![THE DEEP-SEA FLOOR: AN OVERVIEW 31
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Fig. 2.23. Growth (as test volume) of three abyssal xenophyophore
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organisms. Outer scale refers to individuals B and C. Inner scale
refers to individual A. Modified from Gooday et al. (1993).
Copyright: Elsevier Science.
environment (high pressure, low temperature), but by
the meager supply of food.
ACKNOWLEDGEMENTS
The following people answered queries: W. Burnett,
J. Eckman, A. Gooday, P. Jumars, W. Landing,
P. LaRock, G. Paterson, G. Weatherly and G. Wilson.
L. Bouck, S. Ertman, R. Hessler, K. Suderman,
A. Thistle, G. Wilson, and an anonymous reviewer
commented on the manuscript. I am grateful for this
kind help. The chapter was written while the author
was partially supported by ONR grant N00014-95-1-
0750 and NSF grant OCE-9616676.
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Allen, J.A. and Sanders, H.L., 1966. Adaptations to abyssal life as
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Allen, J.A. and Sanders, H.L., 1996. The zoogeography, diversity and
origin of the deep-sea protobranch bivalves of the Atlantic: the
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Altenbach, A.V
., Heeger, T., Linke, P., Spindler, M. and Thies, A.,
1993. Miliolinella subrotunda (Montague), a miliolid foraminifer
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Anonymous, 1989. Seawater: Its Composition, Properties and
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Bagley, P.M. and Priede, I.G., 1997. An autonomous free-fall acoustic
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Barnett, P.R.O., Watson, J. and Connelly, D., 1984. A multiple corer
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Bett, B.J. and Rice, A.L., 1993. The feeding behaviour of an abyssal
echiuran revealed by in situ time-lapse photography. Deep-Sea Res.
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Bett, B.J., Vanreusel, A., Vincx, M., Soltwedel, T., Pfannkuche, O.,
Lambshead, P.J.D., Gooday, A.J., Ferrero, T.J. and Dinet, A., 1994.
Sampler bias in the quantitative study of deep-sea meiobenthos.
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Billett, D.S.M., Lampitt, R.S., Rice, A.L. and Mantoura, R.F.C., 1983.
Seasonal sedimentation of phytoplankton to the deep-sea benthos.
Nature, 302: 520−522.
Blake, J.A. and Hilbig, B., 1994. Dense infaunal assemblages on
the continental slope off Cape Hatteras, North Carolina. Deep-Sea
Res. II, 44: 875−899.
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flow velocity, food concentration and particle flux on growth rates
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Ecol., 129: 45−60.
Cannariato, K.G., Kennett, J.P. and Behl, R.J., 1999. Biotic response
to late Quaternary rapid climate switches in Santa Barbara Basin:
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Carey Jr., A.G., 1972. Food sources of sublittoral, bathyal and abyssal
asteroids in the northeast Pacific Ocean. Ophelia, 10: 35−47.
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Carney, R.S. and Carey Jr., A.G., 1982. Distribution and diversity of](https://image.slidesharecdn.com/2058731-250604222541-148767a9/85/Ecosystems-Of-The-Deep-Oceans-Paul-A-Tyler-46-320.jpg)
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all his company but one man whom he could fully trust; and this was
Tonty. He and Hennepin were on indifferent terms. Men thrown
together in a rugged enterprise like this quickly learn to know each
other; and the vain and assuming friar was not likely to commend
himself to La Salle's brave and loyal lieutenant. Hennepin says that it
was La Salle's policy to govern through the dissensions of his
followers; and, from whatever cause, it is certain that those beneath
him were rarely in perfect harmony.
FOOTNOTES:
Hennepin, Description de la Louisiane (1683), 19; Ibid., Voyage Curieux
(1704), 66. Ribourde had lately arrived.
Lettre de La Motte de la Lussière, sans date; Relation de Henri de
Tonty écrite de Québec, le 14 Novembre, 1684 (Margry, i. 573). This
paper, apparently addressed to Abbé Renaudot, is entirely distinct
from Tonty's memoir of 1693, addressed to the minister
Ponchartrain.
Hennepin, Nouvelle Découverte (1697), 8.
Ibid., Avant Propos, 5.
Ibid., Voyage Curieux (1704), 12.
Hennepin, Voyage Curieux (1704), 18.
Ibid. Avis au Lecteur. He elsewhere represents
himself as on excellent terms with La Salle; with
whom, he says, he used to read histories of travels
at Fort Frontenac, after which they discussed
together their plans of discovery.
This was the Racines Agnières of Bruyas. It was published by Mr. Shea
in 1862. Hennepin seems to have studied it carefully; for on several
occasions he makes use of words evidently borrowed from it, putting
them into the mouths of Indians speaking a dialect different from that of
the Agniers, or Mohawks.
Compare Brodhead in Hist. Mag., x. 268.](https://image.slidesharecdn.com/2058731-250604222541-148767a9/85/Ecosystems-Of-The-Deep-Oceans-Paul-A-Tyler-56-320.jpg)
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Une enterprise capable d'épouvanter tout autre que moi.—
Hennepin, Voyage Curieux, Avant Propos (1704).
Je vous proteste ici devant Dieu, que ma Relation est fidèle
et sincère, etc.—Ibid., Avis au Lecteur.
The nature of these fabrications will be shown
hereafter. They occur, not in the early editions of
Hennepin's narrative, which are comparatively truthful,
but in the edition of 1697 and those which followed. La
Salle was dead at the time of their publication.
This place is laid down on a manuscript map sent to France by the
Intendant Duchesneau, and now preserved in the Archives de la Marine,
and also on several other contemporary maps.
Hennepin's account of the falls and river of Niagara—especially his
second account, on his return from the West—is very minute, and
on the whole very accurate. He indulges in gross exaggeration as to
the height of the cataract, which, in the edition of 1683, he states at
five hundred feet, and raises to six hundred in that of 1697. He also says that
there was room for four carriages to pass abreast under the American Fall
without being wet. This is, of course, an exaggeration at the best; but it is
extremely probable that a great change has taken place since his time. He
speaks of a small lateral fall at the west side of the Horse Shoe Fall which does
not now exist. Table Rock, now destroyed, is distinctly figured in his picture. He
says that he descended the cliffs on the west side to the foot of the cataract,
but that no human being can get down on the east side.
The name of Niagara, written Onguiaahra by Lalemant in 1641,
and Ongiara by Sanson, on his map of 1657, is used by Hennepin
in its present form. His description of the falls is the earliest
known to exist. They are clearly indicated on the map of
Champlain, 1632. For early references to them, see The Jesuits in
North America, 235, note. A brief but curious notice of them is
given by Gendron, Quelques Particularitez du Pays des Hurons,
1659. The indefatigable Dr. O'Callaghan has discovered thirty-nine
distinct forms of the name Niagara. Index to Colonial Documents
of New York, 465. It is of Iroquois origin, and in the Mohawk
dialect is pronounced Nyàgarah.
Tonty, Relation, 1684 (Margry, i. 573).](https://image.slidesharecdn.com/2058731-250604222541-148767a9/85/Ecosystems-Of-The-Deep-Oceans-Paul-A-Tyler-57-320.jpg)
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Near the town of Victor. It is laid down on the map of Galinée,
and other unpublished maps. Compare Marshall, Historical Sketches
of the Niagara Frontier, 14.
Lettre de La Salle à un de ses associés (Margry, ii. 32).
Description de la Louisiane (1683), 41. It is
characteristic of Hennepin that, in the editions of his
book published after La Salle's death, he substitutes, for
anybody but him, anybody but those who had formed
so generous a design,—meaning to include himself, though he lost nothing by
the disaster, and had not formed the design.
On these incidents, compare the two narratives of Tonty, of
1684 and 1693. The book bearing Tonty's name is a compilation
full of errors. He disowned its authorship.
Lettre de La Salle, 22 Août, 1682 (Margry, ii. 212).
Lettre de La Motte, sans date.](https://image.slidesharecdn.com/2058731-250604222541-148767a9/85/Ecosystems-Of-The-Deep-Oceans-Paul-A-Tyler-58-320.jpg)
![THE NIAGARA
PORTAGE.
CHAPTER X.
1679.
THE LAUNCH OF THE GRIFFIN.
The Niagara Portage.—A Vessel on the Stocks.—Suffering and Discontent.
—La Salle's Winter Journey.—The Vessel launched.—Fresh Disasters.
A more important work than that of the
warehouse at the mouth of the river was now to be
begun. This was the building of a vessel above the
cataract. The small craft which had brought La Motte and Hennepin
with their advance party had been hauled to the foot of the rapids at
Lewiston, and drawn ashore with a capstan, to save her from the
drifting ice. Her lading was taken out, and must now be carried
beyond the cataract to the calm water above. The distance to the
destined point was at least twelve miles, and the steep heights
above Lewiston must first be climbed. This heavy task was
accomplished on the twenty-second of January. The level of the
plateau was reached, and the file of burdened men, some thirty in
number, toiled slowly on its way over the snowy plains and through
the gloomy forests of spruce and naked oak-trees; while Hennepin
plodded through the drifts with his portable altar lashed fast to his
back. They came at last to the mouth of a stream which entered the
Niagara two leagues above the cataract, and which was undoubtedly
that now called Cayuga Creek.[124]
Trees were felled, the place cleared, and the master-carpenter set
his ship-builders at work. Meanwhile, two Mohegan hunters,
attached to the party, made bark wigwams to lodge the men.
Hennepin had his chapel, apparently of the same material, where he
placed his altar, and on Sundays and saints' days said mass,](https://image.slidesharecdn.com/2058731-250604222541-148767a9/85/Ecosystems-Of-The-Deep-Oceans-Paul-A-Tyler-59-320.jpg)
![SUFFERING AND
DISCONTENT.
preached, and exhorted; while some of the men, who knew the
Gregorian chant, lent their aid at the service. When the carpenters
were ready to lay the keel of the vessel, La Salle asked the friar to
drive the first bolt; but the modesty of my religious profession, he
says, compelled me to decline this honor.
Fortunately, it was the hunting-season of the Iroquois, and most
of the Seneca warriors were in the forests south of Lake Erie; yet
enough remained to cause serious uneasiness. They loitered sullenly
about the place, expressing their displeasure at the proceedings of
the French. One of them, pretending to be drunk, attacked the
blacksmith and tried to kill him; but the Frenchman, brandishing a
red-hot bar of iron, held him at bay till Hennepin ran to the rescue,
when, as he declares, the severity of his rebuke caused the savage
to desist.[125] The work of the ship-builders advanced rapidly; and
when the Indian visitors beheld the vast ribs of the wooden monster,
their jealousy was redoubled. A squaw told the French that they
meant to burn the vessel on the stocks. All now stood anxiously on
the watch. Cold, hunger, and discontent found imperfect antidotes in
Tonty's energy and Hennepin's sermons.
La Salle was absent, and his lieutenant
commanded in his place. Hennepin says that Tonty
was jealous because he, the friar, kept a journal,
and that he was forced to use all manner of just precautions to
prevent the Italian from seizing it. The men, being half-starved, in
consequence of the loss of their provisions on Lake Ontario, were
restless and moody; and their discontent was fomented by one of
their number, who had very probably been tampered with by La
Salle's enemies.[126] The Senecas refused to supply them with corn,
and the frequent exhortations of the Récollet father proved an
insufficient substitute. In this extremity, the two Mohegans did
excellent service,—bringing deer and other game, which relieved the
most pressing wants of the party, and went far to restore their
cheerfulness.](https://image.slidesharecdn.com/2058731-250604222541-148767a9/85/Ecosystems-Of-The-Deep-Oceans-Paul-A-Tyler-60-320.jpg)
![THE SHIP
FINISHED.
La Salle, meanwhile, had gone down to the mouth of the river,
with a sergeant and a number of men; and here, on the high point
of land where Fort Niagara now stands, he marked out the
foundations of two blockhouses.[127] Then, leaving his men to build
them, he set out on foot for Fort Frontenac, where the condition of
his affairs demanded his presence, and where he hoped to procure
supplies to replace those lost in the wreck of his vessel. It was
February, and the distance was some two hundred and fifty miles,
through the snow-encumbered forests of the Iroquois and over the
ice of Lake Ontario. Two men attended him, and a dog dragged his
baggage on a sledge. For food, they had only a bag of parched corn,
which failed them two days before they reached the fort; and they
made the rest of the journey fasting.
During his absence, Tonty finished the vessel,
which was of about forty-five tons' burden.[128] As
spring opened, she was ready for launching. The
friar pronounced his blessing on her; the assembled company sang
Te Deum; cannon were fired; and French and Indians, warmed alike
by a generous gift of brandy, shouted and yelped in chorus as she
glided into the Niagara. Her builders towed her out and anchored
her in the stream, safe at last from incendiary hands; and then,
swinging their hammocks under her deck, slept in peace, beyond
reach of the tomahawk. The Indians gazed on her with amazement.
Five small cannon looked out from her portholes; and on her prow
was carved a portentous monster, the Griffin, whose name she bore,
in honor of the armorial bearings of Frontenac. La Salle had often
been heard to say that he would make the griffin fly above the
crows, or, in other words, make Frontenac triumph over the Jesuits.
They now took her up the river, and made her fast below the swift
current at Black Rock. Here they finished her equipment, and waited
for La Salle's return; but the absent commander did not appear. The
spring and more than half of the summer had passed before they
saw him again. At length, early in August, he arrived at the mouth of
the Niagara, bringing three more friars; for, though no friend of the](https://image.slidesharecdn.com/2058731-250604222541-148767a9/85/Ecosystems-Of-The-Deep-Oceans-Paul-A-Tyler-61-320.jpg)
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Jesuits, he was zealous for the Faith, and was rarely without a
missionary in his journeyings. Like Hennepin, the three friars were
all Flemings. One of them, Melithon Watteau, was to remain at
Niagara; the others, Zenobe Membré and Gabriel Ribourde, were to
preach the Faith among the tribes of the West. Ribourde was a hale
and cheerful old man of sixty-four. He went four times up and down
the Lewiston heights, while the men were climbing the steep
pathway with their loads. It required four of them, well stimulated
with brandy, to carry up the principal anchor destined for the
Griffin.
La Salle brought a tale of disaster. His enemies, bent on ruining
the enterprise, had given out that he was embarked on a
harebrained venture, from which he would never return. His
creditors, excited by rumors set afloat to that end, had seized on all
his property in the settled parts of Canada, though his seigniory of
Fort Frontenac alone would have more than sufficed to pay all his
debts. There was no remedy. To defer the enterprise would have
been to give his adversaries the triumph that they sought; and he
hardened himself against the blow with his usual stoicism.[129]
FOOTNOTES:
It has been a matter of debate on which side of the Niagara the first
vessel on the Upper Lakes was built. A close study of Hennepin, and a
careful examination of the localities, have convinced me that the spot was
that indicated above. Hennepin repeatedly alludes to a large detached
rock, rising out of the water at the foot of the rapids above Lewiston, on the
west side of the river. This rock may still be seen immediately under the
western end of the Lewiston suspension-bridge. Persons living in the
neighborhood remember that a ferry-boat used to pass between it and the cliffs
of the western shore; but it has since been undermined by the current and has
inclined in that direction, so that a considerable part of it is submerged, while
the gravel and earth thrown down from the cliff during the building of the
bridge has filled the intervening channel. Opposite to this rock, and on the east
side of the river, says Hennepin, are three mountains, about two leagues below
the cataract. (Nouveau Voyage (1704), 462, 466.) To these three mountains,](https://image.slidesharecdn.com/2058731-250604222541-148767a9/85/Ecosystems-Of-The-Deep-Oceans-Paul-A-Tyler-62-320.jpg)
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as well as to the rock, he frequently alludes. They are also spoken of by La
Hontan, who clearly indicates their position. They consist in the three
successive grades of the acclivity: first, that which rises from the level of the
water, forming the steep and lofty river-bank; next, an intermediate ascent,
crowned by a sort of terrace, where the tired men could find a second resting-
place and lay down their burdens, whence a third effort carried them with
difficulty to the level top of the plateau. That this was the actual portage, or
carrying place of the travellers, is shown by Hennepin (1704), 114, who
describes the carrying of anchors and other heavy articles up these heights in
August, 1679. La Hontan also passed the Falls by way of the three mountains
eight years later. La Hontan (1703), 106. It is clear, then, that the portage was
on the east side, whence it would be safe to conclude that the vessel was built
on the same side. Hennepin says that she was built at the mouth of a stream
(rivière) entering the Niagara two leagues above the Falls. Excepting one or two
small brooks, there is no stream on the west side but Chippewa Creek, which
Hennepin had visited and correctly placed at about a league from the cataract.
His distances on the Niagara are usually correct. On the east side there is a
stream which perfectly answers the conditions. This is Cayuga Creek, two
leagues above the Falls. Immediately in front of it is an island about a mile
long, separated from the shore by a narrow and deep arm of the Niagara, into
which Cayuga Creek discharges itself. The place is so obviously suited to
building and launching a vessel, that, in the early part of this century, the
government of the United States chose it for the construction of a schooner to
carry supplies to the garrisons of the Upper Lakes. The neighboring village now
bears the name of La Salle.
In examining this and other localities on the Niagara, I have
been greatly aided by my friend O. H. Marshall, Esq., of Buffalo,
who is unrivalled in his knowledge of the history and traditions of
the Niagara frontier.
Hennepin (1704), 97. On a paper drawn up at the instance of the
Intendant Duchesneau, the names of the greater number of La Salle's
men are preserved. These agree with those given by Hennepin: thus, the
master-carpenter, whom he calls Maître Moyse, appears as Moïse Hillaret;
and the blacksmith, whom he calls La Forge, is mentioned as—(illegible) dit la
Forge.
This bad man, says Hennepin, would infallibly have debauched our
workmen, if I had not reassured them by the exhortations which I made
them on fête-days and Sundays, after divine service. (1704), 98.](https://image.slidesharecdn.com/2058731-250604222541-148767a9/85/Ecosystems-Of-The-Deep-Oceans-Paul-A-Tyler-63-320.jpg)
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Lettre de La Salle, 22 Août, 1682 (Margry, ii. 197); Relation de Tonty,
1684 (Ibid., i. 577). He called this new post Fort Conti. It was burned
some months after, by the carelessness of the sergeant in command, and
was the first of a succession of forts on this historic spot.
Hennepin (1683), 46. In the edition of 1697, he says that it was of sixty
tons. I prefer to follow the earlier and more trustworthy narrative.
La Salle's embarrassment at this time was so great that he
purposed to send Tonty up the lakes in the Griffin, while he went
back to the colony to look after his affairs; but suspecting that the
pilot, who had already wrecked one of his vessels, was in the pay of
his enemies, he resolved at last to take charge of the expedition himself, to
prevent a second disaster. (Lettre de La Salle, 22 Août, 1682; Margry, ii. 214.)
Among the creditors who bore hard upon him were Migeon, Charon, Giton, and
Peloquin, of Montreal, in whose name his furs at Fort Frontenac had been
seized. The intendant also placed under seal all his furs at Quebec, among
which is set down the not very precious item of two hundred and eighty-four
skins of enfants du diable, or skunks.](https://image.slidesharecdn.com/2058731-250604222541-148767a9/85/Ecosystems-Of-The-Deep-Oceans-Paul-A-Tyler-64-320.jpg)
![CHAPTER XI.
1679.
LA SALLE ON THE UPPER LAKES.
The Voyage of the Griffin.—Detroit.—A Storm.—St. Ignace of
Michilimackinac.—Rivals and Enemies.—Lake Michigan.—Hardships.—A
Threatened Fight.—Fort Miami.—Tonty's Misfortunes.—Forebodings.
The Griffin had lain moored by the shore, so near that Hennepin
could preach on Sundays from the deck to the men encamped along
the bank. She was now forced up against the current with tow-ropes
and sails, till she reached the calm entrance of Lake Erie. On the
seventh of August, La Salle and his followers embarked, sang Te
Deum, and fired their cannon. A fresh breeze sprang up; and with
swelling canvas the Griffin ploughed the virgin waves of Lake Erie,
where sail was never seen before. For three days they held their
course over these unknown waters, and on the fourth turned
northward into the Strait of Detroit. Here, on the right hand and on
the left, lay verdant prairies, dotted with groves and bordered with
lofty forests. They saw walnut, chestnut, and wild plum trees, and
oaks festooned with grape-vines; herds of deer, and flocks of swans
and wild turkeys. The bulwarks of the Griffin were plentifully hung
with game which the men killed on shore, and among the rest with a
number of bears, much commended by Hennepin for their want of
ferocity and the excellence of their flesh. Those, he says, who will
one day have the happiness to possess this fertile and pleasant
strait, will be very much obliged to those who have shown them the
way. They crossed Lake St. Clair,[130] and still sailed northward
against the current, till now, sparkling in the sun, Lake Huron spread
before them like a sea.](https://image.slidesharecdn.com/2058731-250604222541-148767a9/85/Ecosystems-Of-The-Deep-Oceans-Paul-A-Tyler-65-320.jpg)
![ST. IGNACE.
For a time they bore on prosperously. Then the
wind died to a calm, then freshened to a gale, then
rose to a furious tempest; and the vessel tossed
wildly among the short, steep, perilous waves of the raging lake.
Even La Salle called on his followers to commend themselves to
Heaven. All fell to their prayers but the godless pilot, who was loud
in complaint against his commander for having brought him, after
the honor he had won on the ocean, to drown at last ignominiously
in fresh water. The rest clamored to the saints. St. Anthony of Padua
was promised a chapel to be built in his honor, if he would but save
them from their jeopardy; while in the same breath La Salle and the
friars declared him patron of their great enterprise.[131] The saint
heard their prayers. The obedient winds were tamed; and the
Griffin plunged on her way through foaming surges that still grew
calmer as she advanced. Now the sun shone forth on woody islands,
Bois Blanc and Mackinaw and the distant Manitoulins,—on the forest
wastes of Michigan and the vast blue bosom of the angry lake; and
now her port was won, and she found her rest behind the point of
St. Ignace of Michilimackinac, floating in that tranquil cove where
crystal waters cover but cannot hide the pebbly depths beneath.
Before her rose the house and chapel of the Jesuits, enclosed with
palisades; on the right, the Huron village, with its bark cabins and its
fence of tall pickets; on the left, the square compact houses of the
French traders; and, not far off, the clustered wigwams of an Ottawa
village.[132] Here was a centre of the Jesuit missions, and a centre
of the Indian trade; and here, under the shadow of the cross, was
much sharp practice in the service of Mammon. Keen traders, with
or without a license, and lawless coureurs de bois, whom a few
years of forest life had weaned from civilization, made St. Ignace
their resort; and here there were many of them when the Griffin
came. They and their employers hated and feared La Salle, who,
sustained as he was by the governor, might set at nought the
prohibition of the King, debarring him from traffic with these tribes.
Yet, while plotting against him, they took pains to allay his distrust
by a show of welcome.](https://image.slidesharecdn.com/2058731-250604222541-148767a9/85/Ecosystems-Of-The-Deep-Oceans-Paul-A-Tyler-66-320.jpg)
![RIVALS AND
ENEMIES.
The Griffin fired her cannon, and the Indians yelped in wonder
and amazement. The adventurers landed in state, and marched
under arms to the bark chapel of the Ottawa village, where they
heard mass. La Salle knelt before the altar, in a mantle of scarlet
bordered with gold. Soldiers, sailors, and artisans knelt around him,
—black Jesuits, gray Récollets, swarthy voyageurs, and painted
savages; a devout but motley concourse.
As they left the chapel, the Ottawa chiefs came to bid them
welcome, and the Hurons saluted them with a volley of musketry.
They saw the Griffin at her anchorage, surrounded by more than a
hundred bark canoes, like a Triton among minnows. Yet it was with
more wonder than good-will that the Indians of the mission gazed
on the floating fort, for so they called the vessel. A deep jealousy
of La Salle's designs had been infused into them. His own followers,
too, had been tampered with. In the autumn before, it may be
remembered, he had sent fifteen men up the lakes to trade for him,
with orders to go thence to the Illinois and make preparation against
his coming. Early in the summer, Tonty had been despatched in a
canoe from Niagara to look after them.[133] It was high time. Most
of the men had been seduced from their duty, and had disobeyed
their orders, squandered the goods intrusted to them, or used them
in trading on their own account. La Salle found four of them at
Michilimackinac. These he arrested, and sent Tonty to the Falls of
Ste. Marie, where two others were captured, with their plunder. The
rest were in the woods, and it was useless to pursue them.
Anxious and troubled as to the condition of his
affairs in Canada. La Salle had meant, after seeing
his party safe at Michilimackinac, to leave Tonty to
conduct it to the Illinois, while he himself returned to the colony. But
Tonty was still at Ste. Marie, and he had none to trust but himself.
Therefore, he resolved at all risks to remain with his men; for, he
says, I judged my presence absolutely necessary to retain such of
them as were left me, and prevent them from being enticed away
during the winter. Moreover, he thought that he had detected an](https://image.slidesharecdn.com/2058731-250604222541-148767a9/85/Ecosystems-Of-The-Deep-Oceans-Paul-A-Tyler-67-320.jpg)
![POTTAWATTAMIE
S.
intrigue of his enemies to hound on the Iroquois against the Illinois,
in order to defeat his plan by involving him in the war.
Early in September he set sail again, and passing westward into
Lake Michigan,[134] cast anchor near one of the islands at the
entrance of Green Bay. Here, for once, he found a friend in the
person of a Pottawattamie chief, who had been so wrought upon by
the politic kindness of Frontenac that he declared himself ready to
die for the children of Onontio.[135] Here, too, he found several of
his advance party, who had remained faithful and collected a large
store of furs. It would have been better had they proved false, like
the rest. La Salle, who asked counsel of no man, resolved, in spite of
his followers, to send back the Griffin laden with these furs, and
others collected on the way, to satisfy his creditors.[136] It was a
rash resolution, for it involved trusting her to the pilot, who had
already proved either incompetent or treacherous. She fired a
parting shot, and on the eighteenth of September set sail for
Niagara, with orders to return to the head of Lake Michigan as soon
as she had discharged her cargo. La Salle, with the fourteen men
who remained, in four canoes deeply laden with a forge, tools,
merchandise, and arms, put out from the island and resumed his
voyage.
