorsal depression of the anterior part of
-the cephalothorax, the absence of a rostrum, and the absence or very
-rudimentary condition of the eyes (fig. 18).
-
-Their nearest relations seem to be certain genera of Crustacea that are
-found in jurassic strata, in the lias, and more particularly in the
-lithographic slates of Solenhofen.
-
-They have a very wide bathymetrical range extending from a depth of 250
-fathoms (_Polycheles crucifera_) to a depth of 2,000 fathoms
-(_Willemoesia_).
-
-[Illustration: FIG. 18.—_Polycheles baccata_, one of the Eryonidæ. The
-eyes and eye-stalks are absent, and the margin and sides of the carapace
-armed with spines. (After Spence Bate.)]
-
-But there are many other curious forms of the macrurous crustacea that
-deserve a passing mention. The graceful _Nematocarcinus gracilipes_,
-distinguished by the extraordinary length of the antennæ and last four
-pairs of legs, these appendages being three or four times the length of
-the body, is by no means rarely met with in depths of over 400 fathoms.
-
-The genus _Glyphus_ captured by the ‘Talisman’ is remarkable for the
-development of a peculiar pouchlike arrangement on the abdomen for the
-protection of the larvæ during the younger stages of their existence.
-
-The proof of the existence of a peculiar cray-fish, _Thaumastocheles
-zaleuca_, at a depth of 450 fathoms, was one of the most important
-contributions to carcinology made by the ‘Challenger’ Expedition. The
-chelæ of this remarkable form are of great but unequal length and armed
-with long tooth-like spines giving it an appearance not unlike that of
-the jaws of some carnivorous fish. The shell is soft and the abdomen
-broad and flattened. There are no eyes nor even eye-stalks, but ‘in
-front of the carapace,’ as Sir Wyville Thomson remarks, ‘between the
-anterior and upper edge and the insertions of the antennæ, in the
-position of the eyes in such forms as _Astacus fluviatilis_, there are
-two round vacant spaces, which look as if the eye-stalks and eyes had
-been carefully extirpated and the space they occupied closed with a
-chitinous membrane.’ The deep-sea prawn, _Psalidopus_, recently taken in
-500 fathoms of water by the ‘Investigator,’ affords us an example of a
-common bathybial character, the whole body being covered with an
-extraordinary array of sharp needle-like spines.
-
-Among the crabs many curious forms have been found in deep water
-extending down to depths of over 2,500 fathoms. They are nearly all
-characterised by blindness and a remarkable development of tooth-like
-spines covering the carapace and limbs.
-
-The remarkable _Lithodes ferox_, from a depth of from 450 to 800
-fathoms, is perhaps the most perfectly armed crab—in the way of
-spines—that exists. Every part of the body and limbs is so covered with
-spines that one has to be extremely careful in handling even a dead
-specimen.
-
-This is only one of the many examples that might be given to illustrate
-this curious feature of the deep-sea Crustacea. Among the crabs alone we
-have such forms as _Galathodes Antonii_, _Pachygaster formosus_,
-_Dicranodromia mahyeuxii_ covered with a fierce armature of spines or
-bristles; but there are nevertheless some species in which this
-character is not particularly noticeable, and in these we usually find
-some other protection against their enemies. An interesting example of
-this has been described by A. Agassiz in a crab allied to the Maiadæ,
-‘in which the dorsal face appears like a bit of muddy area covered by
-corals, with a huge white arm resembling a fragment of an Isis-like
-gorgonian.’ It is evident that this is a case in which the animal is
-protected by its resemblance to the surroundings.
-
-The hermit crabs of the abyss, too, are not usually characterised by any
-very great development of spines. They find their protection in the
-shells they inhabit. Some of the deep-sea hermit crabs carry about with
-them on their shells a sea anemone, as we find to be frequently the case
-among the shallow-water species. _Pagurus abyssorum_, from a depth of
-3,000 fathoms, is an example of this.
-
-In cases where there is a scarcity of gasteropod shells the hermit crabs
-are obliged to find some other form of protection for their bodies. The
-‘Blake’ found in the West Indies a hermit crab that had formed for
-itself a case of tightly compressed sand, and another curious form,
-named _Xylopagurus rectus_, makes its home in pieces of bamboo or in the
-holes in lumps of water-logged wood.
-
-The last group of the Arthropoda we need refer to is that of the
-Pycnogonida, those curious creatures seemingly made up entirely of legs,
-and by some naturalists considered to be related to the Crustacea and by
-others to the scorpions and spiders.
-
-Like the Brachiopoda the Pycnogonida are not usually found in greater
-depths than 500 fathoms. Out of the twenty-seven known genera, only five
-extend into the abyss, and not one of these can be called a true
-deep-sea genus.
-
-There are three genera, _Nymphon_, _Collosendeis_, and _Phoxichilidium_,
-that show a very wide distribution over the floor of the ocean, and are
-capable of existing at the greatest depths, and of these the species of
-the genus Nymphon have a truly remarkable range extending from the shore
-to a depth of 2,225 fathoms.
