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BRITISH ANTARCTIC EXPEDITION 1907-9
UNDER THE COMMAND OF SIR E. H. SHACKLETON, C.V.O.
REPORTS ON THE SCIENTIFIC INVESTIGATIONS
VOL. I
BIOLOGY
EDITOR - - JAMES MURRAY
PARTS I TO IV
I. ONT COLLECTING AT CAPE ROYDS BY JAMES MURRAY
II. ON MICROSCOPIC LIFE AT CAPE ROYDS BY JAMES MURRAY
III. ANTARCTIC ROTIFERA BY JAMES MURRAY
IV. MUSCI BY JULES CARDOT
(WITH THIRTEEN PLATES AND THREE
FIGURES IN THE TEXT)
LONDON
PUBLISHED FOR THE EXPEDITION 15V WILLIAM HEINEMANX
•21 HKnKOKD STUKKT. W.f.
1910
PRICK TWKI.VK SIHLUN'tiS AND SIX I'KN'CK N K I
THESK PAJITK nr. ALSO ISSI'F.D SF.I'
Ons
/U
PART I
ON COLLECTING AT- -(> a-
4 JAMES MURRAY
from the clear transparent ice, through which on our first arrival we could see the
vegetation grosving on the bottom at a depth of several feet. The largest of all was
Blue Lake, which most nearly deserved the name of " lake." It was nearly half
a mile in length, and filled about half of the valley separating the promontory of
Cape Royds from Mount Erebus. The lake was divided into two portions by a very
narrow strait, known as the "Narrows," in which the depth was only about three
feet. The northern half was deeper, and was found to have a few feet of water under
twenty-one feet of ice. The southern half was frozen to the bottom.
The accompanying sketch-map (Fig. 1) shows the positions of some of the principal
geographical features in the immediate vicinity of the camp. The principal lakes are
marked, and the highest point of land on the promontory, known as " High Peak."
The lower hills are not indicated.
Beyond the valley occupied by Blue Lake rise the lower slopes of Mount Erebus.
This is a region of rocky ridges and moraines like those of Cape Royds promontory,
extending for several miles up the mountain, on the sides of which the moraines have
been traced to a height of 1100 feet. Many small lakes occur up to a height of at
least 500 feet. Northward and southward, at the distance of a few miles, the rock
and moraine give place to crevassed glacier. Beyond the limits of the small area thus
briefly described, snowfield and glacier stretch for many miles, offering no support for
any living thing, unless it be some of those lowliest organisms which can exist on
the surface of the snow itself.
At Cape Barne, two miles south of Cape Royds, there is a district of hill and
valley similar to those of Cape Royds, but the hills are higher and the valleys deeper.
In this region there are many interesting lakes. In one place there are two
concentric curved gullies, both ends of which open to the shore. These are occupied
by lakes, one of which, in the gully nearest the shore, is considerably below sea-
level.
The bay immediately east of the hut was our customary dredging ground. All
the dredging was done within a mile of the Cape, in shallow water, nowhere more
than 100 fathoms in depth. Farther out the Sound deepened to 300 fathoms or
more, but that region remained for ever inaccessible.
Collecting on Land. — It is difficult to imagine a more unpromising field for
biological study than Cape Royds appears on a first examination. Nothing is to
be seen but a succession of ridges of black lava (shattered into loose blocks by the
. intense cold), moraines and snowdrifts, all apparently equally barren.
Near the shore the monotony is relieved by the busy Penguin Rookery, the Skua
Gulls, the Weddell Seals, and an occasional Giant or Snowy Petrel. All these
animals are only summer visitors, except the Weddell Seal, which comes ashore at
all seasons. Apart from these there is no conspicuous life, animal or vegetable, in
the whole area. Indeed, leaving the lakes out of account, the collecting which could
be done on land was of the most meagre description.
ON COLLECTING AT CAPE ROYDS 5
On the triangular promontory no moss or lichen wai ever found. Higher up
among the moraines on the slopes of Erebus dwarfed tufts of moss were fairly
abundant, but so feeble is their growth that M. Cardot,* who examined them,
describes them as "sickly plants, struggling painfully against exceptionally hard
conditions." Lichens were more abundant, though scarcely more vigorous in their
growth, and there were more kinds of them. The commonest was a pretty orange-
coloured kind, closely encrusting the rocks, and crowded with apothecia. The Tripe
de Roche, familiar in books of Arctic travel for its role in prolonging the lives of
starving explorers, occurred in two forms. Its little scales, some half-inch in diameter,
and attached by a single point on the under surface, were scarcely so abundant as to
fulfil the same life-saving role in the Antarctic. The largest-growing lichen was a
tufted kind, growing to an inch or more in height, and resembling Usnea in possess-
ing a fragile cortex, which breaks on pulling the stalks and exposes a tougher elastic
medullary portion. The Tripe de Roche and the tufted lichen could be picked off the
rocks, but most of the other kinds could only be collected by selecting conveniently
small stones on which they were growing. When the snow completely melted from
some of the moraines in the height of summer, patches of the tufted species were
found a yard or more in diameter.
In the valley bottoms, where water had run or at least percolated in summer,
there was a faint scum of green, which consisted generally of dried -up green algae.
On some parts of the moraines, where there was some depth of soft soil, considerable
masses occurred of a plant which seems to be identical with that which abounds in
the lakes, and which will be more fully studied when dealing with them. It is of a
whitish, yellowish, or pink colour, and is in sheets like thin paper, superposed to form
masses of considerable thickness. In some eskers on Blue Lake it was traced to a
depth of several feet, and on some flat moraines it was discovered occupying pockets
underground. In all cases when found on land the plant was friable and had a
bleached appearance. From this fact it is doubtful if it ever grew in the kind of
situation it now occupies. It may have been preserved from a time when these
were parts of lake bottoms.
Collecting in the Lakes. — Soon after landing at Cape Royds, on walking across
the lake afterwards called Green Lake, some thin films of vegetation of a dull green
colour were seen projecting above the surface of the ice. Shortly afterwards Wild
found pieces of a similar plant, but of pink or brown colour, exposed on the surface
of Clear Lake. At the margin of Clear Lake, where the ice was transparent, the
same plant was seen at a depth of about a foot, of much brighter colour than that
exposed at the surface. Pieces of this were cut out with an ice-pick, and taken
home and melted, when several microscopic animals were found.
This was the introduction to the most prolific source of fresh-water life in the
district. The plant was found embedded in the ice of nearly all the lakes, and when
* " Musci," by J. Cardot, in Part IV. of this volume.
6 JAMES MURRAY
the smaller ones melted in summer it was seen that it formed continuous sheets over
the whole bed of some of them. Everywhere microscopic life swarmed on this weed.
The method of collecting during the winter was very simple. A few pieces of ice
containing plants were chipped out and taken to. the house to thaw. When quite
melted the weed was put into a coarse silk net, which was again put inside a very
fine silk net, and the whole immersed in a bucket of water. When the nets were
violently shaken in the water the microscopic organisms, animal and vegetable, were
washed off the weed, and strained through the coarse net into the fine one, from
which they could be easily transferred to a bottle. Thus were obtained multitudes
of living things for study.
In summer the collecting was still simpler. The weed could be washed in the lake
water without the need for preliminary thawing.
In this way we collected continually from the shallow lakes. There were some
deeper lakes, which, as it proved, did not melt in summer. In Clear Lake a hole
was dug through the ice. We came on water at a depth of about four feet. Here
we dredged on the lake bottom, at a depth of seventeen feet. There was a quantity
of vegetation brought up, but it was discoloured and dead, and there was no living
thing upon it.
Late in the winter, Priestley sunk a shaft in the southern portion of Blue Lake,
for the purpose chiefly of observing the temperatures of the ice. At a depth of nine
feet some scraps of weed were got, and when thawed a number of living animals
were found on it. At a depth of fifteen feet we came on the bottom of the lake,
which was here composed of angular fragments. These were covered by a continuous
film of yellow weed, and on this also there were numerous living things.
In autumn an attempt was made to use the tow-net in some of the lakes. The
lakes being at this time covered by a thick sheet of ice, the net could not be drawn
through the water. A hole was dug and some gallons of water taken up with a
bucket and poured through the net. At this time the temperatr"0 of the air was
about Zero Fahrenheit, and the net was soon filled with ice. When thawed out
there was no living thing found in it. In summer, when the lakes were melted, and
the air temperature was about freezing-point, the tow-net could be easily used, and
good collections were got.
The vegetation of these lakes is so important a feature in the biology of the district
that it merits some attention. Portions of it have been submitted to expert botanists,
but no report has yet been received, so it cannot be stated to what group of the
vegetable kingdom it belongs. Its appearance and method of growth will be described.
In Green Lake and many other lakes it is in the form of sheets, from a few inches
to many yards in extent, sometimes continuously covering the bed of a pond from
side to side. It varies in thickness, from one-eighth to half an inch or more, and is
of a consistency like sodden paper, so that it was not possible to lift up sheets of any
considerable size without breaking them. The upper surface is of a bright orange
ON COLLECTING AT CAPE ROYDS
FIG. 2.
DIAGRAMMATIC SECTION OF BLUE LAKE
The part shaded with short vertical lines is the uppermost " prismatic " layer of ice. The
convex upper surface of the lake is indicated, and the irregular thickness of the prismatic
layer. The area marked by little arrows is the clear transparent ice which fills the whole
lake to the bottom. The bed of the lake is of angular fragments, and the dark line over
them is meant to represent the film of vegetation which covers the bottom. At the bottom
of the shaft is seen the geologist chipping out some gravel and vegetation.
8 JAMES MURRAY
colour, and is coarsely and irregularly wrinkled. It is composed of few or many
layers, like superposed sheets of paper. The lower surface of the sheet is of a dirty
green colour, and is composed of a tangled mass of many different algse, green and
blue-green. The whole mass was slimy to the touch.
The mode of growth differs in different lakes. The broad sheets above described
are the commonest form. In Clear Lake it does not form flat sheets, but is coarsely
lobed and undulate, and can be seen through the clear ice growing up from the
bottom. In Coast Lake the lobed character is carried further, and little dendroid
masses of fine lobes can be seen embedded in the ice near the surface. When one of
these is cut out and thawed the plant loses its dendroid character and falls down to
form flat sheets. When the ablation of the ice of Coast Lake goes so far as to expose
part of the lake bed, it is seen to be covered with a deposit of small flakes of the
plant, in colour and appearance not unlike used tea-leaves.
In a lake near Cape Barne the ablation of the ice exposed small masses of the
weed in which the successive superposed layers made up a thickness of six inches.
The layers were very thin and the colour a fine pink.
Large fragments dredged from the bottom of Clear Lake and dried on blotting-
paper had a glossy surface and ash-grey colour like some of the lichens of the genus
Peltigera. In most other samples the surface was dull when dried.
Under the microscope the brown weed is seen to be composed of a felt of very
fine fibres, crossing one another irregularly in all directions. Usually no definite
structure can be detected in the fibres, but Mr. Scourfield noticed some in which an
obscure division into cells could be seen.
In some ponds we found another weed of very similar colour and appearance, but
in very small quantity. This was definitely composed of moniliform rows of cells of
some blue-green alga, very probably of the large olive-green laminae which we got in
some streams and ponds in summer. The similarity of the two suggests that the
commoner brown weed has in like manner originated in the blue-green filamentous
algaa generally associated with it. Plausible though the suggestion is, it requires
expert investigation before we can decide upon it. The filaments (Oscillatoria ?)
seem inadequate to the production of such masses, being to a large extent in the
form of longer or shorter rods (as shown in Plate IV. Fig. 15).
The method of trenching a lake in order to get at the sheet of vegetation at the
bottom is illustrated in the accompanying diagram (Fig. 2).
The shallow lakes were very easily trenched, a few hours' work sufficing to reach
the bottom. The trenching of the deeper lakes, Clear Lake and Blue Lake, was a
more laborious undertaking. The trench is marked out by a draft cut with the ice-
pick, enclosing an area of about six feet long by three feet wide. The whole surface
is then picked over to a depth of a few inches, and the chips are shovelled out. A
very little chipping seems to make a great depth of chips, and frequent clearing out
is necessary, or it becomes impossible to get at the solid ice on account of the loose
ON COLLECTING AT CAPE ROYDS 9
stuff. Thus alternately picking and shovelling, a few inches at a time, the trench
gradually deepens to five or six feet. Up to this time everything is easy. The debris
is easily shovelled out with force enough to carry it well clear of the hole. With
every increase in depth this becomes more difficult, till at last the chips come shower-
ino- back on the worker below. It then becomes necessary to have another man to
pull up the chips in a bucket, or if one man attempts to do everything, he must inter-
rupt his work below every little while, climb out of the hole and pull up a load of
chips. Beyond a depth of five feet it is necessary to construct a stairway. It was
usual to have a few broad steps near the top, and lower down to cut niches for the
feet alternately at one side and the other.
Blizzards are apt to interfere with the work, filling the trenches with snow.
After some experience we learnt wisdom, and covered the trenches with sacking,
which was secured with ice-picks whenever we had to go away for a time. The most
laborious part of the picking was the squaring of the corners. Some of the latest
shafts were made round, and of just the diameter at which a human arm can con-
veniently wield an ice-pick. In these the minimum of material had to be removed.
At depths of fifteen feet and more progress becomes very slow. It is necessary
to have a ladder to get down. If there are two men the ladder can be drawn up out
of the way after one man has gone down. If there is only one the ladder is very
much in the way. The man below is in some danger when the bucket is being
hauled up, as the breaking of the line would let it fall upon him.
Collecting in the Sea. — Mr. Hodgson has given an account * of his collecting at
Hut Point, and he mentions some of the difficulties which attend the collector in
polar regions. Though our location at Cape B/oyds was only twenty miles north of
the Discovery winter quarters the local conditions differ very considerably. The
temperature appears to be usually ten degrees or more (Fahrenheit) higher than at
Hut Point. Being close to the spot where McMurdo Sound opens into the Ross Sea
we had open water close by throughout the year. In fact, even in winter the edge
of the permanent ice was never more than a mile from our camp. Beyond the fast
ice the Sound frequently filled with pack-ice stretching as far as eye could see.
Sometimes, in a period of calm, the pack was cemented into a solid field by new ice,
but this was broken up by every storm, and it was therefore always unsafe to go out
on it. The marginal zone of even the permanent winter ice was liable to be broken
off in a severe storm.
From this cause marine dredging was confined within a very small area. We
could work steadily during winter only in a little bay between Cape Royds and Cape
Barne, where the ice formed early and stayed late. Here, as early as the beginning
of May, the ice was strong enough to allow us to cut holes and put down traps. The
traps were baited, and brought up Amphipods and Molluscs. Some pieces of a den-
droid sponge were entangled in the net and from these we got a number of minute
* " On Collecting in Antarctic Seas." By T. V. Hodgson. " National Antarjt. Exped. 1901-4," vol. iii.
BRIT. ANTARCT. EXPED. 1907-9. VOL. I. B
10 JAMES MURRAY
molluscs and other animals. A storm early in May broke up the ice and our dredging
apparatus went out with it.
On May 11 ice again formed in the Bay, and proved to be permanent, remaining
fast till February of the following year. As soon as the ice was strong enough
dredging was begun. The first dredging-line was put down while the Bay was open,
from the edge of a small area of fast ice which remained near the head of the Bay.
Afterwards we had to take advantage of tide-cracks in order to get lines put down.
It rarely happened that we found the cracks open and could get the line down with-
out labour. Usually they were filled with new ice to a depth of 6 inches or a foot,
and it was by hard labour with ice-pick and crowbar that we got a sufficient length
open to serve for dredging. Foot by foot as the crack was cleared the rope was forced
through, for with the low temperatures new ice quickly forms in the part we have
opened. When the rope was through for a sufficient length it was secured at the
two ends to bamboo poles, enough slack being paid out to allow the ends to hang
nearly vertically, thus avoiding the danger of the rope being frozen in. It was then
necessary to dig holes in the ice at the two ends of the rope, through which the
dredge could be lowered and drawn up.
The holes were from fifty to one hundred yards apart, but the effective dredging
distance was less than that on the ice, as the dredge would always leave the bottom
some considerable time before arriving directly under the opening in the ice. Each time
that we wished to dredge the holes had to be reopened with pick and crowbar. They
would be frozen over with ice from a few inches to a foot or more in thickness,
according to the temperature and the length of time they had been left undisturbed.
In cold weather it was not well to leave them for more than a day, and Priestley
sometimes opened them, although there was no intention of dredging, in order to lessen
the labour next time. The Weddell seals were of assistance in keeping the holes
open. They found them useful as breathing-holes and visited them frequently, some-
times arriving in an apparently exhausted condition, to judge from their laboured
breathing.
In order to avoid dredging too frequently over the same ground it was necessary
to cut trenches in the ice alongside the ends of the rope and at right angles to the
line joining the two ends. In these trenches the rope could be shifted a yard or so
at each time of dredging and so the dredge covered entirely fresh ground. Sometimes
the rope was left too long and got frozen in too solidly to be cut out in the usual way.
A new hole was then cut close by the old one, and the line was fished up by means of
a hook on the end of a long bamboo pole.
The arrangement of the apparatus and method of using it are illustrated in
Fig. 3, in which the Bay where we dredged is shown in section.
The dredge was fixed to the middle of the line so that it could be used in either
direction. It was found that it often caught nothing when travelling downhill, so it was.
usual to haul it downhill and then back again uphill before bringing it to the surface
ON COLLECTING AT CAPE ROYDS
11
FIG. 3.
DIAGRAM ILLUSTRATING METHOD OF DREDGING
The ice (which is supposed to be five or six feet in thickness) and the bed of the sea are
shown in section. The vertical lines mark the sea-ice, and their cessation indicates the
positions of the two holes kept open for the purpose of dredging. One man is shown
hauling the dredge, and another is paying out the spare line to lessen the strain which
tends to lift the dredge off the ground. A few feet in front of the dredge a weight is seen,
which series to keep the dredge down, and at the same time by the length of its attaching
cord maintains it in the right position. The weight was sometimes a lump of kenyte tied in
a cloth, but generally some discarded parts of a motor-car were used.
12 JAMES MURRAY
When the dredge reached the surface both men went to the opening, the one who
had been hauling keeping the line taut to prevent it sinking again. The contents of
the dredge, consisting of a thick black mud in which only the larger objects such as
sponges, shells, blocks of kenyte, and sea-anemones could be distinguished, were
emptied into a bucket. This was made from a 4-gallon kerosene tin provided with
two wire handles. The bucket was filled nearly to the top with sea-water in the
hope of enabling the animals to remain alive for some time. At first it was conveyed
home, a distance of about a quarter of a mile, by slinging it on a bamboo pole carried
on the shoulders of two men. This caused too much splashing, and thej bucket was
thenceforward carried by the handles.
Generally all the water in the bucket was frozen into a kind of soft sludge before
we reached home. It was placed behind the stove to thaw, but so cold was it on the
floor of the hut that it often took a day, or even two days, before it^was ready for
examination.
Any large objects visible on the top of the mud were first taken out. To get out
the smaller organisms the thick coherent mud was taken by a handful at a time and
put in a small silk net having a mesh of about one-sixteenth of an inch. This was
shaken in clean sea-water till the fine mud was all washed away. What remained in
the net was emptied into plates and picked over. The larger pieces -of kenyte and
shells, sea-urchins, &c., were first separated. Then came the task (very trying in the
dim gas-light, which alone penetrated to the biological lab., or the equally dim light of
a hurricane lamp) of picking out the smaller things, minute Crustacea, shells, &c.,
requiring the use of a lens to detect them. All these various objects were sorted
according to their size and kinds, and stored in bottles.
When we had enough jars filled to occupy one of the compartment boxes provided
for the purpose, they had to be removed, as there wasn't room in the hut for them.
The first was put under the floor of the hut, as likely to be warmer than the outside
air, and to escape filling with snow during blizzards. The air-lock under the house
was so very difficult of access that the boxes were afterwards put outside, to take
their chance of cold and snow. A few jars were broken by the cold, but there was no
help for it. The formalin which was used for certain kinds of animals suffered a
change from the low temperature, becoming milky, and did not again regain its
clearness.
When not in use the dredge was left at the bottom of the sea. This kept the
rope soft, , and the dredge was ready for use whenever the holes were opened. The
one-inch lines used lasted throughout the season. This may be partly attributable to
leaving them in the sea and never allowing them to freeze solid. The part never
immersed was air-dried and flexible.
The Bay, which was the only place where dredging was possible during almost
the entire season, was very shallow, the depth varying from seven to eighteen
fathoms. The sea bottom was everywhere covered with a deep layer of very fine
ON COLLECTING AT CAPE ROYDS 18
black mud, in which there were many pebbles of kenyte. While this mud was
favourable to certain forms of life it was unfavourable to others, and thus though
life was abundant it was restricted to a comparatively small number of species.
Shells of the large mud-lo ving Mollusc (Anatind) were very plentiful (thoughwe rarely
found the living animal), as well as ofPecten Colbecki. The large predaceous Gastropod
(Neobucdnum) crowded to any bait put down. Dendroid sponges and a large kind
of yellow Sea-anemone adhered to the shells of Anatina or to the pebbles of kenyte,
and rarely large turnip-like Tunicates came up. Ugly little fishes with enormous
heads (Notothenia) were grubbing closely among the other organisms. In the mud
were numerous worms of many kinds, echini, and multitudes of minute Crustacea, &c.
This shallow muddy dredging-ground was used constantly throughout the winter,
and at intervals afterwards till February 1909. It was not till the beginning of July
that there was an opportunity to dredge over fresh ground. At this time a crack
opened which stretched from the cliff of Cape Royds away towards Cape Barne.
This crack appeared to be caused by the contraction of the main icefield in McMurdo
Sound. At any rate it only opened in cold spells and closed in warmer weather.
When the cracks were open the dredge could be put down and dragged a long way
without any need to cut through the ice.
Near the Cape this new ground was quite as shallow as the Bay but it was quite
free from mud, and the collections differed a good deal in their composition. There were
many loose stones near the shore, but as we extended our operations farther and
farther out we reached greater depths and there were no more stones. The sea
bottom here appears to be covered by a continuous carpet of living things.