The parting was not auspicious. The lake, glassy
and calm in the afternoon, was convulsed at night
with a sudden storm, when the canoes were
midway between the island and the main shore. It was with difficulty
that they could keep together, the men shouting to each other
through the darkness. Hennepin, who was in the smallest canoe with
a heavy load, and a carpenter for a companion who was awkward at
the paddle, found himself in jeopardy which demanded all his nerve.
The voyagers thought themselves happy when they gained at last
the shelter of a little sandy cove, where they dragged up their
canoes, and made their cheerless bivouac in the drenched and
dripping forest. Here they spent five days, living on pumpkins and
Indian corn, the gift of their Pottawattamie friends, and on a Canada](https://image.slidesharecdn.com/2058731-250604222541-148767a9/85/Ecosystems-Of-The-Deep-Oceans-Paul-A-Tyler-68-320.jpg)
![porcupine brought in by La Salle's Mohegan hunter. The gale raged
meanwhile with relentless fury. They trembled when they thought of
the Griffin. When at length the tempest lulled, they re-embarked,
and steered southward along the shore of Wisconsin; but again the
storm fell upon them, and drove them for safety to a bare, rocky
islet. Here they made a fire of drift-wood, crouched around it, drew
their blankets over their heads, and in this miserable plight, pelted
with sleet and rain, remained for two days.
At length they were afloat again; but their prosperity was brief. On
the twenty-eighth, a fierce squall drove them to a point of rocks
covered with bushes, where they consumed the little that remained
of their provisions. On the first of October they paddled about thirty
miles, without food, when they came to a village of Pottawattamies,
who ran down to the shore to help them to land; but La Salle,
fearing that some of his men would steal the merchandise and
desert to the Indians, insisted on going three leagues farther, to the
great indignation of his followers. The lake, swept by an easterly
gale, was rolling its waves against the beach, like the ocean in a
storm. In the attempt to land, La Salle's canoe was nearly swamped.
He and his three canoe-men leaped into the water, and in spite of
the surf, which nearly drowned them, dragged their vessel ashore
with all its load. He then went to the rescue of Hennepin, who with
his awkward companion was in woful need of succor. Father Gabriel,
with his sixty-four years, was no match for the surf and the violent
undertow. Hennepin, finding himself safe, waded to his relief, and
carried him ashore on his sturdy shoulders; while the old friar,
though drenched to the skin, laughed gayly under his cowl as his
brother missionary staggered with him up the beach.[137]
When all were safe ashore, La Salle, who distrusted the Indians
they had passed, took post on a hill, and ordered his followers to
prepare their guns for action. Nevertheless, as they were starving,
an effort must be risked to gain a supply of food; and he sent three
men back to the village to purchase it. Well armed, but faint with toil
and famine, they made their way through the stormy forest bearing](https://image.slidesharecdn.com/2058731-250604222541-148767a9/85/Ecosystems-Of-The-Deep-Oceans-Paul-A-Tyler-69-320.jpg)
![FOREBODINGS.
[
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on acorns by the way. Happily, the men left behind, excepting two
deserters, succeeded, a few days after, in rejoining the party.[138]
Thus was one heavy load lifted from the heart of
La Salle. But where was the Griffin? Time
enough, and more than enough, had passed for
her voyage to Niagara and back again. He scanned the dreary
horizon with an anxious eye. No returning sail gladdened the watery
solitude, and a dark foreboding gathered on his heart. Yet further
delay was impossible. He sent back two men to Michilimackinac to
meet her, if she still existed, and pilot her to his new fort of the
Miamis, and then prepared to ascend the river, whose weedy edges
were already glassed with thin flakes of ice.[139]
FOOTNOTES:
They named it Sainte Claire, of which the present name is a perversion.
Hennepin (1683), 58.
There is a rude plan of the establishment in La Hontan,
though in several editions its value is destroyed by the reversal
of the plate.
Relation de Tonty, 1684; Ibid., 1693. He was overtaken
at the Detroit by the Griffin.
Then usually known as Lac des Illinois, because
it gave access to the country of the tribes so called.
Three years before, Allouez gave it the name of Lac
St. Joseph, by which it is often designated by the
early writers. Membré, Douay, and others, call it Lac Dauphin.
The Great Mountain, the Iroquois name for the governor of Canada.
It was borrowed by other tribes also.
In the license of discovery granted to La Salle, he is expressly
prohibited from trading with the Ottawas and others who brought
furs to Montreal. This traffic on the lakes was, therefore, illicit. His](https://image.slidesharecdn.com/2058731-250604222541-148767a9/85/Ecosystems-Of-The-Deep-Oceans-Paul-A-Tyler-73-320.jpg)
![[
1
3
7]
[
1
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[
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9]
enemy, the Intendant Duchesneau, afterwards used this against him. Lettre de
Duchesneau au Ministre, 10 Nov., 1680.
Hennepin (1683), 79.
Hennepin (1683), 112; Relation de Tonty, 1693.
The official account of this journey is given at length in the
Relation des Découvertes et des Voyages du Sieur de la Salle,
1679-1681. This valuable document, compiled from letters and
diaries of La Salle, early in the year 1682, was known to
Hennepin, who evidently had a copy of it before him when he wrote his book,
in which he incorporated many passages from it.](https://image.slidesharecdn.com/2058731-250604222541-148767a9/85/Ecosystems-Of-The-Deep-Oceans-Paul-A-Tyler-74-320.jpg)
![LA SALLE'S
ADVENTURE.
CHAPTER XII.
1679, 1680.
LA SALLE ON THE ILLINOIS.
The St. Joseph.—Adventure of La Salle.—The Prairies.—Famine.—The
Great Town of the Illinois.—Indians.—Intrigues.—Difficulties.—Policy
of la Salle.—Desertion.—Another Attempt to poison La Salle.
On the third of December the party re-embarked,
thirty-three in all, in eight canoes,[140] and
ascended the chill current of the St. Joseph,
bordered with dreary meadows and bare gray forests. When they
approached the site of the present village of South Bend, they
looked anxiously along the shore on their right to find the portage or
path leading to the headquarters of the Illinois. The Mohegan was
absent, hunting; and, unaided by his practised eye, they passed the
path without seeing it. La Salle landed to search the woods. Hours
passed, and he did not return. Hennepin and Tonty grew uneasy,
disembarked, bivouacked, ordered guns to be fired, and sent out
men to scour the country. Night came, but not their lost leader.
Muffled in their blankets and powdered by the thick-falling snow-
flakes, they sat ruefully speculating as to what had befallen him; nor
was it till four o'clock of the next afternoon that they saw him
approaching along the margin of the river. His face and hands were
besmirched with charcoal; and he was further decorated with two
opossums which hung from his belt, and which he had killed with a
stick as they were swinging head downwards from the bough of a
tree, after the fashion of that singular beast. He had missed his way
in the forest, and had been forced to make a wide circuit around the
edge of a swamp; while the snow, of which the air was full, added to
his perplexities. Thus he pushed on through the rest of the day and](https://image.slidesharecdn.com/2058731-250604222541-148767a9/85/Ecosystems-Of-The-Deep-Oceans-Paul-A-Tyler-75-320.jpg)
![where the oozy, saturated soil quaked beneath their tread. All
around were clumps of alder-bushes, tufts of rank grass, and pools
of glistening water. In the midst a dark and lazy current, which a tall
man might bestride, crept twisting like a snake among the weeds
and rushes. Here were the sources of the Kankakee, one of the
heads of the Illinois.[141] They set their canoes on this thread of
water, embarked their baggage and themselves, and pushed down
the sluggish streamlet, looking, at a little distance, like men who
sailed on land. Fed by an unceasing tribute of the spongy soil, it
quickly widened to a river; and they floated on their way through a
voiceless, lifeless solitude of dreary oak barrens, or boundless
marshes overgrown with reeds. At night, they built their fire on
ground made firm by frost, and bivouacked among the rushes. A few
days brought them to a more favored region. On the right hand and
on the left stretched the boundless prairie, dotted with leafless
groves and bordered by gray wintry forests, scorched by the fires
kindled in the dried grass by Indian hunters, and strewn with the
carcasses and the bleached skulls of innumerable buffalo. The plains
were scored with their pathways, and the muddy edges of the river
were full of their hoof-prints. Yet not one was to be seen. At night,
the horizon glowed with distant fires; and by day the savage hunters
could be descried at times roaming on the verge of the prairie. The
men, discontented and half-starved, would have deserted to them
had they dared. La Salle's Mohegan could kill no game except two
lean deer, with a few wild geese and swans. At length, in their
straits, they made a happy discovery. It was a buffalo bull, fast
mired in a slough. They killed him, lashed a cable about him, and
then twelve men dragged out the shaggy monster, whose ponderous
carcass demanded their utmost efforts.
The scene changed again as they descended. On either hand ran
ranges of woody hills, following the course of the river; and when
they mounted to their tops, they saw beyond them a rolling sea of
dull green prairie, a boundless pasture of the buffalo and the deer, in
our own day strangely transformed,—yellow in harvest-time with](https://image.slidesharecdn.com/2058731-250604222541-148767a9/85/Ecosystems-Of-The-Deep-Oceans-Paul-A-Tyler-77-320.jpg)
![THE ILLINOIS
TOWN.
HUNGER
RELIEVED.
ripened wheat, and dotted with the roofs of a hardy and valiant
yeomanry.[142]
They passed the site of the future town of
Ottawa, and saw on their right the high plateau of
Buffalo Rock, long a favorite dwelling-place of
Indians. A league below, the river glided among islands bordered
with stately woods. Close on their left towered a lofty cliff,[143]
crested with trees that overhung the rippling current; while before
them spread the valley of the Illinois, in broad low meadows,
bordered on the right by the graceful hills at whose foot now lies the
village of Utica. A population far more numerous then tenanted the
valley. Along the right bank of the river were clustered the lodges of
a great Indian town. Hennepin counted four hundred and sixty of
them.[144] In shape, they were somewhat like the arched top of a
baggage-wagon. They were built of a framework of poles, covered
with mats of rushes closely interwoven; and each contained three or
four fires, of which the greater part served for two families.
Here, then, was the town; but where were the
inhabitants? All was silent as the desert. The
lodges were empty, the fires dead, and the ashes
cold. La Salle had expected this; for he knew that in the autumn the
Illinois always left their towns for their winter hunting, and that the
time of their return had not yet come. Yet he was not the less
embarrassed, for he would fain have bought a supply of food to
relieve his famished followers. Some of them, searching the deserted
town, presently found the caches, or covered pits, in which the
Indians hid their stock of corn. This was precious beyond measure in
their eyes, and to touch it would be a deep offence. La Salle shrank
from provoking their anger, which might prove the ruin of his plans;
but his necessity overcame his prudence, and he took thirty minots
of corn, hoping to appease the owners by presents. Thus provided,
the party embarked again, and resumed their downward voyage.](https://image.slidesharecdn.com/2058731-250604222541-148767a9/85/Ecosystems-Of-The-Deep-Oceans-Paul-A-Tyler-78-320.jpg)
![On New Year's Day, 1680, they landed and heard mass. Then
Hennepin wished a happy new year to La Salle first, and afterwards
to all the men, making them a speech, which, as he tells us, was
most touching.[145] He and his two brethren next embraced the
whole company in turn, in a manner, writes the father, most
tender and affectionate, exhorting them, at the same time, to
patience, faith, and constancy. Four days after these solemnities,
they reached the long expansion of the river then called Pimitoui,
and now known as Peoria Lake, and leisurely made their way
downward to the site of the city of Peoria.[146] Here, as evening
drew near, they saw a faint spire of smoke curling above the gray
forest, betokening that Indians were at hand. La Salle, as we have
seen, had been warned that these tribes had been taught to regard
him as their enemy; and when, in the morning, he resumed his
course, he was prepared alike for peace or war.
The shores now approached each other; and the Illinois was once
more a river, bordered on either hand with overhanging woods.[147]
At nine o'clock, doubling a point, he saw about eighty Illinois
wigwams, on both sides of the river. He instantly ordered the eight
canoes to be ranged in line, abreast, across the stream,—Tonty on
the right, and he himself on the left. The men laid down their
paddles and seized their weapons; while, in this warlike guise, the
current bore them swiftly into the midst of the surprised and
astounded savages. The camps were in a panic. Warriors whooped
and howled; squaws and children screeched in chorus. Some
snatched their bows and war-clubs; some ran in terror; and, in the
midst of the hubbub, La Salle leaped ashore, followed by his men.
None knew better how to deal with Indians; and he made no sign of
friendship, knowing that it might be construed as a token of fear. His
little knot of Frenchmen stood, gun in hand, passive, yet prepared
for battle. The Indians, on their part, rallying a little from their fright,
made all haste to proffer peace. Two of their chiefs came forward,
holding out the calumet; while another began a loud harangue, to
check the young warriors who were aiming their arrows from the](https://image.slidesharecdn.com/2058731-250604222541-148767a9/85/Ecosystems-Of-The-Deep-Oceans-Paul-A-Tyler-79-320.jpg)
![ILLINOIS
HOSPITALITY.
farther bank. La Salle, responding to these friendly overtures,
displayed another calumet; while Hennepin caught several scared
children and soothed them with winning blandishments.[148] The
uproar was quelled; and the strangers were presently seated in the
midst of the camp, beset by a throng of wild and swarthy figures.
Food was placed before them; and, as the Illinois
code of courtesy enjoined, their entertainers
conveyed the morsels with their own hands to the
lips of these unenviable victims of their hospitality, while others
rubbed their feet with bear's grease. La Salle, on his part, made
them a gift of tobacco and hatchets; and when he had escaped from
their caresses, rose and harangued them. He told them that he had
been forced to take corn from their granaries, lest his men should
die of hunger; but he prayed them not to be offended, promising full
restitution or ample payment. He had come, he said, to protect them
against their enemies, and teach them to pray to the true God. As
for the Iroquois, they were subjects of the Great King, and therefore
brethren of the French; yet, nevertheless, should they begin a war
and invade the country of the Illinois, he would stand by them, give
them guns, and fight in their defence, if they would permit him to
build a fort among them for the security of his men. It was also, he
added, his purpose to build a great wooden canoe, in which to
descend the Mississippi to the sea, and then return, bringing them
the goods of which they stood in need; but if they would not
consent to his plans and sell provisions to his men, he would pass on
to the Osages, who would then reap all the benefits of intercourse
with the French, while they were left destitute, at the mercy of the
Iroquois.[149]
This threat had its effect, for it touched their deep-rooted jealousy
of the Osages. They were lavish of promises, and feasts and dances
consumed the day. Yet La Salle soon learned that the intrigues of his
enemies were still pursuing him. That evening, unknown to him, a
stranger appeared in the Illinois camp. He was a Mascoutin chief,
named Monso, attended by five or six Miamis, and bringing a gift of](https://image.slidesharecdn.com/2058731-250604222541-148767a9/85/Ecosystems-Of-The-Deep-Oceans-Paul-A-Tyler-80-320.jpg)
![FRESH
INTRIGUES.
knives, hatchets, and kettles to the Illinois.[150] The chiefs
assembled in a secret nocturnal session, where, smoking their pipes,
they listened with open ears to the harangue of the envoys. Monso
told them that he had come in behalf of certain Frenchmen, whom
he named, to warn his hearers against the designs of La Salle,
whom he denounced as a partisan and spy of the Iroquois, affirming
that he was now on his way to stir up the tribes beyond the
Mississippi to join in a war against the Illinois, who, thus assailed
from the east and from the west, would be utterly destroyed. There
was no hope for them, he added, but in checking the farther
progress of La Salle, or, at least, retarding it, thus causing his men to
desert him. Having thrown his fire-brand, Monso and his party left
the camp in haste, dreading to be confronted with the object of their
aspersions.[151]
In the morning, La Salle saw a change in the
behavior of his hosts. They looked on him askance,
cold, sullen, and suspicious. There was one
Omawha, a chief, whose favor he had won the day before by the
politic gift of two hatchets and three knives, and who now came to
him in secret to tell him what had taken place at the nocturnal
council. La Salle at once saw in it a device of his enemies; and this
belief was confirmed, when, in the afternoon, Nicanopé, brother of
the head chief, sent to invite the Frenchmen to a feast. They
repaired to his lodge; but before dinner was served,—that is to say,
while the guests, white and red, were seated on mats, each with his
hunting-knife in his hand, and the wooden bowl before him which
was to receive his share of the bear's or buffalo's meat, or the corn
boiled in fat, with which he was to be regaled,—while such was the
posture of the company, their host arose and began a long speech.
He told the Frenchmen that he had invited them to his lodge less to
refresh their bodies with good cheer than to cure their minds of the
dangerous purpose which possessed them, of descending the
Mississippi. Its shores, he said, were beset by savage tribes, against
whose numbers and ferocity their valor would avail nothing; its](https://image.slidesharecdn.com/2058731-250604222541-148767a9/85/Ecosystems-Of-The-Deep-Oceans-Paul-A-Tyler-81-320.jpg)
![LA SALLE AND
THE INDIANS.
waters were infested by serpents, alligators, and unnatural
monsters; while the river itself, after raging among rocks and
whirlpools, plunged headlong at last into a fathomless gulf, which
would swallow them and their vessel forever.
La Salle's men were for the most part raw hands,
knowing nothing of the wilderness, and easily
alarmed at its dangers; but there were two among
them, old coureurs de bois, who unfortunately knew too much; for
they understood the Indian orator, and explained his speech to the
rest. As La Salle looked around on the circle of his followers, he read
an augury of fresh trouble in their disturbed and rueful visages. He
waited patiently, however, till the speaker had ended, and then
answered him, through his interpreter, with great composure. First,
he thanked him for the friendly warning which his affection had
impelled him to utter; but, he continued, the greater the danger, the
greater the honor; and even if the danger were real, Frenchmen
would never flinch from it. But were not the Illinois jealous? Had
they not been deluded by lies? We were not asleep, my brother,
when Monso came to tell you, under cover of night, that we were
spies of the Iroquois. The presents he gave you, that you might
believe his falsehoods, are at this moment buried in the earth under
this lodge. If he told the truth, why did he skulk away in the dark?
Why did he not show himself by day? Do you not see that when we
first came among you, and your camp was all in confusion, we could
have killed you without needing help from the Iroquois? And now,
while I am speaking, could we not put your old men to death, while
your young warriors are all gone away to hunt? If we meant to make
war on you, we should need no help from the Iroquois, who have so
often felt the force of our arms. Look at what we have brought you.
It is not weapons to destroy you, but merchandise and tools for your
good. If you still harbor evil thoughts of us, be frank as we are, and
speak them boldly. Go after this impostor Monso, and bring him
back, that we may answer him face to face; for he never saw either
us or the Iroquois, and what can he know of the plots that he
pretends to reveal?[152] Nicanopé had nothing to reply, and,](https://image.slidesharecdn.com/2058731-250604222541-148767a9/85/Ecosystems-Of-The-Deep-Oceans-Paul-A-Tyler-82-320.jpg)
Ecosystems Of The Deep Oceans Paul A Tyler
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- 6. ECOSYSTEMS OF THE WORLD 28 ECOSYSTEMS OF THE DEEP OCEANS
- 7. ECOSYSTEMS OF THE WORLD Editor in Chief: David W. Goodall Centre for Ecosystem Studies, Edith Cowan University, Joondalup, W.A. (Australia) I. TERRESTRIAL ECOSYSTEMS A. Natural Terrestrial Ecosystems 1. Wet Coastal Ecosystems 2. Dry Coastal Ecosystems 3. Polar and Alpine Tundra 4. Mires: Swamp, Bog, Fen and Moor 5. Temperate Deserts and Semi-Deserts 6. Coniferous Forests 7. Temperate Deciduous Forests 8. Natural Grasslands 9. Heathlands and Related Shrublands 10. Temperate Broad-Leaved Evergreen Forests 11. Mediterranean-Type Shrublands 12. Hot Deserts and Arid Shrublands 13. Tropical Savannas 14. Tropical Rain Forest Ecosystems 15. Wetland Forests 16. Ecosystems of Disturbed Ground B. Managed Terrestrial Ecosystems 17. Managed Grasslands 18. Field Crop Ecosystems 19. Tree Crop Ecosystems 20. Greenhouse Ecosystems 21. Bioindustrial Ecosystems II. AQUATIC ECOSYSTEMS A. Inland Aquatic Ecosystems 22. River and Stream Ecosystems 23. Lakes and Reservoirs B. Marine Ecosystems 24. Intertidal and Littoral Ecosystems 25. Coral Reefs 26. Estuaries and Enclosed Seas 27. Ecosystems of the Continental Shelves 28. Ecosystems of the Deep Ocean C. Managed Aquatic Ecosystems 29. Managed Aquatic Ecosystems III. UNDERGROUND ECOSYSTEMS 30. Subterranean Ecosystems
- 8. ECOSYSTEMS OF THE WORLD 28 ECOSYSTEMS OF THE DEEP OCEANS Edited by P.A. Tyler School of Ocean and Earth Science, Southampton Oceanography Centre, European Way, Southampton SO14 3ZH, United Kingdom 2003 ELSEVIER Amsterdam – Boston – London – New York – Oxford – Paris San Diego – San Francisco – Singapore – Sydney – Tokyo
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- 10. PREFACE The deep ocean floor covers over 50% of the surface of the earth. It is often said that we know more about the surface of the moon than we do about the deep ocean floor and the water column above it. While this is not strictly true, we do know remarkably little, as a proportion of the total, of the deep ocean environment. Paradigms are continually changing, and we know now that the deep sea is an ecosystem of high species diversity, that it may have seasons as seen in temperate land ecosystems, and that in certain areas turbulence can be a great as anything seen in coastal shallow waters. Last, but by no means least, the originally perceived idea that the deep sea was an oligotrophic environment in which all environmental processes were gentle and physiological processes slow is no longer valid. We know now that the deep sea is essentially a heterotrophic system fuelled by organic carbon from surface waters, with the notable exception of hydrothermal vents and cold seeps where substantial ecosystems are fuelled by chemosynthetic processes. The continuing theme of this volume is how this energy input affects the deep-sea ecosystem. All science has its eras of exploration, observation and experimentation. Exploration in deep-sea biology is often considered to have come to a finale with the Galathea cruise of 1950 to 1952. Subsequent discoveries of hydrothermal vents and cold seeps show that the deep-sea age of exploration is still with us and will continue. The 1960s saw the first change in our perception of the deep-sea with the introduction of more sophisticated sampling gear. This has been used from then and still continues to be used for much observation work. The introduction of submersibles, and, more recently, remote operated vehicles and landers, has allowed us to conduct manipulative experimentation on the deep sea bed and in the water column. This volume is a review of where our knowledge stands at this point. All the chapters are written by authorities on their respective subjects, all of whom are still practicing deep-sea biologists. The volume is divided into sections covering the environment of the deep sea, specific deep-water seas and oceans, and lastly a review of the processes that occur there. All the chapters have been peer-reviewed by other experts in deep-sea biology, to all of whom I extend my thanks for their care and advice. I wish to say a special thank you to all the authors. As I have said above, all are active research scientists, often working for extended periods at sea. I know their scientific lives are full, and I am delighted they were willing to write chapters and put up with my impatient prodding. I would also like to thank the series editor David Goodall for his advice, enthusiasm, patience and his unremitting courtesy when I failed to answer his requests! Lastly, I would like to express my sincere thanks to Lida de Maaijer Hoek of Isys Prepress Services for her patience, good humour and exceptional care in the desk editing of this volume. Paul A. Tyler Editor v
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- 12. LIST OF CONTRIBUTORS M.V. ANGEL Southampton Oceanography Centre University of Southampton Southampton SO14 3ZH United Kingdom A. CLARKE Biological Sciences Division British Antarctic Survey High Cross Madingley Road Cambridge CB3 0ET United Kingdom A.W.J. DEMOPOULOS Department of Oceanography University of Hawaii at Manoa 1000 Pope Road Honolulu, HI 96822 USA R.J. ETTER Department of Biology University of Massachusetts 100 Morrissey Boulevard Boston, MA 02125 USA J.D. GAGE Scottish Association for Marine Science PO Box 3 Oban Argyll PA37 1QA United Kingdom A.J. GOODAY DEEPSEAS Benthic Biology Group George Deacon Division for Ocean Processes Southampton Oceanography Centre European Way Southampton SO14 3ZH United Kingdom S.K. JUNIPER GEOTOP Université de Québec á Montréal Montreal, Quebec H2X 3Y7 Canada L.A. LEVIN Integrative Oceanography Division Scripps Institution of Oceanography La Jolla, CA 92093-0218 USA M.A. REX Department of Biology University of Massachusetts 100 Morrissey Boulevard Boston, MA 02125 USA M. SIBUET DERO/EP IFREMER Centre de Brest 29280 Plouzane France C.R. SMITH Department of Oceanography University of Hawaii at Manoa 1000 Pope Road Honolulu, HI 96822 USA C.T. STUART Department of Biology University of Massachusetts 100 Morrissey Boulevard Boston, MA 02125 USA vii
- 13. viii LIST OF CONTRIBUTORS H. THIEL Poppenbuettler Markt 8A Hamburg, 22399 Germany D. THISTLE Department of Oceanography Florida State University Tallahassee, FL 32306-4320 USA V. TUNNICLIFFE Department of Biology University of Victoria Victoria, BC V8W 2Y2 Canada P.A. TYLER School of Ocean and Earth Science and DEEPSEAS Benthic Biology Group Southampton Oceanography Centre University of Southampton Southampton SO14 3ZH United Kingdom C.M. YOUNG Oregon Institute of Marine Biology University of Oregon, PO Box 5389 Charleston, OR 97420 USA
- 14. CONTENTS PREFACE . . . . . . . . . . . . . . . . . . . . . . . v LIST OF CONTRIBUTORS . . . . . . . . . . . . . . vii Chapter 1. INTRODUCTION by P.A. TYLER . . . . . . . . . . . . . . 1 Chapter 2. THE DEEP-SEA FLOOR: AN OVERVIEW by D. THISTLE . . . . . . . . . . . . . . 5 Chapter 3. THE PELAGIC ENVIRONMENT OF THE OPEN OCEAN by M.V . ANGEL . . . . . . . . . . . . . . 39 Chapter 4. REDUCING ENVIRONMENTS OF THE DEEP-SEA FLOOR by V . TUNNICLIFFE, S.K. JUNIPER and M. SIBUET . . . . . . . . . . . . . . . . 81 Chapter 5. THE DEEP ATLANTIC OCEAN by L.A. LEVIN and A.J. GOODAY . . . . 111 Chapter 6. THE DEEP PACIFIC OCEAN FLOOR by C.R. SMITH and A.W.J. DEMOPOULOS 179 Chapter 7. THE DEEP INDIAN OCEAN FLOOR by A.W.J. DEMOPOULOS, C.R. SMITH and P.A. TYLER . . . . . . . . . . . . . . . . 219 Chapter 8. THE POLAR DEEP SEAS by A. CLARKE . . . . . . . . . . . . . . 239 Chapter 9. THE PERIPHERAL DEEP SEAS by P.A. TYLER . . . . . . . . . . . . . . 261 Chapter 10. LARGE-SCALE SPATIAL AND TEMPORAL PATTERNS OF DEEP-SEA BENTHIC SPECIES DIVERSITY by C.T. STUART, M.A. REX and R.J. ETTER . . . . . . . . . . . . . . . . . . . . . . . . 295 Chapter 11. FOOD INPUTS, UTILIZATION, CARBON FLOW AND ENERGETICS by J.D. GAGE . . . . . . . . . . . . . . . 313 Chapter 12. REPRODUCTION, DEVELOPMENT AND LIFE-HISTORY TRAITS by C.M. YOUNG . . . . . . . . . . . . . 381 Chapter 13. ANTHROPOGENIC IMPACTS ON THE DEEP SEA by H. THIEL . . . . . . . . . . . . . . . 427 Chapter 14. EPILOGUE: EXPLORATION, OBSERVATION AND EXPERIMENTATION by P.A. TYLER . . . . . . . . . . . . . . 473 GLOSSARY . . . . . . . . . . . . . . . . . . . . . . 477 SYSTEMATIC LIST OF GENERA . . . . . . . . . . 479 AUTHOR INDEX . . . . . . . . . . . . . . . . . . . 483 SYSTEMATIC INDEX . . . . . . . . . . . . . . . . 507 GENERAL INDEX . . . . . . . . . . . . . . . . . . 517 ix
- 15. To my wife Amanda for her love and support over the years, and her patience with the amount of time I spent at sea!