-
-‘As a rule,’ says Hoek, ‘the deep-sea species are slender, the legs very
-long and brittle, and the surface of the body smooth.’ They have
-further, either no eyes at all or rudimentary eyes without pigment, and
-in many cases—as, for example, _Collosendeis_—they are distinguished for
-reaching to a gigantic size compared with their shallow-water relatives.
-
-The Tunicata is the group of animals that includes all those curious
-vegetable-like organisms found upon our coasts that are familiarly known
-as sea-squirts, or Ascidians, besides the salps, pyrosomas, and the
-microscopic appendicularias of the pelagic plankton.
-
-[Illustration: FIG. 19.—_Collosendeis arcuatus_, from a depth of 1,500
-metres. (After Filhol.)]
-
-Notwithstanding the apparent simplicity of their adult structure,
-naturalists are now agreed that they must be removed from the Mollusca,
-with which they have hitherto been most frequently associated, and
-placed in the group of the Vertebrata. It is the study of embryology
-that has led to this unexpected conclusion, for we find, when we study
-the larval forms, that they possess both a notochord and gill-slits, two
-features that are characteristic of the group of the Vertebrata.
-
-The species of the group Perennichordata, which includes all those
-Tunicates that possess a notochord persistent through life, are chiefly
-pelagic in habit, the little creatures, rarely more than two or three
-millimetres in length, swimming or drifting about with the sagittas,
-copepods, ctenophores, and medusæ that compose the pelagic plankton. Fol
-has recently described a gigantic form belonging to this group, reaching
-a size of thirty millimetres in length, called _Megalocercus abyssorum_,
-which he dredged from a depth of 492 fathoms; and other species have
-been recorded down to a depth of 710 fathoms in the Mediterranean Sea.
-
-Among the simple Ascidians we find no family that is peculiar to deep
-water; but the Cynthiidæ and Ascidiidæ both contain genera that are
-abysmal, and the Molgulidæ have one species, _Molgula pyriformis_, that
-extends into the abysmal zone to a depth of 600 fathoms.
-
-In the genus _Culeolus_ and in _Fungulus cinereus_ and _Bathyoncus_, all
-deep-water Ascidians, there is a very curious modification of the
-branchial sac, the stigmata being apparently not formed, in consequence
-of the suppression of the fine inter-stigmatic vessels. This peculiar
-feature is only found in the deep-sea simple Ascidians and, as we shall
-see presently, in one species of the deep sea compound Ascidians, but it
-is not apparently an essential character of those living in the abysmal
-zone, notwithstanding the fact that it is found in such widely separated
-genera; for _Corynascidia_, _Abyssascidia_, and _Hypobythius_, living in
-depths lying between 2,000 and 3,000 fathoms below the surface, have
-branchial sacs of the ordinary type. Professor Herdman is of opinion
-that this simple form of branchial sac is not a primitive form, but most
-probably a modification of a more complicated type.
-
-In _Culeolus Murrayi_ there is a remarkably abundant supply of
-blood-vessels to the tunic, and these send special branches to a number
-of small papilliform processes on its outer surface. This system of
-highly vascular processes probably constitutes, as Professor Herdman
-suggests, an additional or complementary respiratory apparatus. All
-these modifications of the branchial system are of particular interest,
-for we find so many instances of a similar kind among the inhabitants of
-very deep water. I need only refer here to the modifications of this
-system in the Isopod _Bathynomus_ already referred to (p. 129), and to
-the reduction in the number of the gills of many of the deep-sea fishes.
-Why there should be such modifications is a question upon which the
-physical and natural history investigations of the conditions of life in
-the great depths of the ocean at present throw no light.
-
-In a previous chapter I have referred to the fact that many of the
-bathybial animals are characterised by being stalked. Among the simple
-Ascidia we find many examples of stalked kinds living in deep water,
-such as _Culeolus_ and _Fungulus_, but also several exceptions, such as
-_Bathyoncus_, _Styela bythii_, and _Abyssascidia_, that are sessile. It
-is a noteworthy fact, however, that the genus that has the most deep-sea
-species—namely, _Culeolus_—is a genus that is provided with a very long
-stalk. Furthermore, the only known stalked forms of the very large
-family Ascidiidæ are the abysmal genera _Corynascidia_ and
-_Hypobythius_.
-
-[Illustration: FIG. 20.—_Hypobythius calycodes._ G, nerve ganglion; H,
-heart; M, the position of the atriopore. The large opening on the upper
-side is the mouth. (From a drawing by Professor Moseley in Herdman’s
-‘Tunicata of the “Challenger” Expedition.’)]
-
-The most remarkable character of the genus _Hypobythius_ is the simple
-condition of its branchial sac, reminding one of the structure of this
-organ in the shallow-water genus _Clavelina_. ‘There are no folds and
-there are no internal bars,’ to quote the description given by Professor
-Herdman; ‘only a single system of vessels can be recognised, branching
-and anastomosing so as to form a close network, the small rounded meshes
-of which are the stigmata. The tentacles and dorsal lamina cannot be
-made out.’