The sponges were much more numerous and the siliceous kinds were first obtained.
There were Sea-spiders (Pycnogonids), Lace-corals (Polyzoa), Holothurians, File-shells
(Lima), Alcyonarian Corals, Star-fishes and Brittle-stars, pretty milky white
Nudibranchs (Tritoniella) , and many other things.
The greatest depth at which we dredged in this crack was about eighty fathoms.
The dredge was not left down en the bottom when not in use as we did in the Bay,
because the crack was apt to close at any time, and when it did so the one side of the
floe was often caused to override the other, which would have snapped the line and
lost the dredge.
On July 6, a crack opened from the Penguin Rookery westward out into the
Sound, in which we were able to dredge once at 100 fathoms. Nothing strikingly
different was obtained. In one haul pretty near the shore, the dredge was filled
with the common red Star-fish, and there was almost nothing else.
Collecting Plankton. — Very little plankton collecting could be done in McMurdo
Sound. As the Nimrod passed through the Ross Sea the tow-net was used several
times, being kept clear of the ship by a long boom projecting about ten feet from the
side. The last of these tow-nettings was made as we entered McMurdo Sound. The
boom had then to be unshipped to prepare for the vessel lying alongside the ice-foot.
14 JAMES MURRAY
While she remained for some days moored to the fast ice in the middle of the Sound,
a number of hauls were made with the tow-net used vertically. The net was let
down to a depth of 100 or 200 fathoms and hauled steadily to the surface. These
hauls brought in a good many things, including two kinds of Pteropods, a small
hyaline species, and a pretty red one about an inch in length.
No further collecting could be attempted till we were settled down on shore at
Cape Royds and the ship had gone away. There was at this time open water all
round, but the coast was too rugged to allow of any good tow-netting by throwing in
the nets, and, moreover, we were too busy in other ways. When the ice formed for
the winter it was always unsafe to go to the edge which bordered the open sea.
There were no lanes where the net could be used. The only feasible method was to
attach the net to the dredging-line and pull it from one of the holes to the other,
under the ice. When this was attempted the net came up full of sludge, which
probably came in early in the haul and so prevented much water passing through.
The sludge took a long time to thaw, and when it was reduced to water there was
almost nothing in it. In one such haul we got two phosphorescent Copepods, one
Diatom, and one Peridinium.
The method of taking vertical hauls at the dredging-holes might have been
practised had the depth been sufficient, but at our habitual dredging-ground the
depth was only from seven to eighteen fathoms.
Early in July a crack opened to the west of the Cape. At a distance of a quarter
of a mile from shore the depth was 100 fathoms. Our first concern was to dredge on
this fresh ground, and as the temperature was very low fresh ice filled the crack
before there was time to use the tow-net.
After the Nimrod returned on her second trip south, when the ice had broken
up to beyond Glacier Tongue (twelve miles south of Cape Royds), we noticed a great
many brown bodies, from one to six inches in length, floating in the sea. While the
ship was moored to the Glacier Tongue some members of the ship's crew fished up a
few of them in a tow-net. Some of them, at least, were Ctenophores. The ship's
company also picked up a large medusa (of sorts), but as there were no conveniences
available for preserving it, it was allowed to freeze in a jar of sea-water, and by the
time it could be put in spirit it was in such a condition as to be unrecognisable.
Occasionally shoals of Shizopods (mistaken by the sailors for fish fry) came alongside
the ship.
The Fish-trap. — At the end of August, Day made a fish-trap, at Priestley's
instigation, from Joyce's design, and with the object not so much of adding to
scientific knowledge as replenishing the larder. It was constructed of copper wire,
and was originally of the shape of a water-melon, with a small opening at each of the
poles. From the openings several wires projected into the inside, converging inwards,
so that the fish could easily push them aside to get in, but could not get out again.
The trap was baited and put down in about twenty-five fathoms. After an hour or
ON COLLECTING AT CAPE ROYDS 15
two it was drawn up and two dozen fish found in it. Put down on the same spot for
a time there was nothing caught except star-fish and gigantic red worms.
The trap was used several times, and when put down at a fresh place always
caught plenty of fish the first time used, but few or none afterwards. As the fish
were not appreciated when cooked the trap fell into disuse. The big-headed fish
caught in the trap were good collectors for the Expedition, for in their mouths we
found Isopods and Opisthobranchs, larger and finer than any that ever came in the
dredge, and frequently small fish, to all appearance of the same species as the big
ones.
The fish- trap was the best collector of the giant worms, which came up hanging
from it like long ribbons. As they contracted on feeling the cold, and thickened at
the end inside as well as outside the trap, we could not get them out as complete
specimens. The largest of our star-fish, nine inches across, came into the trap.
,
PART II
MICROSCOPIC LIFE AT CAPE HO YDS
BY JAMES MURRAY
THE finding of an abundant microscopic fauna and flora at Cape Royds came somewhat
as a surprise. It is true that the most northerly lands hitherto carefully examined
(Spitsbergen and Franz Josef Land, in about latitude 80° 0' N.) have a rich microscopic
fauna, but in these lands the higher summer temperature allows of a flora of the
higher plants, and a luxuriant growth of mosses, among which so many microscopic
animals have their haunts. In Grant Land at an equally high latitude there has
recently been found a rich moss flora, which would undoubtedly be found to harbour
plenty of animal life, though this has not yet been reported.
Cape Royds, though at a much lower latitude (77° 30' S.) and close by the open
sea, has a much lower summer temperature. The mean temperature of a summer day
rarely rises above freezing-point, and there is no vegetation higher than mosses. As
contrasted with the northern lands the moss fauna is a very poor one. We found only
four species, and from the whole of Victoria Land there are but eight species known.
At Cape Royds they are very scarce, and are stunted and sickly in growth. The
micro-fauna which they support is very meagre, a few water-bears and rotifers, one
rhizopod, and little else. In some tufts of moss the individual animals were
numerous ; in others no life could be detected.
The kinds of animals which are usually to be found among mosses have at Cape
Royds a shelter of another sort, which, judging from their numbers, appears to suit
them better. This is furnished by the foliaceous vegetation which grows so
abundantly in the lakes and ponds. On the surface and between the layers of this
plant they abound both summer and winter. In summer, when the ponds are
melted, they enjoy for some weeks a warm climate, the temperature rising as high as
60° F. in some ponds. There they are sheltered from the air, which would freeze
them every day if they lived among the mosses. In winter again they are frozen in
the ice for many months, in some of the deeper lakes for many years. While the
mosses appear to be dwarfed by the cold, the microscopic animals are not at all
troubled by the rigours of the climate. When the cold comes they curl up and go to
sleep, it may be for years, and when the thaw comes they go merrily on as though
nothing had happened. Indeed, since the cold does not harm them, the ice
BEIT. ANTAECT. EXPED. 1907-9. VOL. I. PAET 2, ISSUED APEIL 1910 0
18 JAMES MURRAY
preserves them, secure against all other dangers, except only the advent ot
explorers. Their numbers prove how completely they are adapted to the local
conditions. I have never anywhere seen Bdelloid rotifers so plentiful as are the two
dominant species at Cape Royds (Philodina gregaria and Adineta grandis).
Among the higher Invertebrata the Rotifers are easily first in numbers, both of
individuals and species. The Water-Bears are of only a few kinds, but one of them
(Macrobiotus arcticus) is extremely abundant. There are Nematode Worms of two
or more kinds, Mites of several kinds, and two Crustacea belonging to the
Entomostraca. The Ciliate Infusoria are very numerous, there are a good many
Flagellata, but only two Rhizopods were observed.
The vegetation consists solely of Algae, Blue-green and Green, in filaments,
colonies, and single cells. The Diatoms are few and very small, and the Desmids
very rare, only two filamentous kinds being seen.
In this paper a preliminary account will be given of the microscopic life, illustrated
with photographs from life taken on the spot, and with drawings. Some of the
groups, as the Rotifera and Tardigrada, will be worked up and reported upon in
separate papers. Other groups, especially the Infusoria, cannot be worked up in a
scientific manner. Such animals must be studied by a specialist on the spot. Most
of them cannot be preserved in recognisable condition, or to thus preserve them
requires special training and experience. Even were these objections inoperative
there is not time on such an expedition, with a limited scientific staff, to overtake
work of this kind. It is best that the specialist should give as much as possible of
his available time to the line of work which he is best qualified to deal with, and
outside of that do what he can.
Such information as we were able to collect upon the Antarctic Infusoria, &c., in
the form of notes and sketches, will be here presented, without supposing that it can
be of much value to scientific students of these groups. A specialist looking at the
sketches might make a guess at the genera of some of the animals. Lack of know-
ledge prevented observations being made on various organs (mouth, nucleus, &c.)
which are important in determining species.
Photographing the Microscopic Organisms. — The first microphotographs were
made by Mawson, who had some previous experience. Mawson, however, could not
spare much time for this work, since he had all the physical observations to attend
to, as well as the preparations for a long sledge journey. After a few lessons from
Mawson the biologist was able to continue the work when Mawson had left on his
journey.
It was not easy to find an opportunity for microphotography in our crowded
hut. During the day the tramping of feet caused too much vibration. We had
to wait for the hours of the night-watch before anything could be done, and
Mawson gave an entire night-watch to experimenting with different exposures and
plates.
MICROSCOPIC LIFE AT CAPE ROYDS 19
The photographs were all made by acetylene light. This light was quite
unsuitable for ordinary work with the microscope, at any rate with the lenses we
used. We could never get good definition with it. Oil lamps were used, and
though they were of a very poor quality for such purposes, being ordinary stable
lamps with very inferior glass, they sufficed for the work. Photographs were tried
by the light of the oil lamps and also by daylight, but the acetylene proved better
than either for photography. The little daylight which filtered through the frost-
encrusted window was insufficient for microphotography, though it sufficed for
ordinary photography.
In addition to difficulties with regard to light we had to contend against
dust. In our small hut, with its large stove requiring frequent stirring up, its
crowded sleeping accommodation and infinite paraphernalia of fifteen men, cleaning
could be of but a perfunctory character, and there was always a great deal of dust
settling.
The exposure varied, according to light, subject, and especially the degree of
magnification, from two seconds to half an hour. With higher magnifications we got
no satisfactory results, and we ordinarily used a magnification of about 100 diameters,
at the eye, which would give about 200 at the full length of the camera. The eye-
piece was sometimes removed, but the results were not so good. Two plates
were exposed on each occasion and immediately afterwards developed, in order to
check the time of exposure. This varied so much with different subjects that we
could have no general rule. We wished to shorten the time to the utmost, as nearly
all the animals to be photographed were alive, and liable to change their positions.
They were under the influence of a mild narcotic which quietened them down a
little, but which had to be so weak as to permit them to feed in a natural way.
Exposures of five, and even of two seconds were used, but they were too short, and
the average was about half a minute. As the whole of a subject could never be in
focus at one time it was customary when using long exposures to gradually alter
the focus, with the view of giving every depth a chance. This was done too much
by guess to be a conspicuous success, but in some instances it gave a clearer
outline for the whole body of a thick animal than would have been possible
otherwise.
The photographs, as such, are very poor, but since they have an interest as
pictures from life of subjects which cannot be seen anywhere else but in the
Antarctic Region, their shortcomings will perhaps be overlooked. Unfortunately,
through some mischance the best negatives have been mislaid, and are not available
for reproduction.
Crustacea. — In ice from Blue Lake, shortly after we landed, there were found
immature examples of a Calanid of small size. It was "hot in condition to be iden-
tified, and we hoped to find the same species alive and mature when the lake melted,
so the specimen was not mounted. As it happened the lake never melted and we
20 JAMES MURRAY
never saw another example. It is possible that the species is not a native of the lake,
which is situated only a few hundred yards to the north of a bay of the sea, and
it is conceivable that in a severe summer gale microscopic Crustacea and other
organisms might get caught up with the spray and blown into the lake. Against
this is the fact that Blue Lake is the freshest of all the lakes, and the water from it
is as fresh as condensed water. No organism definitely recognisable as marine
was ever got in it.
The only other Crustacean found, a small Harpacticid like Canthocamptus, was
also got in Blue Lake. It was never found in the shallow part of the lake, called the
Narrows, but only at a depth of from 9 to 1 5 feet below the surface in a shaft sunk
by Priestley in July 1908. On July 13, we found a skin on some scraps of weed at
a depth of 9 feet. On July 17, on a film of weed covering the gravel of the bed of
the lake at a depth of 15 feet, another skin was got. There is more probability of
this Crustacean being a native of the lake than the Calanid, as its relatives are not
free-swimmers, but creep about on various plants. No drawings or photographs of
the Crustacea were obtained.
Mites. — In Coast Lake, Blue Lake, Clear Lake, and Deep Lake (near Cape
Barne) skins of small mites were got. During our stay in the Antarctic, none were
seen alive, but after our return to England a living mite was obtained from vegeta-
tion brought from Deep Lake. It had probably hatched from an egg. In Coast
Lake they were abundant, and of several species. No drawings or photographs were
made, but a specimen of one kind was mounted, and it is hoped that enough
examples will yet be got to enable us to report upon them.
Insect. — On Plate IV., Fig. 16, is the photograph of the only Antarctic insect
which we obtained, a parasite on Maccormick's Skua. Two examples were got by
Joyce on one Skua, and it appears to be rare, as very careful search failed to find
any others. On an Emperor Penguin also a small louse was seen, but the specimen
was lost.
Other Arthropods. — In the lakes we occasionally found fragments of Arthropods,
but whether of Crustacean, Insect, or Acarid we did not find out. A probable explana-
tion of such occurrences is that they were parts of small marine Crustacea, brought by
penguins as food for their chicks, and blown into the lakes.
Worms. — In addition to the Kotifera we found worms belonging to several other
groups — Gastrotricha, Turbellaria, and Nematoda.
Gastrotricha. — This small group, supposed to stand near the Rotifera, was repre-
sented by a single example of Chcetonotus found among weed from the Deep Lake
at Cape Barne.
Turbellaria. — Microscopic Turbellaria were found in Coast Lake and Blue Lake.
In the former they were at one time very abundant.
Nematoda. — The Nematodes of the lakes were microscopic and free-living. Two
MICROSCOPIC LIFE AT CAPE ROYDS 21
kinds were common, one of which is figured on Plate IV., Fig. 13. Another kind
had the skin minutely annulate.
Injusoria. — In dealing with the lowest and simplest forms of life, which are
easily carried about in the form of dust, there must always be some doubt as to
whether many of the species are native or introduced by the expedition making the
observations.
When we arrived at Cape Eoyds the season was well advanced towards autumn,
and nearly all the lakes were already frozen. When we cut out blocks of ice
containing portions of the vegetation which so abounds in these lakes we found many
kinds of Infusoria, some of them of large size, dead and embedded in the ice. These
were undoubtedly native. Afterwards, when the lakes melted and living Infusoria
appeared in them, we were able to recognise many of them as of the same kinds which
we had previously found frozen into the ice. Some of them were encysted and
probably alive when found in the ice, but we never observed any of them leaving the
cysts while under observation.
One of the puzzling organisms which we first observed consisted of clusters of
whitish elliptical bodies, in which no definite organs could be seen. They were
supposed to be some kind of eggs. Long afterwards they were accidentally dis-
covered to be Vorticellids. On treating with formalin an "infusion" in which a kind
of Vorticellid abounded, it was found that they contracted into the puzzling egg-like
bodies.
There is little doubt that most, if not all, of the Infusoria and other organisms
hereafter figured were true natives of the lakes of Cape Royds. The number of kinds
seen was much greater than the number noted and figured. Very many were seen
at times when important observations were in progress, which allowed of no time
being given to side issues. The Flagellata, on account of the greater difficulty
attending their study, were generally passed over without note.
Rhizopoda, — The paucity of Rhizopods at Cape Royds was surprising, after it
became known that so many other kinds of microscopic life abounded in the lakes.
In the lakes only two testaceous species were observed : the well-known Difflugia vas,
and a very small kind which appears to be a Quadrula. Among the moss there was
another species, not identified.
On one occasion, when the ice of a lake was melted, we found numbers of an
amorphous granular organism, each with a round nucleus, which were probably
Amoebae, but being only seen dead nothing could be made of them.
When our material is submitted to a specialist it is expected that he will find
other forms which we have overlooked.*
Heliozoa, — In Coast Lake in April 1908, there occurred an animal which appears
* Dr. Penard has, in fact, detected about a dozen species, which will be reported upon in a subsequent
paper.
82 JAMES MURRAY
to be a Heliozoan (Acanthocystis ?). The spines are all of one kind, and they are few
in number and of clavate form (Plate VIII. , Fig. 23).
Microscopic Plants. — The Algae, the majority of which are microscopic, will be
reported upon in a separate paper.
Bacteria. — Many kinds abounded in the lakes in summer ; but in this group it is
impossible to discriminate between native and introduced kinds.
Organisms of Doubtful Position. — Many organisms occurred, of whose nature
we had no guess. Certain of these are figured on Plates VII. and VIII. Perhaps
the sight of these sketches may suggest some of their names to other naturalists.
Acinetaria. — No tentaculiferous Infusoria were definitely recognised, but while
these pages are in the press, Mr. A. W. Sheppard, F.R.M.S., has identified the
organism figured on Plate VII., Fig. 19, as the encysted condition of a species of
Podophrya.
PLATE I
PLATE I
MICROPHOTOGRAPHS OF A ROTIFER
PhUodina gregaria, sp. n.
FIGURE 1. — Portion of the field of the microscope, crowded with the Rotifers, under
a low power. These examples were brought by Priestley from a lake in the
west. They were frozen solid when brought in. As soon as they were thawed
some were transferred by a pipette to a slide and photographed. They were
not narcotised, and were beginning to stretch themselves and creep about, so
the exposure had to be very short. Many of them have moved and are blurred
in the photograph.
FIGURE 2. — One of the Rotifers feeding. It is under the influence ot Eucaine,
but its form has been little affected by it, and is nearly normal. The dark
column in front of the head is formed of an accumulation of minute particles
which have been swept towards the mouth by the cilia of the discs. The
little clouds of particles on each side of the neck are those which have been
rejected by the selectors in the gullet, while those chosen for food have passed
down to the jaws.
FIGURE 3. — Side view of an extremely large example. It has been narcotised and
the constrictions bounding the turgid central trunk have been much deepened
by the influence of the Eucaine on the muscles. (Photograph by Mawson.)
FIGURE 4. — A Rotifer under slight pressure, to show some of the internal organs
and the well-grown young. The dark central tract was coloured deep red,
and the eyes, being also red, show more conspicuously than in life.
BRIT. ANTARCT. EXPED. 1907-9.
VOL. I. PLATE I.
MURRAY: MICROSCOPIC LIFE
n-: 1.
FIGURE 2.
FIOUIIE 3.
FIGURE 4-.
PLATE II
BRIT. ANTARCT. EXPED. 1907-9. VOL. I.
PLATE II
MICROPHOTOGRAPHS OF ROTIFERS
Adineta, Hydatina,' Diaschiza
FIGURE 5. — Adineta grandis under the influence of Eucaine. The trunk and anterior
part of the body are of the normal form. The Eucaine has caused the foot to be
drawn in, which would not usually be the case in the natural condition. The two
examples, of which portions are seen at the foot of the photograph, show the
deep constrictions of the neck which are caused by the influence of the Eucaine
on the muscles of the trunk.
FIGURE 6. — A very large example of Adineta grandis. It is under slight pressure,
so that the foot could not be drawn in. The head and neck have been moved
while the photograph was being taken, but the central trunk shows clearly the
six contained young.
FIGURE 7. — Hydatina senta, side view. This is the only free-swimming Rotifer which
we obtained in the Antarctic. It appeared in Coast Lake in summer. The
photograph is taken from a living example, under the influence of Eucaine, which
causes the cilia to move very slowly. The natural shape was not affected by the
narcotic in this instance, though it frequently causes some distortion.
FIGURE 8. — A small Rotifer, probably Diaschiza tenuior, seen from the side. It was
narcotised, and was showing the natural form very well when the plate was put
in, but at the very moment of exposure the body has become turgid, which is an
indication that death is imminent.
Close under the head of the Rotifer can be seen one of the small Diatoms which
are common in the lakes.
BRIT. ANTARCT. EXPED. 1907-9.
VOL. I. PLATE II.
MURRAY: MICROSCOPIC LIFE
r
FIGURE 5.
FIGURE 6.
v
FIGURE 7.
FIGURE 8.
PLATE III
PLATE III
MICROPHOTOGRAPHS OF WATER-BEARS
FIGURE 9. — Side view of the common Antarctic species of Water-bear, Macrobiotus
arcticus, which is also found in the Arctic Region. In the photograph may also
be seen a Eotifer and some filaments of Algae.
FIGURE 10. — Dorsal view of another example of the same species. Four eggs at an
early stage of development can be seen in the body. To the left of the animal is
one of the eggs of the species, but it is so encumbered by debris that its
characteristic peculiarities cannot be seen.
FIGURE 11. — A different species of Water-bear (Macrobiotus oberhduseri] found among
moss at Cape Royds. It has one excessively long claw on each foot, and the body
is marked with a warm brown colour, forming longitudinal and transverse bands,
indicated in the photograph.
Mb />*' i> i^*£
FIGURE 12. — Another example of Macrobiotus arcticus, containing six eggs not very
far advanced in development. In the anterior part of the body the fat-cells in
the body fluid can be distinguished.
BRIT. ANTARCT. EXPED. 1907-9.
VOL. I. PLATE III.
MURRAY: MICROSCOPIC LIFE
FlClL-RH 10.
'*'
FIGURE 11.
FicniK U.
PLATE IV
PLATE IV
MICROPHOTOGRAPHS OF NEMATODE, ALG^E, AND LOUSE
FIGURE 13. — Part of the field of the microscope under a low power, showing many
kinds of organisms. The largest is a Nematode of the commonest Antarctic kind.
The pear-shaped pharynx can be seen. There is a Water- bear and the cast skin
of another, a contracted Rotifer, and some Algae in round colonies. Near the
upper right-hand corner is a large egg, which is probably that otHydatina senta.