- 16. Chapter 1 INTRODUCTION Paul A. TYLER The largest single ecosystem on earth is the deep sea. The sea surface occupies ~70% of the surface of the earth, and 50% of the surface of the earth is covered by more than 3000 m of ocean, with a mean depth of ~3800 m. It is the very remoteness of the deep sea and the difficulties encountered in its exploration that have resulted in it being one of the least understood environments on earth. At the present time there is detailed information about specific areas of the deep sea, but these are mere pinpricks in the vastness of this environment. The understanding of the deep-sea ecosystem is entwined with some of the most exciting aspects of scientific exploration and with the development of technologies for sampling and penetrating this environment. This volume is a status report, at the beginning of the 21st century, on current knowledge of the deep sea, on how perceptions of it have changed and where the exciting scientific discoveries will be made in the future. CHANGING PARADIGMS Explorers and commercial interests have used the sea as a means of transport for millennia. However, they always looked to the horizon, and it was only in the latter part of the 19th century that scientists went to sea with the specific aim of looking downwards into the impenetrable depths. One of the first was Forbes (1844), who sampled down to a depth of 600 m in the Aegean. Today one would consider this choice of sampling station as unfortunate, since this region of the Mediterranean deep sea is faunistically very poor, and the lack of animals in Forbes’s samples led to the ‘azoic theory’ that little or no life existed below 600 m. The establishment of such a paradigm was in direct opposition to observations of the ophiuroid Astrophyton being brought up on a sounding line from a depth of 1800 m in Baffin Bay (Tyler, 1980), and the pioneering work of Michael and G.O. Sars in Norwegian fjords (Sars, 1864, 1868). Establishing the presence of a fauna in the deep sea presented irresistible challenges to a small group of scientists led by Charles Wyville Thomson. Wyville- Thomson used HMS Porcupine to sample the ocean to the northwest of Scotland and to the west of Ireland in the late 1860s, and found a fauna at depths exceeding 4000 m (Thomson, 1873). This series of cruises established the first ecological observation in the deep sea by showing that there was a marked temperature difference associated with faunal change as one moved across what is now called the Scotland– Faroes–Iceland Ridge from the warm deep North At- lantic to the cold deep Norwegian Sea (see Chapter 6). The results of the Porcupine sampling programme led directly to the HMS Challenger expedition of 1872 to 1876. This expedition traversed the oceans of the globe and demonstrated a widespread and varied fauna in the deep sea, as well as taking numerous physical and chemical measurements. The results of this cruise, now considered the forerunner of modern oceanography, were published in a series of detailed volumes edited by, and at the expense of, John Murray. A readable account of the Challenger expedition has been published by Linklater (1972). The Challenger expedition led directly to the ‘heroic’ age of deep-sea exploration, with expeditions sampling many areas of the world’s oceans (Menzies et al., 1973; Mills, 1983). The heroic age culminated in the Danish Galathea expedition of 1950 to 1952, which demonstrated that life could be found in the deepest of all the oceans, in the ocean trenches. One of the main outcomes of this age of exploration was 1
- 17. 2 Paul A. TYLER the publication of descriptions of the fauna collected on these voyages. Taking stock of deep-sea ecology at this point in time would have led to the establishment of the following paradigms: • The deep sea was species-poor. • It was a tranquil quiescent environment. • There was a slow rain of material from surface to the deep (although see Moseley, 1880). • No primary production occurred within the deep sea. The 1960s heralded a new approach to deep-sea ecology, driven by technology. Quantification became the name of the game, and to get accurate data it was necessary to replace the coarse-meshed qualitative sampling gear of the heroic age with more refined quantitative gear. This was initially achieved by Howard Sanders and Robert Hessler from the Woods Hole Oceanographic Institution, who used an anchor dredge (later an anchor box dredge: Gage and Tyler, 1991) to sample a series of stations down to a depth of 5000 m between Gay Head, Massachusetts and Bermuda. The fine mesh of the anchor dredge collected a wide variety of species, many new to science, which had been missed by the coarse dredges of the heroic age. Thus the concept of high biodiversity in the deep sea was established, although the absolute diversity is still very much subject to debate (see Chapter 10); but it is now believed that the deep oceans are as diverse as tropical rain forests. Although known to be diverse, it was assumed that the deep-sea system was heterotrophic, relying on the slow sinking of material from surface waters to provide an energy source for the inhabitants. The 1970s and 1980s provided evidence that this environment was more dynamic than originally thought. The first example was the discovery of hydrothermal vents along the Galapagos Ridge in 1977 (see Chapter 4). For the first time there was evidence that primary production could take place within the deep sea, and an ecosystem independent of sunlight had been discovered. This discovery led to one of the most active programmes in deep-sea biology, and the discovery of hydrothermal vents continues to this day. There can be few people interested in the natural environment who have not seen photos or videos of these spectacular environments. Subsequently, a second type of primary- production environment was observed in the form of cold seeps (see Chapter 4). Both hydrothermal vents and cold seeps are driven by the availability of reduced chemicals such as hydrogen sulphide and methane, the main difference being the temperature of emission. In terms of energy availability a parallel, but no less important, revolution was occurring in understanding the input of material from surface primary production. The concept of the slow rain of surface primary pro- duction to the seabed was challenged by technological advances, particularly in the use of sediment traps to collect the sinking material. Such sediment traps, together with other techniques (see Chapters 2 and 11) showed that, particularly at temperate latitudes, surface production sank rapidly to the seabed – on average, at a rate of ~100 m d−1 . As a result, the signal of seasonal surface production was transmitted to the seabed, and it is now known that a number of organisms on the deep- sea bed respond seasonally to this input. This theme is explored in many chapters in this volume. This seasonal perturbation is mild in comparison to the last major shift in paradigms. Over certain areas of the seabed, especially under areas of high surface- eddy kinetic energy, benthic storms are created by the input of energy to the seabed. These storms are analogous to the blizzards of Antarctica. They create strong currents transporting sediment, which is then deposited in drifts on the seabed, smothering the local fauna (see Chapter 2). Lastly, technology has allowed humans to penetrate this ‘remote’ environment. SCUBA diving is limited to the top 30 m of the water column; but the development of submersibles has allowed scientists to dive to the deep-sea bed and conduct manipulative experiments as though they were working at the laboratory bench. Current knowledge of hydrothermal vents and cold seeps would be insignificant if it were not for the submersible. Submersibles are still used today; but the Remote Operated Vehicle (ROV) allows similar access from the comfort of the surface tender without the potential dangers of manned submersibles. Today one may summarize the paradigms for the deep-sea environment as: • High species diversity. • Periods of benthic storms perturbing an apparently gentle environment. • Seasonal input of surface-derived energy for het- erotrophic organisms. • Primary production at vents and cold seeps. The change in understanding of the deep sea has been a function of an increase in the ability of scientists to gain knowledge from this environment. Despite recent recognition of the above paradigms, all of them
- 18. INTRODUCTION 3 are natural phenomena. As yet the deep sea is exploited only to a very limited extent, but this may change in the future. Disposal of waste has become prominent on the political agenda, particularly as land-based disposal areas become saturated. The deep sea has already been used for the disposal of low-level radioactive waste, pharmaceuticals and dredge spoil (see Chapter 13). Possibly more insidious is the use of the deep sea in relation to climate change. There is evidence of ‘natural’ decadal-scale changes in the fauna of the northeast Atlantic, possibly related to climate change. The deep sea has also been suggested as a repository for the excess carbon dioxide causing the so-called ‘greenhouse effect’. The vastness of the deep ocean aids its very stability, but in localized areas this is already being challenged. The public outcry over the ‘Brent Spar’ (see Chapter 13) demonstrates that public awareness of this environment is increasing rapidly. Finally, with the decline of continental-shelf fisheries, fishing fleets are moving into deeper and deeper water, and there is evidence that at least one deep-sea fish, the orange roughy (Hoplostethus atlanticus), is already overexploited. THE DEEP SEA TODAY What is the deep-sea? Ask virtually any deep-sea biologist and you get a slightly different answer. For most, it is the region below 200 m, representing the transition from the continental shelves to the continental slope. This is the boundary that has been selected for this volume (see Chapter 2). Definitions based on light penetration, depth of the mixed surface layer, or temperature may be just as valid (see Gage and Tyler, 1991). The approach to this volume has been to examine the deep sea from a number of facets, and differs from the approach of most previous volumes in this series. The linking theme between all the chapters is the availability of energy for organisms in the water column and at the deep-sea floor. Chapters 2, 3 and 4 examine environmental aspects of the deep sea – specifically the deep-sea floor, the water column and reducing environments. Chapters 5, 6, 7, 8, and 9 examine the ecology of the major oceans and those seas peripheral to the main ocean that have waters of oceanic depth. Chapters 10, 11 and 12 examine some of the specific processes that occur within the deep-sea ecosystem; and Chapter 13 explores the anthropogenic impact that has taken place or that may occur in the future. ACKNOWLEDGEMENTS I would like to take this opportunity to thank all the authors who have contributed to this volume. I may be editor but it has been a collective enterprise by a series of world-class scientists whose passion is for the marine environment and the deep sea in particular. I would also like to thank David Goodall for his forbearance throughout its production. REFERENCES Forbes, E., 1844. Report on the Mollusca and Radiata of the Aegean Sea, and their distribution, considered as bearing on geology. Report (1843) to the 13th meeting of the British Association for the Advancement of Science, pp. 30–193. Gage, J.D. and Tyler, P.A., 1991. Deep-Sea Biology: A Natural History of Organisms at the Deep Sea Floor. Cambridge University Press, Cambridge, 504 pp. Linklater, E., 1972. The Voyage of the Challenger. Murray, London. Menzies, R.J., George, R.Y. and Rowe, G.T., 1973. Abyssal Ecology and Environment of the World Ocean. Wiley Interscience, New York. Mills, E.L., 1983. Problems of deep-sea biology: an historical perspective. In: G.T. Rowe (Editor), The Sea, Vol. 8. Wiley Interscience, New York, pp. 1–79. Moseley, H.N., 1880. Deep-sea dredging and life in the deep sea. Nature, 21: 543−547, 569−572, 591−593. Sars, M., 1864. Bemaerkninger Over det Dyriske Livs Udbredning I Havets Dybder, Christiana. Videnskabs-Selskabs Forhandlinger for 1864. Sars, M., 1868. Fortsatte Bemaerkninger Over det Dyriske Livs Udbredning I Havets Dybder, Christiana. Videnskabs-Selskabs Forhandlinger for 1868. Thomson, C.W., 1873. The Depths of the Sea. MacMillan, London, 527 pp. Tyler, P.A., 1980. Deep-sea ophiuroids. Oceanogr. Mar. Biol. Ann. Rev., 18: 125−153.
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- 20. Chapter 2 THE DEEP-SEA FLOOR: AN OVERVIEW David THISTLE INTRODUCTION This chapter provides a general introduction to the ecosystem of the deep-sea floor, beginning with a description of the physical environment of the deep sea. A section on how information is obtained about the deep-sea-floor ecosystem follows, because knowledge of this ecosystem is greatly influenced by the effective- ness of the available technology. Introductions to the fauna of the deep sea where the substratum is sediment (soft bottoms) and where it is not (hard bottoms) follow. The chapter concludes with a section on the pace of life in the deep sea. The geographic extent of the deep-sea-floor ecosystem The deep sea is usually defined as beginning at the shelf break (Fig. 2.1), because this physiographic feature coincides with the transition from the basically shallow-water fauna of the shelf to the deep-sea fauna (Sanders et al., 1965; Hessler, 1974; Merrett, 1989). The shelf break is at about 200 m depth in many parts of the ocean, so the deep sea is said to begin at 200 m. The deep-sea floor is therefore a vast habitat, cov- ering more than 65% of the Earth’s surface (Sverdrup et al., 1942). Much of it is covered by sediment, but in some regions (e.g., mid-ocean ridges, seamounts) bare rock is exposed. In the overview of environmental conditions that follows, the information applies to both hard and soft bottoms unless differences are noted. The ecosystems of hydrothermal vents and cold seeps are special cases and are described in Chapter 4. Environmental setting The deep-sea floor is an extreme environment; pressure is high, temperature is low, and food input is small. It has been characterized as a physically stable envi- ronment (Sanders, 1968). Below I review the major environmental variables and indicate circumstances under which these environmental variables constitute a biological challenge. I also show that the image of the deep-sea floor as monotonous and stable 0 1 2 3 4 5 6 Distance from shore Depth (km) Continental shelf Continental rise Abyssal plain Shelf break Continental slope Bathyal zone Abyssal zone Fig. 2.1. Diagrammatic cross section of the ocean showing the major physiographic features and major depth zones. The sublittoral zone (0–200 m) is not labeled, and the hadal zone (6000–10 000+ m) is not shown. Modified from Gage and Tyler (1991). Copyright: Cambridge University Press 1991. Reprinted with the permission of Cambridge University Press. 5
- 21. 6 David THISTLE must be tempered for some variables and some locations. Pressure Pressure increases by one atmosphere (105 Pascals) for every 10-m increase in water depth, so pres- sure varies from 20 atm at the shelf-slope break to 1000 atm in the deepest parts of the trenches. Pressure can affect organisms physiologically. For example, high deep-sea pressures oppose the secretion of gas. Many bottom-associated deep-sea fishes that use a gas-filled swim bladder to regulate their buoyancy (Merrett, 1989) overcome this problem, in part, by increasing the length of the retia mirabilia (Marshall, 1979), a component of the system that secretes gas into the swim bladder. Pressure also affects an organism biochemically because the performance of proteins (e.g., enzymes) and lipid structures (e.g., membranes) changes with pressure. For example, any biochemical reaction that involves an increase in volume at any step in the transition from reactants to products will proceed more slowly as pressure increases (Hochachka and Somero, 1984). A species that lives in the deep sea must have adaptations that reduce or eliminate the pressure effects on reaction rates. Such adaptations include modifications of the enzymatic machinery (e.g., changes to the amino-acid sequence of an enzyme) to reduce or eliminate volume changes during catalysis (Siebenaller and Somero, 1978). These adaptations come with a cost; pressure-insensitive enzymes are not as efficient at shallow-water pressures as are those of shallow-water species (Hochachka and Somero, 1984). This requirement for molecular-level adaptations has been postulated to constitute an evolutionary barrier that must have been overcome by those species that successfully entered the deep sea. Bottom-water temperature Bottom-water temperatures generally decrease with increasing depth, reaching ~2ºC on the abyssal plain, but the pattern varies with latitude and region (Mantyla and Reid, 1983; Fig. 2.2). Above about 500 m in mid- latitude, temperature varies seasonally, but with dimin- ishing amplitude with increasing depth (Figs. 2.2, 2.3). It should be noted that, at high latitudes, the vertical gradient in bottom-water temperature is small (Sver- drup et al., 1942). A small vertical temperature gradient also occurs in regions where the bottom water is warm (e.g., the Mediterranean Sea and the Red Sea). Fig. 2.2. Typical profiles of mean temperature versus depth for the open ocean. Modified from Pickard and Emery (1990). Reproduced by permission of Butterworth Heinemann. 0 200 400 600 800 1000 1200 1400 24 22 20 18 16 14 12 10 8 6 4 2 0 -2 Temperature ( C) o Depth (m) Minimum temperature Maximum temperature Fig. 2.3. Annual temperature variation in the western North Atlantic illustrating the diminishing amplitude of seasonal variation with depth. Modified from Sanders (1968). Reproduced by permission of the University of Chicago Press. Copyright 1968 by the University of Chicago. In summary, most of the water overlying the deep- sea floor is cold compared to that over most shallow- water habitats. At depths below ~800 m, temperature is remarkably constant (Fig. 2.3). In the abyss, temporal variation is measured in the second decimal place and occurs, for example, because internal tides and waves cause the oscillation of isothermal surfaces. Hydrothermal vents are exceptions; they occur in the cold deep sea, but temperatures near them are elevated and variable (see Chapter 4). The low temperatures have consequences for deep- sea-floor organisms because the cold reduces chemical reaction rates and shifts reaction equilibria toward reactants and away from products (Hochachka and Somero, 1984). To metabolize at reasonable rates, deep-sea species must have biochemical machinery that compensates. For example, low temperatures decrease
- 22. THE DEEP-SEA FLOOR: AN OVERVIEW 7 enzyme flexibility and, therefore, catalytic rates. This effect can be offset over evolutionary time by changes in the amino-acid sequence of an enzyme to reduce the number of weak interactions (e.g., hydrogen bonds) that stabilize its three-dimensional structure (Hochachka and Somero, 1984). The necessity for such adaptation to low temperatures, like that to high pressure, may constitute a barrier which a warm-water shallow-water lineage must overcome evolutionarily to colonize the cold deep sea. Salinity In shallow, coastal waters, salinity can affect benthic species. For example, in estuaries, the organisms must be adapted physiologically to live in water that changes salinity with the tides. In most of the deep sea, on the other hand, the salinity of the bottom water is fully marine (c. 35‰). Exceptions include the Mediterranean and Red Sea (39‰) and hypersaline basins such as the Orca Basin in the Gulf of Mexico (c. 300‰: Shokes et al., 1976). At most locations in the deep sea, salinity varies little with time, and that variation appears to be irrelevant to the ecology of deep-sea organisms. Oxygen Oxygen enters the ocean by exchange with the atmosphere and as a by-product of photosynthesis by marine plants in the euphotic zone. The dissolved gas is carried to the deep-sea floor by the descent of surface waters. The water overlying most of the deep- sea floor is saturated with oxygen or nearly so (5– 6 ml °−1 ), and the variation in space and time of oxygen concentration on the scale of an individual organism is small in absolute terms and does not constitute an environmental challenge for organisms living in the near-bottom water or on the seabed. Two major conditions reduce oxygen concentration to levels that are problematic for organisms. First, organic material (e.g., fecal pellets) that falls from the euphotic zone is decomposed by aerobic bac- teria and is consumed by zooplankton as it sinks. The decomposition and animal respiration reduce the oxygen concentration, producing an oxygen-minimum layer in mid-water, usually between 300 m and 1000 m depth (Fig. 2.4). Where this layer intersects the deep-sea floor, the bottom fauna can be reduced or eliminated (Sanders, 1969). For example, the water bathing Volcano 7 (in the eastern tropical Pacific) above 750 m has an oxygen concentration of 0.08– 1000 2000 3000 4000 0.25 0.20 0.15 0.10 0.05 7 6 5 4 3 2 1 0 A B C Oxygen (ml l ) -1 Oxygen (mol m ) -3 Depth (m) Fig. 2.4. The vertical distribution of dissolved oxygen illustrating the oxygen minimum zones in different regions: (A) south of California, (B) the eastern part of the South Atlantic, and (C) the Gulf Stream. Modified from Anonymous (1989). Reproduced by permission of Butterworth Heinemann. 0.09 ml °−1 , and the mean abundance of sediment- dwelling animals caught on a 0.300-mm mesh sieve is 1854 individuals m−2 . Just below 750 m, the oxygen concentration is slightly higher (0.11–0.16 ml °−1 ), and the mean abundance quadruples to 8457 m−2 . The pattern for the hard-bottom fauna on Volcano 7 is similar (Wishner et al., 1990). The second circumstance concerns basins where the bottom water does not freely exchange with that of the surrounding region, for example, because of a topographic barrier. The reduced exchange decreases the oxygen-supply rate to the bottom waters of the basin. Organic material settles into the basin and is decomposed by microbes. Depending on the balance between the rate at which oxygen is supplied and the rate at which it is consumed, the oxygen concentration in the bottom waters can be much less than that of the surrounding region, or even zero. Such conditions can reduce or eliminate the aerobic benthic fauna. It should be noted that oxygen conditions need not be constant; for instance, the Santa Barbara Basin has alternated between oxic and reduced-oxygen conditions many times in the last 60 000 years (Behl and Kennett, 1996; Cannariato et al., 1999). The ecological effects of low oxygen concentra- tion in the overlying water are complex. For the macrofauna1 , diversity begins to decline at oxy- 1 Macrofauna, meiofauna: see Table 2.1, p. 11.