-
-Among the compound Ascidians only four families extend into the abysmal
-zone, namely, the Botryllidæ, Polyclinidæ, Didemnidæ, and Cœlocormidæ,
-and of these only one species, _Pharyngodictyon mirabile_, of the family
-Polyclinidæ, extends into water of greater depth than 1,000 fathoms. In
-_Pharyngodictyon_ we find the same curious simplification of the
-branchial sac that we have just referred to in the genera of simple
-Ascidians, _Culeolus_, _Fungulus_, and _Bathyoncus_. _Cœlocormus
-Huxleyi_ from a depth of 600 fathoms is a very peculiar form and the
-type of a separate family, the Cœlocormidæ.
-
-The free-swimming Tunicata included in the group _Ascidiæ salpiformes_,
-which contains the genus _Pyrosoma_, and the order Thaliacea containing
-the salps, are in all probability mainly confined to the surface waters.
-A few specimens of _Pyrosoma_ were captured by the ‘Challenger’ dredges
-which came up from very deep water, but it is doubtful at what point in
-the journey to the surface the specimens entered the net.
-
-The most remarkable form of free-swimming Tunicate that has come to
-light is _Octacnemus bythius_, a form that is probably allied to
-_Salpa_. It was found twice, once in the dredge that came from a depth
-of 1,070 fathoms, and once from 2,160 fathoms. The tunic of the animal
-is gelatinous and hyaline, but the most important feature it possesses
-is an imperforate membrane separating the branchial sac from the
-peribranchial cavity. Octacnemus, in other words, possesses no true
-stigmata, these structures being represented only by little pits in the
-walls of the branchial sac. This curious and extremely interesting
-modification of the respiratory organs points very strongly to the
-conclusion that Octacnemus is truly a deep-sea animal.
-
-
-
-
- CHAPTER VIII
- THE FISH OF THE DEEP SEA
-
-
-Of all the groups of animals that constitute the deep-sea fauna, the
-fish show the greatest number of peculiarly abysmal characters. Being
-much more highly differentiated than the invertebrates, they possess
-more organs liable to undergo modifications of colour, size, and
-structure, and consequently we are able to point to a great many more
-features characteristic of deep-sea fish than we can do in any other
-group of animals.
-
-The first point that calls for remark in the consideration of the fish
-fauna of the deep sea is the almost entire absence of ancient and
-primitive types. The Elasmobranchii, including the Sharks, Rays, and
-Chimæra, constituting the order that from anatomical embryological
-grounds is always regarded by naturalists as the most primitive order of
-this class, is represented in very deep water by only one species. _Raia
-hyperborea_ and _Chimæra monstrosa_, it is true, just enter into the
-abysmal zone, but _Chimæra affinis_ is the only Elasmobranch that
-extends to depths of over 1,000 fathoms.
-
-The Ganoidei too, the order that in palæozoic and mesozoic times was so
-rich in genera and species, is entirely absent from the abysmal zone,
-not a single representative having been found at any time by any of the
-deep-sea expeditions.
-
-The Dipnoi, that remarkable order including the three fresh-water
-genera, _Ceratodus_ from Australia, _Lepidosiren_ from Brazil, and
-_Protopterus_ from West Africa, has no representative and no ally in the
-deep waters of the ocean.
-
-The fishes of the deep sea, in fact, with only one or two exceptions,
-all belong to the Order Teleostei, the most modern and most highly
-differentiated order of the class, the families that are most fully
-represented being the Macruridæ and then the Ophidiidæ and Gadidæ, and
-the Berycidæ.
-
-At the limits of the katantic and abysmal zones, a large number of
-families of Teleosteans entirely disappear, and as we approach the
-deepest parts of the ocean, the number of fish that are found is
-considerably reduced. As Dr. Günther very wisely remarks, ‘this
-diminution in the number may be due to the difficulty of capturing
-fishes at great depths, a difficulty which increases in proportion to
-the depths at which the dredge is worked. But it must also be regarded
-as evidence of the actually diminished variety of fishes.’
-
-It may be interesting to the reader to give Dr. Günther’s table of the
-number of species found at different depths, as it shows, among other
-things, the marked change that occurs in the character of the fauna in
-passing from the katantic to the abysmal zone.
-
- Between 100–300 fathoms, 232 species
- Between 300–500 fathoms, 142 species
- Between 500–700 fathoms, 76 species
- Between 700–1,500 fathoms, 56 species
- Between 1,500–2,000 fathoms, 24 species
- Between 2,000–2,900 fathoms, 23 species
-
-As regards the general character presented by the deep-sea fishes, I
-have already pointed out in the chapter dealing with the general
-characters of the deep-sea fauna, the peculiarities in the size of the
-eyes, the colours and markings of the body, and the texture of the bones
-and muscles. There are, however, a few more characters of which mention
-must be made.
