=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.