- 23. 8 David THISTLE gen concentrations of ~0.45 ml °−1 (Levin and Gage, 1998). In terms of abundance, standing stocks at some low-oxygen sites are very low (Sanders, 1969; Levin et al., 1991), whereas at others they are remarkably high (Levin et al., 2000). Sites of high abundance seem to occur where oxygen concentration exceeds ~0.16 ml °−1 (Levin et al., 2000) and the flux of organic carbon is high (Sanders, 1969). Where abundances are high, the number of species that constitute the fauna tends to be low relative to that at comparable, high-oxygen sites, suggesting that only a few species have solved the physiological problems presented by the low oxygen concentration and that the ecological reward for those that have is substantial. Interestingly, the identity of the successful species varies from site to site, suggesting that adaptation to low oxygen concentrations has occurred many times. In general, tolerance of reduced oxygen increases from crustaceans to molluscs to polychaetes, but some exceptions are known (Levin et al., 2000). Meiofauna1 are also sensitive to reduced oxygen. In oxygen-minimum zones, the diversity of benthic foraminiferan faunas tends to be reduced, and most individuals belong to a small number of species (Sen Gupta and Machain-Castillo, 1993). Experimental evidence from shallow water reveals that tolerance to oxygen stress generally decreases from benthic copepods to nematodes and soft-shelled foraminifers to hard-shell foraminifers (Moodley et al., 1997). These taxon-specific differences in tolerance imply that as oxygen-stress increases the meiofauna will change in composition. Oxygen concentration also varies with depth in the sediment. Oxygen enters the pore water of deep- sea sediments by diffusion and by the activities of organisms that pump or mix water into the sediment. Oxygen is consumed by animal and microbial res- piration and by chemical reactions in the sediment. Where the deposition rate of labile organic matter is relatively high and the oxygen concentration in the bottom water is low, as in the basins of the California Continental Borderland, free oxygen disappears within the first centimeter (Reimers, 1987). Where organic- matter deposition rates are low and the bottom water is well oxygenated, as beneath the oligotrophic waters of the central North Pacific, abundant free oxygen is present several centimeters into the seabed (Reimers, 1987). The depth of oxygen penetration into the sediment limits the vertical distribution of organisms that require it, such as most metazoans. Light Light intensity decreases exponentially with depth in the water column because incident photons are absorbed or scattered. Particles suspended in the water (sediment particles, phytoplankton cells) increase both absorption and scattering, but even in the clearest ocean water no photosynthetically useful light reaches the sea floor below about 250 m (Fig. 2.5). Therefore, the deep-sea floor (except the shallowest 50 m) differs from more familiar ecosystems in that plant primary production does not occur. Except for hydrothermal- vent and cold-seep communities, the food of deep-sea- floor organisms must be imported (see Chapter 11). The paucity of food reaching the deep-sea floor has profound consequences for the ecology of organisms living there. The decrease of light intensity with increasing depth has other consequences for deep-sea species. For example, in shallow water most isopods have eyes. As depth increases, the proportion of isopod species without eyes increases until, at abyssal depths, eyes are absent (Hessler and Thistle, 1975; see Thurston and Bett, 1993, for amphipods). The implication of this pattern is that vision is of decreasing importance for some animal groups as depth increases. Its role in the ecology of these species (in prey location, in mate location, in movement) must be taken over by other senses such as chemoreception and mechanoreception. Also, the blindness suggests that they do not use bioluminescence, which is important to many animals of the deep water column (Chapter 3). Demersal fishes (e.g., Macrouridae) show a parallel pattern. They can have eyes, even at great depth, but eyes are smaller in deeper-living species (Marshall, 1979). Near-bottom flow In much of the deep sea, the near-bottom water moves slowly compared to that in shallow-water environments. Speeds in the bathyal zone tend to be less than 10 cm s−1 at 1 m above the bottom, those in the abyssal zone less than 4 cm s−1 . Speeds in both environments vary little from day to day at a location (Eckman and Thistle, 1991). Because the horizontal flow speed must decrease to zero at a solid boundary (Vogel, 1981), the horizonal speeds just above the seabed will be much less than those 1 m above. These flows are benign in that they are too slow to erode sediment or benthic organisms. The flow does move
- 24. THE DEEP-SEA FLOOR: AN OVERVIEW 9 0 200 400 600 800 1000 1200 10-11 10-9 10-7 10-5 10-3 10-1 101 103 105 Light intensity ( W cm ) m -2 Limit of phytoplankton growth Clear coastal water Depth (m) Clearest ocean water Euphotic Aphotic Disphotic Limit of crustacean phototaxis Detection limit for deep-sea fishes Fig. 2.5. The attenuation of light under different conditions of water clarity. Modified from Parsons et al. (1977). Reproduced by permission of Butterworth Heinemann. some material, in particular phytodetritus (flocculent material of low specific density consisting of phyto- plankton cells in an organic matrix, Billett et al., 1983), which accumulates in depressions (Lampitt, 1985). The water is never still, because tidal forces move water at all ocean depths. As a result, the water bathing all sessile sea-bed organisms slowly changes, bringing food and removing wastes. Near-bottom velocities are not slow everywhere in the deep sea. At a site at the base of the Scotian Rise (North Atlantic), near-bottom flows 5 m above the bottom can approach 30 cm s−1 (Gross and Williams, 1991). During periods of fast flow, the sediment can be eroded. These “benthic storms” occur several times each year and have consequences for the fauna. The fast flows can have positive effects. For example, the increase in the horizontal food flux benefits some species (Nowell et al., 1984). In contrast, surface-living crustaceans can be significantly less abundant than at quiescent deep-sea sites (Thistle and Wilson, 1996). Many soft-bottom regions experience erosive flows (see Fig. 1 of Hollister and Nowell, 1991). Such flows also prevent sediment settling from above from covering the horizontal surfaces of some deep-sea hard bottoms. The soft-bottom seafloor Deep-sea sediments consist, in part, of particles derived from the weathering of rock on land (= ter- rigenous particles), which are transported to the sea by wind and in rivers. In consequence, the supply of terrigenous particles is highest near the continents. The rate of supply and the size of the particles decrease with distance from land. Deep-sea sediments also contain particles produced by planktonic organisms in the overlying water. Di- atoms, radiolarians, and silicoflagellates make silica shells; foraminifers, coccolithophores, and pteropods make calcium carbonate shells. As depth increases, the rate of silica and calcium carbonate dissolution increases, but at a given depth, calcium carbonate dissolves more rapidly. The contribution of shells to the sediment depends on the rate at which they are produced in the overlying water and the rate at which they dissolve in the water column and at the seafloor. If shells constitute more than 30% by volume of the deposit, the sediment is called a biological ooze (Gage and Tyler, 1991). The balance between the rates of supply of terrestrial and biological particles and the rate of dissolution of biological particles controls the local sediment composition. For example, only a small amount of terrigenous material reaches the areas farthest from land, but the productivity of the overlying waters in these areas (oceanic central gyres) is so small that the
- 25. 10 David THISTLE few shells that are produced and fall to the seafloor are dissolved away. As a result, the sediment (abyssal red clay) consists of terrigenous particles. Accumulation rates are low, c. 0.5 mm per thousand years. Where productivity is high, the production rate of both siliceous and calcium carbonate shells is high. If the water is deep, the calcium carbonate shells that reach the seafloor dissolve. The sediment will be composed of terrigenous and siliceous particles, a diatomaceous or a radiolarian ooze. For example, a radiolarian ooze occurs under the band of high produc- tivity along the equator in the Pacific. Some productive regions occur where the underlying water is relatively shallow. In these regions, the rate of calcium carbonate dissolution is much reduced, and foraminiferan and coccolithophorid oozes occur (e.g., along most of the Mid-Atlantic Ridge) because production by these plankters is greater than that by those producing silica shells. Biological oozes accumulate at a relatively rapid rate of centimeters per thousand years. Near continents, the supply of terrestrial particles overwhelms that of biological particles, and biological oozes do not form. Accumulation rates vary, but they are higher than for biological oozes. A substantial portion of the surface area of soft bottoms can be occupied by pebble- to cobble-sized manganese nodules. Manganese nodules are accretions of metals (mostly iron and manganese) that grow slowly (~1 mm per 10 000 y). They occur in a few regions of the deep Atlantic, but widely in the deep Pacific, particularly beneath the central gyres. At their most abundant, nodules can almost completely cover the surface of the seabed. Large-scale processes control sediment composition, so it tends to be uniform over hundreds of square kilo- meters. At the spatial scale at which most individual organisms experience their environment (millimeters to meters), the seafloor is made heterogeneous by two processes. The organisms themselves structure the seafloor by building tubes, tests, and mudballs in which to live (Fig. 2.6). These structures are used by other organisms as habitat (Thistle and Eckman, 1990). The second process is small-scale disturbance that creates patchiness in the deep-sea floor – in, for example, species composition, sediment texture, and food con- tent (Grassle and Sanders, 1973; Grassle and Morse- Porteous, 1987). Where they occur, manganese nodules Fig. 2.6. Some representative organism-constructed structures from deep-sea soft-bottom habitats. A. Empty test of the foraminiferan Oryctoderma sp., which is inhabited by a polychaete. B. and C. Foraminifers (the dashed line indicates the surface of the sediment). Scale lines equal 1.0 mm. Modified from Thistle (1979). Reproduced with permission of Plenum Press. impose a third type of small-scale heterogeneity on the surrounding soft bottom. Environmental variation in geologic time The preceding description of physical conditions in the deep sea applies to the modern ocean, but an understanding of modern deep-sea communities cannot be achieved without the incorporation of a historical perspective, because environmental changes at many time scales have helped to shape the present fauna. For example, since the early Eocene (~54 Ma BP)2 , deep- water temperatures have decreased from about 12ºC to their present values (Flower and Kennett, 1994) in four major cooling phases, in the early Middle Eocene, Late Eocene, Late Miocene, and Plio-Pleistocene (Lear et al., 2000). These abrupt temperature changes have been correlated with changes in the deep-sea fauna. For example, the sharp drop at the Eocene– Oligocene boundary (~38 Ma BP) is correlated with large changes in the benthic foraminifer (Kennett, 1982) and ostracod (Benson et al., 1984) assemblages. Within the Pliocene (2.85–2.40 Ma BP), bottom-water temperatures varied by 2ºC on a 40 000-yr time scale in the North Atlantic, as glaciers advanced and 2 1 Ma = 106 years.
- 26. THE DEEP-SEA FLOOR: AN OVERVIEW 11 retreated because of variation in the Earth’s axis of rotation. These temperature changes are correlated with changes in ostracod diversity (Cronin and Raymo, 1997). In the last 60 000 years, global warming and cooling cycles on a 1000-yr time scale are correlated with changes in foraminifer assemblages in the deep sea off California (Behl and Kennett, 1996). Summarizing, in much of the deep sea the variability in temperature, salinity, and oxygen over ecological time at a location is not important, and current velocities are nonerosive. In this sense, the deep-sea- floor environment is physically stable (Sanders, 1968). Even in regions with these physical characteristics, the sediment is heterogeneous at the millimeter-to- meter scale because of the modifications made by the organisms, small-scale disturbances, and manganese nodules. In contrast to these physically quiescent areas, some deep-sea locations experience erosive currents (Hollister and Nowell, 1991; Levin et al., 1994). OBTAINING INFORMATION ABOUT THE DEEP-SEA-FLOOR ECOSYSTEM By definition, 200 m or more of seawater separates deep-sea ecologists from the environment that they study. They, therefore, depend totally on technology to obtain information. Any shortcomings of their sampling devices must be understood, because defects can distort perceptions of the deep-sea-floor ecosystem. For example, the deep-sea floor was thought to be a species-poor environment until Hessler and Sanders (1967) showed that this erroneous view resulted from the inadequacies of older samplers. No single device can sample the entire size range of deep-sea organisms (from bacteria ~1 mm to fish 50 cm) quantitatively and efficiently. Fortunately, the sizes of deep-sea organisms are not spread evenly over this range but tend to fall into a small number of size classes (Mare, 1942; Schwinghamer, 1985; Table 2.1, Fig. 2.7). Sampling techniques have been developed for each. The size classes have the additional advantage that major taxa tend to occur primarily in a single size class, at least as adults. For example, polychaetes, bivalves, and isopods are macrofauna; nematodes and copepods are meiofauna. The technologies in current use differ in their suitability for the study of the various size classes. Table 2.1 Published size categories of deep-sea benthic organisms Category Lower size limit Sampler Representative taxa Megafauna centimeters trawls, photographs fishes, sea urchins Macrofauna 250–500 mm corers polychaetes, bivalves Meiofauna 32–62 mm corers nematodes, harpacticoids Microbiota microns corers protists Fig. 2.7. Size–abundance relationships in the benthos showing the gaps in the distribution that underlie the use of size classes. Equivalent spherical diameter is the diameter of a hypothetical sphere having a volume equal to that of the organism. Gray regions indicate the variability in the size-class boundaries used by different workers. Megafauna are those organisms that are visible in photographs of the seabed taken at more than about one meter off the bottom. Modified from Jumars (1993). Copyright 1993 by Oxford University Press, Inc. Used by permission of Oxford University Press, Inc. Cameras Cameras, mobile or stationary, are used to study the deep-sea-floor megafauna (Owen et al., 1967). Most deep-sea cameras use film, although video cameras and recorders are becoming more common. Because the deep sea is dark, a light source is paired with the camera. Circuitry to control the camera and light source and a source of power (batteries) complete the system. All components are housed in pressure-resistant cases. Megafaunal organisms (e.g., demersal fishes, brittle stars) are sparse, and some are highly mobile and can avoid capture by mechanical sampling devices (see below). Because mobile cameras can be used to survey kilometer-scale transects relatively unobtrusively (but
- 27. 12 David THISTLE see Koslow et al., 1995), they have been crucial in es- timating the abundance and biomass of such organisms and in discerning their distribution patterns (Hecker, 1994). For surveys, vertically oriented cameras have been suspended above the seabed from a ship’s trawl wire to photograph the seabed as the ship moves (Rowe and Menzies, 1969; Huggett, 1987). Cameras have also been mounted obliquely on towed sleds (Thiel, 1970; Rice et al., 1982; Hecker, 1990) and on research submarines (Grassle et al., 1975). Cameras have also been important in documenting the behavior of deep-sea megafauna, and in the dis- covery of rates of some deep-sea processes. For these purposes, cameras are mounted in frames (vertically or obliquely) and left for times ranging from hours to months, taking photographs at preset intervals (Paul et al., 1978). At the appropriate time, ballast weights are released, and the buoyant instrument package rises to the surface for recovery. This “free-vehicle” approach (Rowe and Sibuet, 1983) has been used, for example, to document the date of appearance of phytodetritus on the seafloor (Lampitt, 1985), the rates of mound-building by an echiurid (Smith et al., 1986), and megafaunal activity rates (Smith et al., 1993). Stationary cameras with bait placed in the field of view have been crucial to the discovery and study of food- parcel-attending species in the deep sea (Hessler et al., 1972). Cameras cannot provide information about smaller epibenthic organisms or organisms of any size that are inconspicuous or evasive or that live below the sediment-water interface and make no conspicuous indications of their presence on the sediment surface. Further, cameras return no specimens, so they are not useful for work that requires biological material such as physiological or taxonomic studies (but see Lauerman et al., 1996). Trawls, sledges, and sleds Some devices (trawls and sledges) have been used to collect megafauna. They consist of a mesh collecting bag and a means of keeping the mouth of the bag open (Fig. 2.8). A sledge has runners upon which the device rides; a trawl does not. Both are pulled along the seabed, collecting megafaunal invertebrates and fishes living on or very near the seabed. Smaller organisms are lost through the openings in the mesh. For some purposes, these devices have an advantage over cameras because they collect specimens, but they Fig. 2.8. Some deep-sea trawls (drawn roughly to scale). A, 3-m- wide Agassiz trawl; B, 6-m-wide beam trawl; C, a semiballoon otter trawl. Modified from Gage and Tyler (1991). Copyright: Cambridge University Press 1991. Reprinted with the permission of Cambridge University Press. sample much less area per unit time than cameras and fail to collect agile species that detect the approach of the device and escape. Much effort has been expended toward improving these samplers (Rice et al., 1982; Christiansen and Nuppenau, 1997), but the best that has been achieved is a device that collects all individuals of a few species, a constant proportion of others, and none or a varying proportion of others. The simultaneous use of camera and trawl or sledge surveys may be the best approach to quantification of the megafauna. The epibenthic sled (Hessler and Sanders, 1967) is a type of sledge designed to collect macrofauna from the sediment surface and from the top few centimeters of seabed (Fig. 2.9). The collecting bag has a smaller mesh than that used in a trawl or sledge. As a sled is towed along the seabed, an (adjustable) cutting blade slices under the upper layer of sediment, Fig. 2.9. The epibenthic sled used to collect large, non-quantitative samples of deep-sea infauna and epifauna. For scale, each runner is 2.3 m long by 0.3 m wide. The right-hand figure illustrates the operation of the sled. Modified from Hessler and Sanders (1967). Copyright: Elsevier Science.
- 28. THE DEEP-SEA FLOOR: AN OVERVIEW 13 which moves into the collecting bag. Sleds collect macrofauna in large numbers, supplying specimens for research in which properties of each individual must be determined – for example, studies of reproductive biology, biomass distribution, and taxonomy. Sleds do not collect every individual in their path in the layer to be sampled because the mouth of the bag clogs with sediment as the sled moves along the seabed (Gage, 1975), so sleds are inappropriate for quantitative studies. They can also damage delicate specimens (e.g., the legs of isopods tend to be broken off) and cannot sample macrofauna living at greater depths than 1– 2 cm. The deep-sea-floor ecosystem extends into the near- bottom water because some animals living in or on the seabed make excursions into the near-bottom water, and some animals living in the water just above the seabed interact with the seafloor. Hyperbenthic sledges (see also Rice et al., 1982) have been developed to sample the near-bottom water. Such sledges consist of runners and a frame supporting a vertical array of opening–closing nets (Dauvin et al., 1995; Fig. 2.10). A B C D E F Fig. 2.10. The hyperbenthic sled, a device for collecting deep-sea animals in the waters just above the seabed. The device is 1.51 m tall. A, Attachment point for the cable to the ship; B, frame; C, mouth of a sampler; D, net; E, sample container; F, runner. Modified from Dauvin et al. (1995). Copyright: Elsevier Science. The usual limitations of plankton nets apply to these samplers (e.g., bias in collections owing to differences in avoidance behavior among species, variable filtering efficiency resulting from net clogging). In addition, the frame may put animals from the seabed into suspension and thus cause them to be caught, particularly in the lowest net. Despite their limitations, these samplers provide access to an understudied component of the deep-sea fauna (see also Wishner, 1980). Despite their limitations, most of the taxonomic, systematic, and biogeographic research on the deep- sea fauna has been based on the large collections that trawls, sledges, and sleds provide (Hessler, 1970). This research has resulted in discoveries regarding, for example, the high diversity of the deep-sea-floor fauna (Hessler and Sanders, 1967) and the systematics and phylogeny of major invertebrate groups (Wilson, 1987). Also, such samples taken repeatedly from the same area have provided information on temporal phenomena, in particular reproductive periodicity in the deep sea (Rokop, 1974; Tyler et al., 1982). Corers Corers are used to sample macrofauna, meiofauna, and microbiota. Two types are presently in common use. Box corers, in particular the USNEL-Sandia 0.25-m2 box corer (Hessler and Jumars, 1974; Fig. 2.11), are Fig. 2.11. An advanced version (Hessler–Sandia) of the USNEL box corer (shown in the closed position), a device for collecting quantitative samples of deep-sea macrofauna. The width of the sample box is 0.5 m. A, The detachable spade; B, vent flaps in the open position for descent; C, vent flaps in the closed position for ascent; D, cable to the ship. Some details omitted. Modified from Fleeger et al. (1988). lowered on a ship’s trawl wire. About 100 m above bottom, the rate of descent is slowed to 15 m min−1 until the corer penetrates the bottom. This relatively high entry speed is necessary to minimize multiple touches and pretripping. As the corer is pulled out of the seabed, the top and bottom of the sample box are closed. The advantages of a box corer are that it takes a sample of known area to a depth (20 cm) that encompasses the bulk of the vertical distribution of deep-sea organisms. Box corers are not strictly quantitative. They occa- sionally collect megafaunal individuals, but megafauna are too rare to be effectively sampled. Further, the
- 29. 14 David THISTLE pressure wave that precedes the corer (even in the most advanced designs only about 50% of the area above the sample box is open) displaces material of low mass (e.g., the flocculent layer, phytodetritus; Jumars, 1975; Smith et al., 1996), if any is present (Thistle and Sherman, 1985). Therefore, box-corer samples usually underestimate abundances of organisms that live at the sediment surface or in the upper millimeters. The bias becomes worse as animals decrease in size and mass (see Bett et al., 1994). Deliberate corers (Craib, 1965; Fig. 2.12) are alter- Fig. 2.12. Scottish Marine Biological Laboratory multiple corer, a device for collecting quantitative samples of deep-sea meiofauna, phytodetritus, and other materials that would be displaced by the pressure wave preceding a box corer. A, Sampling tubes; B, supporting frame; C, hydraulic damper; D, cable to the ship. Some details omitted. Modified from Barnett et al. (1984). Copyright: Elsevier Science. natives to box corers. These devices consist of a frame, one or more samplers carried on a weighted coring head hanging from a water-filled hydraulic damper, and mechanisms to close the top and bottom of the sampler(s) during recovery (Soutar and Crill, 1977; Barnett et al., 1984). The corer is lowered on the ship’s trawl wire. At the seabed, the frame takes the weight of the coring head. When the wire slackens, the hydraulic damper allows the coring head to descend slowly, which forces the sampler(s) into the seabed. As a consequence, the pressure wave is minimal. When the trawl wire begins to wind in, the coring head rises, allowing the top and bottom closures to seal the sampler(s). The advantage of deliberate corers is that they can sample quantitatively material that would be displaced by the bow wave of a box corer (Barnett et al., 1984). The disadvantage is that the surface area sampled tends to be smaller; also, stiff sediments are not penetrated as well as when box corers are used. Thus, despite the superior sampling properties of deliberate corers (Bett et al., 1994; Shirayama and Fukushima, 1995), box corers are still used because, for some taxa (e.g., polychaetes) in some environments (e.g., areas of the abyss with low standing stocks), deliberate corers collect too few individuals to be useful. Corers have also been developed for use with research submarines and remotely operated vehi- cles (ROVs). Tube corers are plastic cylinders (~34 cm2 in cross section), each fitted with a removable head that carries a flapper valve and a handle by which the sampler is gripped. To sample, the mechanical arm of the research submarine or ROV presses the corer into the seabed. The corer is then removed from the seabed and transferred to a carrier that seals its bottom. With this method of coring, samples can be taken from precisely predetermined locations, allowing the sampling of particular features or previously emplaced experimental treatments (Thistle and Eckman, 1990). Even though these corers enter the seabed slowly, the water in the corer tube must be displaced for the sediment to enter, so that there is a bow wave, but its effect has not yet been measured. Also, because the bottom of the corer is not sealed during the transfer to the carrier, these cores can only be used in deposits where the subsurface sediment seals the corer, i.e., cohesive muds. Modified Ekman corers are also commonly used by research submarines and ROVs. These corers consist of a metal box of surface area typically between 225 cm2 and 400 cm2 , with a handle for a mechanical arm to grasp and with mechanisms to close the top and bottom after a sample has been taken. These corers have the advantages that they can be deliberately positioned; they take larger samples than do tube corers; and, because they are sealed at the bottom as the sample is taken, they can be used in fluid muds or in sands. A disadvantage is that they sample a much smaller area than a box corer because of handling and payload constraints on their size. Also, despite the low speed at which they are inserted into the seabed, light surface