-
-Notwithstanding the fact that all the abysmal fishes are carnivorous and
-must consequently be capable—in the great number of cases—of rapid and
-vigorous movement, the muscles of the trunk and tail are usually thin,
-and the fascicles loosely connected with one another.
-
-Deep-sea fish are not characterised by an absence of the swimming
-bladder. This organ occurs just as frequently and in the same families
-as in the shallow-water fauna, but we do not know whether it possesses
-any special peculiarities or not, as it is usually so ruptured and
-destroyed by the change of pressure it undergoes in being brought to the
-surface, that it is impossible to make any thoroughly accurate
-investigation of its anatomy and relations.
-
-The extraordinary development of glands in the skin which secrete mucus,
-and the presence in many forms of very highly specialised organs for
-emitting phosphorescent light, are characters of the deep-sea fish
-fauna, to which I have referred in a previous chapter.
-
-As with the Tunicates, some of the Crustacea and other groups, the fish
-of the abysmal zone show curious modifications of the respiratory
-system. The gill laminæ of these animals are not only reduced in number,
-but appear to be short and shrunken. It is possible, of course, that
-during life they may end in fine delicate points which are broken off or
-ruptured during their capture, but still the horny rods that support
-them are shorter than they are in shallow-water forms, and the general
-evidence of their structure tends to show that they have undergone
-profound modifications in the change to the conditions of deep-sea life.
-
-An extremely common and almost general character of deep-sea fishes is
-the black coloration of some of the body cavities; this is limited to
-the pharynx in many of the fishes that live about the hundred fathoms
-limit, but the colour is more intense and spread all over the oral,
-branchial, and peritoneal cavities in typical deep-sea forms. It may
-seem very difficult at first to account for this remarkable development
-of black pigment in parts of the body that are not usually, and, in some
-cases, cannot at any time be exposed to view. It is obvious that it
-cannot be functional as a hiding colour, either in offence or defence.
-But it is quite possible that it is due to some modification of the
-function of excretion. It is well known that in many cases of disease or
-injury to the kidneys in vertebrates, the colour of the skin is
-affected, and every one recognises now the fact that in many
-invertebrates the colour of the skin is greatly dependent upon the
-function of the secretion of the urates.
-
-It would at least be interesting to know if this dark coloration of the
-mucous membranes is in any way correlated with any modification of the
-structure or function of the kidneys. At present we have no recorded
-observations on this point, but it is to be hoped that, when we have a
-sufficient number of specimens brought home from the deep water, a
-systematic investigation of this subject will be made.
-
-Lastly, it should be pointed out that our knowledge of the abysmal fauna
-has not, at present, brought to light any evidence that the fish are of
-an extraordinarily large size. In many groups of animals, as I have
-frequently pointed out in the last few chapters, the large and gigantic
-species or specimens are only found in the abyss. This may also be the
-case with fishes, but we have no evidence that it is so. The only
-methods that have been used at present for the investigation of the
-fauna living on or near the floor of the deep oceans, are not of a kind
-to lead to the capture of really large fish. That they may exist is
-highly probable, but all that we know at present is, that the fish with
-which we are acquainted living at great depths are not in any way
-remarkable for their great size.
-
-Of the only two Elasmobranchs, one, namely _Raia hyperborea_, has been
-found in water extending from 400 to 608 fathoms in depth. Only four
-specimens have yet been taken, one by the Norwegian expedition off
-Spitzbergen and three by the ‘Knight Errant’ off the northern coasts of
-Scotland. It is interesting to find that this, the only deep-sea species
-of the Rays, shows some striking peculiarities. ‘The teeth are
-remarkably slender,’ says Günther, ‘small, irregularly and widely set,
-different from those of other British Rays. In young specimens at any
-rate those of the male do not differ from those of the female. The
-mucous membrane behind the upper jaw forms a pad with a lobulated
-surface. The mucous cavities of the head are extremely wide, and finally
-the accessory copulatory organs have a spongy appearance, and are
-flexible, the cartilage by which they are supported being a simple
-slender rod.’
-
-The other Elasmobranch, that extends into very deep water, is _Chimæra
-affinis_, a species which can hardly be distinguished from the better
-known _Chimæra monstrosa_, a fish that itself very frequently wanders
-within the limits of the abysmal zone.
-
-Among the Teleostei, the family Berycidæ has several representatives in
-the deep water. They are small fish rarely exceeding four inches in
-length, with large heavy heads, with functional but small eyes, and an
-abundant supply of large mucous glands on the skin.
-
-_Melamphaes beanii_, belonging to this family, has been captured at the
-enormous depth of 2,949 fathoms.
-
-_Bathydraco antarcticus_, belonging to the family Trachinidæ, from a
-depth of 1,260 fathoms, is an example of a true abysmal fish possessing
-very large eyes.