(Photograph by Mawson.)
FIGURE 14. — Fine filamentous Alga from Pony Lake. This is probably an Oscillatoria,
but is different from the kind most commonly found in the other lakes.
FIGURE 15. — A short rod of the common blue-green Alga (probably Oscillatoria),
highly magnified. Beside it are some narrower filaments of another kind.
FIGURE 16. — Photograph from life of the only parasite which we obtained from the
feathers of Antarctic birds. Two examples were got on a Skua Gull (Megalestris
Maccormicki). Though not so purely microscopic as the other animals photo-
graphed, it is a very small species.
BUIT. ANTARCT. EXPE1). 1907-9.
VOL. I. PLATE IV.
MURRAY: MICROSCOPIC LIFE
FlGI'KK 13.
FIGURE 14.
FIG IT UK
FlCiUUK 1(>.
PLATE V
PLATE V
CiLIATE INFUSORIA ; VOET1CELLA AND SIMILAR FORMS
FIGURE 1. — Vorticella, a narrowly pyriform species, with slender contractile stalk,
which can extend to five or six times the length of the body. The whole body
was covered by papillae, as in V. monilata Tatem, but the shape of the body is
different.
FIGURE 2. — Vorticella, a species of conical form, widest at the mouth, and with a con-
striction separating a basal portion, one-third of the total length. There is only
a short broad stalk, but it may have been broken off.
FIGURE 3. — Vorticella, mouth not expanded.
FIGURK 4. — Vorticellid, narrowly oblong ; free-swimming example, with posterior
ring of cilia and no stalk.
FIGURE 5. — Vorticella, long narrowly oval species, with narrow mouth.
FIGURE 6. — Vorticellids, two contracted individuals on one stalk.
FIGURE 7. — An animal of doubtful position, resembling the Vorticellids in possessing
only a circlet of cilia round the mouth.
FIGURE 8. — Vorticellid, another free-swimming example.
Brit Antarct. Exped 1907-9.
MURRAY; MICROSCOPIC LIFE.
Vol. 1. Plate V
J. Murray del ad nat.
PLATE VI
BRIT. ANTARCT. EXPED. 1907-9. VOL. I. E
PLATE VI
CILIATE INFUSORIA
FIGURE 9a. — A large reddish brown animal, common in all the lakes. It is covered
all over with short cilia, which appear to continue beneath the surface as short
rods, making a thick skin. A bundle of pbaryngeal rods forms a kind of narrow
cage, tapering downwards, as in Nassula, &c.
FIGURE 96. — Probably the same animal as Fig. 9a, at a different stage in its life-
history. The cage of pharyngeal rods is of the same form, and projects far above
the surface. The skin is thin, and not ciliated. Many of these colourless
examples were found at the same time.
FIGURE 10. — A narrow curved animal, narrowing to one end. It is marked longitu-
dinally with faint ridges, which are seen in profile to be finely undulate. It is
ciliated on all the ridges, the cilia being long and not very close together.
They gradually elongate to the broad end. All are motile but only those on a
limited area move actively. No mouth could be detected.
FIGURE 11. — A ciliate with projecting crest ; no other details noted.
FIGURE 12. — Ciliate with a beak, a series of strong cilia at the mouth, and another
series round half the circumference of the body.
FIGURE 13. — Large oval ciliate, ciliated all over, and with a vibrating membrane
occupying an elliptical area. The cilia appear to be continued under the skin
as rods (as in the species figured in 9a).
FIGURE 14. — A dark brown ciliate, with cilia all over the body. It is elliptical in
form, with the poles produced into rounded processes. No mouth or other
organs could be seen. As it travels along in the direction of its long axis it
at the same time revolves round it.
FIGURE 15. — An oval ciliate, marked with longitudinal ridges. There are four
strong setse at each end, a row of motile setae half-way down one side, and
a single seta on the same side.
Brit Antarct. Expect 1907-9.
MURRAY: MICROSCOPIC LIFE.
Vol. 1. Plate VI
9. a
10.
15
M.rray del ad nat.
PLATE VII
PLATE VII
FLAGELLATA; AND VARIOUS ORGANISMS OF DOUBTFUL POSITION
FIGURE 16. — An oval flagellate with green granular contents. One strong
flagellum was seen.
FIGURE 17. — A cluster of four cells, with one flagellum visible on each. The cluster
rotated rapidly with an irregular motion. The contents were green.
FIGURE 18. — A colourless flagellate, of ovate form with a long tail: one flagellum
seen : contents granular.
FIGURE 19. — A hyaline, almost spherical body, set on a slender stalk. The body is
marked with ten prominent annular ridges, about equidistant from one another,
except near the pole to which the stalk is attached, where they are crowded.
The protoplasmic contents form a globular mass much smaller than the envelope.
Identified as an encysted Podophrya, by Mr. A. W. Sheppard, F.R.M.S.
(see p. 22).
FIGURE 20. — A soft orange -coloured ciliated body issuing from a small opening in
a hyaline shell.
FIGURE 21. — A cluster of oval bodies of sienna-brown colour, in a darker brown
matrix.
FIGURE 22. — A soft thick-skinned animal, finely ciliate all over. Little internal
structure could be seen except two cavities occupying the greater part of the
interior. The granular part between the cavities is sienna-brown in colour.
Two vibrating membranes were seen, like those which are found in various
sorts of worms.
Brit Antarct. Exped 1907-9.
MURRAY MICROSCOPIC LIFE
16
17
Vol. 1. Plato VII
20
19
21
22
J. Murray del ad nat.
PLATE VIII
PLATE VIII
VARIOUS ORGANISMS OF DOUBTFUL POSITION
FIGUEE 23. — Possibly a heliozocm (Acanthocystis ?). The spines are few and of one
kind only, clavate in form. Plentiful in Coast Lake.
FIGURE 24. — Oval bodies of a dingy purple colour, always occurring in clusters, as
many as eight together. They always included many round pellet-like bodies.
FIGUBE 25. — Apparently a bit of vegetable tissue. Such a plant must have been
introduced, but if so it must have been previous to our expedition, as it was
found in the ice of a lake which was frozen when we arrived.
FIGURE 26. — An elliptical hyaline body, made up of eight rings.
FIGURE 27a. — A brown discoid body with finely reticulate surface. One pole is pro-
duced with a little point. There is no opening of any kind. Fig. 27b shows a
bit of the reticulate surface more highly magnified.
FIGURE 28. — Round vegetable cells, in irregular clusters and strings. A few were
filled with green contents, like two of those figured ; most had a little shrivelled
dark mass as in four of the cells figured ; the empty cells were open, as shown
in the two cells at the ends of the series.
Brit Antarct Exped 1907-9.
MURRAY MICROSCOPIC LIFE.
Vol. 1. Plate Vlll.
23
25
27
26
27 fe
ad nat.
PART III
ANTARCTIC KOTIFERA
BY JAMES MURRAY
AMONG the various forms of freshwater life the Rotifera were most conspicuous.
They were found among mosses and in the lakes and ponds. In the former situation
they were relatively scarce, while in the lakes they were extremely abundant, and
were often associated in such numbers as to affect the colour of the water.
Some sixteen species were distinguished, representing three of the recognised
orders, and five or six distinct families. The predominance of the Rotifera was
entirely due to the Bdelloida, of which a dozen species were found. In the other
orders there were only one or two members of each family present.
Five of the Bdelloids are species previously unknown. Most of the others differ
more or less from the types of their species. Only two non-Bdelloids species were
recognised (Hydatina senta and Diaschiza tenuior), the others were only assigned to
their genera. No Rotifera were found in the sea.
I have been able to find no record of any species of rotifers found on the Antarctic
Continent or on any of the islands which, though lying outside the Antarctic Circle,
possess a polar climate. The German Expedition found Rotifers on the Gaussberg,
just on the Antarctic Circle (Richters, 31) ; * the Swedish Expedition obtained them
at Snow Hill Island, not far from the Circle (Richters, 32), and the Scottish Expedi-
tion at the South Orkneys, a little farther north (Murray, 26). t All of these were
unrecognisable.
The only species doubtfully identified from this whole great area is Callidina
papillosa, Thomp. (40). Richters found an egg in moss from the Gaussberg which
resembled the egg of C. papillosa figured by Jansou (20). The identification of this
egg can never be certain, as there are several other Bdelloids which have similar eggs,
covered with blunt processes.
To find definite records of Rotifera in the southern hemisphere we must go far
* Figures in heavy type enclosed in brackets refer to list of books at the end of the paper.
t The Rotifer vulgaris casually recorded in that paper must be regarded with doubt. As all the others
•were dead, it had probably been accidentally introduced into the bottle.
BRIT. ANTARCT. KXPED. 1907-9. VOL. I. PART 3, ISSUED APRIL 1910 F
42 JAMES MURRAY
beyond the Antarctic Region. A few species have been found in Gough Island
(Murray, 29), and Amsterdam Island (Richters, 31). Hilgendorf (19) has published
a list of over forty species from New Zealand, in about latitude 43° S., and these are-
the nearest neighbours to the Antarctic Rotifers of which I can find any notice.
Order— BDELLOIDA
Family — PHILODINAD^E
Genus — Philodina
Philodina gregaria, sp. n. (Plate XL, Figs. 7a—7e)
Specific characters. — Large, corona large, narrower than trunk, wider than collar,
space between discs wide, the rounded lobes springing from collar large, the interspace
with two smaller convexities ; collar very prominent, but scarcely marked off from
pedicels; rostrum short and broad; antenna short; eyes large, pale brown; teeth 2+1/1+2;-
foot of four joints, spurs slender, acute, moderately divergent, separated by broad flat
interspace; viviparous; found aggregated in great numbers, forming larger or smaller
patches.
Detailed description. — Size variable ; smallest about •£$ inch long, in the feeding
attitude (= 416 M) ; longest measured ^ inch (= 800 /*) when fully extended creep-
ing : a fairly large example had the following measurements, diameter of corona y^
inch (= 166 yu), of collar y^ (inch = 132 /*), of neck -^ inch (= 106 /*), of trunk
yyT inch (= 213 ft), length of jaw ^fa inch (=* 47 M). of spur ygVs inch (=25 /*).
Central setae were not detected on the discs. The very prominent collar passes
insensibly into the pedicels, the junction marked only by the line of fine cilia
(secondary wreath). The rostrum bears short lamellae, a brush of active cilia, and
four stout straight setae, corresponding to the tactile setae of P. brevipes (25) and
some other species, but they were not seen in motion. The length of the antenna is
about equal to half the diameter of the neck segment which bears it.
The brain is fairly large. Each jaw bears two strong teeth and one thin one, and
in addition the usual fine striae. There is a prominent hook at the back of the ramus.
The stomach is very voluminous, and of a very bright deep ruby or crimson colour,
due to the presence of coloured granules and globules. The central trunk is the
broadest part of the body, and varies in size with the number of young carried. It is-
regularly plicate, the folds shallower on the back, deeper on the sides.
The reproduction is viviparous. Usually there are two or three young carried at a
time. Whatever the number, they appear to be all at the same stage of development.
At any rate they are all of the same apparent size, and after the jaws are developed
they appear to be alike in all. It is very different with the genus Rotifer, in which.
ANTARCTIC ROTIFERA 43
the young are obviously of very different ages, one being fully developed while another
shows no detail at all. The yolk mass contains eight nuclei.
The spurs vary greatly in size, generally most closely resembling those of P. acuti-
cornis (25). The interspace is relatively broader. Occasionally they are very long
and apparently two-jointed. The last foot-joint is long. The ventral toes are very
large, and the dorsal ones very small.
The vibratile tags are short, and broadly spindle-shaped. Three pairs have been
seen.
Habitat. — In lakes and ponds at Cape Royds, Cape Barne, and at the Stranded
Moraines on the west side of McMurdo Sound. It was absent from the very saline
Green Lake, but was in nearly all the other waters examined. It was much the most
abundant species in the district. Its abundance in Coast Lake and in the lake at
the Stranded Moraines was remarkable. In winter it was got by digging out ice
containing plants from the lakes. As soon as these were thawed the rotifers were
found active in great plenty. In summer blood-red patches began to form on the
stones at the margin of Coast Lake. These attained to a diameter of an inch.
Similar patches were on the plants, but these were more difficult to detect owing to
the orano-e-red colour of the plants. In the lake at the Stranded Moraines, Priestley
reports that the patches reached to six or eight inches in diameter, and were of
appreciable thickness. These patches were formed solely of P. gregaria, which were
fixed side by side, as close as they could stand.
To obtain them a handful of weed was taken and washed in a bucket of water,
being vigorously shaken in order to detach all the adherent microscopic organisms.
The sediment thus obtained was strained through a coarse silk net, in order to remove
the larger particles, and the fine sediment was then bottled and allowed to settle. At
first it was of a dull green colour, from the preponderance of blue-green Algae. After
an hour or two a red film, like blood, appeared on the surface of the mud. The rotifers
have crept out of the mud. After a time they leave the mud and creep up the sides
of the bottle into the clear water above ; eventually they reach the surface of the water
and there form a ring of red round the bottle. They may then be collected in
thousands with a brush and put into clean water. This process may be continued
with one lot for days, an hour or two being sufficient time for new hosts of the endless
procession to reach the surface.
Habits. — P. gregaria is ordinarily rather a restless animal. It is ready enough
to feed, and remains fixed in one spot for a long time, but it swings about continually
so that it is not easy to get a good portrait. Though normally anchoring itself, in
company with its myriad neighbours, it occasionally casts off" and goes swimming. In
Coast Lake it was got in the tow-net, and a few might be seen if a bottle of the lake
water was held up to the light. According to Priestley it was much more plentiful
in the water of the lake at the Stranded Moraines.
It was particularly amenable to treatment with mild narcotics. When a very
44 JAMES MURRAY
dilute solution of Eucaine was added to the water, it at once changed both habits and
appearance. It ceased its restless swaying about and went on feeding so steadily
that it could easily be photographed. The circular muscles were slightly contracted,
thus deepening the constrictions between the principal divisions of the body. The
foot was inclined to be further retracted than in the normal condition. Otherwise
there was little change in form, and the corona was quite unaltered. The effect of the
narcotic was much less than on P. laticeps. When that species is treated with Eucaine
it partially retracts the foot, expands the usually narrow central trunk, and reduces
the width of the corona, in fact it so completely alters its proportions that it is not
recognisable for the same species, and might be mistaken for some species of the central
group (P. citrina, P. brevipes, &c.) unless particular attention were paid to the spurs.
Affinities. — P. gregaria belongs to the central group of the genus, possessing eyes
and tapering spurs of moderate size. There is nothing distinctive in the general
form. The viviparous reproduction distinguishes it from all the species of that group.
The red colour is differently distributed from that of P. roseola, in which the red is
diffused. In this species it is limited to the stomach. The slender spurs, with broad
interspace, are like those of P. laticeps and P. acuticornis. The short antenna
separates it from both. It is of larger size than any of the other species in the
central group, except perhaps P. citrina.
The large size, red stomach, viviparous reproduction, and slender spurs set far
apart, will distinguish it from all known species of Philodina. The absence of a
groove between the prominent collar and the pedicels is also a good character.
Natural history. — As the dominant species in the lakes of Cape Royds the
natural history of P. gregaria received a good deal of attention, and many experi-
ments were made to elucidate it. These will not be detailed here, but a short
summary of the facts will be given.* Its extraordinary abundance must indicate that
it is possessed of great powers of resistance to all the adverse influences which would
be supposed to attend upon it in such a rigorous climate, or else that it is of remark-
able fecundity. It appears to triumph in both ways.
It is perhaps inaccurate to call it "gregarious." It is found in large " flocks," but
it is doubtful if they ever " flocked " together. The great crowds in which they occur
appear to arise from the rapidity with which they reproduce themselves. Several
young, probably sometimes as many as six or eight, are produced at a time, and they
seem to stay and fix themselves where they are born. Thus the patches increase till
they reach inches in diameter, and as there is not foothold for all, they stand on one
another's heads (so to speak) till a layer of measurable thickness is produced.
They withstood all the tests applied to Adineta grandis except the heating, which
was not tried on them. They are normally frozen in the ice of the lakes for the
greater part of the year, and revive at any time that the ice is thawed. When dried
and exposed to the lowest air temperatures for a long time, they were not killed, nor
* The detailed account of the observations and experiments will be found in a paper on " Life under
Polar Conditions," in a later number of this series of Reports.
ANTARCTIC ROTIFERA 45
did they die when alternately thawed and refrozen at weekly intervals for several
months. They lived for a month in sea-water and in a much more saline solution, and
became active again immediately on being transferred to fresh water. They were
dried while in the Antarctic by exposing to the air till all the ice passed off by ablation,
' and were then conveyed by a long voyage through the tropics to England, where they
revived within an hour of being moistened and could be seen alive in London a year
after they were collected.
In England they were subjected to a temperature of — 78° Cent, for many hours,
by Mr. J. H. Priestley, of Bristol, and survived.
Philodina antarctica, sp. n. (Plate X., Figs. 5a-5c)
Specific characters. — Large, elongate : corona of moderate breadth, wider than
the prominent collar, discs with central papilla?, each bearing several fine setae ;
antenna long : teeth 2/2 : foot four-jointed, long, slender ; spurs with broad triangular
basal portion, and narrow apical portion of about equal length (Fig. 5b) ; last joint
of foot short, dorsal toes nearly as large as the ventral ones : pale brown eyes.
Detailed description. — The size is variable. The shortest measured was 380 /* in
length, in the feeding attitude, and with the foot well drawn in. The longest
measured 714 n, fully extended, creeping. In a large example the diameter of the
corona was 96 n, of the collar 73 M, of the neck 60 p., of the trunk 106 n : the length
of the spur was 30 /x.
The stomach is of a deep ruby-red colour, the anterior part of the body of a
faint brown, and the foot clear and hyaline. The longitudinal folds of the trunk
are deep at the sides and shallow on the back. The discs are separated by a space
equal to half the diameter of a disc. Into this come the low rounded lobes terminat-
ing the collar. They are slightly separated, and the part between is convex. The
collar appears two-lobed in dorsal view, and is distinctly marked off from the pedicels.
The rostrum is short and broad. The length of the antenna is equal to the diameter
of the neck segment bearing it. The reproduction is unknown. Neither eggs nor
embryos were ever seen. The yolk mass has the eight nuclei usual in the order.
The vibratile tags were not detected.
It was an extremely difficult animal to study, on account of its restless disposition.
It went wriggling and twisting and creeping about, often stopping to feed for a
moment, but never still. There were no narcotics available when it was first found
in considerable numbers, and it afterwards proved to be a rare species, and very
uncertain in its occurrence. For this reason no photographs were obtained, and
no specimens could be preserved.
Habitat. — Among plants in the ice of several lakes at Cape Royds and Cape
Barne. It was never got except by thawing the ice of the lakes. Most of the lakes
in which it lived did not melt in either of the two summers we spent in the district.
Affinities. — Belonging also to the central group of species, there is little in its general
proportions and characters to distinguish it from several other species. It is more
46 JAMES MURRAY
elongate than most of them, and in that respect comes nearest to P. erythrophthalma
Ehr. (15). The characteristic spurs, consisting of a narrow blunt apical portion
springing abruptly from a broad conical base, will separate it from that and all other
known species when well developed. Sometimes the narrow part is considerably
reduced. No other member of the genus has such spurs, but very similar spurs,
differing only in being sharper pointed, are possessed by Callidina hexodonta, Ber-
gendal (3), formerly regarded (from the possession of cervical eyes) as belonging to
the genus Philodina.
The slender foot, which can be elongated much more than the drawing (Fig. 5a)
shows, the lack of interspace between the spurs, and the deep red stomach are
also good characters.
Philodina alata, sp. n. (Plate X., Figs. 4o-4/)
Specific characters. — Size moderate : corona broad, diameter about equal to that
of the trunk (exclusive of the processes) in ordinary extension ; collar inconspicuous,
the lobes going to the upper lip not reaching beyond the line of the bridge joining
the pedicels : interspace between discs equal to diameter of disc ; central setae on
discs : teeth 2/2 : trunk plicate, bearing two large rounded lateral processes (one on
each side) a little way in front of the widest part of the central segments : rump
short, with a rounded boss in the middle of the preanal segment : foot short, four-
jointed, spurs diverging, tapering, conical, rather blunt. Eyes brown.
General description. — Length 300 /t, in the feeding attitude. The stomach is of
the same deep red as in P. gregaria, the colour being seated in small granules and
globules. The rostrum is short and rather narrow. The length of the antenna is
about equal to the diameter of the neck. On the fairly large brain are seated the
pale brown eyes. The jaws are rather small, and bear two teeth each. The trunk
is regularly plicate, the dorsal folds being wider and shallower than the lateral ones.
The trunk is sometimes decidedly viscous, and has extraneous matter adhering to it,
but this is not always so. It is sometimes quite clear.
The lateral processes are not thickenings, like most of the trunk processes of
Bdelloids. They are large, hollow, approximately conical protuberances, with skin
no thicker than that of the trunk, unless at the extreme apex, where it is a little
thicker. They are controlled by special muscles, by which the apex may be more or
less pulled in and inverted, making the form truncate, as shown in Fig. 4a. Some-
times, when creeping or feeding, the processes are so far inverted that they are
scarcely visible in dorsal view, but this is not always the case, as shown in Fig. 4c,
where they are fully extended when the animal is creeping. In complete con-
traction of the animal they have their greatest projection, and are then more
directed forward than at any other times.
No suggestion as to the function of these curious processes has been offered. In
other Bdelloids having warts on the body they are supposed to have a defensive
ANTARCTIC ROTIFER A 47
function. It can hardly be so with P. alata, as the processes are, from their form,
more vulnerable than the trunk wall would be.
The boss on the preanal segment is of another nature. It is a thickening, and
may have a protective function.
The vibratile tags are short and somewhat broadly spindle-shaped (Fig. 4e).
Habitat. — Among weeds from the Narrows between the two portions of Blue
Lake, pretty numerous, March 27, 1908. Afterwards found in several lakes at Cape
Royds and Cape Barne, always scarce and uncertain. A few could usually be got
when wanted from the Narrows of Blue Lake.