- 30. THE DEEP-SEA FLOOR: AN OVERVIEW 15 material can be displaced from the periphery of the sample (Eckman and Thistle, 1988). A variety of corers have been used historically to sample macrofauna, meiofauna, and microbiota (gravity corers, Smith–McIntyre grabs). The sampling properties of these devices were not as good as those of the box corer, deliberate corers, or submarine/ROV samplers (see below). In particular, the bow wave was more severe. Therefore, the data obtained with such samplers must be interpreted with caution. Finally, the collection of subsurface megafauna remains an unsolved problem, but acoustical approaches (Jumars et al., 1996) seem likely to be useful for some types of measurements. Research submarines and remotely operated vehicles (ROVs) A research submarine is comparable in size to a delivery truck. Those in service typically carry a pilot and one or two scientists in a pressure sphere about 2 m in diameter. Surrounding the sphere is equipment for life support, propulsion, ascent and descent, and scientific purposes (manipulator arms, cameras, specialized payload in a carrying basket) (Heirtzler and Grassle, 1976). Research submarines bring the ecologist into the deep sea and thereby confer large benefits by correcting the tunnel vision that deep-sea scientists acquire from the study of deep- sea photographs. Further, research submarines permit a wide range of ecological experiments. For example, trays of defaunated sediment have been placed on the seabed for study of colonization rates (Snelgrove et al., 1992), and dyed sediment has been spread and subsequently sampled for estimates of sediment mixing rates (Levin et al., 1994). Research submarines have limitations. For example, positioning the vehicle and then removing the device to be used (e.g., a corer) from its carrier, performing the task, and returning the device to its carrier require a substantial amount of time, so relatively few tasks can be done during a dive. Also, because the vehicle is large, maneuvering can be awkward, and experiments are occasionally run over and ruined. Because of their cost, few research submarines are in service, so dives are rare. Much more research needs to be done than can be accommodated. Remotely operated vehicles (ROVs) are self-propelled instrument packages. Some operate at the end of a cable that provides power and hosts a two-way communications link; others are untethered, carrying their own power and recording images and data. The instrument package consists of a propulsion unit, sensors (particularly television), and, in some cases, manipulator arms. Some ROVs are designed to “fly” over the seabed. These ROVs tend to be used for large-scale surveys, but some can be maneuvered with precision and can inspect or sample centimeter-scale targets (e.g., the MBARI ROV: Etchemendy and Davis, 1991). Other ROVs are bottom crawlers (e.g., the Remote Underwater Manipulator: Thiel and Hessler, 1974) and are more suitable for seabed sampling and experimentation. The great advantage that ROVs have over research submarines is endurance. Because the investigators are on the support ship rather than in the vehicle, the ROV does not have to be recovered each day to change crew as does a research submarine. The time savings result in far more ROV bottom time than research submarine bottom time for each day at sea. Limitations of ROVs include slow sampling and cumbersome maneuvering. Also, there are substantial benefits to allowing deep-sea scientists to come as close as possible to experiencing the deep-sea environment. Scientists who have made dives relate how their conception of the deep sea was substantially changed by the experience, improving their science. Sensors Knowledge of the chemical milieu in which deep- sea-floor organisms live has increased markedly since the introduction of microelectrode sensors. These devices measure chemical parameters (oxygen, pH) with a vertical resolution measured in millimeters. Early measurements were made on recovered cores, but free-vehicle technologies have been developed so that measurements can be made in situ (see Reimers, 1987). Other technologies The devices discussed above are those that are in common use. Many other devices have resulted in important work but have not become common (see Rowe and Sibuet, 1983). It is beyond the scope of this chapter to present all these devices, but two are conspicuous. The free-vehicle respirometer (Smith et al., 1976), which measures oxygen utilization by the benthic community, has been important in
- 31. 16 David THISTLE studies of deep-sea community energetics, which have implications for global carbon cycling. Free-vehicle traps have been crucial to the study of food-parcel- attending species in the deep sea (Hessler et al., 1978). Costs and benefits Good techniques are available with which to sample, and reasonable techniques are available with which to do experiments in the deep sea, but the expense is substantial. Both sampling and experimentation require the use of large, and therefore expensive, ships. Research submarines and ROV’s add additional costs. For soft bottoms, separating the animals from the sediment and identifying the diverse fauna (Grassle and Maciolek, 1992) are time-consuming, so sample processing is costly. These expenses are among the reasons why relatively few data have been collected from this vast ecosystem and why few ecological experiments have been performed. Despite these costs, scientists persist in the study of the deep sea, and their research provides a va- riety benefits for society. For example, research on hydrothermal-vent animals led to the discovery of DNA polymerases that work at high temperatures, which are crucial tools in pure and applied molecular biology. Safe repositories for human waste, such as dredge spoils, sewage sludge, industrial waste, and radioactive materials, are needed. Ongoing ecological work will help determine whether wastes dumped in the deep sea make their way back into contact with humans, and the effects of these wastes on the functioning of natural ecosystems in the ocean (Van Dover et al., 1992). The deep-sea floor contains mineral resources; for example, economically important amounts of cobalt and nickel occur in manganese nodules. The work of deep-sea ecologists is helping to determine the environmental consequences of deep-ocean mining (Ozturgut et al., 1981). More generally, the deep-sea benthos provides critical ecological services (e.g., recycling of organic matter to nutrients: Snelgrove et al., 1997). THE SOFT-BOTTOM FAUNA OF THE DEEP-SEA FLOOR Taxonomic composition At high taxonomic levels (i.e., phylum, class, and order), the soft-bottom, deep-sea fauna is similar to that of shallow-water soft bottoms (Hessler, 1974; Gage, 1978). For example, the megafauna consists primarily of demersal fishes, sea cucumbers, star fishes, brittle stars, and sea anemones. The macrofauna consists primarily of polychaetes, bivalve mollusks, and isopod, amphipod, and tanaid crustaceans. The meiofauna consists of primarily of foraminifers, nematodes, and harpacticoid copepods. At lower taxonomic levels (family and below), however, the similarities disappear. In particular, the species that live in the deep sea are not, in general, found in shallow water. Gage and Tyler (1991) have reviewed the natural history of deep-sea taxa. Many taxa that have large numbers of species in shallow water have a few members that penetrate into the deep sea. For example, of 300 stomatopod (mantis shrimp) species, only 14 occur below 300 m (Manning and Struhsaker, 1976). The decapod crustacean fauna in shallow water (200 m) consists of more than 200 species, predominantly brachyuran crabs. Below a depth of 1500 m, there are fewer than 40 species of decapods, and brachyurans make up ~10% of this total. The proportion of the bivalve mollusks that are eulamellibranchs decreases as well (Sanders et al., 1965). In contrast, the proportion of isopod species that are asellotes (Hessler and Wilson, 1983) and of bivalve mollusk species that are protobranchs increases with depth. Finally, some taxa inhabit the deep sea exclusively; for instance, the protist group of xenophyophores have not been found above ~500 m (Tendal, 1996). Variation of biomass and numbers with depth The general pattern of the distribution of the biomass of organisms on the deep-sea floor is known (Fig. 2.13) and appears to be controlled by the rate at which food is supplied to the seabed (Rowe, 1971). The basic pattern is set by the productivity of the surface waters. For example, primary productivity is highest nearest the continents, and the deep-sea floor near continents tends to have the highest biomass. The depth of the overlying water modifies this pattern. As food particles sink, a portion of each particle is lost to decay, and some particles are consumed by mid- water organisms. The deeper the water, the longer it takes particles to reach the seabed, and the greater the loss from these processes. Given two regions with identical primary productivities in the overlying water, the deeper location will have the lesser food input
- 32. THE DEEP-SEA FLOOR: AN OVERVIEW 17 Fig. 2.13. The correspondence between water-column primary production and deep-sea benthic biomass. A, Distribution of benthic biomass (g wet weight m−2) in the Pacific; B, zones of primary productivity in the Pacific. Values 1–4 are 100, 100–150, 150–250, and 250– 650 mg C m−2 d−1, respectively. Modified from Hessler (1974). Reproduced by permission of the Oregon State University Press. Fig. 2.14. Biomass from 709 deep-sea quantitative samples plotted against depth showing the logarithmic decline in biomass with increasing depth. Modified from Rowe (1983), who gives the sources of the data. Numerals in the figure indicate the number of co- occurring points. Copyright: 1983, John Wiley and Sons. Reprinted by permission of John Wiley and Sons, Inc. and, therefore, the smaller benthic biomass. Food is also supplied to the deep-sea benthos as organic debris (e.g., pieces of seagrass and macroalgae) that moves across the seabed from shallow to deep water (Carey, 1981). This phenomenon appears to explain why those trenches that are near continents have higher biomasses than would be expected for their depths (Belyaev, 1989). The trench traps and concentrates the organic debris that would otherwise be spread over a wider area (Rowe, 1983). Because shallow water is adjacent to the continents in most regions, higher productivity, shorter food- particle settling times, and larger seabed fluxes of organic debris are all correlated, resulting in the generalization that the abundance of life in the deep sea decreases with depth and distance from a major land mass (Murray, 1895). More simply, the biomasses of megafauna (Lampitt et al., 1986), macrofauna (Rowe, 1983; Fig. 2.14), and meiofauna (Shirayama, 1984; Tietjen, 1992) decrease as depth increases. The number of animals per unit area of sea floor also decreases with depth (Hessler, 1974; Thiel, 1979). For example, macrofaunal abundance decreases significantly with depth in the Gulf of Mexico (Rowe and Menzel, 1971) and in the northwest Atlantic (Rowe et al., 1982), as does that of meiofauna in the western Pacific (Shirayama, 1984) and Mediterranean (de Bovée et al., 1990; see also Thiel, 1979). Trophic composition of the deep-sea-floor fauna Ecologists often find it useful to combine species into groups whose members are similar in selected attributes, to facilitate the search for generalizations
- 33. 18 David THISTLE (Jumars and Fauchald, 1977; Sokolova, 1997). Deep- sea workers have frequently grouped species by feeding mode. This approach has led to interesting results, but few direct observations of the feeding of deep- sea species have been made. Although some gut- content studies have been done (Sokolova, 1994), most inferences about how a deep-sea species feeds have been based on knowledge of the feeding of its shallow- water relatives. Deposit feeders A deposit feeder ingests sediment. During gut passage, the animal converts a portion of the organic material contained in the sediment into a form that can be assimilated. Deposit feeding is the dominant feeding mode in the deep sea (Thiel, 1979). For example, at an oligotrophic site in the abyssal Pacific, 93% of the macrofauna were deposit feeders (Hessler and Jumars, 1974; see also Flach and Heip, 1996). The dominance of deposit feeding may arise because the rain of organic material into the deep sea consists primarily of small particles of little food value. Deposit feeders apparently can collect and process this material profitably despite the costs of manipulating the mineral grains that they simultaneously ingest. Adaptations to deep-sea deposit feeding include an increase in gut volume (Allen and Sanders, 1966). The larger volume is thought to allow the rate of sediment processing to increase without a decrease in gut residence time or to allow gut residence time to increase without a decrease in the rate of sediment processing (Jumars and Wheatcroft, 1989). Either adjustment would increase the rate of food assimilation by organisms feeding on the relatively food-poor deep- sea sediment as compared with that which could be achieved with the gut morphology of a closely related shallow-water species. Deposit feeders can be grouped by the sediment horizon at which they feed and by their mobility (Jumars and Fauchald, 1977). Sessile surface-deposit feeders remain in a fixed location and feed from the sediment surface. Discretely motile surface-deposit feeders move infrequently but must be stationary to feed efficiently (echiuran worms: Ohta, 1984; Bett and Rice, 1993). Both sessile and discretely motile surface deposit feeders extend structures (a proboscis, palps, tentacles) over the sediment surface to collect material. Motile surface-deposit feeders (holothurians such as Scotoplanes globosa) ingest sediment as they move over the sediment surface. Subsurface deposit feeders tend to be motile and feed as they burrow through the sediment. Among deposit feeders, some ecologically inter- esting patterns have been observed. The decrease in the average size of macrofaunal deposit feeders as depth increases (and the rate at which food reaches the deep-sea floor decreases) was described above. In addition, as depth increases from about 400 m to that of the abyss, the proportion of sessile forms among deposit-feeding polychaetes decreases (Jumars and Fauchald, 1977; see also Rowe et al., 1982). Jumars and Fauchald (1977) suggested that this pattern could arise if the maximum feeding radius of sessile surface-deposit feeders were fixed (e.g., because of mechanical limitations to the length of polychaete tentacles). Therefore, as food flux decreases, fewer sessile deposit feeders are able to reach a large enough area to survive. Because the foraging areas of motile polychaete deposit feeders do not have such mechanical limits, they would not be as much affected by the decrease in food flux. The rules can be different in areas that experience strong near-bottom flows. For example, at such a site at a depth at which sessile deposit-feeding polychaetes should be rare, the dominant polychaete is a sessile deposit feeder (Thistle et al., 1985). This species digs a pit around itself approximately 1 cm deep and 4 cm in diameter. As the near-bottom flow encounters the pit, the streamlines of the flow expand and its speed decreases (by the principle of continuity: Vogel, 1981). When the speed of the flow decreases, its capacity to transport particles (including food particles) is reduced, which increases the flux of food particles to the bed. The worm harvests these particles (Nowell et al., 1984) and thus can occur in large numbers at a depth where sessile feeding on deposits would not be expected to function well. Exploiters of large food parcels Not all of the food that enters the deep sea does so as small particles of little food value. For example, the carcasses of fishes and whales reach the sea floor. These high-quality food parcels are rare (Smith et al., 1989) but attract a subset of the fauna. These “parcel-attending species” include necrophages, which consume the carcass directly, and species that benefit indirectly from the food fall. The parcel attenders include certain species of demersal fishes (Dayton and Hessler, 1972; Smith, 1985), amphipods of the family Lysianassidae (Hessler et al., 1978; Thurston, 1979),
- 34. THE DEEP-SEA FLOOR: AN OVERVIEW 19 decapod shrimps (Thurston et al., 1995), gastropods (Tamburri and Barry, 1999), and brittle stars (Smith, 1985). Whether any species depends exclusively on car- casses has not yet been shown (Jumars and Gallagher, 1982; Ingram and Hessler, 1983), but some omnivorous species include carcasses adventitiously in their diets (Smith, 1985; Priede et al., 1991). The response of the parcel attenders to carcasses placed on the seafloor has revealed much about their ecology. Minutes to hours after a bait parcel is placed on the seafloor, swimming parcel attenders begin to arrive; nonswimmers arrive more slowly. Both approach predominantly from down current (Dayton and Hessler, 1972; Thurston, 1979; Smith, 1985), attracted by a current-borne cue, probably odor (Sainte- Marie, 1992). These animals feed voraciously until their guts are full. Satiated individuals leave the carcass but remain in the vicinity, perhaps to optimize digestive efficiency (Smith and Baldwin, 1982) or to return to the carcass after the gut is partially emptied (Smith, 1985). At peak abundance around a fish carcass, tens of fishes, hundreds of amphipods, and hundreds of brittle stars may be present (although these peaks are not simultaneous) (Smith, 1985). These abundances are many times greater than abundances in the background community, so carcasses cause local concentrations of individuals. As the amount of flesh decreases, the parcel attenders disperse. Some species depart while some flesh remains; others remain weeks after the flesh has been consumed (Smith, 1985). Dispersal distances may be a few meters for walkers, such as brittle stars; but Priede et al. (1990) have shown that food-parcel-attending fishes disperse more than 500 m. Of the parcel attenders, amphipods are best known biologically (but see Tamburri and Barry, 1999, for other taxa). According to Smith and Baldwin (1982), these crustaceans survive the long periods between food parcels by greatly reducing their metabolic rate while retaining an acute sensitivity to the arrival of carcasses at the seafloor. When they detect the odor from a carcass, they rapidly increase their metabolic rate and begin a period of sustained swimming toward the bait. To maximize consumption at the food parcel, they feed rapidly, filling their extensible guts. At satiation, the gut fills most of the exoskeleton, which can be greatly distended (Shulenberger and Hessler, 1974; Dahl, 1979). The ingested material is rapidly digested (95% in 1–10 days), making space in the gut for more food and increasing the flexibility of the body for swimming (Hargrave et al., 1995). Younger stages need to feed more frequently than adults, but all can survive for months between meals (Hargrave et al., 1994). Differences in behavior and morphology suggest that groups of parcel attenders have different strategies. For example, some parcel-attending amphipods have shearing mandibles. They consume bait rapidly and probably combine scavenging and carnivory in their feeding strategy. Other parcel-attending amphipods have triturating mandibles and combine scavenging with detritivory (Sainte-Marie, 1992). Jones et al. (1998) have reported that the former arrive first at the carcass and are replaced by the latter over time. Ingram and Hessler (1983) found that the populations of three species of small-bodied, parcel-attending amphipods were concentrated about 1 m above the bottom and that the population of a larger-bodied species was concentrated about 50 m above the bottom. Turbulent mixing in the bottom boundary layer causes the chemical signal from a carcass to widen and to increase in vertical extent with increasing distance from a carcass, while it simultaneously decreases in concentration. Ingram and Hessler (1983) therefore suggested that the two groups of species exploited the carcass resource differently. The high-hovering species surveys a wide area and detects primarily large carcasses. The low-hovering species detect the full range of carcass sizes but from a smaller area. These ideas are suggestive, but depend on the untested assumptions that carcasses produce chemical signals in proportion to their sizes, and that the threshold concentrations at which a signal can be detected are approximately the same for the two guilds (Jumars and Gallagher, 1982). Also, differences between guilds in swimming speed and ability to sequester food are likely to be necessary to explain why the optimal foraging height for the small-bodied species is lower than that for the large-bodied species (see also Sainte-Marie, 1992). After leaving the carcass, necrophages transfer calories and nutrients to other deep-sea soft-bottom organisms by defecating (Dayton and Hessler, 1972). Smith (1985) estimated that about 3% of the energy required by a bathyal benthic community can be provided in this way (see also Stockton and DeLaca, 1982). The concentration of potential prey that a carcass attracts may itself be a resource. Jones et al. (1998) reported that none of the fish species attending cetacean carcasses that they placed in the abyssal Atlantic
- 35. 20 David THISTLE consumed the carcass. Rather, they preyed on the parcel-attending amphipods. Suspension feeders Suspension feeders (Fig. 2.15) feed on material they collect from the water column, intercepting epibenthic plankton, particles raining from above, and particles that have been resuspended from the seabed. The food particles captured vary in size from microns to millimeters, depending on the suspension feeder. The smaller particles include bacteria, pieces of organic matter, microalgae, and silt- and clay-sized sediment particles with microbial colonies. Larger particles include invertebrate larvae and the organic aggregates known as “marine snow” (Shimeta and Jumars, 1991). Examples of suspension feeders on deep-sea soft bottoms are sea anemones (Aldred et al., 1979), sea pens (Rice et al., 1992), sponges (Rice et al., 1990), and stalked barnacles (personal observation). Fig. 2.15. Representative suspension feeders. A, Glass sponge; B, horny coral. Modified from Gage and Tyler (1991). Cambridge University Press 1991. Reprinted with the permission of Cambridge University Press. Particles can be collected from seawater in five basic ways (Levinton, 1982). In mucous-sheet feeding, an animal secretes a mucous sheet that particles encounter and stick to, which the animal (e.g., members of the polychaete genus Chaetopterus) collects and consumes. In ciliary-mucus feeding, the feeding current passes over rows of mucus-covered cilia. The mucus and the embedded particles are moved by the cilia to the mouth. This approach to suspension feeding is used by ascidians (Monniot, 1979), sabellid polychaetes, brachiopods, bryozoans, and some bivalve mollusks (Levinton, 1982). In setose suspension feeding, a limb is drawn through the water, and suspended particles are captured by setae on the limb. The collected particles are scraped from the limb and transferred to the mouth. Suspension-feeding crustaceans feed in this manner, in particular barnacles and suspension- feeding amphipods. In sponges, water enters through pores and is drawn along internal canals to flagellated chambers by the pumping action of the flagellated cells. The entrained particles encounter the collars of the flagellated cells. Particles that are retained are phago- cytized or transferred to phagocytic amebocytes, where digestion also occurs (Barnes, 1987). In suspension- feeding by foraminifers (e.g., Rupertina stabilis: Lutze and Altenbach, 1988), suspended particles encounter and stick to pseudopodia extended into the near-bottom water. Active suspension feeders expend energy to cause water to flow over their feeding structures; for example, barnacles move their cirri through the water, and sponges pump water over the collars of their flagellated cells. Passive suspension feeders – for example, some foraminifers, crinoids, some ophiuroids, some holothurians, some octocorals, and some ascidians – depend on external flows to move water over their feeding structures. For both active and passive suspen- sion feeders, the rate of particle capture (and to a first approximation their rate of energy acquisition) depends on the product of the flow rate over their feeding apparatus and the concentration of food particles in the filtered water (= the particle flux). Passive suspension feeders depend on the local particle flux, whereas active suspension feeders depend only on the local particle concentration because they control the speed of the flow over their feeding apparatus (Cahalan et al., 1989). For a passive suspension feeder to survive at a location, the particle flux must be sufficient to meet its metabolic requirements; thus, not all locations in the deep sea are suitable. Rather, the interaction of local flow with topography will create a finite number of appropriate sites. Because both average particle concentration and average flow velocity decrease with depth, the number of sites suitable for passive suspen- sion feeders decreases with depth. Similarly, suspended particle concentration varies locally, so only a finite number of sites will be suitable for active suspension feeders, and this number will decrease with depth as the suspended-particle concentration decreases. For passive suspension feeders, the minimum particle concentra- tion for survival can be lower than for active suspension feeders because the animal expends no energy filtering; and, up to some limit, more rapid ambient flow can increase the effective concentration for passive
- 36. THE DEEP-SEA FLOOR: AN OVERVIEW 21 but not for active suspension feeders. Therefore, the number of suitable locations (and therefore abundance) should decrease more rapidly with depth for active suspension feeders than for passive suspension feeders. This pattern has been observed (Jumars and Gallagher, 1982). Given the low suspended-particle concentrations in the deep sea, maximizing the particle-capture rate may be particularly important. In particular, passive suspen- sion feeders should orient their collecting surfaces to maximize the flux of particles that they intercept. Data from the deep sea with which to test this prediction are sparse, but some types of behavior are suggestive. For example, the sea anemone Sicyonis tuberculata bends its body in such a way that its feeding surface faces into the current as the current direction rotates with the tide (Lampitt and Paterson, 1987). Also, under the West-African upwelling, the vertical flux of food particles is large and near-bottom currents are slow, so the vertical flux of food particles greatly exceeds the horizontal flux. There, the passive suspension-feeding sea anemone Actinoscyphia aurelia orients its collector upwards, as expected (Aldred et al., 1979). Some passive suspension feeders increase particle capture rates by exploiting the increase in horizontal speed of the near-bottom water as distance from the seabed increases. For example, the deep-sea foraminifer Miliolinella subrotunda builds a pedestal 1–6 mm tall on which it perches to suspension-feed (Altenbach et al., 1993). Other passive suspension feeders occur on topographic features or the stalks of other organisms, such as glass sponges, thus placing their feeding apparatus in regions of more rapid flow. A sea anemone moved ~30 cm up the side of an experimental cage in ~5 days to perch at the highest point (personal observation). Some shallow-water polychaetes can switch feeding modes (Taghon et al., 1980; Dauer et al., 1981). When the flux of suspended particles is large enough, these species suspension-feed. When it is not, they deposit-feed. Many deep-sea polychaetes are thought to have this capability (G. Paterson, personal communica- tion, 1997). Carnivores/predators Carnivores select and consume living prey. For example, in the deep sea, kinorhynchs have been found with their heads embedded in the sides of nematodes (personal observation). Such direct evidence of feeding on live prey is difficult to obtain from the deep sea. Gut-content analysis, both by visual inspection (Langer et al., 1995) and by immunological methods (Feller et al., 1985), has been used; but this approach cannot always distinguish carnivores from scavengers. As a result, feeding mode is often inferred from the feeding patterns of similar, shallow-water species. For example, a group of deep-sea nematodes with teeth in their buccal cavities (Fig. 2.16) are thought to be carnivores because shallow-water species with such armature are carnivorous (Jensen, 1992). The proportion of the deep-sea fauna that is carnivorous is not well known. Jumars and Gallagher (1982) estimated that carnivores constituted between 2% and 13% of the polychaetes at four Pacific sites. Tselepides and Eleftheriou (1992) reported that 49–52% of the polychaetes between 700 and 1000 m depth off Crete were carnivorous. Fig. 2.16. Examples of deep-sea nematodes that are thought to be carnivores because their buccal cavities have teeth as do carnivorous nematodes in shallow water. Only the anterior portion of each worm is shown. Modified from Jensen (1992). Reproduced by permission of the Station Biologique de Roscoff. In the food-poor deep sea, prey are rare, so the time between encounters with prey will be long compared to that needed to subdue and ingest a prey item once encountered. Under these circumstances, optimal- foraging theory predicts that diets should be gener- alized to shorten the time between prey encounters, increasing the food-acquisition rate (MacArthur, 1972). The step from feeding on live prey to including carrion in the diet is a small one, so organisms that might be predators in shallow water are likely to consume both