-
-The Pediculati, the family of the anglers, is represented at depths of
-over 2,000 fathoms by the interesting form _Melanocetus Murrayi_. The
-eyes are very small indeed, the mouth huge and armed with long uneven
-rasp-like teeth. At the end of the fishing-rod tentacle hanging over the
-mouth, there is an organ that has been supposed to be capable of
-emitting a phosphorescent light. This curious modification of the red
-worm-like bait of the common shallow-water angler into a will-o-the-wisp
-lantern attracting little fishes to their destruction in the deadly jaws
-of the _Melanocetus_ is one of the most interesting adaptations that
-have been brought to light by our study of the deep-sea fauna.
-
-Several species of the family Lycodidæ occur in the abysmal zone, but
-they do not possess any features that call for special mention in this
-place.
-
-[Illustration: FIG. 21.—_Melanocetus Murrayi_, 1,850–2,450 fathoms.
-(After Günther.)]
-
-The family Ophidiidæ contributes very largely to the fish fauna of the
-abyss. Some of the deep-water genera, such as _Neobythites_, have a wide
-bathymetrical distribution extending from 100 fathoms to depths of over
-2,000 fathoms, but others, such as _Bathyonus_, _Typhonus_, and
-_Aphyonus_, only occur in depths of over 1,000 fathoms.
-
-The body is usually elongate and slender, ending in a pointed tail, the
-head large and heavy, and the eyes, in the genera confined to the
-abysmal zone, usually so far degenerated that they are not visible at
-all from the outside.
-
-The Macruridæ form a family that contributes very largely to the
-deep-sea fauna; no fewer than twenty-six different species are known to
-occur within the limits of the abysmal zone. Not only do the Macruridæ
-contribute a large number of different species, but they probably occur,
-in some districts at any rate, in vast numbers.
-
-During the voyage of the ‘Talisman,’ for example, the French naturalists
-caught in one haul of the dredge off the coast of Morocco in 500 fathoms
-of water no fewer than 134 fish, of which number 95 belonged to the
-family Macruridæ.
-
-They are usually small fish, measuring from a few inches to two feet in
-length, with a body terminating in a long compressed tapering tail and
-covered with spiny, keeled, or striated scales.
-
-The Pleuronectidæ or flat fish are not, as a rule, found in the abysmal
-zone; one species, however, _Pleuronectes cynoglossus_, was found by the
-American ship ‘Blake’ to extend into 732 fathoms of water.
-
-The families Sternoptychidæ and Scopelidæ are of particular interest to
-us, as almost all the genera they contain belong either to the pelagic
-or abysmal zones, and lend support to the view enunciated by Moseley,
-that the deep-sea fauna has, partly at any rate, been derived from the
-fauna of the pelagic zone. They are nearly all small slender fish with
-delicate and frequently semi-transparent bodies, large gaping mouths
-armed with numerous long irregular teeth, and frequently provided upon
-the head and sides of the trunk with rows of eye-like phosphorescent
-organs.
-
-These families, and others that have still to be referred to, belong to
-the group of Teleostei that is called Physostomi, the name referring to
-the open communication that usually exists in all these families between
-the swimming bladder and the alimentary canal. It is a remarkable fact
-that in none of the deep-sea representatives has this open communication
-been discovered. It is true that many specimens are, when examined, so
-lacerated by the diminution in pressure as to render anatomical study a
-matter of difficulty, but still a fair number of uninjured
-well-preserved specimens have now been examined and the duct has not
-been found.
-
-Of the family Sternoptychidæ, _Gonostoma microdon_ has a most remarkable
-distribution. It has been found at numerous stations in both the Pacific
-and Atlantic Oceans at depths ranging from 500 to nearly 3,000 fathoms
-of water.
-
-The Scopelidæ are represented by some very extraordinary types. The
-genus _Bathypterois_, for example, occurring in depths ranging from 500
-to 2,500 fathoms, is characterised by the development of enormously long
-pectoral fins to serve probably as organs of touch. ‘The rays of the
-pectoral fin,’ says Dr. Günther, ‘are much elongated. The ventral fins
-abdominal, with the outer rays prolonged, eight-rayed.... Gill rakers
-long.’ They are further characterised by the absence of any true
-phosphorescent organs and the smallness of their eyes.
-
-There can be little doubt, I think, that in these fishes the sense of
-touch or taste to a great extent takes the place of the sense of sight
-in other Scopelids. Not being provided with well-developed eyes or
-phosphorescent organs to attract their prey, the pectoral fins and the
-outer rays of the pelvic fins have become elongated and provided with
-special sense organs for searching for their food in the fine mud of the
-floor of the ocean.