It was living, among scraps of weed, at a depth of 1 1 feet in Blue Lake.
Habits. — A quiet animal and slow in its movements. When feeding, which it
rarely did, it could be easily studied. Under the influence of a narcotic (Eucaine)
it behaved in a very unsatisfactory manner. It kept the corona expanded and
continued feeding, but contorted itself till it was not recognisable except by the
side processes. No good photographs of it could be obtained, but under pressure
some were made which showed the internal structure and the characteristic processes.
Affinities. — P. alata, like the other two Antarctic members of the genus, belongs
to the central group of species. The lateral processes, which are unique in the
order, serve to distinguish it from all other Philodince and Bdelloids. The boss on
the preanal is an uncommon character. Without these processes it would be very
difficult to characterise.
Philodina, sp. (Plate X., Fig. 6)
Description. — Size moderate. Corona narrow, about equalling the prominent
collar, and less than the trunk. Upper lip with two rounded lobes meeting in the
middle line. Two pale or colourless eyes. Teeth 2/2. Foot short (number of
joints ?) ; spurs broad short cones slightly diverging and with no interspace.
This species, which is undoubtedly distinct from all the other Antarctic species,
was only once seen, and is insufficiently studied. The form of the upper lip and
spurs are like those of P. plena (4). There is little to separate it from that species
except the possession of eyes. It is doubtful if the presence or absence of eyes
constitute characters of even specific value, as several species are variable in this
respect (P. rugosa, P. macrostyla, &c.).
It had not the red stomach of nearly all the Antarctic Bdelloids, but that might
merely indicate youth.
Habitat. — Among weed from the Narrows of Blue Lake.
Genus — Callidina
Note. — The genus Callidina contains a host of species, many of them not at all
closely related, and offering material for many genera. C. constricta and C. angularis
48 JAMES MURRAY
in our list should le in a different genus from C. habit a. As the whole order requires
revision by a competent authority,* the old names are retained here.
Callidina constricta, Duj. (14) (Plate XII. , Figs. 13a, 136)
Dujardin's meagre description would fit equally well a considerable number of
those CallidincB which mould the food into pellets, and which are distinguished from
one another by characters requiring more careful examination than the Bdelloids
usually receive. The commonest of those species which have numerous teeth on the
jaws, the corona somewhat less in diameter than the widest part of the head, the
discs close together, and the spurs short divergent cones, may be taken as the type
of C. constricta.
To establish C. constricta, Duj., and C. elegans, Ehr. (15), and to distinguish them
from the numerous related species, it would be necessary that both should be re-
described by a competent authority. Janson (20) briefly describes them, but scarcely
with the detail necessary to firmly establish them, neglecting, for example, the form
of the upper lip, a character of the utmost importance in all Philodinadse. Janson
gives C. constricta as having fewer teeth than C. elegans (8/8 instead of 10 or 11) and a
shorter jaw. There is nothing so difficult as to make certain of the number of teeth
in those pellet-making Callidince which have many teeth. They can be clearly enough
seen, but the anterior ones are usually thickest, and the others diminish successively
till they cannot be distinguished from the fine striae which are found on the rami of
all Bdelloids. Very commonly there are four strong teeth, the fourth (counting
from the anterior end) much thinner than the first; then there is an abrupt transition
to finer teeth which are still thicker than the striae, but merge gradually into them.
The Cape Royds Callidina, which I identify as C. constricta (see Plate XII.,
Fig. 13), is a small animal, about ^-Q inch in length (250 /u). It is not much enlarged
in the central trunk, which is faintly plicate and not obviously stippled. The small
discs are touching, and the whole diameter of the corona is distinctly less than the
widest part of the head. The jaws are long and narrow, and each bears four distinct
teeth, diminishing in thickness backwards, succeeded by finer teeth which continue
to diminish in thickness to the posterior end of the jaw. The rump is clearly
marked off from the central trunk, but its two segments are only seen as the animal
extends itself in creeping. The foot is short, of three joints, and the spurs are very
short, quickly tapering, and widely divergent.
Habits. — It is a quiet sedate animal, moving steadily and readily feeding. It
often swims free, rotating on its long axis.
Habitat. — In most of the lakes at Cape Royds. Common, but not abundant. It
is one of the two Bdelloids found in Green Lake.
s- — In company with C. constricta two different forms of egg have been
* Mr. D. Bryce is at present engaged on such a revision.
ANTARCTIC ROTIFERA 49
found, which may belong to this species or to C. angularis (Plate XII., Figs. 12c-12c£).
Each of these eggs was found with the jaws of the contained young so well grown
that the numerous teeth could be seen. As there are only the two species known at
Cape Royds which have many teeth, it is practically certain that these eggs belong to
them, but as they were never found in the bodies of the rotifers it cannot be determined
to which each belongs.
Callidina angularis, sp. n. (Plate XII., Figs. 12a— I2d)
Specific characters. — Small, yellow, much widest in central trunk ; strongly
nodose from the lateral projection of certain segments, from the neck to the preanal :
trunk strongly plicate, and stippled (except head, neck, and foot). The head is widest
at the cheeks, and the corona is distinctly smaller, and about equal in diameter to the
neck. Rump narrower than the fourth central segment, its two segments distinct.
Foot short, of three segments. Spurs small, divergent, acuminate and acute. Food
moulded into pellets. Jaws narrow, teeth many.
General description. — Length when feeding about ^^ inch, rather greater
when creeping. The diameter of the corona is about -§^ inch (50 ju), and of the
widest part of the trunk about ^<y inch (100 M). The rostrum is short and broad
and the antenna short. There are from eight to ten teeth in each jaw, and they
diminish rapidly in thickness towards the posterior end of the jaw. The anterior
edges of the three segments succeeding the neck are strongly produced outward,
giving a nodose appearance to the anterior trunk. The mid-trunk is also laterally
produced into an angle. The fourth central and preanal segments are also produced
at their posterior edges. The trunk has few broad clearly marked folds. The
preanal has two folds near the middle line. The three toes are short and obscure.
The upper lip is a somewhat triangular area. The central portion forms a pointed
arch, or may be rounded or slightly cleft. The collar is very inconspicuous. The
voluminous stomach is red or brown or pale yellow. The pellets are very obscure.
Habitat. — In most of the lakes at Cape Royds.
Reproduction. — The absence of living young makes it practically certain that the
species is oviparous. Associated with it were eggs (already referred to under
C. constricta) which could only belong to one of these species.
Affinities. — C. angularis is undoubtedly derived from C. constricta. There are so
many points in common that they were for long considered to be identical. The out-
line of a Bdelloid rotifer can vary so much during the different movements that it was
considered inadvisable to put much importance on the angular outline of this species.
It was only when the peculiar form was found to be always associated with a stippled
skin that it came to be regarded as distinct from G. constricta.
Eggs. — As to the eggs of this species, see remarks under C. constricta.
BRIT. ANTARCT. EXPED. 1907-9. VOL. I. G
50 JAMES MURRAY
Callidina habita, Bryce (4) Variety (Plate XL, Figs. 8a-8e)
Description. — Large, stout, hyaline or yellowish, much widest in central trunk,
which is plicate with broad folds. Length when feeding, up to 570 M, diameter of
trunk 125 M and upwards, of corona 95 /u- The corona is much wider than the collar,
and that is much wider than the neck. The discs are large and bear central setae.
They are separated by a space equal to half the diameter of the disc. The collar is
prominent and appears two-lobed in dorsal view. It is continued on to the upper
lip as two large rounded lobes which meet in the middle line.
The rostrum is stout, and the lamellae appear to be quite separated. The
antenna is equal to one-third the diameter of the neck. The jaws are large and
have a thickened border, sometimes coloured brown. The teeth are two in each jaw,
witha thinner one.
The voluminous stomach is of a deep crimson colour, and the walls are filled with
larger and smaller globules. The yolk-mass contains eight nuclei. The two
segments of the rump are obscurely distinguishable. The foot is four-jointed, but
there is often difficulty in making out more than three. The spurs are stout, almost
cylindrical, quickly tapering to the acute points, divergent. The first foot-joint has
a thickening forming a more or less distinct boss.
The egg is elliptical and is produced at each pole into a rounded prominence
(Fig. 8d).
This variety, which is typical in every other respect, differs conspicuously in" the
form of the spurs. Those of the type are broad and very acuminate, the lower edge
making an ogee curve. The highly coloured stomach is an important feature, but it
cannot be considered of much specific value.
Another variety. — Rather smaller, the two lobes of the upper lip less distinct,
separated only by a small notch. Spurs in form like the type, but smaller, and the
curvature of the lower edge less marked. The foot-boss more distinct. Otherwise
like the type (Plate XL, Fig. 86).
Both forms were fairly common in most of the lakes at Cape Royds. They did
not occur in Green Lake. In Blue Lake at a depth of fifteen feet.
Fa m ily — ADINET AD^E
'Genus — Adineta
The genus Adineta, the only one yet described in the family, is a small one. It
contains at present some eight recognised species. It is better represented in the
Antarctic Region than any of the other genera of Bdelloids.
Five species were recognised, of which one (A. grandis) is new to science.
ANTARCTIC ROTIFER A 51
None of them, except A. grandis, is very common. That species swarms in numbers
- only inferior to those of Philodina gregaria.
Adineta grandis, sp. n. (Plate XIL, Figs. Wa-Wd)
Specific characters. — Very large, stout, rostrum philodinoid, with lamellae and
brush of cilia as in Philodina ; posterior margin "of mouth pectinate : spurs short
broad cones, separated by straight interspace, reproduction viviparous.
Detailed description. — Large examples measure 750 M in length. The colour is
light brown or yellowish, darker in the alimentary tract. It has the graceful form
usual in the genus, very broad in the central region of the trunk and tapering to
narrow extremities. The trunk is regularly, and not very deeply, plicate. The
neck is slightly constricted below the head, then there is an expansion at the normal
level of the mastax.
The head is ovate, and rounded in front. The rostrum is short and stout,
and is quite like that of a typical philodine, except that it is not retractile.
In ventral view the lamella?, which are relatively smaller than is usual in the genus,
appear to meet in the middle line. Beneath them is the brush of cilia, looking as
they do in a philodine when contracted, but the tip was never seen everted so as to
make the brush project. On each side, close to the edge of the lamellae, is a longer
cilium, moving like the tactile setae occurring in the same situation in Philodina
macrostyla, &c. Only one could be distinguished at each side. The antenna is
short and broad.
The pectinate part of the mouth appears to be a' fold within the margin proper-
The processes are flat plates, with the spaces between about equal to the thickness of
the plates. They are rounded at the ends. The number varies from six to ten on
each side. At the inner end of each series the terminal process is drawn out into a
longer narrow rod, which appears to be attached to the flat surface of the corona.
The furred surface is uninterrupted from side to side, but at the anterior end a tooth-
like process projects backwards in the median line. The jaws are normal, with the
usual two teeth on each.
The stomach is large and of a warm brown colour. The intestine is elliptical.
The two segments can be distinguished in the rump, which tapers gradually in line
with the stout four-jointed foot. The spurs are stout and subacute. They diverge
widely and are separated by an interspace about equal to the diameter of the spur at
its base. The terminal joint of the foot is of moderate length and the three toes are
small and short.
The stoutness of the trunk varies greatly according to the number of young
carried, and their degree of development." It becomes enormous when it contains
six or seven young almost full-grown. The largest number observed was seven, but
the usual number was three. It was noticed that when the young were well enough
52 JAMES MURRAY
giown to show definite structures (such as the teeth) all of them were in the same
degree of advancement, as far as could be perceived. This is different from what is
the case in the genus Rotifer, in which all the species are viviparous. There the
two or three embryos are at different stages. The yolk-mass, which could rarely be
seen, contained eight nuclei. On one occasion (March 27, 1908) a yolk-mass was.
seen divided up into six portions, each containing a nucleus, no doubt a brood at an
early stage. Three pairs of narrow spindle-shaped vibratile tags have been seen.
Habits. — A. grandis has not the restlessness which is characteristic of the genus.
It is sufficiently active, but it creeps steadily, and "right side up," like many of the
Philodinadse, and can therefore be more easily studied. It feeds on minute organic
particles, among which there is rarely any recognisable organism.
Habitat. — Among the brown vegetation in the lakes at Cape Royds. It was the
most generally distributed of all the Antarctic rotifers, and occurred in saline lakes
from which most of the other species were absent.
Natural history. — A detailed study of Adineta grandis will be made in another
paper.* A summary of the ascertained facts will be here given. The species is the
only rival to Philodina gregaria in abundance, and in this connection it is important
to note that it shares with it the viviparous mode of reproduction. It is found in a
greater number of lakes than P. gregaria, but it never appears to be in such
prodigious numbers. This may be due merely to its less conspicious colouring, which
is almost identical with that of the plant on which it lives.
It is not extremely abundant in the freshwater lakes, where it has many com-
petitors, but in the very saline Green Lake, from which all the other species but
Callidina constricta are absent, it is almost as abundant as P. gregaria is in Coast
Lake, and may be collected in the same way. When the fine debris washed from
the weed of Green Lake is allowed to stand undisturbed for some hours a pale brown
layer appears on the surface. This may be taken up by the pipette and is found to
be pure Adineta grandis, without admixture of other organisms. The rotifer has
crept out of the mud to the surface. This habit allows quantities to be got for study
and makes it easy to make simple experiments upon them. It did not usually creep
up the sides of the bottle, as P. gregaria does, but on one occasion, when a moderate
number were mixed with that species, they also crept to the surface of the water.
They could be obtained at any time during the winter by melting some ice from
Green Lake enclosing some of the brown weed. They usually began to move as soon
as released.
In its power to endure extreme changes of temperature and other adverse
conditions, Adineta grandis is the most interesting of the Antarctic rotifers. From
its large size and great abundance, it was the species selected for most of the experi-
ments made with the object of finding the limits to the vitality of Rotifers. It
survived the lowest temperatures experienced at Cape Royds ( - 40° F.), and repeated
* On " Life under Polar Conditions."
ANTARCTIC ROTIFER A 53
freezing and thawing. It was the only species subjected to the heating experiment,
in which a proportion of them lived after the bottle containing them (in the dry con-
dition) was immersed in boiling water for a short time. It was one of the rotifers
which was to be seen alive and active in London in September 1909, after being dry
for about a year, and spending some months in tropical and subtropical climates. It
revived in about an hour after being moistened. It was immersed in sea-water, and
in the much more saline fluid obtained from under ihe ice of Green Lake (of which it
is a native), and kept there for one month, after which it revived quickly when trans-
ferred to fresh water. The diameter of the contracted animal in fresh water was
225 p. — in the brine of Green Lake it contracted to 150 p, or two -thirds of the normal
diameter.
It is indifferent to the interruption of its active vital functions by freezing, often
for long periods, and quickly resumes activity when thawed. The development of the
young likewise does not appear to suffer from its interruption at any stage. In con-
sequence of interruption development may often take many years for its completion,
but in the periods when the lakes are melted it probably only occupies a few days,
and many generations may be completed in the few weeks of summer.
Affinities. — A. grandis shows most resemblance to A. vaga Davis (12). The
pectinate margin to the mouth is a character which may readily be overlooked in
species of this genus, as the structures of the head of an Adineta are very difficult to
see, on account of the restless contortions of the animal. The pectinate border is
easily seen in A. grandis owing to its large size.
The species may be a derivative of A. vaga, but of very long standing, as the
profound changes show. A different mode of reproduction has been adopted, and
the form of spurs and rostrum, &c., changed. The ciliated surface of the face of
A. vaga is described as divided into two parts by an unciliated band. This is not
the case with A. grandis, in which there is no perceptible interruption of the furred
surfaee from side to side.
Adineta barbata, Janson? (20) (Plate XII. , Figs. 9a-9c)
Description. — Of moderate size. Head ovate ; rostral tip produced laterally into
little sharp points ; lamellae long, slender, curved, rounded, accompanied by several
long bristles ; spurs divergent, curved, tapering, acuminate.
While corresponding in general to the type of A. barbata, the animal found at
Cupe Royds differs in two particulars : the spurs taper from the base and are
acuminate ; in the type the thickness is maintained to near the tip, and the tapering
is then abrupt ; the processes on the rostrum which correspond to the lamellae of
the Philodinadse are very long and slender, and in dorsal view look like antennae of
insects.
Jansou does not describe the lamellae as of this narrow elongate form, and his
figure shows them quite moderate. The difference is of specific value, but as the
54 JAMES MURRAY
Antarctic species was not at the time studied critically enough, and the knowledge of
the peculiar lamellae is only gathered from my sketches, I prefer to leave it meanwhile
as a form of A. barbata, to which it appears to be closely related.
The pectinate border of the mouth was difficult to see, owing to the restless
movements of the animal, but it was clearly seen in a well-grown young in the egg
(Fig. 9c).
The first eggs found confirmed the belief that the animal is distinct from
A. barbata. They were elliptical, and lacked the knobs characteristic of the egg of
A. barbata. Later on a different egg was found, which had the knobs as in A. barbata,
but much reduced in size (Fig. 9c).
Adineta vaga, Davis (12)
The type of this species (which is the var. minor of Bryce) was found in several
lakes soon after we landed at Cape Royds. It was not seen later, and was thus never
subjected to a very critical examination.
Habitat. — Blue Lake, Clear Lake, and Coast Lake.
Adineta gracilis, Janson (20)
Habitat. — Blue Lake, Clear Lake, and a little pond on the lower slopes of Mount
Erebus. The examples appeared to be quite typical.
Adineta longicornis, Murray ? (27)
Description. — Small, head ovate, rostral part divided into two large, evenly rounded
lobes. The spurs are long, slightly divergent, and taper to acute points.
Habitat. — Among moss from the High Moraines, Cape Royds, January 1909, one
example seen.
It is with hesitation that this animal is referred to A. longicornis. The spurs are
shorter than in the type, and the head is of slightly different shape. As the rostral
structures were not clearly made out, it cannot be confidently separated from that
species.
NOTE
Additional Species. — In April 1910, just as we go to press, an additional Bdelloid
has been found alive, in moss collected by Priestley at the Stranded Moi-aines. It is
a pellet-maker having three teeth in each jaw, and may be identified as Callidina
(Macrotrachela) tridens, Milne. That species is very similar to C. constricta, except
in the very distinct teeth. A more detailed description of it is wanted.
This addition, with the identification of the Floscidaria as F. cornuta, brings up
the list of recognised Antarctic Rotifera to 16 species.
ANTARCTIC ROTIFERA 55
Order— EHIZOTA
Family — FLOSCULARIAD^E
Genus — Floscularia
Floscularia, sp. (Plate XIII., Fig. 15) *
On January 18, 1909, among weed from a pond between Cape Barne and Cape
Royds, a species of Floscularia was found in some abundance. When examined after
reaching the hut they were moving languidly and showing an inclination to expand.
The long setoe, which exceeded the trunk in length, projected as a long brush. The
lobes were not unfolded, and there was little time to wait for it.
Although there is no hope of identifying the species, a sketch of it in the partly
contracted condition is given, in order to complete our records of the rotifera of the
region.
In this state the length, exclusive of the setae, was 284 /j.. The foot is transversely
wrinkled in the usual manner, and terminates in a conical portion, tapering to a
narrow apex (probably the adhesive disc unattached at the time).
Neither tube nor eggs were observed. The teeth and the various viscera were
seen, but detailed studies under pressure could not be made.
Order— PLOIMA
Family — HYDATINAD^E
Genus — Hydatina
Hydatina senta, Ehr. (Photograph from life. Plate II., Fig. 7)
No description need be given, as the Antarctic examples appear to be quite typical.
The species was identified with certainty by Mr. Rousselet from preserved examples.
It was found in great abundance in Coast Lake when it was tow-netted for the first
time on January 2, 1909. It might have been found much earlier if the lake had
been examined in a suitable manner, as there was some open water from the end of
November 1908, and the temperature of the water, as early as December 4, was
+ 47° Fahr. When the Hydatina was first found the temperature had gone down
to +40°. On January 18, 1909, the temperature had gone up again to +45°, and
the Hydatina was much less plentiful.
Large oval smooth brown eggs were known, which proved to be those of
Hydatina.
Under the influence of Eucaine Hydatina behaved well, retaining the normal form
* An example having the corona fully expanded has s-ince been found, which allows of its identifi-
cation as F. cornuta, Dobie.
56 JAMES MURRAY
perfectly, but remaining so still that it could easily be photographed while living and
feeding.
Hydatina was only seen in Coast Lake. Its restriction to one lake led Mr.
Rousselet to suggest the possibilit}r of its recent introduction, say by the Discovery
Expedition. It is not impossible, though hardly probable. Hydatina is a common
animal in ponds around farmhouses, and eggs might readily enough adhere to farm
produce such as hay or straw. When Captain Scott and Dr. Wilson camped for a
short time at Cape Koyds, close by the spot where we afterwards had our hut, they
may have had some straw packing among their gear. In this connection it is worth
noting that we dredged in Back-door Bay a stalk of old sodden straw on which a
sponge had grown. This straw could hardly be of older date than the Discovery
Expedition. Against the theory of the recent origin of Hydatina is the fact that
Coast Lake is a mile from Captain Scott's camp, and that several suitable lakes lie
nearer the camp. Still, one egg is enough for the introduction, and the straw bear-
ing the one egg may have blown to Coast Lake.
Family — NOTOMMATAD^E
Genus — Pleurotrocha
Pleurotrocha, sp. ? (Plate XIII. , Figs. 14a-14c)
Description. — Of large size, 520 yu in length. In lateral view greatly elevated
just behind the middle of the trunk. Head long, mouth narrow. Jaws very like
those of P. grandis, Western, and some related species. Toes longer than those of
P. grandis. Stomach large, orange-coloured.
This large active animal was found in Blue Lake early in the season, before the
requisites for narcotising and preserving were available. It was fairly abundant then
but was not obtainable afterwards, when it could have been photographed and pre-
served. We are therefore dependent for our knowledge of the animal on a few
sketches by onelittle acquainted with the order to which it belongs. Mr. Rousselet
has examined the sketches critically, taking such points as could be best trusted, and
compared them with the same points in the species coming nearest to our one.