- 37. 22 David THISTLE Fig. 2.17. The proportion of asteroid feeding types at increasing depth, showing the shift to omnivory as depth increases. Modified from Carey (1972). Reproduced by permission of Ophelia Publications. living and dead material in the deep sea. For example, Carey (1972) reports a trend for the proportion of predaceous asteroids to decrease and the proportion of omnivorous asteroids to increase with increasing depth in the deep sea (Fig. 2.17). Although prey are rare, they may be more detectable in the deep sea than in shallow water. Flow in the benthic boundary layer is slower and more orderly in the deep sea than in shallow water, so chemical gradients should be more persistent and provide better information for prey location. Also, pressure waves produced by prey (Ockelmann and Vahl, 1970) should be more easily detected in the deep-sea benthic bound- ary layer because of its lower turbulence. Background acoustic noise is also lower in the deep sea, making weak acoustic signals produced by prey relatively easy to detect. These physical attributes of the deep sea also facilitate the transmission of information to the prey about the approach of a predator, so that sensory capabilities of the prey may be evolving in parallel with those of the predators (Jumars and Gallagher, 1982). The general decrease in food input with increasing depth in the deep sea appears to affect predators dis- proportionately. For example, Rex et al. (1990) found that abundance of predaceous gastropods decreased at a greater rate with depth than did that of deposit-feeding gastropods (Fig. 2.18). One possible explanation for this pattern is that, as the distances between prey increase with depth, the energy spent in searching 0 1 2 3 0 1000 2000 3000 4000 Depth (m) Number per 0.09 m 2 Neogastropoda Opisthobranchia Mesogastropoda Archaeogastropoda Fig. 2.18. Logarithm of abundance of major taxonomic groups of gastropods at different depths in the North Atlantic, showing that predators decline more rapidly than do deposit feeders. Neogastropoda and Opisthobranchia are predators. Modified from Rex et al. (1990). Copyright: Elsevier Science. increases, but the energetic return per prey item found remains the same. Therefore, as depth increases, fewer gastropod species (and perhaps fewer species of other taxa) can make an energetic profit as predators. Croppers In the food-poor deep sea, there should be strong selection to digest and assimilate any organic material encountered, living or dead. Dayton and Hessler (1972) proposed the term “cropper” for an animal that ingests live prey, whether exclusively or in combination with dead prey or inorganic materials. Deep-sea croppers include species of holothurians, echinoids, ophiuroids, asteroids, cephalopods, and some polychaetes, de- capods, and demersal fishes. Most deposit feeders in the deep sea are croppers because they feed on living and dead material. Given the large proportion of deposit feeders among deep-sea-floor animals, much of the living prey may be consumed by deposit feeders (Dayton and Hessler, 1972). There are several corollaries of this view. Deposit feeders vary in size from fishes to nematodes. Given that food is in short supply, the size of the prey ingested should be limited only by the size of the deposit feeder’s mouth. Therefore, the smaller the prey organism, the greater its predation risk because the number of mouths large enough to ingest it increases as its size decreases. Thus larvae, juveniles, and meiofauna of all life stages should experience more intense predation than do macrofaunal and megafaunal adults. This increase in predation pressure with decreasing size should decrease the probability of competitive exclusion among smaller animals, and allow larger overlaps in their utilization of resources. In particular, as an animal’s size decreases,
- 38. THE DEEP-SEA FLOOR: AN OVERVIEW 23 its diet should become increasingly broad (Dayton and Hessler, 1972). Finally, the environment of the deep sea is less physically variable than that in shallow water and is likely to impose less mortality on deep-sea organisms than the physical environment of shallow water imposes on shallow-water organisms. As a consequence, the mortality imposed by croppers may be crucial to the organization of deep-sea communities (Jumars and Gallagher, 1982). Size structure The size of the average macrofaunal individual de- creases with increasing depth in the deep sea (Fig. 2.19). 0 20 40 0 20 40 60 80 0 40 60 80 0 1.00 0.50 0.25 0.13 0.06 20 40 Mesh size (mm) Percent of total biomass A. 3 m B. 295 m C. 2000 m Fig. 2.19. Macrofaunal biomass of different size groups in three depth zones, showing the decrease in average size with depth. Note that size decreases from left to right. Modified from Shirayama and Horikoshi (1989). Reproduced with the permission of Wiley-VCH Verlag. An early indication of this pattern was that workers who wished to retain the individuals of macrofaunal taxa from deep-sea samples quantitatively had to use sieves with smaller mesh openings than would be necessary to collect those taxa in shallow water (Sanders et al., 1965; see Table II in Rowe, 1983). More formally, Rowe and Menzel (1971) showed that the proportional decrease in macrofaunal biomass with depth was greater than the proportional decrease in macrofaunal numbers in the Gulf of Mexico, a result that has been found in other areas (Shirayama and Horikoshi, 1989; but see Polloni et al., 1979) and for the meiofauna (Soetaert and Heip, 1989). Direct measurements of animal sizes along depth gradients reinforce these conclusions. For example, Soltwedel et al. (1996) reported that the length of nematodes declined with depth; Vanaverbeke et al. (1997) found that nematode biomass per individual decreased with depth. For macrofauna, the miniaturization appears to occur by species replacement rather than by decreases in the average body size within species (Gage, 1978). Hessler and Jumars (1974) have presented a hypoth- esis to explain the decrease in the size of macrofauna with depth (see also Thiel, 1975; Gage, 1977). Food arrives at the deep-sea floor from above. As food supply decreases with increasing depth, fewer animals are present per unit area, so food should not be mixed as deeply by bioturbation. Therefore, the layer in which the food is concentrated at the sediment surface becomes increasingly thin. For organisms that feed by ingesting sediment, Hessler and Jumars (1974) argued that those with small mouths should be best at restricting their ingestion to the food-rich layer, minimizing the costs of feeding by reducing the amount of food-poor sediment inadvertently ingested. On the assumption that mouth size and body size are correlated, the size of successful deposit feeders should decrease as food supply decreases with depth. Because the bulk of the macrofauna in the deep sea are deposit feeders, this explanation could account for the decrease in their average size. Suspension feeders also decrease in size with increasing depth. Ascidians (sea squirts) decrease from 1 cm to 2.5 mm in size and by a factor of 25 in mass from the upper slope to the abyssal plain (Monniot, 1979). Monniot (1979) argued that the dwarfing of these active suspension feeders is an adaptation to the decrease in food for suspension feeders with increasing depth. That is, an individual ascidian uses energy to pump water through its filtering apparatus and gains energy by ingesting the particles it collects. The energy harvested must exceed the energetic costs of the pumping and the energetic costs of maintaining the individual. The amount of energy harvested decreases as food-particle concentration decreases with depth, but the energetic costs of pumping a unit of water remain constant. As a consequence, Monniot (1979) suggested that the maximum mass that can be supported decreases with depth, resulting in dwarfing. In contrast to the trend toward miniaturization, gigantism (= species many times the size of their near
- 39. 24 David THISTLE relatives) occurs in the deep sea. Among the giants are species of scavenging amphipods (Hessler et al., 1972), surface-deposit-feeding holothurians (Gage and Tyler, 1991), and isopods (Wolff, 1956). The giants are not closely related evolutionarily and have different life styles, so it is unlikely that gigantism arises for a single reason. Some abyssal ascidians are three to ten times larger than other abyssal members of their families. These giants have greatly modified filtering structures and appear to have abandoned active suspension feeding for passive suspension feeding. Monniot (1979) argued that the elimination of the energetic costs of pumping allowed individuals of greater mass to be supported from a given concentration of suspended particles than could be supported by active suspension feeding; but these giants can occur only in locations where the flux of food particles is relatively high. The changes in size with depth reported above were for entities that contained many species, such as macro- fauna and nematodes. Rex and Etter (1998) argued that such patterns should be investigated within individual species, because only within species will it be possible to understand the ecological and evolutionary forces that created the patterns. When they did so for several gastropod species, they found that the trend was for individuals to increase rather than decrease in size with depth (see also Wilson, 1983, and Macpherson and Duarte, 1991). Biogeography Introduction Geographical patterns in the distribution of species (or higher taxa) and the causes of those patterns are not well known for animals of the deep-sea floor. This situation arises, in part, because of the great mismatch between the vastness of the habitat and the small amount of sampling that has been done. In addition, large numbers of species are present in the deep sea, most of which are undescribed. Further, the number of specialists who can provide identifications or taxonomic descriptions is small and is decreasing. The shape of the ocean floor sets the stage for deep- sea biogeography. Briefly, within major ocean basins at slope depths, the habitat is more or less continuous along isobaths, interrupted by relatively small features, such as submarine canyons. The major oceans and most of the secondary seas are connected at these depths. Below about 2500 m, the mid-ocean ridges and submarine mountain ranges divide the major oceans into regions, for instance, the eastern Atlantic. Below about 3500 m, the deep-sea floor consists of isolated basins (Fig. 2.20). Fig. 2.20. The Atlantic Ocean showing the location of the deep- ocean (3500 m) basins (black) and the mid-Atlantic Ridge (dashes). Modified from Allen and Sanders (1996). Copyright: Elsevier Science. Patterns along isobaths The distribution of higher taxa is unusually homoge- neous in the deep sea. For example, of the 143 genera of asellote isopods known from the World Ocean, all but nine are found in the Atlantic. This level of similarity is much greater than expected on the basis of shallow-water isopods (Hessler and Wilson, 1983). Some deep-sea species are widespread, but many more have restricted distributions. For example, along 180 km of the 2100-m isobath off the northeastern coast of the United States, 10 abundant macrofaunal species occurred at all stations (Grassle and Maciolek, 1992), but 43% of the peracarid crustacean species, 34% of the polychaete species, and 21% of the bivalve species occurred at only one. The constraints (ecological or historical) that cause large numbers of species to be endemics in this environment are unknown, but the differences among higher taxa in the proportion of species with localized distributions may provide a point of departure for further research. Because the widespread species constitute the bulk of the individuals at each station, the faunas at either end
- 40. THE DEEP-SEA FLOOR: AN OVERVIEW 25 of the 180-km transect are relatively similar (of the species expected in a random sample of 200 individuals at each of the two localities, 79% were shared) (Grassle and Maciolek, 1992). Despite this similarity, the endemics are a source of faunal heterogeneity among stations. The pattern of a few widespread species and many endemics also occurs in the ocean basins. For example, at the scale of the Atlantic, 11 of 109 species and subspecies of protobranch bivalves have been found in six or more of the 15 deep basins, but 48 have been found in only one, and the remaining species tend to occur in two adjacent basins (Allen and Sanders, 1996). At least for protobranchs, the distributions of most species tend to be much smaller than pan-Atlantic, and the protobranch faunas of adjacent basins tend to be similar – for instance, the Sierra Leone and Angola Basins share 20 of a total of 40 species. The Norwegian Basin is an exception. It shares no protobranch species with any other Atlantic basin. The reason for this difference is not known, but the Norwegian Basin became anoxic during the last glaciation (Schnitker, 1979) and lost most of its metazoan fauna. It is unclear why some species occur in several basins and others occur in only one, but ecological differences among the basins, dispersal limitations, and history are all likely to play a role. Patterns with depth The faunal break at the shelf/slope transition has been confirmed repeatedly (Sanders and Hessler, 1969; Haedrich et al., 1975; Rex, 1977; Carney and Carey, 1982). Below that depth, regions of relatively slow faunal change (= zones) are separated by bands of more rapid faunal change. The depths of zone boundaries vary among taxa and locations, but in the North Atlantic appear to be at about 0.5, 1, 1.5, and 2 km (Gage and Tyler, 1991). Below 2 km, the rate of change of the fauna slows, and zonation does not appear to be as marked, perhaps in part because of the smaller sampling effort at these depths. Although the depth range of an individual species does not often extend from 200 m into the trenches or even from 200 m to the abyssal plain, some species have extensive ranges. For example, the brittle star Ophiomusium lymani off North Carolina has been found from 1372 to 3987 m (Grassle et al., 1975). Species are not equally abundant throughout their ranges, tending to be rare at the extremes as expected, if conditions become less suitable as their range boundaries are approached (Grassle et al.: their table 6). The depth range of a species may vary along isobaths. For example, in contrast to its depth range off North Carolina, O. lymani was found only between 1705 and 2170 m on a transect from Massachusetts to Bermuda (Schoener, 1969). Many species appear to have very small depth ranges, having been collected at only one station along a transect. Because many transects have had stations at depth increments of a few hundreds of meters, the depth ranges of such species must be less. Contrary to expectation, representatives of deep-sea taxa are found on the shelf in the Arctic, Antarctic, and Mediterranean seas and in Scandinavian fjords. For example, species of asellote isopods of deep- sea genera have been found at shelf depths in the Bay of Naples (Schiecke and Fresi, 1969) and off Sweden (Hessler and Strömberg, 1989); foraminifer species of deep-sea genera have been found at SCUBA- diving depths in Antarctica (Gooday et al., 1996); and a sponge from a bathyal–abyssal subphylum has been found in a cave at 18 m in the Mediterranean Sea (Vacelet et al., 1994). This pattern could arise if representatives of taxa that had evolved in the deep sea entered shallow water. Alternatively, the shallow- water representatives of these taxa could be relics in the original environment of groups that invaded the deep sea. At least for the shelf representatives of asellote isopods, the former appears to be the case (Hessler and Thistle, 1975). The environments where representatives of deep-sea taxa are found in shallow water have deep- sea temperatures at shelf depths (cold at the poles but warm in the Mediterranean), suggesting that ordinarily the temperature gradient (or a correlate) separates the deep-sea fauna from that of the shelf. Combined along- and across-isobath patterns Relatively few deep-sea studies have had sample coverage adequate to address along- and across-isobath patterns in the distribution of species. For protobranchs at the scale of the Atlantic, Allen and Sanders (1996) found that the similarity of the fauna among stations was much greater along isobaths than across isobaths. In particular, the faunas of stations separated by 16 500 km along an isobath were as similar as those of stations separated by 0.8 km across isobaths. The pattern of greater similarity along than across isobaths has also been found at the 100-km, within-ocean- basin scale (Carney et al., 1983; Grassle and Maciolek, 1992). At the 100-km scale, recent work on benthic decapods from the Mediterranean slope has revealed
- 41. 26 David THISTLE discrete regions of high abundance within the range of a species which extend both along and across isobaths (Maynou et al., 1996). Therefore, the distribution of a species within its depth range may be a series of patches (see Blake and Hilbig, 1994). These results have been based on the distribution of macrofauna. Theory suggests (Fenchel, 1993) that animals of smaller size (meiofauna, microfauna) may have larger species ranges, so many species may be cosmopolitan. Relatively little information is available with which to evaluate this idea in the deep sea, but many common deep-sea formaminiferal species appear to be cosmopolitan (Gooday et al., 1998). Finally, deep-sea biogeography is based almost en- tirely on morphological species, which is not surprising given the difficulty of working in this environment. Substantial genetic variation has been discovered within some nominal species (Etter et al., 1999), raising the possibility that some morphological species may be complexes of cryptic species, a situation that would make deep-sea biogeography even more difficult. Factors controlling the depth range of a species The physiological limits of a species set the ultimate bounds of its range; shallow-water species could there- fore be excluded from the deep sea and vice versa, and the ranges of deep-sea species restricted. For example, mitosis is inhibited when shallow-water sea urchins are exposed to deep-sea pressures (Marsland, 1938, 1950). Also, larvae of a bathyal sea urchin require bathyal temperatures to develop properly, setting a physiological limit to the minimum and maximum depths at which the larvae can develop (Young and Cameron, 1989). Similarly, when eggs of an asteroid that lives between 1000 and 2500 m were exposed to pressures corresponding to 0 and 3000 m, virtually no normal development occurred (Young et al., 1996). Ecological limitations could be imposed in a variety of ways. As depth increases, sediment composition can change. Because some species are restricted to particular sediment types, sediment changes can limit their ranges. For example, on a depth transect off North Carolina, sands give way to sandy silts and then to clayey silts. Nematode species were restricted to these sediment-defined depth bands – 17, 5, and 49 species, respectively (Tietjen, 1976). The depth distribution of a species may also be controlled by food availability. For example, the decrease in suspended food concentration with depth limits the penetration of certain suspension-feeding species into the deep sea (see above, pp. 20–21). In contrast, on the Carolina slope unusual conditions occur such that food is supplied to the slope at rates comparable to those on the shelf, and some species ordinarily confined to the shelf are present on the slope in large numbers (Blake and Hilbig, 1994). Examples of other environmental variables that can affect the ranges of species include low oxygen concentration in the near-bottom water (Wishner et al., 1990), strong near-bottom flows (Rice et al., 1990; Paterson and Lambshead, 1995), and correlates of the permanent thermocline (Gage, 1986). The depth distribution of a species may also be controlled by ecological interactions with other species. For example, Rex (1977) found that groups made up largely of croppers and predators such as the epibenthic macrofauna and gastropods had smaller species ranges on average than did infaunal deposit feeders. To explain the difference, he noted that the former were at a higher trophic level than the latter and that studies in other environments have found that the higher the trophic level the greater the competitive interactions among species. He suggested that the increased probability of competition among species of epibenthic macrofauna and gastropods resulted in lower average range sizes than in the infauna, whose populations were much less likely to interact competitively because of the intense predation upon them (but see Carney et al., 1983). THE HARD-BOTTOM HABITAT Examples of deep-sea hard-bottom habitats include the exposed portions of rocks and manganese nodules, mollusk shells, new oceanic crust, regions of steep topography where sediment does not accumulate, and locations where the sediment has been washed away by currents (e.g., portions of some seamounts). Despite this variety, two generalizations can be made. The hard- bottom fauna differs in taxonomic composition and life-style from that of deep-sea soft bottoms (Table 2.2), and near-bottom flow tends to be more important to this fauna than to that of the deep-sea soft bottoms. The fauna Soft bottoms are three-dimensional, and although the sediment surface has great ecological importance to many of the species present, few animals live on it. Hard bottoms are two-dimensional. Most of the animals live on the surface of the substratum, and infaunal
- 42. THE DEEP-SEA FLOOR: AN OVERVIEW 27 Table 2.2 Comparison of soft-bottom and hard-bottom faunas in the deep sea Feature Soft-bottom habitat Manganese- nodule habitat Other hard-bottom habitats Dominant taxa polychaetes, nematodes foraminifers sponges, black corals, horny corals Dominant size macrofaunal and meiofaunal macrofaunal and meiofaunal megafaunal Dominant mobility mode mobile sessile sessile Dominant feeding mode deposit suspension and surface deposit suspension organisms are essentially absent. One exception to this generalization is “cauliflower”-type manganese nodules, where sediment accumulates in the crevices and harbors a more or less conventional infauna (Thiel et al., 1993). The dominant taxa of deep-sea soft bottoms are polychaetes, nematodes, and foraminifers. The domi- nant taxon on manganese nodules is the agglutinating foraminifers (Mullineaux, 1987). On other hard bot- toms, sponges, horny corals (gorgonians), and black corals (antipatharians) dominate (Genin et al., 1986, 1992; Tyler and Zibrowius, 1992), and xenophyophores may be present (Levin and Thomas, 1988). Soft bottoms are dominated by macrofauna and meiofauna. The dominant hard-bottom taxa on manganese nodules and mollusk shells are macrofauna and meiofauna, but on other hard bottoms megafauna appear to dominate. Soft-bottom animals tend to be mobile and to deposit- feed; whereas on manganese nodules and mollusk shells, most organisms are sessile (Mullineaux, 1987; Voight and Walker, 1995). They suspension-feed or collect particles from the surface of the substratum (Mullineaux, 1987, 1989), a process analogous to surface-deposit feeding in soft bottoms. On other hard- bottom habitats, the dominant animals are sessile or sedentary, and they suspension-feed (Genin et al., 1986, 1992; Tyler and Zibrowius, 1992). Distribution Regularities in the distribution of hard-bottom organ- isms are apparent at several scales. For example, on the Bahamian slope, sponges occur in greater numbers on vertically oriented surfaces than on horizontal surfaces, creating marked patchiness on scales of meters. The difference in exposure to siltation, which adversely affects sponges, may underlie the difference (Maldonado and Young, 1996). At scales of hundreds of meters along a depth transect, there are changes in species composition, and there may be changes in the general characteristics of the fauna. For example, sponge species diversity is high on the shelf and upper slope of the Bahamas. As depth increases, the diversity decreases as the species of the lower reefs disappear. Deeper on the slope, deep- sea sponges occur, resulting in a secondary peak in diversity and abundance. Branched erect sponges are common in shallow but not in deep water, so there is also a change in the morphological composition of the fauna with depth (Maldonado and Young, 1996). The importance of near-bottom flow The hard-bottom fauna includes many sessile suspen- sion feeders. These animals depend on near-bottom flow to transport their propagules to settlement sites. To a first approximation, the flux (flow velocity x propagule concentration) of propagules determines the initial distribution of these species. These animals also depend on the near-bottom flow to supply them with food. To a first approximation, the flux of food determines survivorship. Evidence of such effects on the hard-bottom fauna has been found at several scales. For example, on a wide peak on the summit of Jaspar Seamount (Pacific Ocean), individuals of a species of black coral are more abundant on the edges of the peak than in the center. This pattern matches that of variation in the velocity field (Genin et al., 1986). Also, where the Western Boundary Undercurrent encounters the Blake Spur (Atlantic Ocean), the cliffs are bathed by flows of ⩾ 30 cm s−1 (much faster than the 3 cm s−1 of typical deep-sea flows). There, massive sponges and horny corals are unusually abundant (Genin et al., 1992). In these examples, Genin et al. assumed that the concentration of propagules (or food) was the same in the water that impinged on the high- and low- flow environments. This assumption may not always be valid, particularly near the seabed. For example, Mullineaux (1988) compared the settlement rate of sessile organisms (predominantly foraminifers) on manganese nodules suspended 20 cm above the seabed to the rate on nodules at the seabed. She found that the
- 43. 28 David THISTLE rate of settlement was significantly greater on the lower nodules despite their exposure to much slower flows. A complete understanding of such flow effects will require simultaneous measurements of velocity and the concentration of propagules (or food). Flow effects also arise because of the vertical gradient in horizontal velocity adjacent to the seabed (Fig. 2.21). At the seabed, the horizontal velocity is Fig. 2.21. An illustration of the decrease in horizontal speed in the bottom boundary layer as the seabed is approached. zero (Vogel, 1981). In the layer immediately above the seabed, molecular viscosity suppresses turbulent motions, and horizontal velocity increases slowly and linearly with height. In the deep sea, this region of slow flow (the viscous sublayer) may be a centimeter thick (Jumars and Gallagher, 1982). Above the viscous sublayer, turbulence is present and horizontal velocity increases logarithmically with height. This “log” layer can be 1–10 m thick in the deep sea (Jumars and Gallagher, 1982). As the horizontal velocity in the log layer increases, the viscous sublayer becomes thinner, and turbulent eddies collide with the seabed more frequently, disrupting or destroying the viscous sublayer. This vertical gradient in horizontal velocity can be expected to influence the settlement location of sessile suspension feeders, because most hard-bottom suspension feeders extend a collecting apparatus into the flow and depend on water motion to bring particles to it. Assuming that there is no vertical gradient in food concentration, the flux of food increases with height above the bottom (and the rate of increase rises rapidly with height as the viscous sublayer gives way to the log layer). Therefore, there should be strong selection for passive suspension feeders on hard bottoms to extend their feeding apparatus as far from the seabed as possible, and there is a large premium on extension above the viscous sublayer. Accordingly, the propagules of passive suspension feeders that are small as adults may be expected to settle at locations where the local horizontal velocity is great enough to disrupt the viscous sublayer or reduce its thickness (Jumars and Gallagher, 1982). For example, a variety of foraminifers attach in such a way that they are millimeters to centimeters above the seabed (Lutze and Altenbach, 1988; Lutze and Thiel, 1989). This selection pressure should also apply to passive suspension feeders that are large as adults, because their juveniles should benefit from extending out of the viscous sublayer (Jumars and Gallagher, 1982). For example, passive suspension feeders tend to be found on the edges of rocks, where the flow can be expected to be locally accelerated and the viscous sublayer thinned. Also, the feeding polyps of horny corals (gorgonians) and sea pens (pennatularians) are absent near the base of the animal, where the flow is slowest, and feather stars (comatulid crinoids) perch on other animals well above the seabed (Tyler and Zibrowius, 1992). Although the flux of propagules and the flux of food may increase monotonically with increasing velocity, the rate of larval settlement and the rate of food capture by a suspension feeder may not. In particular, as near-bottom velocity (more correctly, the vertical gradient of horizontal velocity) increases, the shear (horizontal) force exerted on the seabed increases. To settle, a larva must attach with sufficient strength to resist that force. At some velocity, the frequency of successful settlement will begin to decrease, raising the possibility that settlement success could be greatest at intermediate-velocity locations (Crisp, 1955). Also, when velocity exceeds some limit, the feeding rate of some suspension feeders will begin to decline because their feeding apparatus will be deformed by the flow (Koehl, 1977) and become less effective. The distribution of suspension feeders (predominantly sessile foraminifers) on manganese nodules may reflect such effects. They were more abundant in a band below the summits of manganese nodules than on the tops of the nodules, where the velocity (and shear stress) was greatest (Mullineaux, 1989; see also Mullineaux and Butman, 1990). Flow is not the only variable that influences the settlement location of propagules on hard substrata. Mullineaux and Butman (1990) found that some species would settle on plates coated with ferroman- ganese but would not settle on smooth control plates, indicating that some propagules seek specific substrata or textures (see also Mullineaux, 1988; Bertram and Cowen, 1994, 1999).