-
-These long pectoral rays must have a very curious appearance in the
-living fish. Mr. Murray observes: ‘When taken from the trawl they were
-always dead, and the long pectoral rays were erected like an arch over
-the head, requiring considerable pressure to make them lie along the
-side of the body; when erected they resembled the Pennatulids like
-Umbellula.’ Filhol considers that when the fish is progressing through
-the obscurity of the abyss it probably carries these organs directed
-forward, seeking with them in the mud for any worms or other animals
-upon which it preys, or receiving through them warning of the approach
-of an enemy from whom it is necessary to make an immediate escape. One
-of the most remarkable of the deep-sea fish is closely related to
-Bathypterois, namely _Ipnops Murrayi_, living in depths of over 1,000
-fathoms. It is about five inches long, of a yellowish brown colour, with
-an elongated subcylindrical body covered with large thin deciduous
-scales. There are no phosphorescent organs of the ordinary type met with
-in the Scopelidæ, but upon the upper surface of the head there is found
-a pair of organs somewhat resembling the ordinary eyes of fishes but
-devoid of retina and optic nerve, that, from the researches of Moseley,
-seem to be undoubtedly organs for emitting light. ‘The organs are paired
-expanses, completely symmetrical in outline, placed on either side of
-the median line of the upper flattened surface of the head of the fish,
-extending from a line a little posterior to the nasal capsules nearly to
-a point above the posterior extremity of the cranial cavity.’ They are
-covered by the upper walls of the skull, which is extremely thin and
-completely transparent in the region lying over them. ‘They are
-membranous structures 0·4 mm. in thickness marked by hexagonal areas
-about 0·04 mm. in diameter. When their surface is viewed by reflected
-light the appearance is that of a number of glistening white isolated
-short columns standing up in relief from its basal membrane.’ Each
-hexagonal column is composed of a number of transparent rods disposed
-side by side at right angles to the outer surface of the organ, with
-their bases applied against the concave surface of a large hexagonal
-pigment cell, one of which forms the basis of each hexagonal column. It
-is still very doubtful what are the true homologies of this
-extraordinary phosphorescent organ, but Moseley was of opinion that, ‘on
-the whole, it seems not unlikely that the remarkable head organs of
-Ipnops may be regarded as highly specialised and enormously enlarged
-representatives of the phosphorescent organs on the heads of such allied
-Scopelidæ as _Scopelus rafinesquii_ and _Scopelus metopoclampus_. It may
-be conceived that in _Ipnops_ the supra-nasal and sub-ocular
-phosphorescent organs of these species on either side have united and
-become one with the result of the total obliteration of the eye.’
-
-Most of the species of the genus _Scopelus_ are undoubtedly pelagic in
-habit, descending during the day to depths of semi-darkness but rising
-at night to the surface waters. It is not certain how many of the known
-species occasionally or habitually dwell in very deep water, but there
-seems to be no doubt that two species at least—_S. macrolepidotus_ and
-_S. glacialis_—belong to the abysmal zone. Both of these species were
-found in dredges that had been at work in depths of over 1,000 fathoms
-and showed signs when examined of having been brought from the abyss.
-
-The Stomiatidæ are almost entirely confined to water from 450 to nearly
-2,000 fathoms in depth. They may be distinguished from the Scopelidæ by
-the long hyoid barbel close to the symphysis of the lower jaw, but like
-many of the genera of that family they have wide gaping mouths armed
-with a profusion of vicious looking teeth and a series of luminous spots
-on the sides of the head and body. (_See_ Frontispiece.)
-
-In _Eustomias obscurus_, found in depths of over 1,000 fathoms in the
-Atlantic by the ‘Talisman,’ the barbel is provided with a terminal
-swelling, shaped like a dumb-bell, which may be capable of emitting a
-phosphorescent light and serve the animal as a lure for the attraction
-of its prey. The genus _Malacosteus_, too, presents us with some of the
-most remarkable forms that are found in the abysmal zone. The mouth is
-of enormous size and the integuments of the abdomen present very
-definite longitudinal folds, leaving no doubt that this fish is able,
-like several others living in deep water, to swallow prey of an enormous
-size.
-
-But a perfectly unique structure in this fish ‘is a thin cylindrical
-muscular band which connects the back part of the mandibular symphysis
-with the hyoid bone. It is probably the homologue of a muscular band
-which, in other Stomiatids, stretches on each side from the mandible to
-the side of the hyoid, the two bands coalescing into an unpaired one in
-_Malacosteus_. It is, in the present state of preservation, much
-elongated, like a barbel, but during life it is notably contractile, and
-serves to give to the extremity of the mandible the requisite power of
-resistance when the fish has seized its prey, as without such a
-contrivance so long and slender a bone would yield to the force of its
-struggling victim.’
-
-[Illustration: FIG. 22.—_Saccopharynx ampullaceus_; a deep-sea eel, with
-the head of a large fish, which it has swallowed, showing through the
-thin integuments of the body. (From Günther.)]
-
-Belonging to the family of the Salmons we find one genus _Bathylagus_
-that is undoubtedly an abysmal form. Although there may be some doubt as
-to the exact depth at which the specimens were captured, the thinness of
-the bones, the enormous size of the eyes, and other bathybial characters
-prove that they must live in very deep water. Closely allied to the
-salmon and the herrings is the family of the Alepocephalidæ, a family
-that contributes several forms to the fauna of the deep sea, but they do
-not possess any characters that call for special comment. Their vertical
-distribution varies between 345 and 2,150 fathoms.