It is undoubtedly very near P. grandis (41), which is like it in size and activity.
The jaws differ in being more ovoid in form, diminishing backward from the widest
part. In P. grandis the greatest width is maintained for some distance backward.
The toes are considerably longer and narrower than in P. grandis. The general
outline of the jaws is very similar to that of Diglena permollis, Gosse (16), as shown
in a drawing by Mr. Dixon-Nuttall.
Reproduction. — Two kinds of egg have been seen which contained animals having
jaws exactly like those of the Pleurotrocha (the drawing, Fig. 14c, is made from an
example in an egg). One of the eggs was elliptical and smooth. The other was
ANTARCTIC ROTIFERA 57
also elliptical but was thicker shelled, and was covered with little points projecting
from the surface (Fig.
Genus — Diaschiza
Diaschiza tenuior, Gosse ? (17) (Plate XIII., Figs. 16o-16/)
Of the species described in Dixon-Nuttall and Freeman's Monograph of the genus
(13) the Antarctic Diaschiza can only be D. tenuior. It differs only in being somewhat
laterally compressed, and in carrying the toes more ventrally than usual. The
length of the toes easily separate it from D. caeca, Gosse:
With it was associated a much smaller animal, which may be the male of the same
species (Figs. 14c-14d). It has a large head and comparatively insignificant body.
The corona has the large strong cilia common to most males, and it swam powerfully.
The dorsal cleft is seen on the trunk. The segment between the trunk and the toes
is well marked. The toes are small, conical, divergent and decurved. Unlike most
males it possesses teeth. The male organs were not definitely seen, nor were any
viscera distinguished.
In the female the seise at the base of the toes were not made out, but we were
working with very poor light.
The elliptical, smooth, thin-shelled egg was seen, containing the young. A
smaller egg, with thick shell, through which obscure lines passed obliquely in various
directions (Fig. 16/), contained an animal with jaws like those of the supposed male.
REMARKS ON THE ANTARCTIC ROTIFER FAUNA
Its Composition. — The Rotifer Fauna of Cape Royds, comprising not much more
than a fiftieth part of the known species, is very remarkable in its composition. When
the number of species is so small it is curious that all the orders of Rotifera are repre-
sented [the Scirtopoda being, as a result of Beauchamp's studies (2), reduced from
ordinal rank].
The great preponderance of the small order Bdelloida (with twelve species) over the
Plo'ima (with four species) is not surprising in view of the well-known remarkable
vitality and facility of distribution of these animals.
The proportions in which the various genera of Bdelloids occur in the fauna are
very curious and interesting. The four species of Philodina are all unknown else-
where. The large genus Callidina, which elsewhere contains half, or more than half
of the species in the entire order, has only three species at Cape Royds. Two of these
are known species and one is new. The small genus Adineta, of which only seven
species have been described, has no fewer than five species at Cape Royds. Only one
of these is new to science. The genus Rotifer is absent.
BRIT. ANTARCT. EXPED. 1907-9. VOL. I. H
58 JAMES MURRAY
The sixteen species ofRotifera are thus distributed : eight known species, five new
species, three not identified. While there are only five new species named, one of the
others (Philodina, sp.) though not sufficiently studied to be named, is certainly new,
and several of the known species differ more or less from their types, and may be
incipient species resulting from long isolation in peculiar conditions.
Peculiarities. — The distinctive characters of the new species are not very remark-
able. They consist in the forms and proportions of the spurs, head, upper lip, &c., or in
the possession of peculiar processes (Philodina alata). There are no peculiar types.
Development has gone on lines similar to those it might have followed in any region.
The rotifers may have acquired peculiar physiological properties, enabling them to
resist the rigours of the Antarctic climate, but such adaptation (if it has taken place)
is not correlated with any peculiarity in outward form.
Reviewing the rotifer fauna as a whole, its most notable features are the general
prevalence of red colour among the Bdelloids, and the viviparous reproduction
acquired by some of them belonging to groups which are rarely viviparous.
Colour of Bdelloids. — All the Philodince, except the unnamed species, have the
voluminous stomach coloured of a vivid deep crimson. This colour is shared in equal
degree by Callidina habita. Callidina constricta and C. angularis have the stomach
sometimes red, sometimes brown, and occasionally pale yellow. None of the
Adinetce have red stomachs. A. grandis has the alimentary tract of a warm brown
colour, the others are pale yellow or colourless.
The prevalent red colour may be related to the nature of the food. Callidina
habita and C. constricta are not normally red in other countries. All the species
live among the same plants, which are of a warm orange colour, but associated with
this are numerous green and blue-green Algae, so that there is a variety of food
available, and the different species may select different foods. We have no exact
information as to the kinds of food actually eaten by the Antarctic species. Though
they can be readily watched feeding, they rarely swallow organic matter in recognis-
able condition, but fine, flocculent material, the result of decomposition. Bdelloids
have the power of selecting their food. A pair of knuckle-like processes in the
gullet can be brought together, and these are opened or closed according as the
particles swept down the funnel are acceptable or not.
There were no red-coloured rotifers in Green Lake, which was very saline. The
only two Bdelloids which occurred there were Adineta grandis and Callidina con-
stricta. A. grandis was never red in any lake, but in freshwater lakes C. constricta
had sometimes a red stomach.
Reproduction. — Of the Antarctic rotifers seven are oviparous, two are viviparous^
and the reproduction of the remaining seven is unknown. The genus Rotifer, all
the members of which are viviparous, is unknown in the region.
The two viviparous species, Philodina gregaria and Adineta grandis, prepon-
derate enormously in numbers over all the other species. This mode of reproduction
ANTARCTIC ROTIFERA 59
appears therefore to be best adapted to secure success in the struggle for existence
under the severe conditions experienced at Cape Royds.
Dispersal. — Most of the Bdelloid Rotifers were found generally distributed
among the lakes of the district. Dispersal from one to another is therefore prob-
ably easy. The most striking exception is Green Lake, where there are only two
of the species. There is little doubt that the qualities of the water are responsible
for the limitation in that lake, as there are other lakes close by from which rotifers
might be derived.
Hydatina is only known in Coast Lake. It is the only normally free-swimming
rotifer in the region, and very likely its large eggs fall to the bottom in the deeper
part of the lake, and are not likely to be in the marginal zone which is exposed by
the ablation of the ice. Even so it is to be expected that some eggs would get
among the weeds which are thus exposed.
No rotifers were found in Pony Lake, but the proximity of the penguin rookery
renders the water so foul that it would be surprising if there were rotifers in it,
always excepting Hydatina, which frequents such situations.
There are only two probable means by which rotifers can be transferred from one
lake to another. One is by the agency of birds, and the other is by means of the
wind. The skua gulls are the only birds which frequent the fresh water. They are
very fond of bathing, and stand in the shallow parts of the lakes, where the water is
a few inches deep and the bottom usually covered with weed and rotifers, and there
they splash the water about with great gusto. Fragments of weeds may adhere
to their feet or may get on their feathers as the water splashes over them. The
skuas may visit several ponds in the course of their flights and thus distribute the
rotifers over the countryside.
At the margins of the lakes there is generally a zone of dried weed, which
increases in breadth as the winter advances, as a result of the ablation of the ice
surface. This zone varies in colour from brown to dull green. The weed is strongly
wrinkled and very light. Pieces of it might easily be torn off by the wind and blown
about the country till the next summer, when it might happen to get into another
body of water. Fragments of weed were often seen blowing about. In Coast Lake
the ablation of the ice exposed the weed on the bottom, which was in small flakes.
Showers of these flakes were blown by the wind over the shore and out to sea.
Dispersal by the wind and conveyance from lake to lake in a small district like
Cape lioyds is easily understood. The prevalent strong winds in the region are all
from the south, and it is therefore difficult to imagine that the plants and their living
freight could be transferred in this way over long distances where there are not
intermediate resting-places, except from south to north. It is, however, possible that
northerly gales occur at long intervals, which would help to disperse the species in
the opposite direction.
Distribution. — Of the distribution of the Rctifera in the Antarctic Region there is
60
JAMES MURRAY
nothing to be said, as there are no previous records, except the doubtful egg of
Callidina papillosa mentioned by Richters (31). Rlchters gives details of the jaws
and teeth of four species from the Gaussberg, and figures them, as well as other details
of two of the species. These are insufficient for certain identification, though if the
rotifer fauna of the Gaussberg could be studied in life we might have a good guess as
to what those four were. As our knowledge at present stands we cannot connect any
of them with the species found at Cape Royds, though it may be suggested that the
figures five and six on his Plate XVII. may be Callidina constricta, which was the
only moss-dwelling species identified at Cape Royds.
As the thirteen species found at Cape Royds appear to be the first recorded for
the continent, we can only study the distribution over the world of those of the
species which are not new to science. The distribution is most clearly seen when the
facts are given in tabular form. It is not claimed that the records tabulated are
complete, but I have given all which I could find.
TABLE SHOWING THE GENERAL DISTRIBUTION OF ALL RECORDED ANTARCTIC
ROTIFERA.
o
OD "3
u
g
I
31
1
i
3
3
B
£
0
H
s
a s
5
s
s
5
H
ISB
§
<]
•<
"1
<l
•5
•<
< ^
fc
02
Philodina gregaria, sp. n.
X
_
_
_
„ antarctica, sp. n.
X
—
—
—
—
—
—
—
„ alata, pp. n.
X
—
—
—
—
—
—
—
Callidina angularis, sp. n.
X
—
—
—
—
—
—
—
Adineta grandis, sp. n.
X
—
—
—
—
—
—
—
Callidina conslricta, Duj.
X
X
X
—
X
X
X
X
,, habita., Bryce '( .
X
X
X
X
X
X
X
X
„ papillosa, Thomp.
?
X
X
X
—
X
—
X
Adineta vaga, Davis
X
X
X
X
X
X
X
X
„ barbata, Janson ? .
X
X
X
—
X
X
—
X
„ gracilis, Janson . .
X
X
X
—
X
X
X
X
,, longicornis, Murray ?
X
X
—
X
X
—
—
—
Hydatina senta, Ehr. . .
X
X
X
X
X
?
X
Diaschiza tennior, Gosse .
?
X
X
—
1 — '
—
—
The facts made use of in the above table are compiled in part from the following
sources : for the Antarctic from a paper by Richters (31) : for Australasia from Hilgen-
dorfs list (19) and from my own unpublished notes which will appear in a later
number of this series : for Europe from papers by Bryce, Janson (20), and others :
for Africa from a paper by Rousselet (33) and unpublished notes : for N. America
from a paper by Jennings (21) and from unpublished notes : for the Arctic Region
from papers by Bryce (5), Bergendal (3) and myself (28).
The new Antarctic species are placed at the top of the list in order that the other
facts may be compactly grouped. The facts of distribution of the nine previously
ANTARCTIC ROTIFERA 61
known species are given under eight headings, being a column for each continent, for
Australasia, and for the Arctic and Antarctic regions. This covers the greater part
of the earth's surface, only excluding oceanic islands. It may fairly be said that an
animal which occurs in all of these broad divisions is " generally distributed."
Callidina habita and Adineta vaga are in every column ; C. constricta, A. gracilis,
and Hydatina senta are absent from one region only ; C. papillosa and A. barbata
from two ; A. longicornis from four ; and Diaschiza tenuior from five. These facts
have very little significance, as some of the regions have been very little worked.
Considered in another way the table shows that all the previously known species
are also in New Zealand, all but one in Europe, all but two in Africa, N. America,
and the Arctic, all but four in S. America, and all but five in Asia. For the reason
given in the preceding paragraph these facts have no special significance. Were
the world more fully explored we would in all probability find that all the species
are ubiquitous.
The Rotifera of the southern hemisphere, from which any migrants to the Ant-
arctic have most probably come, are very imperfectly known. In New Zealand
Hilgendorf s was the only work, and in the Index Faunae Novse-Zealandise (19) he
gives a list of over furty species then known. In Australia short lists were published
by Anderson (1) and Shephard (34 to 37), and Whitelegge (42) in 1889 summed up
the known species, giving a list of 110. In Africa there were short lists by Milne (24)
and Kirkman (22 and 23). Rousselet (33) in 1905 visited S. Africa, and in his
paper summarises all that was known, giving a list of 156 species. In S. America
Schmarda (39) found many rotifers, but his descriptions and figures are such that
most of the species are unrecognisable. Recently Daday wrote upon the Rotifera
of Patagonia (9), Chile (10), and Paraguay (11). He enumerates 106 species,
including some noted by Wierzejski (43). There may be isolated references to
rotifers of the southern hemisphere in works other than those cited, but they
comprise almost all that was known till recently.
All those lists are noticeable for the very subordinate position occupied by the
Bdelloids. This is due to the fact that it was not suspected that mosses supported
a very rich fauna of rotifers and other animals. Some of the early investigators of
these animals, who were more interested in their physiology than in the discrimina-
tion of species, distinguished between the moss-dwellers and water-dwellers.
In papers by Thompson (40) and Bryce (6) such terms as " moss-haunting" are
found in use, and in Germany also the word " Moosbewohner " [Richters (32)] is
recognised. So little was the existence of the moss-fauna known among naturalists
that in some of the latest expeditions the mosses were taken only as botanical
specimens, and were treated with preservatives which killed the fauna.
Much work has in recent years been done on the Rotifera of all parts of the
globe, from specimens procured from moss. No easier method of collecting is possible
than by bringing home dried mosses ; and it is to be hoped that future expeditions
62 JAMES MURRAY
will realise the desirability of collecting some moss for the sake of its fauna. The
moss-fauna of rotifers consists chiefly of Bdelloids, though there are a few others
which have adopted the same habitat.
Origin of the Rotifer Fauna. — In the present state of our knowledge it would
be premature to discuss the origin of the Antarctic Rotifer Fauna with any expecta-
tion of arriving at definite conclusions. We know almost nothing of the geological
history of Antarctica, especially of its recent glaciation ; we know nothing of the dis-
tribution of the Rotifera round the fringe of the continent (the heart we may infer to
be as dead as anything on earth) ; and our knowledge of the Rotifera of the nearest
land-masses and of the scattered islands of the southern hemisphere is inadequate.
It may, however, be permitted to discuss in a tentative manner the bearing of
the known facts. It may safely be assumed that the Antarctic Region has known
periods of greater glaciation or of greater cold than at present, and periods of temperate
or warm climate. Is it possible that the rotifer fauna is aboriginal, and has survived
through the various climatic changes of long geological ages ?
Judging from the summer condition of the Antarctic coast at present, it does not
seem too much to suppose that during the coldest periods there might always be
some rock faces so inclined as to make the most of the northern sun, and so form
little pools where the rotifers might live. The rotifers have been proven to survive a
lower temperature than any yet known under natural conditions on the earth, and
we have seen (under descriptions of Adineta grandis and Philodina gregaria) that
they do not ask for much in the way of luxury. Give them a week or two of warm
weather ( + 40° F. or so) and they are content to be frozen up for years.
Periods of intense cold need not be periods of great precipitation, as we see in the
present low snowfall in South Victoria Land, and thus the bearing of temperature on
the problem falls to be separately discussed from that of glaciation. Has there ever
been a period when the glaciation was such that the very mountain peaks were covered
by a thick ice-cap which would scrape the rotifers, in common with all other living
things great or small, off the surface of the continent? That will not be discussed
here, but short of such a mechanical destruction as extreme glaciation or complete
submergence would achieve, I see no reason why there should not always have been
rotifers on the coast at least of the Antarctic continent.
The peculiarities of the species of rotifers which at present appear to be peculiar
to the Antarctic are not very great, morphologically considered, yet they are great
enough to require us to presuppose a very long isolation if they have been acquired
in their present location. For four of them it would be difficult to suggest a probable
ancestor among known species. Although our list of fourteen species shows five new
and nine previously known, the amount of peculiarity is greater than these figures
would indicate. Three species indicated bjr marks of interrogation in the table
given differ more or less from their supposed types elsewhere. It was doubtful
whether they should be regarded as new species or as varieties. In such cases one has
ANTARCTIC ROTIFERA 63
to decide whether the advance of knowledge will be better served by emphasising the
affinities in using the old names or pointing the differences in bestowing new names.
If the rotifers are not aboriginal but have reached the Antarctic by immigration
in comparatively recent times, since the last period in which the conditions were such
as to prevent the continuance of life on the continent, we must look for some possible
means of immigration.
The Rotifera share with the lowest forms of life that facility for distribution
which makes them, as Jennings (21) puts it, "potentially cosmopolitan." The agent
of distribution is the wind. When some rotifers, and the eggs of others, are dried they
may be blown in the form of dust for long distances. There is no difficulty in
supposing the Antarctic peopled in this way, though there is no region where such
distances of sea must be crossed in the process ; but all round the Antarctic continent
the storm-winds generally blow off the land, and so could play no part in bringing a
rotifer population to the country.
There are numerous small islands scattered over the Antarctic Ocean, and there
are some storms which blow from the north : even in the Antarctic they are known,
though rare. While it is difficult to believe the wind currents of the lowest strata
of the atmosphere adequate to transport rotifers over the wide ocean separating, for
example, New Zealand and South Victoria Land, they might transport them from one
island to another, and thus the rotifers might in long ages work their way by slow
and intermittent steps from the one land to the other.
There is yet another way in which the wind might be supposed to effect the
transference. If a violent local storm were capable of whirling rotifer dust up till it
was caught in those high currents which set to the southward, then they might
conceivably be carried all the way and dropped on land. If, however, it were as easy
as all that, we would expect a much more extensive rotifer fauna.
In one region, south of Cape Horn, the Antarctic continent approaches very near
one of the other continents, and there the rotifer dust might readily be blown across.
From such a point of easy access the animals might get distributed all round the
Antarctic coast by the aid of wind and birds.
The paucity of the rotifer fauna, as far as we know it, points to great difficulty of
access. The varying degrees of peculiarity exhibited by the different species suggest
that some have been longer resident in the Antarctic than others. Hydatina senta
may be a recent immigrant, may even have taken passage with Captain Scott in the
early days of the present century : Callidina habita and C. angularism&y have been
there for a few hundreds or thousands of years : whilst Philodina alata, P. gregaria
and Adineta grandis may be aborigines of immense antiquity.
That great difficulties lie in the way of emigration to the Antarctic, and
difficulties quite apart from the climatic rigours which meet the immigrants, we have
one more indication in the adaptability of many rotifers to various climates. Not
only do a grea.t many species extend from the temperate into the Arctic Region, but
64 JAMES MURRAY
rotifers taken from a subtropical forest in the extreme north of New Zealand, and
subjected to low temperatures at Cape Royds, revived on being moistened. Many
Rotifers, Bdelloids at least, do not require much, if any, extension of their powers of
endurance to fit them for life in the polar regions, so that if species are few in the
Antarctic it is almost certainly because of mechanical difficulties in the way of getting
there.
Vitality of Rotifers. — Observations on the Biology of the Rotifera constituted the
most interesting part of the naturalist's work in the Antarctic. Many experiments,
of a very simple nature, were made upon them in order to ascertain their powers of
resistance to every kind of adverse circumstance which might affect them in the
course of a migration to Antarctica, or from place to place in it.
These observations and experiments will be described in detail in a paper now in
preparation. A short summary of the most interesting points will be found under
the descriptions of Philodina gregaria and Adineta grandis in the preceding pages.
LIST OF WORKS REFERRED TO IN THE TEXT
1 . ANDERSON, H. H. and SHEPHAUD, J., Victorian Rotifers, Proc. Boy. Soc. of Victoria, iv., 1892, pp. C9-80.
2. BEAUCHAMP, P. DE, " Recherches sur les Rotiferes," Arch, de Zool.Experiment. et Gen., t: x., p. 43, 1 909.
3. BERGENDAL, D., " Beitrage zur Fauna Gronlands," K. Fysiograf. Sdllskapets Handl., N.F., 1891-2,
Bd. iii.
4. BEYCE, D., " Further Notes on ' Macrotrachelous Callidinao,'" Journ. Quekett Micr. Club, Ser. II.,
vol. v., 1894, p. 451.
5. ,, ,, - " Contributions to the Non-Marine Fauna of Spitsbergen," Proc. Zool. Soc. London, 1897,
pp. 793-9.
6. „ „ "Moss-dwelling Cathypnadse," Science Gossip, 1892, p. 271.
7. CEKTES, A., " Mission a Cap Horn, 1889.
8. „ „ "Infusoires et Rotiferes," Act. Soc. Sclent. Chili, iv., 1894.
9. DADAY, E. VON, " Mikro-Siisswasserthiere aus Patagonien," Term, rajzifiiz., Bd. 25, 1902, p. 201.
10. „ „ "Susswasser-Mikrofauna von Chile," Term, rajzifuz., Bd. 25, 1902, p. 436.
11. „ ,, "Susswasser-Mikrofauna Paraguays," in Chun's Zoologica, Bd. 18, H. 44, Stutt-
gart, 1905.
12. DAVIS, H., " A New Callidina," &c., Month. Micr. Journ,., ix., 1873, p. 201.
13. DixoN-NuTTALL, F. R., AND FREEMAN, R., " The Rotatorian Genus Diaschiza," Journ. Roy. Micr
Soc., 1903, p. 1.
14. DUJARDIN, F., " Histoire Naturelle des Zoophytes," Suites & Euffon, 1841.
15. EHRENBERQ, C. G., "Organisation der Infusorien/' Abhandl. K. Akad. Wiss. Berlin (1831), 1832.
16. GOSSE, P. H., " Catalogue of Rotifera found in Britain," Ann. Nat. Hist., vol. viii., 1851, p. 197.
17. „ „ In " The Rotifera," Hudson and Gosse, London, 1889, vol. ii., p. 81.
18. HILGENDORF, F. W., " A Contribution to the Study of the Rotifera of New Zealand," Trans, and
Proc. of New Zealand Inst., vol. xxxi. (14 of New Series), 1898, pp. 107-134.
19. ,, -, " Rotatoria," in Index Faunae Novce-Zealandice, 1894.