- 44. THE DEEP-SEA FLOOR: AN OVERVIEW 29 Fig. 2.22. Weight-specific respiration rate of benthopelagic organisms living in shallow water (dashed regression line) and in the deep sea (solid regression line) as a function of individual organic carbon weight (W), illustrating the lower metabolic rates in the deep sea. Modified from Mahaut et al. (1995). Copyright: Elsevier Science. THE PACE OF LIFE IN THE DEEP SEA Because the rate of input of food to the deep sea is small, it has been assumed that energy-conserving strategies (low movement rates, low metabolic rates, long life, late maturity, and small reproductive output) have been selected for, giving rise to the expectation that the pace of life is slower in the deep sea than in shallow water. The results of some early deep-sea rate measurements (Jannasch et al., 1971; Smith and Hessler, 1974; Turekian et al., 1975) reinforced this impression, but some rates in the deep sea may be similar to or only marginally lower than rates in shallow water (Gage, 1991). Locomotion Movement rates of epibenthic megafauna and demersal fishes have been measured, but comparable shallow- water data are not always available. Most individuals most of the time are stationary or are moving slowly. For example, demersal deep-sea fishes swam slowly (8 cm s−1 ) over the seabed in a nomadic search for food (Bagley and Priede, 1997). Deep-sea brittle stars moved at 1–3 cm min−1 (LaFond, 1967), whereas shallow- water brittle stars moved at 15–45 cm min−1 (Broom, 1975). Deep-sea holothurians moved at 1–2 cm min−1 (Gage, 1991), whereas shallow-water holothurians moved at 7 cm min−1 (Parker, 1921). Although ordinary movement is slow, when stimu- lated, many deep-sea animals can move at rates com- parable to those of similar shallow-water animals. For example, when approached by a research submarine, demersal fishes swam away rapidly enough to stir up a cloud of sediment (Grassle et al., 1975). Brittle stars are ordinarily still, and parcel-attending amphipods are still or drift with the currents. When these animals detect a food parcel, they move rapidly toward it (Smith, 1985). Speeds of 7 cm s−1 have been measured for amphipods, which are comparable to speeds measured on shallow- water confamilials (Laver et al., 1985). Metabolic rates To investigate the relative metabolic rates of deep- sea and shallow-water animals, Mahaut et al. (1995) calculated regression lines of metabolic rate (as weight- specific respiration) against mass from published data (Fig. 2.22). The regression line for deep-sea animals fell below that for shallow-water animals, so on the average, weight-specific metabolic rates are lower in the deep sea. The difference decreased with the size of the animal, suggesting that rates for macrofauna and meiofauna in the deep sea are lower but not markedly lower than in shallow water (see also Gage, 1991). The data for the larger deep-sea animals came from measurements on fishes. The metabolism of demersal fishes, even those that swim constantly, appears to be slower than that of shallow-water species. For example, the resting oxygen consumptions for individuals of two fish species from 1230 m were significantly lower
- 45. 30 David THISTLE than rates measured on similar shallow-water, low- temperature fishes (Smith and Hessler, 1974). These low rates appear to be necessary because of low food availability and are achieved by minimization of locomotion and accompanying economies at the cellular level, e.g., lower enzyme concentrations in muscles than in comparable, shallow-water species (Siebenaller and Somero, 1982). In contrast, the weight-specific oxygen consumption of bathyal brittle stars (three species) and a bathyal holothuroid (Scoto- planes globosa) were similar to those in shallow water (Smith, 1983). The collective metabolism of the organisms (bacte- ria, protozoa, meiofauna, and macrofauna) resident in the sediment has been estimated by measurements of sediment-community oxygen consumption (SCOC) per unit area. Smith and Teal (1973) found that the SCOC at a station at 1850 m was two orders of magnitude less than that at several shallow-water stations. Further work (Smith, 1987) has shown that SCOC decreases with increasing depth in the deep sea. On a unit- biomass basis, the 1850-m station had a tenth the SCOC of the shallow-water stations (Smith and Teal, 1973). In contrast, there was no trend in SCOC per unit biomass with depth among the deep-sea stations (Smith and Hinga, 1983; Smith, 1987). If one assumes that the relative metabolic rate among the component groups of organisms does not change markedly as depth increases, these results imply that the metabolic rate of the sediment-dwelling organisms is lower in the deep sea than in shallow water, but within the deep sea, there is no trend with depth. Bioturbation Bioturbation occurs when organisms (primarily the infauna) move sediment. Bioturbation rates decrease with increasing depth in the deep sea, but when bioturbation rates are normalized by the number of animals per unit area, which also decreases with increasing depth, the rate does not change with depth (Gage, 1991). Individual deep-sea animals mix the sediment at about the same rate as shallow-water animals, suggesting that movement rates and metabolic rates of infauna may not be too dissimilar to such rates in shallow water. Growth The growth of demersal fishes appears to be slow in the deep sea (Beamish and Chilton, 1982; Merrett, 1989), as might be expected from their low metabolic rate. The rate of growth of the deep-sea infauna is less clear. Turekian et al. (1975) used a radioisotope technique to study the growth rate of a deep-sea clam. Their best estimate was that the largest size class (8 mm) was about 100 years old and that reproductive maturity was reached in 50–60 years. Although the confidence limits around these estimates were large, the report has been very influential because it supported preconceptions that growth rates would be low and life spans long in the deep sea. The disparity between the results of Turekian et al. (1975) and those from shallow water are not as great as originally thought. For example, the growth rates of some deep-sea bivalve species are comparable to those of shallow-water bivalves (Gage, 1991). Following the work of Turekian et al. (1975), the growth rates of deep- sea benthic organisms have been estimated by other indirect methods. If the assumptions made are correct, the growth rates of deep-sea invertebrates are lower than, but not markedly different from, growth rates of shallow-water species (Gage, 1991). Direct mea- surements from time-lapse photographs have shown that the growth of a deep-sea hard-bottom barnacle was almost as rapid as that of similar, shallow-water barnacles (Lampitt, 1990) and that the volume of three xenophyophore individuals increased by a factor of 3– 10 in 8 months; during these 8 months, periods of rapid growth were interspersed with periods of no growth (Gooday et al., 1993) (Fig. 2.23). Colonization studies suggest that an aplacophoran mollusk can reach adult size in two months (Scheltema, 1987). Food availability appears to be a leading variable in the control of the growth rate of deep-sea animals; where food is abundant, growth is rapid compared to that where food is scarce. For example, food is abundant for tube worms (Vestimentifera) at hydrother- mal vents, and their length can increase by tens of centimeters per year (Lutz et al., 1994). Similarly, pieces of wood (e.g., tree trunks) that wash offshore and sink are a rich source of food for wood-boring bivalves (Xylophaginidae), which reach adult size within months of settlement (Turner, 1973). The growth rate of a gooseneck barnacle increased several-fold when phytodetritus, a likely source of food for this suspension feeder, was present (Lampitt, 1990). It appears that growth rates in the deep sea are limited, not by the physiological challenges of the physical
- 46. THE DEEP-SEA FLOOR: AN OVERVIEW 31 0 10 50 2 4 6 8 10 12 14 16 18 20 8 10 12 A B C 14 16 18 20 22 0 2 4 6 Time (days) Test volume (cm 3 ) 150 200 250 300 0 Fig. 2.23. Growth (as test volume) of three abyssal xenophyophore individuals illustrating that growth can be rapid for deep-sea organisms. Outer scale refers to individuals B and C. Inner scale refers to individual A. Modified from Gooday et al. (1993). Copyright: Elsevier Science. environment (high pressure, low temperature), but by the meager supply of food. ACKNOWLEDGEMENTS The following people answered queries: W. Burnett, J. Eckman, A. Gooday, P. Jumars, W. Landing, P. LaRock, G. Paterson, G. Weatherly and G. Wilson. L. Bouck, S. Ertman, R. Hessler, K. Suderman, A. Thistle, G. Wilson, and an anonymous reviewer commented on the manuscript. I am grateful for this kind help. The chapter was written while the author was partially supported by ONR grant N00014-95-1- 0750 and NSF grant OCE-9616676. REFERENCES Aldred, R.G., Riemann-Zürneck, K., Thiel, H. and Rice, A.L., 1979. Ecological observations on the deep-sea anemone Actinoscyphia aurelia. Oceanol. Acta, 2: 389−395. Allen, J.A. and Sanders, H.L., 1966. Adaptations to abyssal life as shown by the bivalve Abra profundum (Smith). Deep-Sea Res., 13: 1175−1184. Allen, J.A. and Sanders, H.L., 1996. The zoogeography, diversity and origin of the deep-sea protobranch bivalves of the Atlantic: the epilogue. Prog. Oceanogr., 38: 95−153. Altenbach, A.V ., Heeger, T., Linke, P., Spindler, M. and Thies, A., 1993. Miliolinella subrotunda (Montague), a miliolid foraminifer building large dentritic tubes for temporary epibenthic life style. Mar. Micropaleontol., 20: 293−301. Anonymous, 1989. Seawater: Its Composition, Properties and Behaviour. Pergamon Press, Oxford, 165 pp. Bagley, P.M. and Priede, I.G., 1997. An autonomous free-fall acoustic tracking system for investigation of fish behaviour at abyssal depths. Aquat. Living Resourc., 10: 67−74. Barnes, R.D., 1987. Invertebrate Zoology. Harcourt Brace Jovanovich College Publishers, New York, 893 pp. Barnett, P.R.O., Watson, J. and Connelly, D., 1984. A multiple corer for taking virtually undisturbed samples from shelf, bathyal and abyssal sediments. Oceanol. Acta, 7: 399−408. Beamish, J.R. and Chilton, D.E., 1982. Preliminary evaluation of a method to determine the age of sable fish (Anopoploma fimbria). Can. J. Fish. Aquat. Sci., 39: 277−287. Behl, R.J. and Kennett, J.P., 1996. Brief interstadial events in the Santa Barbara Basin, NE Pacific during the past 60 kyr. Nature, 379: 243−246. Belyaev, G.M., 1989. Deep-Sea Oceanic Trenches and Their Fauna. Institute of Oceanology, USSR Academy of Sciences, Moscow. In Russian. [Quoted by Gage and Tyler (1991)]. Benson, R.H., Chapman, R.E. and Deck, L.T., 1984. Paleoceano- graphic events and deep-sea ostracods. Science, 224: 1334−1336. Bertram, M.A. and Cowen, J.P., 1994. Testate rhizopod growth and mineral deposition of experimental substrate from Cross Seamount. Deep-Sea Res., 41: 575−601. Bertram, M.A. and Cowen, J.P., 1999. Temporal variations in the deep-water colonization rates of small benthic foraminifera: the results of an experiment on Cross Seamount. Deep-Sea Res. I, 46: 1021−1049. Bett, B.J. and Rice, A.L., 1993. The feeding behaviour of an abyssal echiuran revealed by in situ time-lapse photography. Deep-Sea Res. I, 40: 1767−1779. Bett, B.J., Vanreusel, A., Vincx, M., Soltwedel, T., Pfannkuche, O., Lambshead, P.J.D., Gooday, A.J., Ferrero, T.J. and Dinet, A., 1994. Sampler bias in the quantitative study of deep-sea meiobenthos. Mar. Ecol. Prog. Ser., 104: 197−203. Billett, D.S.M., Lampitt, R.S., Rice, A.L. and Mantoura, R.F.C., 1983. Seasonal sedimentation of phytoplankton to the deep-sea benthos. Nature, 302: 520−522. Blake, J.A. and Hilbig, B., 1994. Dense infaunal assemblages on the continental slope off Cape Hatteras, North Carolina. Deep-Sea Res. II, 44: 875−899. Broom, D.M., 1975. Aggregation behaviour of the brittle star Ophiothrix fragilis. J. Mar. Biol. Assoc. U. K., 55: 191−197. Cahalan, J.A., Siddall, S.E. and Luchenbach, M.W., 1989. Effects of flow velocity, food concentration and particle flux on growth rates of juvenile bay scallops Argopecten irradians. J. Exp. Mar. Biol. Ecol., 129: 45−60. Cannariato, K.G., Kennett, J.P. and Behl, R.J., 1999. Biotic response to late Quaternary rapid climate switches in Santa Barbara Basin: ecological and evolutionary implications. Geology, 27: 63−66. Carey Jr., A.G., 1972. Food sources of sublittoral, bathyal and abyssal asteroids in the northeast Pacific Ocean. Ophelia, 10: 35−47. Carey Jr., A.G., 1981. A comparison of benthic infaunal abundance on two abyssal plains in the northeast Pacific Ocean. Deep-Sea Res., 28A: 467−479. Carney, R.S. and Carey Jr., A.G., 1982. Distribution and diversity of
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- 54. Chapter 3 THE PELAGIC ENVIRONMENT OF THE OPEN OCEAN Martin V. ANGEL INTRODUCTION The waters of the ocean provide the most voluminous habitat on the planet, since 71% of the Earth’s surface is covered by sea to an average depth of 3800 m. The water in the oceans is estimated to have a volume of 1.368×109 km3 , and to be equivalent to about 0.24% of the Earth’s total mass. The shelf seas 0–200 m deep, that fringe the continental landmasses, extend over about 5% of the Earth’s area. At the outer edge of most shelf seas is the shelf-break where the seabed falls quite steeply to depths of around 3000 m, forming the continental slope and rise; these continental margins account for a further 13% of the Earth’s area. Beyond that, abyssal depths 3000 to 6000 m deep cover about 51% of the Earth’s surface, and the deepest or hadal depths (including many of the ocean trenches) where depths are 6000 m cover less than 2%. The volume of the living space provided by the water of the oceans is 168 times that of terrestrial habitats (Cohen, 1994; see Table 3.1). The oceans have a major influence on climate, since they transfer substantial amounts of heat from equatorial zones to the polar regions. Beneath the clearest of oceanic waters sunlight is detectable only to depths of 1000 to 1250 m. In consequence, the major part of this, the most extensive environment on Earth, is virtually completely dark, lit only by brief flashes of bioluminescence, the light the animals themselves produce. It is a very cool environment; the waters at depths of 1000 m and below have temperatures that range mostly between −0.9º and 5ºC. Their physical and chemical environments hardly vary, and the very limited variation is at scales that are much coarser than those of terrestrial and freshwater habitats. Since uniformity of habitat is unfavourable for the evolution of new species, the total inventory of species inhabiting the waters of the oceans is surprisingly small considering their vast volume. Certainly there are generally far fewer pelagic species in the oceans than occur in terrestrial environments, but estimates of the numbers of benthic species, based on very meagre sampling, range from 0.5 to 2000 million depending on the assumptions made. Table 3.1 Biophysical characteristics of oceans compared to those of continents1 Oceans Continents Surface area (108 km2) 3.6 1.5 Surface area as % Earth’s surface 71% 29% Mean depth of life zone (km) 3.8 0.05 Volume of life zone (109 km3) 1.37 0.0075 Volume % of total 99.5 0.5 Standing crop of plants (1027 kg C)1 ~2 560 Biomass per unit area (103 kg C km−2) 5.6 3700 Biomass per unit volume (103 kg C km−3) 1.5 75.000 Dead matter (1015 kg C) ~2 1.5 Dead organic matter per unit area (106 kg C km2) 5.5 10 Net primary productivity (NPP) y−1 25–44 ~501 NPP per unit area (103 kg C km−2 y−1) 69 330 Carbon residence time in living biomass (years)2 0.08 11.2 1 Adapted from Cohen (1994). 2 Based on Harte (1988). Despite the relative uniformity of the chemical and physical conditions, the oceanic water column is an environment that is physiologically highly challenging to life because of resource limitations. However, before discussing the oceanic water column as a habitat, some 39
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- 56. [ 1 0 4] [ 1 0 5] [ 1 0 6] [ 1 0 7] [ 1 0 8] [ 1 0 9] [ 1 1 0] [ 1 1 1] [ 1 all his company but one man whom he could fully trust; and this was Tonty. He and Hennepin were on indifferent terms. Men thrown together in a rugged enterprise like this quickly learn to know each other; and the vain and assuming friar was not likely to commend himself to La Salle's brave and loyal lieutenant. Hennepin says that it was La Salle's policy to govern through the dissensions of his followers; and, from whatever cause, it is certain that those beneath him were rarely in perfect harmony. FOOTNOTES: Hennepin, Description de la Louisiane (1683), 19; Ibid., Voyage Curieux (1704), 66. Ribourde had lately arrived. Lettre de La Motte de la Lussière, sans date; Relation de Henri de Tonty écrite de Québec, le 14 Novembre, 1684 (Margry, i. 573). This paper, apparently addressed to Abbé Renaudot, is entirely distinct from Tonty's memoir of 1693, addressed to the minister Ponchartrain. Hennepin, Nouvelle Découverte (1697), 8. Ibid., Avant Propos, 5. Ibid., Voyage Curieux (1704), 12. Hennepin, Voyage Curieux (1704), 18. Ibid. Avis au Lecteur. He elsewhere represents himself as on excellent terms with La Salle; with whom, he says, he used to read histories of travels at Fort Frontenac, after which they discussed together their plans of discovery. This was the Racines Agnières of Bruyas. It was published by Mr. Shea in 1862. Hennepin seems to have studied it carefully; for on several occasions he makes use of words evidently borrowed from it, putting them into the mouths of Indians speaking a dialect different from that of the Agniers, or Mohawks. Compare Brodhead in Hist. Mag., x. 268.
- 57. 1 2] [ 1 1 3] [ 1 1 4] [ 1 1 5] [ 1 1 6] [ 1 1 7] [ 1 Une enterprise capable d'épouvanter tout autre que moi.— Hennepin, Voyage Curieux, Avant Propos (1704). Je vous proteste ici devant Dieu, que ma Relation est fidèle et sincère, etc.—Ibid., Avis au Lecteur. The nature of these fabrications will be shown hereafter. They occur, not in the early editions of Hennepin's narrative, which are comparatively truthful, but in the edition of 1697 and those which followed. La Salle was dead at the time of their publication. This place is laid down on a manuscript map sent to France by the Intendant Duchesneau, and now preserved in the Archives de la Marine, and also on several other contemporary maps. Hennepin's account of the falls and river of Niagara—especially his second account, on his return from the West—is very minute, and on the whole very accurate. He indulges in gross exaggeration as to the height of the cataract, which, in the edition of 1683, he states at five hundred feet, and raises to six hundred in that of 1697. He also says that there was room for four carriages to pass abreast under the American Fall without being wet. This is, of course, an exaggeration at the best; but it is extremely probable that a great change has taken place since his time. He speaks of a small lateral fall at the west side of the Horse Shoe Fall which does not now exist. Table Rock, now destroyed, is distinctly figured in his picture. He says that he descended the cliffs on the west side to the foot of the cataract, but that no human being can get down on the east side. The name of Niagara, written Onguiaahra by Lalemant in 1641, and Ongiara by Sanson, on his map of 1657, is used by Hennepin in its present form. His description of the falls is the earliest known to exist. They are clearly indicated on the map of Champlain, 1632. For early references to them, see The Jesuits in North America, 235, note. A brief but curious notice of them is given by Gendron, Quelques Particularitez du Pays des Hurons, 1659. The indefatigable Dr. O'Callaghan has discovered thirty-nine distinct forms of the name Niagara. Index to Colonial Documents of New York, 465. It is of Iroquois origin, and in the Mohawk dialect is pronounced Nyàgarah. Tonty, Relation, 1684 (Margry, i. 573).
- 58. 1 8] [ 1 1 9] [ 1 2 0] [ 1 2 1] [ 1 2 2] [ 1 2 3] Near the town of Victor. It is laid down on the map of Galinée, and other unpublished maps. Compare Marshall, Historical Sketches of the Niagara Frontier, 14. Lettre de La Salle à un de ses associés (Margry, ii. 32). Description de la Louisiane (1683), 41. It is characteristic of Hennepin that, in the editions of his book published after La Salle's death, he substitutes, for anybody but him, anybody but those who had formed so generous a design,—meaning to include himself, though he lost nothing by the disaster, and had not formed the design. On these incidents, compare the two narratives of Tonty, of 1684 and 1693. The book bearing Tonty's name is a compilation full of errors. He disowned its authorship. Lettre de La Salle, 22 Août, 1682 (Margry, ii. 212). Lettre de La Motte, sans date.
- 59. THE NIAGARA PORTAGE. CHAPTER X. 1679. THE LAUNCH OF THE GRIFFIN. The Niagara Portage.—A Vessel on the Stocks.—Suffering and Discontent. —La Salle's Winter Journey.—The Vessel launched.—Fresh Disasters. A more important work than that of the warehouse at the mouth of the river was now to be begun. This was the building of a vessel above the cataract. The small craft which had brought La Motte and Hennepin with their advance party had been hauled to the foot of the rapids at Lewiston, and drawn ashore with a capstan, to save her from the drifting ice. Her lading was taken out, and must now be carried beyond the cataract to the calm water above. The distance to the destined point was at least twelve miles, and the steep heights above Lewiston must first be climbed. This heavy task was accomplished on the twenty-second of January. The level of the plateau was reached, and the file of burdened men, some thirty in number, toiled slowly on its way over the snowy plains and through the gloomy forests of spruce and naked oak-trees; while Hennepin plodded through the drifts with his portable altar lashed fast to his back. They came at last to the mouth of a stream which entered the Niagara two leagues above the cataract, and which was undoubtedly that now called Cayuga Creek.[124] Trees were felled, the place cleared, and the master-carpenter set his ship-builders at work. Meanwhile, two Mohegan hunters, attached to the party, made bark wigwams to lodge the men. Hennepin had his chapel, apparently of the same material, where he placed his altar, and on Sundays and saints' days said mass,
- 60. SUFFERING AND DISCONTENT. preached, and exhorted; while some of the men, who knew the Gregorian chant, lent their aid at the service. When the carpenters were ready to lay the keel of the vessel, La Salle asked the friar to drive the first bolt; but the modesty of my religious profession, he says, compelled me to decline this honor. Fortunately, it was the hunting-season of the Iroquois, and most of the Seneca warriors were in the forests south of Lake Erie; yet enough remained to cause serious uneasiness. They loitered sullenly about the place, expressing their displeasure at the proceedings of the French. One of them, pretending to be drunk, attacked the blacksmith and tried to kill him; but the Frenchman, brandishing a red-hot bar of iron, held him at bay till Hennepin ran to the rescue, when, as he declares, the severity of his rebuke caused the savage to desist.[125] The work of the ship-builders advanced rapidly; and when the Indian visitors beheld the vast ribs of the wooden monster, their jealousy was redoubled. A squaw told the French that they meant to burn the vessel on the stocks. All now stood anxiously on the watch. Cold, hunger, and discontent found imperfect antidotes in Tonty's energy and Hennepin's sermons. La Salle was absent, and his lieutenant commanded in his place. Hennepin says that Tonty was jealous because he, the friar, kept a journal, and that he was forced to use all manner of just precautions to prevent the Italian from seizing it. The men, being half-starved, in consequence of the loss of their provisions on Lake Ontario, were restless and moody; and their discontent was fomented by one of their number, who had very probably been tampered with by La Salle's enemies.[126] The Senecas refused to supply them with corn, and the frequent exhortations of the Récollet father proved an insufficient substitute. In this extremity, the two Mohegans did excellent service,—bringing deer and other game, which relieved the most pressing wants of the party, and went far to restore their cheerfulness.
- 61. THE SHIP FINISHED. La Salle, meanwhile, had gone down to the mouth of the river, with a sergeant and a number of men; and here, on the high point of land where Fort Niagara now stands, he marked out the foundations of two blockhouses.[127] Then, leaving his men to build them, he set out on foot for Fort Frontenac, where the condition of his affairs demanded his presence, and where he hoped to procure supplies to replace those lost in the wreck of his vessel. It was February, and the distance was some two hundred and fifty miles, through the snow-encumbered forests of the Iroquois and over the ice of Lake Ontario. Two men attended him, and a dog dragged his baggage on a sledge. For food, they had only a bag of parched corn, which failed them two days before they reached the fort; and they made the rest of the journey fasting. During his absence, Tonty finished the vessel, which was of about forty-five tons' burden.[128] As spring opened, she was ready for launching. The friar pronounced his blessing on her; the assembled company sang Te Deum; cannon were fired; and French and Indians, warmed alike by a generous gift of brandy, shouted and yelped in chorus as she glided into the Niagara. Her builders towed her out and anchored her in the stream, safe at last from incendiary hands; and then, swinging their hammocks under her deck, slept in peace, beyond reach of the tomahawk. The Indians gazed on her with amazement. Five small cannon looked out from her portholes; and on her prow was carved a portentous monster, the Griffin, whose name she bore, in honor of the armorial bearings of Frontenac. La Salle had often been heard to say that he would make the griffin fly above the crows, or, in other words, make Frontenac triumph over the Jesuits. They now took her up the river, and made her fast below the swift current at Black Rock. Here they finished her equipment, and waited for La Salle's return; but the absent commander did not appear. The spring and more than half of the summer had passed before they saw him again. At length, early in August, he arrived at the mouth of the Niagara, bringing three more friars; for, though no friend of the
- 62. [ 1 2 4] Jesuits, he was zealous for the Faith, and was rarely without a missionary in his journeyings. Like Hennepin, the three friars were all Flemings. One of them, Melithon Watteau, was to remain at Niagara; the others, Zenobe Membré and Gabriel Ribourde, were to preach the Faith among the tribes of the West. Ribourde was a hale and cheerful old man of sixty-four. He went four times up and down the Lewiston heights, while the men were climbing the steep pathway with their loads. It required four of them, well stimulated with brandy, to carry up the principal anchor destined for the Griffin. La Salle brought a tale of disaster. His enemies, bent on ruining the enterprise, had given out that he was embarked on a harebrained venture, from which he would never return. His creditors, excited by rumors set afloat to that end, had seized on all his property in the settled parts of Canada, though his seigniory of Fort Frontenac alone would have more than sufficed to pay all his debts. There was no remedy. To defer the enterprise would have been to give his adversaries the triumph that they sought; and he hardened himself against the blow with his usual stoicism.[129] FOOTNOTES: It has been a matter of debate on which side of the Niagara the first vessel on the Upper Lakes was built. A close study of Hennepin, and a careful examination of the localities, have convinced me that the spot was that indicated above. Hennepin repeatedly alludes to a large detached rock, rising out of the water at the foot of the rapids above Lewiston, on the west side of the river. This rock may still be seen immediately under the western end of the Lewiston suspension-bridge. Persons living in the neighborhood remember that a ferry-boat used to pass between it and the cliffs of the western shore; but it has since been undermined by the current and has inclined in that direction, so that a considerable part of it is submerged, while the gravel and earth thrown down from the cliff during the building of the bridge has filled the intervening channel. Opposite to this rock, and on the east side of the river, says Hennepin, are three mountains, about two leagues below the cataract. (Nouveau Voyage (1704), 462, 466.) To these three mountains,
- 63. [ 1 2 5] [ 1 2 6] as well as to the rock, he frequently alludes. They are also spoken of by La Hontan, who clearly indicates their position. They consist in the three successive grades of the acclivity: first, that which rises from the level of the water, forming the steep and lofty river-bank; next, an intermediate ascent, crowned by a sort of terrace, where the tired men could find a second resting- place and lay down their burdens, whence a third effort carried them with difficulty to the level top of the plateau. That this was the actual portage, or carrying place of the travellers, is shown by Hennepin (1704), 114, who describes the carrying of anchors and other heavy articles up these heights in August, 1679. La Hontan also passed the Falls by way of the three mountains eight years later. La Hontan (1703), 106. It is clear, then, that the portage was on the east side, whence it would be safe to conclude that the vessel was built on the same side. Hennepin says that she was built at the mouth of a stream (rivière) entering the Niagara two leagues above the Falls. Excepting one or two small brooks, there is no stream on the west side but Chippewa Creek, which Hennepin had visited and correctly placed at about a league from the cataract. His distances on the Niagara are usually correct. On the east side there is a stream which perfectly answers the conditions. This is Cayuga Creek, two leagues above the Falls. Immediately in front of it is an island about a mile long, separated from the shore by a narrow and deep arm of the Niagara, into which Cayuga Creek discharges itself. The place is so obviously suited to building and launching a vessel, that, in the early part of this century, the government of the United States chose it for the construction of a schooner to carry supplies to the garrisons of the Upper Lakes. The neighboring village now bears the name of La Salle. In examining this and other localities on the Niagara, I have been greatly aided by my friend O. H. Marshall, Esq., of Buffalo, who is unrivalled in his knowledge of the history and traditions of the Niagara frontier. Hennepin (1704), 97. On a paper drawn up at the instance of the Intendant Duchesneau, the names of the greater number of La Salle's men are preserved. These agree with those given by Hennepin: thus, the master-carpenter, whom he calls Maître Moyse, appears as Moïse Hillaret; and the blacksmith, whom he calls La Forge, is mentioned as—(illegible) dit la Forge. This bad man, says Hennepin, would infallibly have debauched our workmen, if I had not reassured them by the exhortations which I made them on fête-days and Sundays, after divine service. (1704), 98.
- 64. [ 1 2 7] [ 1 2 8] [ 1 2 9] Lettre de La Salle, 22 Août, 1682 (Margry, ii. 197); Relation de Tonty, 1684 (Ibid., i. 577). He called this new post Fort Conti. It was burned some months after, by the carelessness of the sergeant in command, and was the first of a succession of forts on this historic spot. Hennepin (1683), 46. In the edition of 1697, he says that it was of sixty tons. I prefer to follow the earlier and more trustworthy narrative. La Salle's embarrassment at this time was so great that he purposed to send Tonty up the lakes in the Griffin, while he went back to the colony to look after his affairs; but suspecting that the pilot, who had already wrecked one of his vessels, was in the pay of his enemies, he resolved at last to take charge of the expedition himself, to prevent a second disaster. (Lettre de La Salle, 22 Août, 1682; Margry, ii. 214.) Among the creditors who bore hard upon him were Migeon, Charon, Giton, and Peloquin, of Montreal, in whose name his furs at Fort Frontenac had been seized. The intendant also placed under seal all his furs at Quebec, among which is set down the not very precious item of two hundred and eighty-four skins of enfants du diable, or skunks.
- 65. CHAPTER XI. 1679. LA SALLE ON THE UPPER LAKES. The Voyage of the Griffin.—Detroit.—A Storm.—St. Ignace of Michilimackinac.—Rivals and Enemies.—Lake Michigan.—Hardships.—A Threatened Fight.—Fort Miami.—Tonty's Misfortunes.—Forebodings. The Griffin had lain moored by the shore, so near that Hennepin could preach on Sundays from the deck to the men encamped along the bank. She was now forced up against the current with tow-ropes and sails, till she reached the calm entrance of Lake Erie. On the seventh of August, La Salle and his followers embarked, sang Te Deum, and fired their cannon. A fresh breeze sprang up; and with swelling canvas the Griffin ploughed the virgin waves of Lake Erie, where sail was never seen before. For three days they held their course over these unknown waters, and on the fourth turned northward into the Strait of Detroit. Here, on the right hand and on the left, lay verdant prairies, dotted with groves and bordered with lofty forests. They saw walnut, chestnut, and wild plum trees, and oaks festooned with grape-vines; herds of deer, and flocks of swans and wild turkeys. The bulwarks of the Griffin were plentifully hung with game which the men killed on shore, and among the rest with a number of bears, much commended by Hennepin for their want of ferocity and the excellence of their flesh. Those, he says, who will one day have the happiness to possess this fertile and pleasant strait, will be very much obliged to those who have shown them the way. They crossed Lake St. Clair,[130] and still sailed northward against the current, till now, sparkling in the sun, Lake Huron spread before them like a sea.
- 66. ST. IGNACE. For a time they bore on prosperously. Then the wind died to a calm, then freshened to a gale, then rose to a furious tempest; and the vessel tossed wildly among the short, steep, perilous waves of the raging lake. Even La Salle called on his followers to commend themselves to Heaven. All fell to their prayers but the godless pilot, who was loud in complaint against his commander for having brought him, after the honor he had won on the ocean, to drown at last ignominiously in fresh water. The rest clamored to the saints. St. Anthony of Padua was promised a chapel to be built in his honor, if he would but save them from their jeopardy; while in the same breath La Salle and the friars declared him patron of their great enterprise.[131] The saint heard their prayers. The obedient winds were tamed; and the Griffin plunged on her way through foaming surges that still grew calmer as she advanced. Now the sun shone forth on woody islands, Bois Blanc and Mackinaw and the distant Manitoulins,—on the forest wastes of Michigan and the vast blue bosom of the angry lake; and now her port was won, and she found her rest behind the point of St. Ignace of Michilimackinac, floating in that tranquil cove where crystal waters cover but cannot hide the pebbly depths beneath. Before her rose the house and chapel of the Jesuits, enclosed with palisades; on the right, the Huron village, with its bark cabins and its fence of tall pickets; on the left, the square compact houses of the French traders; and, not far off, the clustered wigwams of an Ottawa village.[132] Here was a centre of the Jesuit missions, and a centre of the Indian trade; and here, under the shadow of the cross, was much sharp practice in the service of Mammon. Keen traders, with or without a license, and lawless coureurs de bois, whom a few years of forest life had weaned from civilization, made St. Ignace their resort; and here there were many of them when the Griffin came. They and their employers hated and feared La Salle, who, sustained as he was by the governor, might set at nought the prohibition of the King, debarring him from traffic with these tribes. Yet, while plotting against him, they took pains to allay his distrust by a show of welcome.