-
-The family Halosauridæ contains five species all confined to the abysmal
-zone. They have long bodies tapering to a finely pointed tail, and the
-head is provided with a snout that projects considerably in front of the
-mouth.
-
-Of the family of the Eels there are several representatives in the deep
-sea. They are characterised by a combination of true eel characters with
-special modifications due to a bathybial existence. ‘To enable them to
-seize upon prey more powerful than themselves certain organs have
-undergone a degree of specialisation, as is observed in bathybial
-members of other families with a similar mode of life; the jaws are
-exceedingly elongate and the whole gape, the pharynx and stomach capable
-of enormous distension.’ The head is very large, the eye very small and
-the tail long and tapering (fig. 22).
-
-The lessons we learn from the study of the fishes of the deep sea are
-particularly instructive. It would take far more space than can be
-afforded here to fully illustrate all of the points that seem clear to
-us, but I hope I have said sufficient to show that the fish fauna is
-made up of genera and species belonging to several widely separate
-families of the Teleostei; that some of them show, in a very marked way,
-what may be looked upon as peculiarly bathybial characters, whilst
-others are but slightly modified from their shallow-water
-representatives. These facts by themselves lend support to the view that
-the fauna of the deep sea has been derived from the fauna of shallower
-water by successive migrations at different periods of the world’s
-history. Those that exhibit in a most marked degree the special
-bathybial characters are probably those whose immigration took place
-long ago, whilst those more closely related to shallow-water forms are,
-comparatively speaking, recent importations. The occurrence of Scopelidæ
-and Sternoptychidæ in deep water suggests, as Moseley pointed out many
-years ago, that the fauna is partly derived from the pelagic plankton.
-But while these points may seem clear to us, there are others that still
-require much more investigation and consideration. The whole question of
-the function and use of the phosphorescent organs, the mucous glands,
-the barbels and elongated fin rays, the mode of life, the deposition of
-ova and their development, afford problems which in the present state of
-our knowledge must remain unsolved. Let us hope that in the future there
-may be a new stimulus given to deep-sea research, and these problems may
-be again seriously studied and eventually solved.
-
-
-
-
- INDEX
-
-
- Actiniaria, 36, 93
-
- —— two remarkable genera of, 15
-
- _Aegir_, 15
-
- Agassiz, A.,
-
- —— on colour of Cœlentera, 65
-
- —— on Echinoidea, 101, 103
-
- Agassiz, L., on board the ‘Hassler,’ 12
-
- ‘Albatross,’ American vessel, 15
-
- Alcyonaria, 95
-
- —— phosphorescence of, 81
-
- Amphipoda, 124
-
- Anemones
-
- —— of deep water, 36, 92
-
- —— two remarkable genera of, 15
-
- Annelida, 117
-
- Ascidia compositæ, 146
-
- —— salpiformes, 147
-
- —— simplices, 142
-
- Asteroidea, 104
-
-
- Barriers of temperature, 32
-
- _Bathynomus_, 129
-
- Beddard, F. E., on Isopoda, 127
-
- Benthos, 53
-
- Berycidæ, 154
-
- ‘Blake,’ American vessel, 12
-
- Blue mud, 42
-
- Brachiopoda, 115
-
- Brachyura, 138
-
- _Brisinga_, 9, 105
-
- Buchanan’s experiment, 19
-
-
- Carpenter, P. H., on Crinoidea, 100
-
- _Cephalodiscus_, 113
-
- Cephalopoda, 120
-
- ‘Challenger,’ H.M.S., voyage of, 12
-
- Cirripedia, 130
-
- Cœlentera,
-
- —— colour of, 65
-
- —— of deep sea, 91
-
- Colour
-
- —— of the deep-sea fauna, 59, 66
-
- —— of the deep-sea fish, 60
-
- Copepoda, 124
-
- Corals, 94
-
- Crinoidea, 99
-
- Crustacea, 123
-
- —— colour of, 63
-
-
- Darkness of the abyss, 22
-
- Diatom ooze, 39
-
- Dipnoi, 149
-
-
- Echinoderma, 99