20. JANSON, O., " Rotatorien-Familie der Philodinseen," Marburg, 1893.
21. JENNINGS, H. S., " Rotatoria of the United States," U.S. Fish. Comm. Bull, for 1899, p. 81.
22. KIRKMAN, T., "Rotifera of Natal/' Journ. Roy. Micr. Soc. London, 1901, pp. 229-241.
23. „ „ " Second List of Rotifera of Natal," Journ. Roy. Micr. Soc. London, 1906, pp. 263-268.
ANTARCTIC ROTIFERA 65
24. MILNE, W., "On the Function of the Water- vascular System in the Rotifera," Proe. Phil. Soc.
Glasgow, 1907, pp. 1-12.
25. MURRAY, J., " The Rotifera of the Scottish Lochs," Trans. Roy. Soc. Edin., vol. xlv., 1906, pp. 151-191 .
26. ,t ,, " Scottish National Antarctic Expedition, Tardigrada of the South Orkneys," Trans.
Roy. Soc. Edin., xlv., 1906, p. 323.
27. „ „ " Some Rotifera of the Sikkim Himalaya," Journ. Roy. Micr. Soc., 1906, p. 643.
28. ,, „ "Arctic Rotifers," Proc. Roy. Phys. Soc. Edin., vol. xvii., 1908, pp. 121-7,
29. „ „ "Microscopic Life in Gough Island," Proc. Roy. Phys. Soc. Edin., vol. xvii., 1908, pp.
127-9.
30. „ „ " Some African Rotifers," Journ. Roy. Micr. Soc. London, 1 908, p. 669.
31. RICHTERS, F., " Fauna der Moosrasen des Gaussbergs und Einiger Siidlicher Inseln," Deut. Sudpol.
Exped'.'^QZ, Bd. ix. Zool.,^>?261.
32. „ „ " Moosbewohner," Schwed. Sudpol. Exped., 1901-3, Bd. vi., 1908.
33. "ROUSSELET, 0. F., " Rotifera of South Africa," Journ. Roy. Micr. Soc. London, 1907, pp. 395-414.
34. SHEPHARD, J., "A New Rotifer, Lacinularia elongata," Victorian Naturalist, xiii., No. 2, May 1896,
pp. 22-4.
35. „ „ "A New Rotifer, Lacinularia eUiptica," Victorian Naturalist, October 1897.
36. „ i, " A New Rotifer, Lacinularia striolata," Proc. Soy. Soc. of Victoria, August 1899,
pp. 20-35.
37. SHEPHARD, J., and STRICKLAND, W., " A New Rotifer, Melicerta fimbriata," Victorian Naturalist,
xvi., No. 3, July 1899, pp. 38-40.
38. STRICKLAND, W., " The Rotifer in Melbourne," Victorian Naturalist.
39. SCHMARDA, L., "Neue Wirbellose Thiere," Leipzig, 1859.
40. THOMPSON, P. G., " Moss-haunting Rotifers," Science Gossip, 1892, p. 56,
41. WESTERN, G., " Notes on Rotifers," Journ. Quekett Micr. Club, 1893, pp. 155-160.
42. WHITELEGOE, T., " Invertebrate Fauna of Port Jackson and Neighbourhood," Journ. and Proc. Roy.
Soc. of New South Wales for 1899, pp. 308-17.
43. WIERZEJSKI, A., " Rot. W. Argentinie," Bull. Acad. Cracovie, 1902, p. 158.
NOTE
In the preparation of this paper I have been indebted to several friends for assist-
ance, which I desire here to acknowledge. Mr. D. Bryce examined the drawings of
the Bdelloid rotifers, and gave me the advantage of his opinion on the values of the
species. Mr. C. F. Kousselet did the same for the non-Bdelloid rotifers, and helped
me with the list of literature. Mr. D. J. Scourfield experimented on the dried
rotifers brought from the Antarctic, and demonstrated that some had survived the
many changes of climate which they had experienced. Mr. J. H. Prieetley subjected
some of the rotifers to a temperature of —78° C. for many hours, and showed that
some , were not killed by this, thus completing the simple series of experiments
commenced in the Antarctic.
Mr. Rousselet was also good enough to remount the few specimens of the new
species of Antarctic rotifers, which had been mounted in a temporary way, thus
saving them from destruction.
BRIT. ANTARCT. EXPED. 1907-9. VOL. I. I
PLATE IX
PLATE IX
This plate is designed to illustrate the bright coloration which distinguishes most
of the Antarctic Bdelloids. The brilliant red of the stomach of Philodina gregaria
and Callidina hctbita (Figs. 1 and 3) may give the impression of being overdone.
If we have not caught the precise shade of the red, it is not that it was lees vivid
than we show it. It was a deep, clear, pure crimson or ruby. When seen in large
mass the colour approaches blood-red, but by transmitted light it appears crimson
under the microscope.
When we succeeded in reviving a few examples of Philodina gregaria and showed
them to a naturalist in London, he exclaimed, " Are they so red as all that."
FIGURE 1. — Philodina gregaria, lateral view, showing the red stomach and eyes,
small antenna, and two young.
FIGURE 2. — Adineta grandis, dorsal view, showing the brown colour deepest in the
alimentary tract.
FIGURE 3. — Callidina habita, dorsal view. The distinctive spurs of this variety are
shown, and the prominent foot-boss.
Brit. Antarct. Expecl. 1907-9. Vol. I. Plate IX.
MURRAY: ANTARCTIC R o T i F K R A.
i) P. gregaria.
(2) A. grandis.
Colours of Antarctic Rotifers.
(3) C. habita.
J. Murray, del. ad. nat.
PLATE X
PLATE X
FIGURE 4a. — Philodina alata, sp. n., dorsal view, feeding : the lateral processes, which
distinguish it from all other known species, are in this figure slightly drawn in at
the tips (which is accomplished by special muscles, shown in Fig. 4e).
FIGURE 46. — The same, fully contracted. The lateral processes are then turgid from
the pressure of the body-fluid, and are projected to their fullest extent and
somewhat forward.
FIGURE 4c. — The same, in the creeping attitude : usually the lateral processes are
drawn in, almost out of sight, among the skin-folds when creeping, but in this
instance they were fully extended.
FIGURE 4c£. — Jaw of the same with its two teeth and wide border.
FIGURE 4e. — One of the lateral processes of the same. It is seen to be hollow, and
special muscles enter it and are attached near the top. One of the vibratile tags
is seen at the base of the process. It is unusually short and wide for a Bdelloid.
FIGURE 4f. — Foot and rump of the same seen from the side. The prominent boss of
the preanal segment can also be seen as a curved line in Fig. 4c.
FIGURE 5a. — Philodina antarctica, sp. n. The figure shows the prominent collar, the
eyes on a conspicuous brain, the elongate foot, and the peculiar spurs. The
basal portion of the spurs is badly drawn in this figure (it is correctly shown in
Fig. 56).
FIGURE 56. — Spurs of the same. These, with the broad triangular bases and peg-like
apices, are the best character of the species. The distinction of the two parts is
sometimes obliterated. Even then the absence of interspace will distinguish
this from P. gregaria, and there are many other differences of general propor-
tions and details, requiring, however, careful study.
FIGURE 5c. — Lateral view of the foot of the same. The dorsal toes are almost as thick
as the ventral, though shorter. This is a point of difference from P. gregaria,
in which the dorsal toes are relatively very small.
FIGURE 6. — Philodina, unnamed species.
Brit Antarct Exped 1907-9.
Vol. 1. Plate X.
MURRAY ANTARCTIC ROTIFER A
J. ML
PHILODINAALATA.sp.il 5, P ANTARCTICA sp.n. 6. PHILODINA.
sp.
Murrav
PLATE XI
PLATE XI
FIGURE 7ot. — Philodina gregaria, sp. n. dorsal view of the animal when feeding. The
very prominent collar is only marked off from the pedicels by the cilia of the
wreath. Two well-grown young are seen.
FIGURE 7b. — Spurs and toes of the same. The dorsal toes are seen to be greatly
smaller than the ventral.
FIGURE 7c.— One of the spindle-shaped vibratile tags of the same.
FIGURE Id.— Jaw of the same, with two strong teeth, one thinner tooth, and many
fine striae.
FIGURE 7e. — Lateral view of the jaw of the same, showing the process of the ramus
behind.
FIGURE Sa. — Callidina habita, dorsal view of a variety which differs from the type
in the form of the spurs.
FIGURE 86. — The same, a variety having spurs nearer the typical form, but smaller.
FIGURE 8c. — Lateral view of the foot of the same showing the thickened part of the
first foot-joint which forms the boss.
FIGURE 8d. — Egg of the same, elliptical in form, with each pole produced into a
process.
FIGURE 8e. — Tip of rostrum of the same. The lamellae are widely separated.
Brit Antarct Exped 1907-9.
Vol. 1. Plate XI
MURRAY: ANTARCTIC ROTIKERA.
8.c
7. PHILODINA GREGARIA, sp.n.
Murray del ad nat.
8. CALLIDINA HABITA BRYCE.
PLATE XII
BRIT. ANTARCT. EXPED. 1907-9. VOL. I.
PLATE XII
FIGURE 9a. — Adineta barbata, Janson ?
FIGURE 96. — Egg of the same from Blue Lake ; one pole is produced, and there are
no other processes.
FIGURE 9c. — Egg from Deep Lake, Cape Barne, from which a similar Adineta was
hatched on September 24, after the egg had been dry for about a year, and had
been conveyed from Antarctica to Britain.
FIGURE 10a. — Adineta grandis, large stout example, showing six young, apparently
all in an equally advanced state of development, and with the jaws well grown.
FIGURE 106. — Head of the same, ventral surface, showing the lamellse, brush of
cilia and motile setse, pectinate folds at posterior margin of corona, &c.
FIGURE lOc. — Jaw of the same, showing the projection at back of ramus.
FIGURE Wd. — Yolk-mass of the same, dividing into six parts, each containing one
nucleus. The number of nuclei is normally eight, as in moat Bdelloids. In this
instance the full number may be present, two of them hidden behind the centre
of the mass.
FIGURE lla. — Adineta longicornis ? The rostral part is not of the same form as in
the type, but allowing for a different angle of inclination it may be the same.
FIGURE 116. — Spurs of the same. They are considerably shorter than in the type,
but are still relatively much longer than in any other known species.
FIGURE 12a. — Callidina angularis, sp. n., dorsal view, showing angular outline and
stippled surface.
FIGURE 126. — Jaw of the same, with four principal teeth and many finer ones
diminishing backwards.
FIGURES 12cand I2d. — Eggs found associated with this species and with G. constricta.
FIGURE 13a. — Callidina constricta, Duj., dorsal view, illustrating the differences
between this species and G. angularis ; the outline is not angular and the skin
is not stippled.
FIGURE 136. — Part of one disc of the corona of C. constricta, showing the groups ot
cilia which give the appearance of toothed wheels to the philodinoid corona.
Diagrams elucidating the motions of the cilia which cause the appearance of
teeth have often been published. This is not a diagram, but a drawing from an
example which had just been killed and fixed with osmic acid while in the act
of feeding. Each group has cilia at every stage of the stroke, thus showing the
course followed by each. The specimen was mounted in fluid and preserved.
Unfortunately, as was only to be expected, the slight difference of density between
the fluids within and without the body has resulted in making the disc turgid and
opened out the groups of cilia into one uniform fringe.
Brit Antarct. Exped 1907-9. Vol. 1. Plate Xll
MURRAY: ANTARCTIC ROTIFER A.
HI
10. A
9, ADINETA BARBATA^ 10, A GRANDIS, sp.-n. II, A LONGICORNIS
12, CALLIDINA ANGULARIS, sp n. 13, C. CONSTRICTA.
A I
PLATE XIII
PLATE XIII
FIGURE 14a. — Pleurotrocha, sp. A large active animal resembling P. grandis, but
with longer spurs and some other differences.
FIGURE 1 46. — Elliptical papillose egg of the same.
FIGURE 14c. — Jaws of the same, drawn from an example in the egg.
FIGURE 15. — Floscularia, sp., semi-contracted.
FIGURE 16a. — Diaschiza tenuior, lateral view with the foot drawn well under the
body.
FIGURE 166. — The same, dorsal view.
FIGURE 16c and 16cZ. — Dorsal and lateral views of a supposed male of the same,
having rudimentary jaws.
FIGURE 16e. — Rough sketch of jaws of the female.
FIGURE 16/ — Thick-shelled egg containing jaws similar to those of the supposed
male (Figs. 16c and
Brit Antarct. Exped 1907-9. yo] j pjate Xlll
MURRAY: ANTARCTIC ROTIFERA.
16. b.
14, PLEUROTROCHA, sp 15, FLOSCULARIA, SP. 16, DIASCHIZA TENUIOR
^Murray del ad nat
PART IV
*
MUSCI
NOTE SUB LES MOUSSES RAPPORTEES PAR L'EXPEDITION DU "NJMROD"
PAR JULES CARDOT
LES Mousses de 1'exp^dition du Nimrod proviennent de la me'me region que celles
rapporte"es anterieurement par le Discovery* Elles consistent seulement en
4 especes, dej& connues, mais dont 1'une toutefois est nouvelle pour la flore ant-
arctique. Les echantillons sont extremement rabougris, et tous donnent 1 'impression de
plantes malades, luttant pe"niblement centre des conditions de milieu exceptionelle-
ment rudes.
Alors que nous ne connaissons encore que 8 mousses ve"ge"tant mise'rablement sur
les cotes inhospitalieres de la Terre Victoria, M. Bryhn vient de publier f une liste
de 62 especes de Muscinees (57 Mousses et 5 Hepatiques) rapportees par Peary de
3 localites de la Terre de Grant, par 81°-82° de latitude boreale — chiffre supe"rieur a
celui des especes actuellement connues dans tout le domaine antarctique au sud du
60ime parallele.
La seule comparaison du nombre des especes connues d'une part a la Terre
Victoria, par 77°-78° lat. sud, et d'autre part a la Terre de Grant, sous une latitude
sensiblement plus elevee, fait ressortir d'une fagon frappante combien sont diflerentes
les conditions climateriques auxquelles sont soumises les regions polaires dans les
deux hemispheres.
•
DlCRANACE^:
Dicranella Hookeri
Dicranetta Hookeri (0. Mull.), Card, in Bull. Herb. Boissier, 2* sir., vi., p. 4, et Fl. bryol. Terres Magell.,
etc., p. 60.
Angstrcemia ffookeri (C. Miill.), Syn. II. p. 607.
* Of. National Antarctic Expedition, Nat. History, vol. ii., Musci.
t Ad cognitionem Bryophytorum arcticorum contributiones sparsae (Christiania, Videiifck.-Selhk
Fordhandl. for 1908, No. 5.).
BRIT. ANTARCT. EXPED. 1907-9. VOL. I. PART 4, ISSUED APRIL 1910 I,
78 J. CARDOT
Anisothecium Jamesoni, Mitt. Musci., austro-amer., p. 39, pro parte.
Dichodontium Jamesoni, Sch. Syn. Muse. Europ., ed. i., p. CO, et auct. plurim., pro parte.
Dicranella Jamesoni, Broth, in Nat. Pflanzenfam., Musci, p. 311, jro parte.
"CapeEoyds, 1908."
Echantillon en tres mauvais ^tat, d'un vert noiratre. Les feuilles sont tantot
entieres, comme dans la forme normale de 1'espece, tantot denticule'es vers I'extre'mite.
II me parait ne'anmoins certain que ce specimen appartient bien au D. Hookeri, espece
connue dans le domaine magellanique, a la Ge"orgie du sud, & Kerguelen et & 1'ile
Heard, et qui n'avait pas encore ete signalee dans 1'Antarctide.
II y a quelques tiges de Bryum argenteum, L., en melange.
POTTIACE^E
Sarconeurum glaciate
Sarconeurum glaciale (Hook, fils et Wils.), Card, et Bryhn, in Nat. Antarc. Exped., Musci, p. 3.
Didymodon (?) glacialis (Hook, fils et Wils.), Fl. Antarc., ii. p. 408, t. 152, fig. 6.
Sarconeurum antarcticum, Bryhn, in Nyt. Mag. f. Naturvidensk, B. 40, H. iii., p. 204, tab. i. et ii.
Barbula, sp., Gepp, Report on the coll. voyage Southern Cross, XXI. Cryptog., Musci, p. 319.
"Hut Point, Boss Island, 77° 50' S. lat., Coll. E. Joyce, Nov. 1908 ; Cape Irizar,
S. Victoria Land, Coll. T.W.E. David, Nov. 1908; Cape Barne, Ross Island, Coll.
J. Murray, Nov. 1908 ; High Moraines, Cape Royds, 77° 30' S. lat., Coll. J. Murray,
Jan. 1909."
Les rhizoides forment une masse feutre*e, compacte, me'langee de sable et de terre,
epaisse d'un centimetre environ, et limitee inferieurement par une surface nette. II
est probable que cette surface indique le contact avec la partie du sol qui ne de"gele
jamais et qui arrete ainsi la penetration des rhizoiides.
BRYACE.E
Bryum argenteum
Bryum argenteum, L., Sp. pi. p. 1120.
"High Moraines, Cape Royds, Ross Island, 77° 30' S. lat., Col. J. Murray,
Jan. 1909."
Forme excessivement rabougrie, identique aux dchantillons deja rapportds de la
meme region par le Discovery
" Stranded Moraines, McMurdo Sound, S. Victoria Land, Coll. R. E. Priestley,
1908."
MUSCI 79
Tiges e*mettant de nombreux rameaux tres greles, longs de 2 a 4 millimetres,
garnis de tres petites feuilles espacees, tres concaves, orbiculaires, ou suborbiculaires,
obtuses ou apiculees, rappelant beaucoup les feuilles du B. cephalozioides, Card., mais
les feuilles inferieures sont normales.
Bryum antarcticum
Bryum anlarcticum (Hook, fils et Wila.), Fl. Antarct. n. p. 414, t. 153, fig. 6.
Webera antarctica, Jaeg. Ad. i. p. 599.
" Stranded Moraines, McMurdo Sound, S. Victoria Land, Coll. R. E. Priestley,
1908."
I BRITISH ANTARCTIC EXPEDITION 1907-9
UNDER THE COMMAND OF SIR E. H. SHACKLETON, C.V.O.
REPORTS ON THE SCIENTIFIC INVESTIGATIONS
VOL. I
BIOLOGY
EDITOR JAMES MURRAY
PART V
TARDIGRADA
BY JAMES MURRAY, F.R.S.E., F.Z.S.
(WITH EIGHT PLATES)
LONDON
PUBLISHED FOR THE EXPEDITION BY WILLIAM HEINEMANN
21 BEDFORD STREET, W.C.
1910
ISSUED SEPARATELY. DECEMBER IfllO
PRICE TEN SHILLINGS NET
REPORTS ALREADY ISSUED
VOL. I. PART I., ON COLLECTING AT CAPE ROYDS
BY JAMES MURRAY PRICE Is. 6d.
VOL. I. PART II., ON MICROSCOPIC LIFE AT CAPE ROYDS
By JAMES MURRAY PRICE 5s. Od.
VOL. I. PART III., ANTARCTIC ROTIFERA
By JAMES MURRAY PRICE 5s. Od.
VOL. I. PART IV., MUSCI
BY JULES CARDOT PRICE Is. Od.
PART V
TAEDIGEADA
BY JAMES MURRAY
(WITH PLATES XIV.-XXI.)
CONTENTS
PAGE
INTRODUCTION .83
ON STRUCTURE OF TAHDIGRADA .84
ON NOMENCLATURE 91
ON THE VALUE OF SPECIES OF TARDIGRADA 91
ANTARCTIC TARDIGRADA 95
TARDIGRADA OF NEW ZEALAND .107
AUSTRALIAN TARDIGRADA 123
TARDIGRADA OF SOME PACIFIC ISLANDS 147
CANADIAN TARDIGRADA 159
SUMMARY OF RESULTS 179
ON DISTRIBUTION 179
BIBLIOGRAPHY .... 183
INDEX TO SPECIES 185
PLATES XIV.-XXI. . . .187
BRIT. ANTARCT. EXPED. 1907-9. VOL. I. PART 5, ISSUED DECEMBER 1910
M
INTRODUCTION
TARDIGRADA. were collected not only in the Antarctic, but in all other countries
visited by the Expedition in the course of the voyage round the world. These were
New Zealand, the Macquarie Islands, Australia, Fiji, Hawaii, and Canada. A brief
visit was paid to South Africa, and some moss was collected on Table Mountain by
Drs. Mackay and Michell, but though animals of other sorts were plentiful enough,
no Tardigrada were found.
The great majority of the Tardigrada obtained were of the k'nds which live among
moss. These everywhere over the world greatly exceed in numbers the true water-
dwellers. All, of course, are aquatic animals, in the sense that they must have at least
a film of water to support their active life, but the dwellers among moss have to
endure frequent and often long-continued desiccation, while the others live in ponds
and other waters of a more permanent sort. Though a number of species are common
to both habitats, there is no doubt that most are confined to one or other. The moss-
dwellers are adapted to withstand desiccation, the others may be equally so, but they
are rarely exposed to tests of their powers. By desiccation it is not meant that the
animals can endure the loss of all moisture from their bodies, but that they can live
when completely deprived of the external watery element. When so deprived they
are dormant, and if any physiological change goes on, it must be excessively slow.
Only some half-dozen species are well recognised as habitual pond-dwellers,
though in Scotland a good many others are frequently found in ponds, lakes, and
rivers.
It was only in New Zealand, Australia, and the Antarctic that there was much
opportunity of looking for the purely aquatic kinds. In those countries which were
only visited in passing there was no available method of collecting except by taking
dry moss to be examined afterwards at leisure. Nothing could exceed the facility of
this mode of collecting. The moss is preferably gathered dry ; the animals have
already become dormant in the natural course, and nothing is required for the
examination afterwards, but to moisten the moss, when in half an hour or so the
animals are found to be active. They will not live indefinitely, but for a year at
least they can quickly be revived when wanted, and some have been kept for a
number of years alive.
In this paper, after some introductory paragraphs dealing with structure, nomen-
83
13
84 J. MURRAY
clature, and other matters, the Tardigrada of each country visited will be treated in
a separate section, five altogether.