- 67. RIVALS AND ENEMIES. The Griffin fired her cannon, and the Indians yelped in wonder and amazement. The adventurers landed in state, and marched under arms to the bark chapel of the Ottawa village, where they heard mass. La Salle knelt before the altar, in a mantle of scarlet bordered with gold. Soldiers, sailors, and artisans knelt around him, —black Jesuits, gray Récollets, swarthy voyageurs, and painted savages; a devout but motley concourse. As they left the chapel, the Ottawa chiefs came to bid them welcome, and the Hurons saluted them with a volley of musketry. They saw the Griffin at her anchorage, surrounded by more than a hundred bark canoes, like a Triton among minnows. Yet it was with more wonder than good-will that the Indians of the mission gazed on the floating fort, for so they called the vessel. A deep jealousy of La Salle's designs had been infused into them. His own followers, too, had been tampered with. In the autumn before, it may be remembered, he had sent fifteen men up the lakes to trade for him, with orders to go thence to the Illinois and make preparation against his coming. Early in the summer, Tonty had been despatched in a canoe from Niagara to look after them.[133] It was high time. Most of the men had been seduced from their duty, and had disobeyed their orders, squandered the goods intrusted to them, or used them in trading on their own account. La Salle found four of them at Michilimackinac. These he arrested, and sent Tonty to the Falls of Ste. Marie, where two others were captured, with their plunder. The rest were in the woods, and it was useless to pursue them. Anxious and troubled as to the condition of his affairs in Canada. La Salle had meant, after seeing his party safe at Michilimackinac, to leave Tonty to conduct it to the Illinois, while he himself returned to the colony. But Tonty was still at Ste. Marie, and he had none to trust but himself. Therefore, he resolved at all risks to remain with his men; for, he says, I judged my presence absolutely necessary to retain such of them as were left me, and prevent them from being enticed away during the winter. Moreover, he thought that he had detected an
- 68. POTTAWATTAMIE S. intrigue of his enemies to hound on the Iroquois against the Illinois, in order to defeat his plan by involving him in the war. Early in September he set sail again, and passing westward into Lake Michigan,[134] cast anchor near one of the islands at the entrance of Green Bay. Here, for once, he found a friend in the person of a Pottawattamie chief, who had been so wrought upon by the politic kindness of Frontenac that he declared himself ready to die for the children of Onontio.[135] Here, too, he found several of his advance party, who had remained faithful and collected a large store of furs. It would have been better had they proved false, like the rest. La Salle, who asked counsel of no man, resolved, in spite of his followers, to send back the Griffin laden with these furs, and others collected on the way, to satisfy his creditors.[136] It was a rash resolution, for it involved trusting her to the pilot, who had already proved either incompetent or treacherous. She fired a parting shot, and on the eighteenth of September set sail for Niagara, with orders to return to the head of Lake Michigan as soon as she had discharged her cargo. La Salle, with the fourteen men who remained, in four canoes deeply laden with a forge, tools, merchandise, and arms, put out from the island and resumed his voyage. The parting was not auspicious. The lake, glassy and calm in the afternoon, was convulsed at night with a sudden storm, when the canoes were midway between the island and the main shore. It was with difficulty that they could keep together, the men shouting to each other through the darkness. Hennepin, who was in the smallest canoe with a heavy load, and a carpenter for a companion who was awkward at the paddle, found himself in jeopardy which demanded all his nerve. The voyagers thought themselves happy when they gained at last the shelter of a little sandy cove, where they dragged up their canoes, and made their cheerless bivouac in the drenched and dripping forest. Here they spent five days, living on pumpkins and Indian corn, the gift of their Pottawattamie friends, and on a Canada
- 69. porcupine brought in by La Salle's Mohegan hunter. The gale raged meanwhile with relentless fury. They trembled when they thought of the Griffin. When at length the tempest lulled, they re-embarked, and steered southward along the shore of Wisconsin; but again the storm fell upon them, and drove them for safety to a bare, rocky islet. Here they made a fire of drift-wood, crouched around it, drew their blankets over their heads, and in this miserable plight, pelted with sleet and rain, remained for two days. At length they were afloat again; but their prosperity was brief. On the twenty-eighth, a fierce squall drove them to a point of rocks covered with bushes, where they consumed the little that remained of their provisions. On the first of October they paddled about thirty miles, without food, when they came to a village of Pottawattamies, who ran down to the shore to help them to land; but La Salle, fearing that some of his men would steal the merchandise and desert to the Indians, insisted on going three leagues farther, to the great indignation of his followers. The lake, swept by an easterly gale, was rolling its waves against the beach, like the ocean in a storm. In the attempt to land, La Salle's canoe was nearly swamped. He and his three canoe-men leaped into the water, and in spite of the surf, which nearly drowned them, dragged their vessel ashore with all its load. He then went to the rescue of Hennepin, who with his awkward companion was in woful need of succor. Father Gabriel, with his sixty-four years, was no match for the surf and the violent undertow. Hennepin, finding himself safe, waded to his relief, and carried him ashore on his sturdy shoulders; while the old friar, though drenched to the skin, laughed gayly under his cowl as his brother missionary staggered with him up the beach.[137] When all were safe ashore, La Salle, who distrusted the Indians they had passed, took post on a hill, and ordered his followers to prepare their guns for action. Nevertheless, as they were starving, an effort must be risked to gain a supply of food; and he sent three men back to the village to purchase it. Well armed, but faint with toil and famine, they made their way through the stormy forest bearing
- 70. HARDSHIPS. a pipe of peace, but on arriving saw that the scared inhabitants had fled. They found, however, a stock of corn, of which they took a portion, leaving goods in exchange, and then set out on their return. Meanwhile, about twenty of the warriors, armed with bows and arrows, approached the camp of the French to reconnoitre. La Salle went to meet them with some of his men, opened a parley with them, and kept them seated at the foot of the hill till his three messengers returned, when on seeing the peace-pipe the warriors set up a cry of joy. In the morning they brought more corn to the camp, with a supply of fresh venison, not a little cheering to the exhausted Frenchmen, who, in dread of treachery, had stood under arms all night. This was no journey of pleasure. The lake was ruffled with almost ceaseless storms; clouds big with rain above, a turmoil of gray and gloomy waves beneath. Every night the canoes must be shouldered through the breakers and dragged up the steep banks, which, as they neared the site of Milwaukee, became almost insurmountable. The men paddled all day, with no other food than a handful of Indian corn. They were spent with toil, sick with the haws and wild berries which they ravenously devoured, and dejected at the prospect before them. Father Gabriel's good spirits began to fail. He fainted several times from famine and fatigue, but was revived by a certain confection of Hyacinth administered by Hennepin, who had a small box of this precious specific. At length they descried at a distance, on the stormy shore, two or three eagles among a busy congregation of crows or turkey buzzards. They paddled in all haste to the spot. The feasters took flight; and the starved travellers found the mangled body of a deer, lately killed by the wolves. This good luck proved the inauguration of plenty. As they approached the head of the lake, game grew abundant; and, with the aid of the Mohegan, there was no lack of bear's meat and venison. They found wild grapes, too, in the woods,
- 71. ENCOUNTER WITH INDIANS. and gathered them by cutting down the trees to which the vines clung. While thus employed, they were startled by a sight often so fearful in the waste and the wilderness,—the print of a human foot. It was clear that Indians were not far off. A strict watch was kept, not, as it proved, without cause; for that night, while the sentry thought of little but screening himself and his gun from the floods of rain, a party of Outagamies crept under the bank, where they lurked for some time before he discovered them. Being challenged, they came forward, professing great friendship, and pretending to have mistaken the French for Iroquois. In the morning, however, there was an outcry from La Salle's servant, who declared that the visitors had stolen his coat from under the inverted canoe where he had placed it; while some of the carpenters also complained of being robbed. La Salle well knew that if the theft were left unpunished, worse would come of it. First, he posted his men at the woody point of a peninsula, whose sandy neck was interposed between them and the main forest. Then he went forth, pistol in hand, met a young Outagami, seized him, and led him prisoner to his camp. This done, he again set out, and soon found an Outagami chief,—for the wigwams were not far distant,—to whom he told what he had done, adding that unless the stolen goods were restored, the prisoner should be killed. The Indians were in perplexity, for they had cut the coat to pieces and divided it. In this dilemma they resolved, being strong in numbers, to rescue their comrade by force. Accordingly, they came down to the edge of the forest, or posted themselves behind fallen trees on the banks, while La Salle's men in their stronghold braced their nerves for the fight. Here three Flemish friars with their rosaries, and eleven Frenchmen with their guns, confronted a hundred and twenty screeching Outagamies. Hennepin, who had seen service, and who had always an exhortation at his tongue's end, busied himself to inspire the rest with a courage equal to his own. Neither party, however, had an appetite for the fray. A parley ensued: full compensation was made for the stolen goods,
- 72. and the aggrieved Frenchmen were farther propitiated with a gift of beaver-skins. Their late enemies, now become friends, spent the next day in dances, feasts, and speeches. They entreated La Salle not to advance farther, since the Illinois, through whose country he must pass, would be sure to kill him; for, added these friendly counsellors, they hated the French because they had been instigating the Iroquois to invade their country, Here was another subject of anxiety. La Salle was confirmed in his belief that his busy and unscrupulous enemies were intriguing for his destruction. He pushed on, however, circling around the southern shore of Lake Michigan, till he reached the mouth of the St. Joseph, called by him the Miamis. Here Tonty was to have rejoined him with twenty men, making his way from Michilimackinac along the eastern shore of the lake; but the rendezvous was a solitude,—Tonty was nowhere to be seen. It was the first of November; winter was at hand, and the streams would soon be frozen. The men clamored to go forward, urging that they should starve if they could not reach the villages of the Illinois before the tribe scattered for the winter hunt. La Salle was inexorable. If they should all desert, he said, he, with his Mohegan hunter and the three friars, would still remain and wait for Tonty. The men grumbled, but obeyed; and, to divert their thoughts, he set them at building a fort of timber on a rising ground at the mouth of the river. They had spent twenty days at this task, and their work was well advanced, when at length Tonty appeared. He brought with him only half of his men. Provisions had failed; and the rest of his party had been left thirty leagues behind, to sustain themselves by hunting. La Salle told him to return and hasten them forward. He set out with two men. A violent north wind arose. He tried to run his canoe ashore through the breakers. The two men could not manage their vessel, and he with his one hand could not help them. She swamped, rolling over in the surf. Guns, baggage, and provisions were lost; and the three voyagers returned to the Miamis, subsisting
- 73. FOREBODINGS. [ 1 3 0] [ 1 3 1] [ 1 3 2] [ 1 3 3] [ 1 3 4] [ 1 3 5] [ 1 3 6] on acorns by the way. Happily, the men left behind, excepting two deserters, succeeded, a few days after, in rejoining the party.[138] Thus was one heavy load lifted from the heart of La Salle. But where was the Griffin? Time enough, and more than enough, had passed for her voyage to Niagara and back again. He scanned the dreary horizon with an anxious eye. No returning sail gladdened the watery solitude, and a dark foreboding gathered on his heart. Yet further delay was impossible. He sent back two men to Michilimackinac to meet her, if she still existed, and pilot her to his new fort of the Miamis, and then prepared to ascend the river, whose weedy edges were already glassed with thin flakes of ice.[139] FOOTNOTES: They named it Sainte Claire, of which the present name is a perversion. Hennepin (1683), 58. There is a rude plan of the establishment in La Hontan, though in several editions its value is destroyed by the reversal of the plate. Relation de Tonty, 1684; Ibid., 1693. He was overtaken at the Detroit by the Griffin. Then usually known as Lac des Illinois, because it gave access to the country of the tribes so called. Three years before, Allouez gave it the name of Lac St. Joseph, by which it is often designated by the early writers. Membré, Douay, and others, call it Lac Dauphin. The Great Mountain, the Iroquois name for the governor of Canada. It was borrowed by other tribes also. In the license of discovery granted to La Salle, he is expressly prohibited from trading with the Ottawas and others who brought furs to Montreal. This traffic on the lakes was, therefore, illicit. His
- 74. [ 1 3 7] [ 1 3 8] [ 1 3 9] enemy, the Intendant Duchesneau, afterwards used this against him. Lettre de Duchesneau au Ministre, 10 Nov., 1680. Hennepin (1683), 79. Hennepin (1683), 112; Relation de Tonty, 1693. The official account of this journey is given at length in the Relation des Découvertes et des Voyages du Sieur de la Salle, 1679-1681. This valuable document, compiled from letters and diaries of La Salle, early in the year 1682, was known to Hennepin, who evidently had a copy of it before him when he wrote his book, in which he incorporated many passages from it.
- 75. LA SALLE'S ADVENTURE. CHAPTER XII. 1679, 1680. LA SALLE ON THE ILLINOIS. The St. Joseph.—Adventure of La Salle.—The Prairies.—Famine.—The Great Town of the Illinois.—Indians.—Intrigues.—Difficulties.—Policy of la Salle.—Desertion.—Another Attempt to poison La Salle. On the third of December the party re-embarked, thirty-three in all, in eight canoes,[140] and ascended the chill current of the St. Joseph, bordered with dreary meadows and bare gray forests. When they approached the site of the present village of South Bend, they looked anxiously along the shore on their right to find the portage or path leading to the headquarters of the Illinois. The Mohegan was absent, hunting; and, unaided by his practised eye, they passed the path without seeing it. La Salle landed to search the woods. Hours passed, and he did not return. Hennepin and Tonty grew uneasy, disembarked, bivouacked, ordered guns to be fired, and sent out men to scour the country. Night came, but not their lost leader. Muffled in their blankets and powdered by the thick-falling snow- flakes, they sat ruefully speculating as to what had befallen him; nor was it till four o'clock of the next afternoon that they saw him approaching along the margin of the river. His face and hands were besmirched with charcoal; and he was further decorated with two opossums which hung from his belt, and which he had killed with a stick as they were swinging head downwards from the bough of a tree, after the fashion of that singular beast. He had missed his way in the forest, and had been forced to make a wide circuit around the edge of a swamp; while the snow, of which the air was full, added to his perplexities. Thus he pushed on through the rest of the day and
- 76. THE KANKAKEE. the greater part of the night, till, about two o'clock in the morning, he reached the river again, and fired his gun as a signal to his party. Hearing no answering shot, he pursued his way along the bank, when he presently saw the gleam of a fire among the dense thickets close at hand. Not doubting that he had found the bivouac of his party, he hastened to the spot. To his surprise, no human being was to be seen. Under a tree beside the fire was a heap of dry grass impressed with the form of a man who must have fled but a moment before, for his couch was still warm. It was no doubt an Indian, ambushed on the bank, watching to kill some passing enemy. La Salle called out in several Indian languages; but there was dead silence all around. He then, with admirable coolness, took possession of the quarters he had found, shouting to their invisible proprietor that he was about to sleep in his bed; piled a barricade of bushes around the spot, rekindled the dying fire, warmed his benumbed hands, stretched himself on the dried grass, and slept undisturbed till morning. The Mohegan had rejoined the party before La Salle's return, and with his aid the portage was soon found. Here the party encamped. La Salle, who was excessively fatigued, occupied, together with Hennepin, a wigwam covered in the Indian manner with mats of reeds. The cold forced them to kindle a fire, which before daybreak set the mats in a blaze; and the two sleepers narrowly escaped being burned along with their hut. In the morning, the party shouldered their canoes and baggage and began their march for the sources of the river Illinois, some five miles distant. Around them stretched a desolate plain, half-covered with snow and strewn with the skulls and bones of buffalo; while, on its farthest verge, they could see the lodges of the Miami Indians, who had made this place their abode. As they filed on their way, a man named Duplessis, bearing a grudge against La Salle, who walked just before him, raised his gun to shoot him through the back, but was prevented by one of his comrades. They soon reached a spot
- 77. where the oozy, saturated soil quaked beneath their tread. All around were clumps of alder-bushes, tufts of rank grass, and pools of glistening water. In the midst a dark and lazy current, which a tall man might bestride, crept twisting like a snake among the weeds and rushes. Here were the sources of the Kankakee, one of the heads of the Illinois.[141] They set their canoes on this thread of water, embarked their baggage and themselves, and pushed down the sluggish streamlet, looking, at a little distance, like men who sailed on land. Fed by an unceasing tribute of the spongy soil, it quickly widened to a river; and they floated on their way through a voiceless, lifeless solitude of dreary oak barrens, or boundless marshes overgrown with reeds. At night, they built their fire on ground made firm by frost, and bivouacked among the rushes. A few days brought them to a more favored region. On the right hand and on the left stretched the boundless prairie, dotted with leafless groves and bordered by gray wintry forests, scorched by the fires kindled in the dried grass by Indian hunters, and strewn with the carcasses and the bleached skulls of innumerable buffalo. The plains were scored with their pathways, and the muddy edges of the river were full of their hoof-prints. Yet not one was to be seen. At night, the horizon glowed with distant fires; and by day the savage hunters could be descried at times roaming on the verge of the prairie. The men, discontented and half-starved, would have deserted to them had they dared. La Salle's Mohegan could kill no game except two lean deer, with a few wild geese and swans. At length, in their straits, they made a happy discovery. It was a buffalo bull, fast mired in a slough. They killed him, lashed a cable about him, and then twelve men dragged out the shaggy monster, whose ponderous carcass demanded their utmost efforts. The scene changed again as they descended. On either hand ran ranges of woody hills, following the course of the river; and when they mounted to their tops, they saw beyond them a rolling sea of dull green prairie, a boundless pasture of the buffalo and the deer, in our own day strangely transformed,—yellow in harvest-time with
- 78. THE ILLINOIS TOWN. HUNGER RELIEVED. ripened wheat, and dotted with the roofs of a hardy and valiant yeomanry.[142] They passed the site of the future town of Ottawa, and saw on their right the high plateau of Buffalo Rock, long a favorite dwelling-place of Indians. A league below, the river glided among islands bordered with stately woods. Close on their left towered a lofty cliff,[143] crested with trees that overhung the rippling current; while before them spread the valley of the Illinois, in broad low meadows, bordered on the right by the graceful hills at whose foot now lies the village of Utica. A population far more numerous then tenanted the valley. Along the right bank of the river were clustered the lodges of a great Indian town. Hennepin counted four hundred and sixty of them.[144] In shape, they were somewhat like the arched top of a baggage-wagon. They were built of a framework of poles, covered with mats of rushes closely interwoven; and each contained three or four fires, of which the greater part served for two families. Here, then, was the town; but where were the inhabitants? All was silent as the desert. The lodges were empty, the fires dead, and the ashes cold. La Salle had expected this; for he knew that in the autumn the Illinois always left their towns for their winter hunting, and that the time of their return had not yet come. Yet he was not the less embarrassed, for he would fain have bought a supply of food to relieve his famished followers. Some of them, searching the deserted town, presently found the caches, or covered pits, in which the Indians hid their stock of corn. This was precious beyond measure in their eyes, and to touch it would be a deep offence. La Salle shrank from provoking their anger, which might prove the ruin of his plans; but his necessity overcame his prudence, and he took thirty minots of corn, hoping to appease the owners by presents. Thus provided, the party embarked again, and resumed their downward voyage.
- 79. On New Year's Day, 1680, they landed and heard mass. Then Hennepin wished a happy new year to La Salle first, and afterwards to all the men, making them a speech, which, as he tells us, was most touching.[145] He and his two brethren next embraced the whole company in turn, in a manner, writes the father, most tender and affectionate, exhorting them, at the same time, to patience, faith, and constancy. Four days after these solemnities, they reached the long expansion of the river then called Pimitoui, and now known as Peoria Lake, and leisurely made their way downward to the site of the city of Peoria.[146] Here, as evening drew near, they saw a faint spire of smoke curling above the gray forest, betokening that Indians were at hand. La Salle, as we have seen, had been warned that these tribes had been taught to regard him as their enemy; and when, in the morning, he resumed his course, he was prepared alike for peace or war. The shores now approached each other; and the Illinois was once more a river, bordered on either hand with overhanging woods.[147] At nine o'clock, doubling a point, he saw about eighty Illinois wigwams, on both sides of the river. He instantly ordered the eight canoes to be ranged in line, abreast, across the stream,—Tonty on the right, and he himself on the left. The men laid down their paddles and seized their weapons; while, in this warlike guise, the current bore them swiftly into the midst of the surprised and astounded savages. The camps were in a panic. Warriors whooped and howled; squaws and children screeched in chorus. Some snatched their bows and war-clubs; some ran in terror; and, in the midst of the hubbub, La Salle leaped ashore, followed by his men. None knew better how to deal with Indians; and he made no sign of friendship, knowing that it might be construed as a token of fear. His little knot of Frenchmen stood, gun in hand, passive, yet prepared for battle. The Indians, on their part, rallying a little from their fright, made all haste to proffer peace. Two of their chiefs came forward, holding out the calumet; while another began a loud harangue, to check the young warriors who were aiming their arrows from the
- 80. ILLINOIS HOSPITALITY. farther bank. La Salle, responding to these friendly overtures, displayed another calumet; while Hennepin caught several scared children and soothed them with winning blandishments.[148] The uproar was quelled; and the strangers were presently seated in the midst of the camp, beset by a throng of wild and swarthy figures. Food was placed before them; and, as the Illinois code of courtesy enjoined, their entertainers conveyed the morsels with their own hands to the lips of these unenviable victims of their hospitality, while others rubbed their feet with bear's grease. La Salle, on his part, made them a gift of tobacco and hatchets; and when he had escaped from their caresses, rose and harangued them. He told them that he had been forced to take corn from their granaries, lest his men should die of hunger; but he prayed them not to be offended, promising full restitution or ample payment. He had come, he said, to protect them against their enemies, and teach them to pray to the true God. As for the Iroquois, they were subjects of the Great King, and therefore brethren of the French; yet, nevertheless, should they begin a war and invade the country of the Illinois, he would stand by them, give them guns, and fight in their defence, if they would permit him to build a fort among them for the security of his men. It was also, he added, his purpose to build a great wooden canoe, in which to descend the Mississippi to the sea, and then return, bringing them the goods of which they stood in need; but if they would not consent to his plans and sell provisions to his men, he would pass on to the Osages, who would then reap all the benefits of intercourse with the French, while they were left destitute, at the mercy of the Iroquois.[149] This threat had its effect, for it touched their deep-rooted jealousy of the Osages. They were lavish of promises, and feasts and dances consumed the day. Yet La Salle soon learned that the intrigues of his enemies were still pursuing him. That evening, unknown to him, a stranger appeared in the Illinois camp. He was a Mascoutin chief, named Monso, attended by five or six Miamis, and bringing a gift of
- 81. FRESH INTRIGUES. knives, hatchets, and kettles to the Illinois.[150] The chiefs assembled in a secret nocturnal session, where, smoking their pipes, they listened with open ears to the harangue of the envoys. Monso told them that he had come in behalf of certain Frenchmen, whom he named, to warn his hearers against the designs of La Salle, whom he denounced as a partisan and spy of the Iroquois, affirming that he was now on his way to stir up the tribes beyond the Mississippi to join in a war against the Illinois, who, thus assailed from the east and from the west, would be utterly destroyed. There was no hope for them, he added, but in checking the farther progress of La Salle, or, at least, retarding it, thus causing his men to desert him. Having thrown his fire-brand, Monso and his party left the camp in haste, dreading to be confronted with the object of their aspersions.[151] In the morning, La Salle saw a change in the behavior of his hosts. They looked on him askance, cold, sullen, and suspicious. There was one Omawha, a chief, whose favor he had won the day before by the politic gift of two hatchets and three knives, and who now came to him in secret to tell him what had taken place at the nocturnal council. La Salle at once saw in it a device of his enemies; and this belief was confirmed, when, in the afternoon, Nicanopé, brother of the head chief, sent to invite the Frenchmen to a feast. They repaired to his lodge; but before dinner was served,—that is to say, while the guests, white and red, were seated on mats, each with his hunting-knife in his hand, and the wooden bowl before him which was to receive his share of the bear's or buffalo's meat, or the corn boiled in fat, with which he was to be regaled,—while such was the posture of the company, their host arose and began a long speech. He told the Frenchmen that he had invited them to his lodge less to refresh their bodies with good cheer than to cure their minds of the dangerous purpose which possessed them, of descending the Mississippi. Its shores, he said, were beset by savage tribes, against whose numbers and ferocity their valor would avail nothing; its
- 82. LA SALLE AND THE INDIANS. waters were infested by serpents, alligators, and unnatural monsters; while the river itself, after raging among rocks and whirlpools, plunged headlong at last into a fathomless gulf, which would swallow them and their vessel forever. La Salle's men were for the most part raw hands, knowing nothing of the wilderness, and easily alarmed at its dangers; but there were two among them, old coureurs de bois, who unfortunately knew too much; for they understood the Indian orator, and explained his speech to the rest. As La Salle looked around on the circle of his followers, he read an augury of fresh trouble in their disturbed and rueful visages. He waited patiently, however, till the speaker had ended, and then answered him, through his interpreter, with great composure. First, he thanked him for the friendly warning which his affection had impelled him to utter; but, he continued, the greater the danger, the greater the honor; and even if the danger were real, Frenchmen would never flinch from it. But were not the Illinois jealous? Had they not been deluded by lies? We were not asleep, my brother, when Monso came to tell you, under cover of night, that we were spies of the Iroquois. The presents he gave you, that you might believe his falsehoods, are at this moment buried in the earth under this lodge. If he told the truth, why did he skulk away in the dark? Why did he not show himself by day? Do you not see that when we first came among you, and your camp was all in confusion, we could have killed you without needing help from the Iroquois? And now, while I am speaking, could we not put your old men to death, while your young warriors are all gone away to hunt? If we meant to make war on you, we should need no help from the Iroquois, who have so often felt the force of our arms. Look at what we have brought you. It is not weapons to destroy you, but merchandise and tools for your good. If you still harbor evil thoughts of us, be frank as we are, and speak them boldly. Go after this impostor Monso, and bring him back, that we may answer him face to face; for he never saw either us or the Iroquois, and what can he know of the plots that he pretends to reveal?[152] Nicanopé had nothing to reply, and,
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