-
- —— colour of, 64
-
- Echinoidea, 101
-
- Eels, 165
-
- Elasipoda, 106
-
- Elasmobranchii, 148, 153
-
- Eryonidæ, 135
-
- Eyes of abysmal animals, 67
-
- —— of deep-sea crustacea, 72
- —— of deep-sea fish, 69
-
- —— of deep-sea mollusca, 71
-
- —— of _Genityllis_, 118
-
- —— of _Neobythites_, 69
-
- —— of _Serolis_, 73
-
-
- _Fenja_, 15
-
- ‘Fish Hawk,’ American vessel, 12
-
- Fol and Sarasin’s experiments, 25
-
- Foraminifera, 90
-
- Forbes, on the probable existence of a deep-sea fauna, 2
-
- —— on zones of distribution, 49
-
-
- Ganoidei, 149
-
- Gasteropoda, 119
-
- Gephyrea, 116
-
- Gills of deep-sea fish, 151
-
- Globerigina ooze, discovery of, 5
-
- —— distribution and composition of, 37
-
- Green mud, 42
-
- Gunn, Dr., on the eyes of _Genityllis_, 118
-
- Günther, Dr., on deep-sea fish, 150
-
-
- Hall, Marshall, 12
-
- Halosauridæ, 164
-
- ‘Hassler,’ American ship, 12
-
- Herdman, on Ascidians, 143, 146
-
- Hermit crabs, 139
-
- Hoek, Dr., on Cirripedia, 132
-
- —— on Pycnogonida, 140
-
- Holothuridea, 106
-
- Hoyle, on Cephalopoda, 120
-
- Hydroids, 92
-
-
- ‘Investigator,’ H.M.S., 16
-
- _Ipnops Murrayi_, colour of, 60, 61
-
- _Ipnops Murrayi_, phosphorescent organs of, 160
-
- Isopoda, 127
-
-
- Katantic sub-zone, 50
-
- ‘Knight Errant,’ H.M.S., 12
-
-
- Lamellibranchia, 119
-
- ‘Lightning,’ H.M.S., 8, 9
-
- Lime, scarcity of, in bones of bathybial fish, 83
-
- —— in shells of mollusca, 83
-
- Littoral sub-zone, 49
-
- Lycodidæ, 155
-
-
- Macrura, 135
-
- Macruridæ, 156
-
- Madreporaria, 94
-
- Medusæ, 91
-
- Mollusca, 119
-
- —— colour of, 62
-
- Moore, Capt., 94
-
- Moseley, H. N., on colour of Cœlentera, 65
-
- —— on phosphorescent organs of _Ipnops_, 160
-
- —— on the darkness of the abyss, 22
-
- —— on the phosphorescence of Alcyonarians, 25
-
- Murray, on _Bathypterois_, 159
-
-
- Nekton, 53
-
- Neritic zone, 48
-
- ‘Norma,’ Mr. Hall’s yacht, 12
-
- Norske Nord-havns expedition, 7, 15
-
-
- Ophidiidæ, 156
-
- Ostracoda, 123
-
-
- Packard, on the illumination of the abyss, 23
-
- Pediculati, 155
-
- Pelagic zone, 47
-
- Pennatulidæ, 96
-
- _Phoronis_, 111
-
- Phosphorescence of Alcyonarians, 81
- — of deep-sea Crustacea, 80
- — of Echinoderma, 81
-
- Phosphorescent light in the abyss, 24
- — organs of deep-sea fish, 77
-
- Pigment in mucous membranes of deep-sea fish, 84, 152
-
- Plankton, 52
-
- Pleuronectidæ, 157
-
- Polar currents, 30, 33
-
- Polychæta, 118
-
- ‘Porcupine,’ H.M.S., 8, 9
-
- Porifera, 91
-
- Pourtales, Count, 10
-
- Pressure in the abyss, 19
-
- Protozoa, 88
-
- Pteropod ooze, 39
-
- Pycnogonida, 139
-
-
- Radiolaria, 89
-
- Radiolarian ooze, 39
-
- Red mud, 37
- — — off the Brazilian coasts, 42
-
- _Rhabdopleura_, 111
-
- Ross, Sir James, on the fauna of the deep sea, 3
-
-
- Salmonidæ, 163
-
- Sargasso sub-zone, 48
-
- Sars, 6, 9,
- — on Amphipoda, 125
- — on _Brisinga_, 105
-
- Schizopoda, 133
-
- Scopelidæ, 158
-
- _Serolis_, 127
-
- Siphonophora, 92
-
- Size of deep-sea animals, 85
- — of fish, 153
-
- Smith, Mr., on Lamellibranchia, 119
-
- Spatangoids, 101
-
- Sponges, 91
-
- Stebbing, Rev. T. R., on Amphipoda, 125
-
- Sternoptychidæ, 158
-
- Stomatopoda, 133
-
- Stomiatidæ, 162
-
-
- ‘Talisman,’ French vessel, 12
-
- Teleostei, 149, 154
-
- Temperature of the abyss, 28
-
- Thomson, Sir Wyville, on _Pourtalesia_, 10
- — on _Thaumastocheles_, 137
- — on the darkness of the abyss, 22
- — on the phosphorescence of the sea, 26
-
- Thoracostraca, 133
-
- ‘Travailleur,’ French vessel, 12
-
- ‘Triton,’ H.M.S., 12
-
- Tunicata, 140
-
-
- Vegetable life, absence of, 42
-
- Verrill, on the illumination of the abyss, 23
-
- ‘Vittor Pessani,’ Italian vessel, 12
-
- ‘Vöringin,’ Norwegian vessel, 14
-
-------------------------------------------------------------------------
-
-
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