In the section dealing with each country there will be given first an account 01
how the collecting was done in that country, and a summary of previous knowledge
of the Tardigrada. Then will follow the list of species and notes upon them, includ-
ing descriptions of new species. The peculiarities of the Tardigrade fauna, and its
relation to the rest of the world, will be briefly treated. General questions of
geographical distribution, &c., concerning the entire area over which work has been
done, will be reserved for discussion in the general summary of results.
The information brought together about Tardigrada might have been exhibited in
a more concise form by giving a complete list of all the species collected in all the
countries, in one table, followed by notes on the species, and a discussion of general
questions of distribution. It seems to me, however, so important to the student to
find all the information about each country collected together by itself, that I
have adopted this form, even at the expense of a little repetition, and some redun-
dancy in the tables illustrating the distribution.
The object has been to offer the information in accessible form, so that the
student of one country may readily find what he wants. This has been attempted
by a suitable classification of the facts under various headings, and for further
assistance, there is a " Contents " page, and at the end an alphabetical index to the
species, while the explanations of the plates have references to the pages of the text
where the species figured are described.
We have thus what is practically a series of five papers, each complete in itself,
with Introductory and Recapitulatory chapters, dealing with all of them collectively.
STRUCTURE OF TARDIGRADA
The student is referred to other works for a general account of the structure of
Tardigrada. Here there will be given only such details as are necessary to the
understanding of the descriptions of the species.
Ecliiniscus. — In this genus the important points in description are : the number and
texture of the plates ; the processes on the body ; the claws ; and the skinfold (fringe)
of the last legs. The plates vary in number. The minimum may be taken as nine
and the maximum as twelve. There may often appear to be more than twelve
plates, but I regard these as originating in the subdividing of the normal plates.
The plates are either single or paired. The single plates are of two kinds : first,
large, covering the back arid sides ; second, small, triangular, median plates. The
pairs meet in the middle line, and go over the sides like the large single plates.
The number of segments of an Ecliiniscus have been variously estimated by
different writers. Doyere gives four body-segments, without regarding the head,
which he supposes to show traces of several segments. Schultze's E. heller-
TARDIGRADA 85
manni (43), has nine segments. What they mean by segments can only be
ascertained by studying the figures which these writers have given. Richters'
scheme is perhaps the most useful. It does not aim at accounting for all the
morphological segments of some hypothetical primitive or typical Tardigrade, but is
merely a division of the body for practical use to systematic naturalists.
He reckons six segments, and indicates them by roman numbers, I. to VI. These
numbers are here adopted to distinguish the segments, and as there are two distinct
types of segmentation I give two figures, one of each type, on which the segments
are numbered, and other points of structure indicated (see Plates XVIII. Fig. 34,
and XVI. Fig. 14).
In one group of Echinisci the segments V. and VI. have coalesced so completely
that there is no external mark of their separation (Plate XVI. Fig 14). This
includes the great majority of the species. Segment I. is the head, II., III., and IV.
the body segments bearing the first, second, and third pairs of legs, V. -f VI. bears
the fourth legs. The reckoning of V. as a segment is rather inconvenient, as there
is in most species no visible trace of it.
In the other group, which includes only about a dozen species, segments V.
and VI. remain distinct (Plate XVIII. Fig. 34). When V. is separate it usually
bears a pair of plates, like those of segments III. and IV., but it may be a half-ring,
without trace of division in the middle, as in E. pulclier here figured.
All of the plates are liable in one species or another to be subdivided in various
ways. This usually only affects the surface markings. The divisions may be merely lines
separating different areas, or they may be broad bands devoid of markings. The median
plates are often divided by a transverse line into two parts (E. novcezeelandice, Plate
XV. Fig. 5), or by median lines into pairs (E. pulcher, Plate XVIII. Fig. 34).
E. tessellatus (Plate XVI. Fig. 15) is an extreme case of the division of segments
II. and V. + VI. by transverse and longitudinal bands. E. scrofa, Richters (26),
is an example of subdivision of the median plates into many narrow bands.
The plates of the first and second pairs are commonly divided into two parts by a
transverse plain band (E. duboisi, Plate XVII. Fig. 19, E. spinulosus, Plate XIX.
Fig. 38). This band occupies a furrow. Sometimes the separation is a mere line,
and the markings continue into the furrow. Occasionally there are two lines on each
plate, separating narrow portions at the anterior and posterior borders, without any
cessation of the markings (E. tessellatus, Plate XVI. Fig. 15).
The markings of the plates are of several kinds. As their true character is often
in doubt they are named from their appearances. There are pellucid dots which do not
appear to project from the general surface, actual projecting granules (E. granulatus,
.Doyere, very large in E. tessellatus, Plate XVI. Fig. 15), pits or apparent perfora-
tions, and reticulation of raised lines. The reticulation may be simply the borders of
adjacent depressions (E. intermedius, Plate XVI. Fig. 17). Very peculiar are the
close-set spicules of E. spiculifer, Schaudinn (41), and the reticulation of pearly dots
86 J. MURRAY
of E. islandicus (18). A very few species are said to have no surface markings
whatever on the plates.
The lumbar plate, which covers segment VI., or V. + VI. when these are joined,
is nearly always divided by two lateral slits into a sort of trefoil. Often the middle
and lateral flaps thus produced are bent downwards at an angle to the small median
portion of the plate, which is then said to be faceted. The angle joining the two
slits has caused it to be supposed that the posterior " tail-piece " thus separated is a
distinct segment, and it has also been supposed that the portion in front of the slits
is homologous with segment V. This is undoubtedly not the case, as when V. and
VI. are separate, VI. is trefoliate in the usual way. A number of species have been
figured without a trefoliate "lumbar plate" _E. sintsbergensis, Scorn field (45), and E.
bisetosus, Heinis (7) ]. Personally I have only seen one species in which the lumbar
plate is not trefoliate (E. intermedium, described in this paper, see Plate XVI. Fig. 17,
and Plate XX. Fig. 52). In other cases I do not take the evidence of figures, unless
the authors refer to the character in their descriptions, as in certain positions the slits
are hidden and may be overlooked.
The processes connected with the plates are either setce, longer or shorter spines,
little spicules, or blunt cones or knobs. Certain processes often occur in definite
situations, easily indicated in descriptions. Near the mouth there are four cirri and
two palps, which are in most species of no distinctive value. In a few species they
are conspicuously large, or slightly modified in form [considerably in E. cornutus
(34)]. In one or two species they are stated to be lacking [E. imberbis (38)].
Of most regular occurrence are the five lateral processes and two dorsal processes,
on each side. Richters distinguishes the lateral processes by the letters a, b, c, d, e,
(Plate XVI. Fig. 14)* counting from the head backwards. « is the head seta, the
only process present in every known Echiniscus. It is situated at the base of the
head, between I. and II. The pair commonly curve forward, and look very like
cow's horns, but very thin, except in E. cornutus, where the resemblance is perfect.
At the base of seta a there is generally (perhaps always) a little triangular or
cylindrical palp or " Auricle " as it may be called. It is rarely distinctive but in a
few species is of characteristic shape or large size.
In a few species there are other processes besides these most common ones. The
most frequent are little spicules on the posterior borders of the plates of segments
II., III., and IV. between the lateral and dorsal processes. In such cases there may
be a similar spicule beside the slit in the lumbar plate, close to seta e, if that be
present. There may be more than one spicule between the lateral and dorsal
processes, especially on segment IV. (plates of the second pair), where there may be
two or three [E. oihonnce, Heinis (9)].
The dorsal processes usually spring from the postero-dorsal angle of the paired
plates, that is, at the angle marking the limits of the space occupied by the median
* Seta e is not present in the species figured.
TARDIGRADA 87
plates. Sometimes the dorsal processes are nearer the median line (E. borecdis, &c.),
and when there are any processes on segment V. they are usually near the middle
line (E. islandicus, E. novcezeelandice).
In the group which has segments V. and VI. united there are usually only two
median plates. A well-defined third median is rare, but there is often an area in
the triangular space behind the second pair, dotted exactly like the plates, but
without definite bounding lines.
In the group which has V. and VI. separate there are always three median plates,
although they are often very obscurely separated from the pairs in front of them.
In the majority of species there is a serrate fold of skin on the fourth leg. The
teeth or spines on this fold vary from short and blunt to slender and acute, and are
useful for distinguishing species.
Near the base of the fourth leg, on the outer side, there is usually a blunt palp.
On the first leg there is often a longer or shorter sharp spine. Rarely there are
little spines on other legs (E. perarmatus).
The colour of Echiniscus is usually red, varying from pale pink to deep crimson or
scarlet. A very few species are yellow (E. mutabilis, sylvanus, tessellatus), green
(E. viridis) or colourless (E. intermedius).
The eggs are always smooth, shortly oval, and are laid in the cast skin. From
one to nine have been seen in a single skin. As many as nine is quite exceptional,
the usual number being three or four.
Oreella. — Like Echiniscus in every important detail of structure except that the
body is not covered with plates. It is soft and flexible like Macrobiotus. The only
trifling differences from Echiniscus are that the " Auricle " at the base of seta a is
elongate and acute, and is elevated on a large papilla which bears the seta also ;
and that the anterior cirri at the mouth are replaced by narrowly conical processes.
The generic forms of land Tardigrada do not seem to be very numerous.
Although work has been done in all regions of the globe now, no new genus has
been found since Diphascon (Plate, 1888) till now.
Oreella has no eye-spots. The eggs are unknown.
Milnesium. — Though several species have been described, their differences are
slight and inconstant, and only M. tardigradum is here admitted. It is a large
animal with very wide gullet and elongate pear-shaped pharynx, in which there are
no rods such as Macrobiotus has. There are six cylindrical palps round the mouth,
and a pair farther back. A distinct genus, Acrophanes, of no value, was separated
by Ehrenberg (5) apparently to receive a shrivelled skin of Milnesium.
The claws are very different from those of Macrobiotus. On each foot there are
two claws, usually with two or three, or even four, branches. The three-branched claw
appears to be homologous with a pair of Macrobiotus claws, including the supple-
mentary point. On each foot there are also two very slender, bristle-like claws, each
with a very fine supplementary point.
88 J. MURRAY
The number of points on the branched claws has been used to distinguish species.
Some examples have all these claws alike, and three-branched. In others they vary on
the different legs. I find the commonest form to have fewer points on the front legs
and more on the last. Thus, they will have only one point or two on the front legs ;
two or three on the second and third legs, and three on the fourth legs.
The colour in old animals is a very warm brown. The eggs are laid in the skin,
and are often pretty numerous.
Macrobiotus. — In this genus the characters which lend themselves for descriptive
purposes are : the organs of the mouth (teeth, gullet, and pharynx) ; the claws ; the
skin ; the eyes ; the fat-cells ; and the egg. The teeth are slender or thick, more or
less curved or bent abruptly, and have a larger or smaller furea where the bearer
is attached, the bearer itself, though often mentioned in descriptions, having no
specific importance. The gullet is slender or wide, and at the end in the pharynx
may have merely a slightly enlarged run, or may be bent out into a flange-
more or less wide. The pharynx may be nearly round, or somewhat elongated,
The hard rods or nuts which surround the tube may be two, three, or four in each
row. The last one in the row is often small and obscure, and is called from its
shape the comma. There is often no comma. The more important rods (or nuts
when they are so short that the name " rod " would be inappropriate) are never more
than three in number. When there are three they are generally nearly equal in
length : when there are only two, that nearer the gullet is longer, and often
shows obvious traces of the two rods by the joining of which it may be supposed to
have originated. Besides the rods there are apophyses attached to the end of the
gullet. These are not in line with the rods, but alternate with them. When they
are large, however, they may readily be mistaken for rods, and even recent
descriptions have not always clearly distinguished the two structures.
The claivs vary chiefly in their relative sizes, and in the degree of union of the
pairs. They are always united in pairs, if only at the bases. Several types have
been recognised, but they are connected by intermediate forms which are difficult to
classify. The hufelandi type is commonest. The claws of each pair are united for
some way above the base, usually for half their length or more. There is much
variation within this type. Usually the claws of a pair are unequal, and the larger
one has one or two supplementary points. Sometimes they are equal and placed side
by side, but only one has the extra points. In some species they are very thick, and
the supplementary point is very strong, and the whole thing makes a distinct
approach to the branched claw of Milnesium.
The dispar type (or macronyx type) has the claws of each pair very unequal, the
larger claw strongly curved and bearing a supplementary point, the smaller claw like
a little barb from the base of the larger one. The two pairs are similar, and on the
last legs the lesser claw is relatively larger.
The Diphascon type (found throughout the genus Diwhascon, and in M.
TARDIGRADA 89
oberhciuseri and a few other Macrobioti) has the two pairs dissimilar. One pair has
two claws, of which one is longer, united at the base ; the other pair has one very
long claw (bristle-like in M. oberhanaeri), which is loosely attached to the middle of
the back of a smaller claw.
The echinogenitus type has two similar pairs of claws which are united at the base
only. The claws of each pair are equal or unequal. This type is not so distinct from
the hufelandi type as it seems, and is connected with it by a series of forms
having the claws united more or less above the base.* The typical form has the
claws very widely divergent. Forms having the pairs unequal lead to the
Diphascon type.
The skin varies in texture, &c. It may be smooth and hyaline, papillose, or
more or less pigmented. Often the young are transparent and colourless, and the older
ones increasingly pigmented. The pigment is rarely of specific value, as most of the
large old animals become brown, but in M. oberhciuseri it is characteristic. A few'species
are papillose (M. annulatus, M. nodosus, &c.), and a few bear processes on the skin
(M. ornatus, M. tuberculatus, M. sattleri, M. papillifer, and sometimes M. dispar).
The eyes are of doubtful specific value. They are usually dark brown or black,
occasionally red. Though it is believed that the presence or absence of eye-spots is
characteristic of each species, there is enough variation to make the character
unreliable. It should, however, always be noted in descriptions.
The fat-cells are generally of no specific importance. In a few species (M.
coronifer, M. islandicus, M. rubens, M. occidentalis, &c.) they have a characteristic
colour, red, yellow, orange, or brown.
The egg varies greatly. It is either smooth or covered with processes which are
characteristic for each species. The smooth eggs are never quite spherical, but
shortly oval, rarely narrowly oval. They are left in the skin at the moult. The
ornamented eggs are spherical, with a very few exceptions (M. coronifer, M. pullari,
&c.). The form of the processes should be noted, the distance (if any) separating
them, and the nature of the egg-shell where free from processes. In a number of
forms the shell is areolate. It is thick, and composed of two layers, the space
between which is divided into a number of equal chambers, showing on the surface
as hexagons. From some of these the processes spring, at equal distances apart.
The turgidity of the processes destroys the regularity of the hexagons, and additional
septa in the chambers give rise to a variety of patterns in the " areolation."
The spiny eggs are laid free, not enclosed in the cast skin. An intermediate type
of egg has the shell ornamented with rods, which are embedded in a hyaline substance,
so that the surface is even (M. arcticus, &c.). In M. hastatus these rods have tops
shaped like fleurs-de-lis, which project above the general surface.
* Prof. Richters informs me, in a recent letter, that he has ascertained that M. echinogenitus and related
species have the claws separate, but close together, in embryos in the egg, and that they join afterwards.
The adult M. echinogenitus has claws of the hufelandi type, not V-shaped.
BRIT. ANTARCT. BXPED. 1907-9. VOL. I. N
90 J. MURRAY
Many species of Macrobiotus cannot be distinguished with certainty unless the
egg is seen. The mere presence of eggs beside the adults is of no use, although there
may be a presumptive relationship in such cases. The relation must be demonstrated,
and this can be done in two ways : first, by finding in an egg the young so far
advanced that the claws and pharynx can be recognised ; second, by finding, in the
adult, eggs so far advanced as to show the characteristic processes. The first way is
the more generally useful. The second may be of greater use than the beginner
would suspect. The eggs are soft till they are almost ready for laying, and the
processes, when formed, are still so soft that they are squeezed flat against the shell
by the enclosing membrane. If the membrane can be torn, and the eggs released, the
processes often stand up and can be recognised.
All species of Macrobiotus, and probably other water-bears, may be found in what
is called the " simplex " state. They are then devoid of teeth, of rods in the pharynx,
sometimes of all the mouth parts. Plate founded the genus Doyeria on an animal in
this condition. The simplex state is now explained as a part of the moult, though
it does not occur simultaneously with the casting of the skin. Identification of an
animal in the simplex condition is difficult, as one of the most important characters
is lacking.
Many water-bears, perhaps all, encyst themselves. This appears to be a sort of
hibernation, in which a quiescent period is passed in a special protective case. Some
species, when encysted, undergo a simplification going far beyond the ordinary
" simplex," for they lose all recognisable organs, teeth, pharynx, and claws, and
afterwards acquire them anew.
Diphascon. — The genus differs from Macrobiotus solely in having the gullet
elongated between the teeth and the pharynx. As specimens of Macrobiotus are
sometimes found which have the gullet elongated, the value of the genus is very
slight. Nevertheless, I believe Diphascon to be a natural group. All the species
have claws of one type, known as the Dipliascon type, which is also found, but rarely,
in Macrobiotus. Many of the species have the pharynx very narrow, but others
have it quite round. Most of the known species have no eye-spots.
In distinguishing species the chief points to attend to are the diameter of the
gullet, which varies extremely, the proportions of the pharynx, and the number and
sizes of the rods or nuts in it.
In outward form there is little variety. All are very similar except D. bnllatum,
which is short, nodose, and papillose, and D. augustatum, which has a heavy body,
but tapers to a very narrow head.
As a rule the animals of the genus are small, but some very large forms inhabit
the arctic regions.
The eggs are smooth, and are laid in the cast skin, as in Echiniscus and a large
section of Macrobiotus,
TARDIGRADA 91
NOMENCLATURE
The nomenclature of the Tardigrada is in need of revision. The validity of many
of the accepted names, generic and specific, may be questioned, and some of them
will certainly not hold. Professor Hay (6) has suggested a number of corrections
which must be made. He has shown that the water-bears have no right to bear the
name Tardigrada, which is preoccupied (for a suborder of Edentata).
The recognition by the earlier naturalists of species having two, three, and four
claws may be explained as arising from different interpretations of the claws of
Macrobiotus and Milnesium. Macrobiotun may be said to have two forked claws on
each foot, or to have four claws united in pairs, just as you please. It is curious
that so many good naturalists have recorded a water-bear with three claws, which
nobody finds nowadays.
Milnesium is commonly supposed to be the same as Schrank's Arctiscon, in which
case the earlier name must be used, unless it can be shown to be preoccupied. If
Schultze's Macrobiotus hufelandii is accepted as the common water-bear, and the
same as Miiller's Acarus ursellus, Miiller's specific name should be used for it. All
these points would require a very careful revision of the whole literature of the
group, and a comparison of all the early descriptions and figures.
The revision is so important that it should be made in a work of monographic
character, or at any rate should comprehend the whole group. While, therefore, in
perfect agreement with Professor Hay as to the necessary changes, I shall in this
paper continue to use the familiar names. By this means the present report will be
kept in line with a whole series of others dealing with the water-bears of various
countries under the name " Tardigrada." This report has been announced in the
earlier numbers of this publication under that title.
When changing the name it is desirable that the change should be final, if that
be possible. It would require a careful examination of all writings on the subject
prior to 1835 to satisfy ourselves that Perty's name Xenomorphidce (22) [which
Professor Hay shows to have precedence of Schultze's Arctiscoida (44) ] is really the
earliest applied to the water-bears as a family.
The name Xenomorphidce is a family name, and the group of the water-bears
must be considered as of more than family value. Schultze's name Arctiscoida,
although proposed as a family name, might be adopted as an ordinal name for the
group, and Xenomorpkida for the only family as yet recognised.
ON THE VALUE OF SPECIES OF TARDIGRADA.
In the course of this Expedition a great many diverse forms of Tardigrada were
collected or observed. Some of these are described in the following papers as species,
92 J. MURRAY
one even as of generic rank. Had the early practice been followed, whereby species
of the genus Echiniscus were supposed to be sufficiently characterised if they differed
by one pair of spines or setoe from the related species, or if even a seta were replaced
by a spine or vice versa, a hoat of new species would have been added. I prefer a
definition of species in that genus which excludes such forms, and requires species to
show peculiarities in other points of structure than the spines (such as claws, or plates,
or texture), or to have the spinous armature conspicuously different from that of the
nearest relatives.
No definition of species can be made which will be universally applicable. In
some groups it is necessary to distinguish species by characters so slight that they
would not be admitted in other groups. Species corresponds to no entity in Nature —
it is a human convenience. We may seek to distinguish by it such races of any
organism which have through some degree of isolation and the tendency to vary
(under influences of which we are profoundly ignorant) acquired more or less distinct
marks by which they may be distinguished from the related races of the same stock,
and which we believe to be constant (in the sense that they continually recur, and
that we believe the recurrence to indicate community of origin). The amount of
distinctness considered necessary to constitute specific value gives an opportunity for
the personal element in naturalists to assert itself, and has, incidentally, given occasion
for much disputation.
Every species throws some light on the biological history of the world, and so we
want to discover them all. Such cheap species as the ardent discoverer could so
easily manufacture out of the myriad forms of Echiniscus would yield the minimum
of light on biological problems. In so far as they are permanent, they are only
fortuitous side eddies in the stream of evolution, if such a term can be per-
mitted a popular application in science. I mean that the species having two
spines and those having three, and not otherwise differing, are not instances of
essential adaptation to the conditions of life of the animals, but that one is as
good as the other. Further, the probability is that many of these slight forms are
really not species in even the widest acceptation of the term, but merely individual
peculiarities.
Among Echinisci there are forms known, such as E. granulatus and E. quadri-
spinosus, var. cribrosus. '[see Murray (12) and (18)] which possess several good charac-
teristics, besides their distinctive spines. In colonies of such species, which may be
confidently identified by the concurrence of many characters, it continually happens
that individuals occur in which one or more of the customary spines is lacking.
That it is frequently only one spine of a pair which is absent indicates that these are
:nly errors or peculiarities in the development of the individual. Yet by the accepted
practice one pair of spines or seta) makes a species.
There is next to be considered the case where these slight peculiarities seem to be
constant. Admittedly when example after example turns up, having all the trifling
TARDIGRADA 93
features identical, the idea of the value of the characters grows. This happens most
frequently in colonies got at one place and time. Now a whole brood, or even a
whole colony, may possess some little peculiarity which would not be permanent in a
race of animals having the sexes distinct, and where cross-fertilisation may occur.
The case of the Bdelloid liotifera, which will be discussed in a subsequent paper, is
very different, as only females are known. The reproduction being parthenogenetic
there seems to be no check to the multiplication «f slightly marked species, and the
study of certain groups of forms bears this out.
The group of Tardigrada has hitherto been fortunate in that it has not attracted
the attention of any too-enthusiastic discoverer of species. Professor Richters has
shown a commendable reserve in simply describing and figuring those doubtful
Echinisci, without giving them names.
This appeal's to be the right course to take, in the present state of our knowledge,
and yet these unnamed forms are a trouble to the student. There is nothing to get
hold of, and the memory recognises them not — even reference to them is laborious.
Since species are a human convenience one is sometimes tempted to wonder whether
the convenience of naturalists is better served by describing unnamed forms than by
lightly naming them in the good old way, with, however, its accompaniment of
burdening the synonymy when we come to know better.
In the genus Macrobiotus the species difficulty takes another form. There is little
external variability. As few of the species have superficial processes there is little
difficulty of the sort we have in Echiniscus.
Variability is seen in the coloration — the animal may be colourless when young,
and highly pigmented when old ; in the claws — the amount of union of the pairs
may vary considerably ; in the rods of the pharynx — the first rod, next the gullet,
may be a long one, or divided into two shorter ones, and the " comma " may be
present or absent ; in the eyes, which may be present or absent. Some of these
differences are known to be dependent upon age. The limits and value of others are
not definitely known.
Two forms of Macrobiotus may differ in two of the most important characters used
in discriminating species (claws and pharynx for example), yet the range of variation
of each of these characters, taken separately, is wide enough to embrace both forms.
Whether it is likely that two or more characters would vary simultaneously, to the
extreme extent, in animals which are of the same species, we cannot tell. If it were
so specific distinctions in the genus would to a large extent break down, and we would
have only a few species, representing the principal types of claws recognised
(hufolandi, oberhiiusem, dispar), or of eggs (the hufelandi type, the hastatus type,
and the smooth eggs).
Another difficulty with Macrobiotus is that series of species agree so closely in all
the characters of the adult that they cannot be distinguished with certainty, and it is
necessary to find the egg before a species can be identified. These series of closely
94 J. MURRAY
related species only occur in tliu groups which lay spiny eggs, or at least the eggs
are only of value in that group.
Now the eggs do not show the characteristic processes while they are in the body
of the parent, and once they have been deposited there is nothing to indicate
their relation to one animal or another. The actual act of deposition can rarely be
observed.
While the eggs are still soft and their shape readily distorted by the
pressure of the enclosing membrane, and of adjacent eggs, the processes are often
already formed, and if the membrane be ruptured, the external pressure being
removed, the turgidity of the egg causes it to assume its proper shape, and the
processes then stand out. The opportunities for completing an identification in this
way must be casual and uncertain, and it is oftener possible to find an egg in which
the young is so well grown that it shows the distinctive characters of the adult.
While many species are only distinguishable by their eggs, the converse occurs.
Quite distinct species have eggs exactly similar, or so nearly alike that identification
cannot be certain.
These facts show the necessity there is for making an exhaustive study of each
species whenever the opportunity offers, and the danger there is that species will be
multiplied to no good purpose, if such study is neglected.
Bibliography. — Throughout the text references to the bibliographical list are
made in heavy type enclosed in brackets.
Tables of Species. — In the lists of species given to illustrate distribution there
was some difficulty in presenting the facts concisely. To classify the records of
Tardigrada broadly the world was divided into nine great regions, one for each of
the five continents, for Australasia, and for the Antarctic and Arctic Regions. As
regards these eight divisions the classification is obvious and natural enough. The
ninth division (Oceanic Islands) was introduced to receive a lot of scattered records,
not otherwise provided for. This is not satisfactory, as the islands are so scattered.
Those which lie near continents are included with them, the others are all slumped
together. It is a makeshift which may serve for the moment. Greater accuracy is
not called for till a more comprehensive survey of the distribution of all the Tardi-
grada is attempted.
I have given all the records I could find, but many may have been overlooked.
Some discrepancies may be found between the number of species for any one region
given in different tables. This is due to the fact that many records were added from
time to time after the tables were first compiled, and it could hardly be hoped that
no errors would occur in collating so many tables.
In dealing with each country the local distribution is treated with more detail,
and in addition to the nine main divisions of the earth's surface there may be given a
column for such adjacent regions as may be supposed likely to show some relation-
ship. Thus with the Antarctic a sub-Antarctic area is distinguished.
ANTARCTIC TAEDIGRADA
On collecting Antarctic Tardigrada. — The methods employed in collecting the
fresh-water organisms in general have been detailed in Part I. of this volume, and
need not be repeated here. The water-bears were collected among the vegetation in
the lakes, and also among moss. They came next in importance after the Rotifers,
but not on account of being numerous in species. They were conspicuous in the
lakes from the great abundance of one species (Macrobiotus arcticus), which occurred
in numbers comparable with those of the two Bdelloid Rotifers, Philodina gregarict
and Adineta grandis.
Two species were got in the lakes. The same two occurred among moss, with
two others.
Priestley collected moss on the west side of Macmurdo Sound, at the Stranded
Moraines. It contained three species, one of which was not known at Cape Royds
(M. polaris), and one species found on Ross Island (M. oberhtiuseri) did not occur at
the Stranded Moraines.
All the Tardigrada were collected between 77°30' and 78°S.lat. No Tardigrada were
found among the mosses collected by Professor David about 2 degrees farther north.
Summary of previous knoivledge. — The water-bears are better known than any
other group of Antarctic fresh-water animals. Half a dozen papers on the subject
have appeared, most of them by Professor Richters. The first mention I can find of
an Antarctic Tardigrade is by Richters, who in 1904 (31), in his preliminary report
on the Antarctic Moss Fauna, described a species (Macrobiotus antarcticus) found by
the German " Gauss " Expedition on the Gaussberg, a hill on the Antarctic continent,
and just within the Antarctic Circle. This was the only truly Antarctic species
recorded in that paper, the other ten species being from islands in the southern ocean.
In 1906 there appeared the report on the Tardigrada collected in the South
Orkneys by Bruce's Scottish Expedition [Murray (15)]. These islands are truly polar
in climate, although lying outside the Antarctic Circle and distant from the continent.
Fifteen distinct species were found, but owing to the condition of the specimens it was
only possible to name six of them, three of which were known and three were new.
Richters' full report on the Tardigrada of the Gauss Expedition appeared in 1907
(35). It made no addition to the list of Antarctic species, though it extended the
list of species known from the islands of the southern ocean to seventeen.
95
96 J. MURRAY
In 1907 (36) Richters published also a summary of all that was known of Ant-
arctic Tardigrada, including the results of the work of the German, Scottish, and
Swedish Expeditions (although the report on the last had not then appeared). He
gives a list of twenty-three species recognised, and mentions fifteen others which had
been imperfectly observed, but he does not in that paper discriminate between the
Antarctic and the sub- Antarctic species.
The Tardigrada of the Swedish Expedition were described by Richters in 1908 (38).
In a table at the end of this paper he enumerates ten species from the Antarctic
Region proper. Adding one species collected by the Scottish Expedition and one by
the German, we have a total of twelve Antarctic species recognised and named,
besides many insufficiently studied.
Lastly, in 1909 (40), in a short note in the Zoologischen Anzeiger, Richters
noted three species found in moss collected by the National Antarctic Expedition,
under Captain Scott, in S. Victoria Land. Only one of these was sufficiently studied,
and was found to be a new species, and described as Macrobiotus meridionalis.
There were thus thirteen species recorded for the Antarctic. We collected only
four species, but ,of these three were new to the Antarctic and one was new to
science.
The full list of the Antarctic species, with their relation to sub- Antarctic and
other regions, will be given in tabular form at the end of this paper.
LIST OF SPECIES
Macrobiotus arcticus, Murray.
M. oberhauseri, Doy6re.
M. polaris, sp.n., Murray.
Diphascon alpmum, Murray.
Diphascon or Macrobiotus^).
NOTES ON THE SPECIES
GENUS — Macrobiotus, Schultze (42)
Macrobiotus arcticus, Murray (19) (Plate XIV. Figs. 2a—2f)
Specific characters. — Large : young hyaline, adult pigmented. Teeth strongly
curved ; gullet narrow ; pharynx shortly oval, with two short rods, the second
shorter, and no comma. Claws large, of the Diphascon type, one pair with nearly
equal claws, united for some way above the base, the other pair with a very long
claw springing from the back of a shorter thick one. Egg oval or round, thick-
shelled, studded with short rods which are embedded in a hyaline substance. Dark
eyes.
ANTARCTIC TARDIGRADA 97
As M. arcticus was originally described from an egg containing the young ready
to hatch, and the adult had never been seen, it is here redescribed from adult
examples. The principal differences are that the rods in the pharynx are longer in
the adult (they are nearly round or quadrate in the young), and that it acquires a
brown pigment like that of M. hnjelandii.
Detailed description. — Length 500 /u. and upwards. The young is perfectly
hyaline, and has the long claws as slender and bristle-like as those of M. oberhciuseri.
The long claws become a little thicker in the adult. They are strongly curved
towards the tip, which is not the case with oberhauseri,
The gullet is narrow, and is expanded at the end in the pharynx into a prominent
thickened flange. In the adult the rod nearest the gullet is between three and four
times as long as broad— the second rod is a little shorter, three times as long as
broad, or rather less than that.
The stomach is voluminous : about a dozen of its component cells are visible in
dorsal view. They are filled with large granules of a brownish colour, and there are
deeper brown patches. About the middle of August many were observed which had
the stomach green or deep blue. Macrobiotus with blue stomach has been noticed
in Scottish lochs and in the Arctic.
Four unripe eggs have been seen in the body at once. The eggs are spherical or
very shortly oval, 96 p. in diameter. The thickness of the shell is about 5 or 6 n, but
varies considerably. The eggs often appear to be viscous, and have much adherent
material : some are quite clean and don't seem viscous. It may be that they are
viscous when first laid, and become hard afterwards.
Habitat. — In nearly all the lakes about Cape Royds and Cape Barne, extremely
abundant.
It was the first water-bear which we found. On March 15 we obtained it, and
also eggs, among dried vegetation on the ground. This had doubtless originated in
a lake. It was most abundant in Coast Lake, where we could always get great
quantities.
In Blue Lake on July 17 it was found alive at the bottom, on a thin pellicle of
vegetation under 1 5 feet of ice.
In the experiments made to test the vitality of rotifers, many water-bears of this
species were present with the rotifers. They endured equally well exposure to the
greatest Antarctic cold, and repeated freezing and thawing, at weekly intervals, for
months. They were not subjected to heat.
After being kept dry for a year, and conveyed on a voyage through the Tropics '
to England, no adult animals revived, but the eggs retained their vitality and some jx•
Scourfield.
M. arcticus appeared to be a thorough pond-dweller, and was never found among
mosses at Cape Royds. Nevertheless, in mosses which Priestley brought from the
BEIT. ANTARCT. EXPED. 1907-9. VOL I O
98 J. MURRAY
Stranded Moraines, on the west side of Macmurdo Sound, they were plentiful, and
there were many eggs.
These eggs differ from those got in the lakes, but not enough to cause us to
regard it as a distinct species. They are smaller and much thinner-shelled. The
shell is so thin that the rods which traverse it are scarcely longer than broad. They
do not seem viscous. The animal does not differ at all. The thinner-shelled eggs
may be summer eggs.
The distribution of M. arcticus is peculiar. As far as our meagre knowledge
goes it is confined to the polar regions, but an egg of the thin-shelled form has been
got in a Scottish loch, and a similar egg, but smaller, in tropical Africa. In various
countries visited on the homeward voyage (New Zealand, Australia, Canada) an
animal was found which is almost certainly this species, as it has all the characters
of the adult, but as no eggs were found there is a certain doubt about the
identification.
M. arcticus has only one near relative, M. hastatus, Murray (18), an inhabitant of
peat-bogs. Several Macrobioti are now known which have claws of the Diphascon
type, but only these two species have the peculiar eggs, studded with rods.
Macrobiotus polaris, sp. n. (Plate XIV. Figs, lo-le)
Specific characters.— Large, hyaline, or old examples somewhat pigmented.
Gullet of intermediate width : teeth curved : pharynx shortly oval, with apophysis,
and three short rods and comma in each row. Claws slender, of hufelandi type,
very unequal, and united less than half-way. Eggs spherical : processes from
polygonal bases, varying from round to acuminate, separated by polygons of equal
size to their bases. Dark eyes.
Detailed description. — Length, up to 800 /x- The egg measures 75 M in diameter,
exclusive of the processes, and about 85 M over the rounded processes, or 95 M over
the pointed ones. A young one squeezed out of an egg was 250 /JL in length, which
is a very large animal to come from such a small egg, but it may have been slightly
elongated by pressure. The animal is somewhat narrow and elongated. The gullet
is about 4 n wide, with parallel sides, and bent out to form a narrow flange at the
end in the pharynx. The pharynx measures 36 M by 30 M- The three rods are of
nearly equal length and are rounded at the ends. They vary in length, according
to age, from 1^ to 2|- times as long as broad. The stomach is narrowed at both
ends, and consists of numerous cells, about twenty being in view at one time. The
fat-cells are small, 5 or 6 M in diameter. The body-fluid in old animals is pale
yellow, the stomach sienna-brown.
The claws are unusually slender for a large animal of the hufelandi group. The
lesser claw of each pair is little more than half as long as the larger one. Only one
supplementary point could be seen on the long claw.
ANTARCTIC TARDIGRADA 99
The egg is extremely variable. It is one of the "areolate" type (see M.
areolatus, p. 167). The shell appears to be double, and the enclosed space is
divided into a number of equal cells or chambers by septa which appear on the
surface as a hexagonal reticulation. From certain hexagons, at equal distances
apart, the processes spring. They are separated by one hexagon. They vary
greatly in form. Some are rounded at the top and appear as semicircles in optical
section : others are conical and acute : others again acuminate, with slender points.
The turgidity of the processes causes pressure upon the surrounding hexagons, and
thus interferes with the regularity of the reticulation. Sometimes the processes
appear wrinkled or shrunken at their bases, when they may become smaller than the
intervening polygons. .
Habitat. — Among moss collected by R. E. Priestley at the Stranded Moraines on
the west side of Macmurdo Sound, very abundant, eggs also numerous.
After being dry for more than a year, several adults were found alive when the
moss was moistened on February 15, 1910. Some eggs were hatched, thus completing
the life- history of the animal.
M. polovris has affinities with a great many species forming what may be called
the hufdandi group. Most of the species are characterised by eggs having very
distinctive characters, but it must be admitted that several of them could not be
recognised with certainty, unless the egg were found, and its relation to the animal
demonstrated by hatching or otherwise. It is not one of the species nearest to
M. hufelandii, which have very wide gullets. It is only necessary here to point out
the characters in which it differs from those species which have somewhat similar eggs.
The nearest relative is perhaps M. ineridionalis , Richters (40), with which I at
first identified it. Professor Eichters' figure does not give all the details necessary
to separate the species, but he informs me in a letter that the processes of the egg
are hemispheres with a small discoid projection on top, and that the surface is not
reticulate in the manner described above. This form of egg I know as occurring on
Scottish mountains, and it is quite different from M. polaris.
M. echinogenitus and M. areolatus have the claws joined near the base only : the
former has not a reticulate shell, and the latter has a much larger egg and no comma
in the pharynx.
M. montanus (nee p. 116) has the processes hemispherical, almost touching at
their bases, and the surface not reticulate. M. harmsworthi has the processes
acuminate, and close together on the shell, which is not reticulate.
Macrobiotus oberhciuseri, Doyere (2)
As an Antarctic species this is only known from a single example, found in moss
from the High Moraines at Cape Royds, January 1909. A photograph of the animal
from life is shown in Part 2 of this volume, Plate III. Fig. 11.
100 J. MURRAY
The animal is strongly pigmented, of the characteristic colour, and is smooth all
over, not papillose as in many examples from tropical regions.
GENUS — Diphascon, Plate (23)
Diphascon alpinum, Murray (14) (Plate XIV. Fig. 3)
It has been pointed out by Professor Richters that the southern representative of
D. alpinum is not precisely like the type as found on Ben Lawers in Scotland. The
Antarctic form has a comma in the pharynx, in addition to the three rods. The
comma is not noticed in the original description or figure, but at that time I do not
think the comma had begun to be taken notice of as a serious character.
In the Antarctic examples the three rods do not increase in size so much from
first to third as in the type, and the claws appear to be relatively larger and more
slender. All these differences do not justify the separation of a new species without
fuller study.
At Cape Royds D. alpinum was much less common than M. arcticus. It was
present in a good many of the lakes and ponds, but was never abundant, perhaps
most so in Terraced Lake, Cape Barne.
It was first found in Blue Lake on March 27, 1908, and in that lake, on
July 17, a skin with three smooth oval eggs, very probably of this species, was
got at a depth of 15 feet, on a thin film of vegetation which covered the stones on
the bottom.
One example measured 200 /j. in length, but that was undoubtedly a small one,
though I have no measurements of any larger. The type was 250 /* long, and some
examples from the South Orkneys were 360 /x in length.
At Cape Royds it was never found among moss, but at the Stranded Moraines it
did occur. There also three eggs were found in a skin.
It is now recorded for Spitsbergen and Canada. There are no notes as to
the " comma " in these countries.
Diphascon (?), sp. (Plate XIV. Fig. 4)
A small animal, of which only one example was found, is referred doubtfully to
the genus Diphascon, to which it technically belongs, having a slightly elongated
gullet, and claws of the Diphascon type.
The animal is smooth and hyaline. It measures 300 M in length. The teeth are
small, and curved strongly outwards towards the furca. The gullet is of moderate
width. It is about twice as long as the teeth. The pharynx is shortly oval
and contains two rods in each row of thickenings. The end of the gullet in the
pharynx is expanded into a broad flange, to which the apophyses are attached. The
ANTARCTIC TARDIGRADA 10]
first rod (nearest the gullet) is about three times, the second about twice as long as
broad. There is no "comma."
The claws are of the Diphascon type, but the longest claw is less in excess of
the others than usual. The lesser pair has one claw considerably longer than the
other.
The eggs are unknown ; eyes were not seen ; there are no notes about the
stomach or fat-cells.
The animal is not sufficiently studied for identification. Although technically a
Diphascon, there is some reason to think that it is a sport of a Macrobiotics. It
will be noticed that, apart from the elongated gullet, the teeth and pharynx are very
like those of M. arcticus (Fig. 26).
As that species also has claws of the Diphascon type, there is little to separate the
two. The elongation of the gullet is less than in any Diphascon except angustatum,
and the portion between the bearers and the pharynx did not appear flexible.
A similar slightly elongated gullet is known in an animal having claws of the
dispar type, and therefore certainly no Diphascon.
Found among dried vegetation, probably of lake origin, which also contained
M. arcticus and its eggs.
REMARKS ON THE ANTARCTIC TARDIGRADE FAUNA.
The Antarctic is defined for the purposes of this paper as comprising the whole
Antarctic Continent, not excepting the small part which lies outside the Antarctic
Circle, and such islands lying off the coast as enjoy a similar frigid climate to that of
the continent itself. It is a region where the summer temperature of the air is never
very far above freezing-point. The sea is for the most part permanently of a tem-
perature somewhere between the freezing-point of fresh and that of sea water. It
is a desolate and inhospitable region, presenting a remarkable contrast to the North
Polar region, which has a relatively genial climate in latitudes several degrees higher
than any in which biological studies have been attempted in the south.
So far as known there is no life in the Antarctic except on or very near the coast.
The remotest living thing recorded for the Antarctic is probably the Lichen obtained
by Captain Scott at an altitude of 5000 feet in the Western Mountains. This is a
remarkable occurrence. The summer climate there is like the winter polar climate on
the coast. The occurrence of Penguins 80 miles from the sea is an altogether different
case, as they could not live there. Like the men who saw them they were provisioned
and clothed for a journey to a distance from their base. So hardy are the Lichens
that one wonders whether there may not be some on the bare rocks of Mount Hope or
the Cloudmaker. If one had time on such journeys for such things, and if a keen-
eyed lichenologist were of the party, it is not unlikely that he would detect some
stains which no one else would recognise for Lichens.
102 J. MURRAY
I
Tardigrada we would not expect far inland. As far as moss could exist water-
bears might be expected. At present none is known south of latitude 78°.
So few are the Tardigrada of Victoria Land that it is hardly worth while entering
into any discussion of general questions concerning their origin and relationships.
Taken in conjunction with the Tardigrada of other parts of the Antarctic, altogether
making the modest total of sixteen identified species, it may be better worth while to
discuss such questions.
The whole circuit of the Antarctic Coast has only been touched (for purposes of
biological study) at three points in its many thousands of miles : at Graham Land, at
the Gaussberg, and at South Victoria Land. The Graham Land region, including the
Islands, lies entirely outside the Antarctic Circle ; the Gaussberg is almost on the
Circle ; Victoria Land is far to the south. The region round Graham Land is best
known. Some half-dozen expeditions have visited it, but only the Scottish and
Swedish have published reports on the Tardigrada.
It is hoped that Dr. Charcot's recent expedition will afford material for further
study of the Tardigrada of this region.
There is undoubtedly a fairly rich Tardigrade fauna in the region lying south of
Cape Horn. The other two points of the continent examined are very poor in
Tardigrada. In Victoria Land, Ross Island is reckoned as part of the continent.
Though technically an island, permanent ice or snow joins it with the mainland.
In the following table a list is given of all known Antarctic species, with their distri-
bution in the three Antarctic localities, and their general distribution over the world.
The list includes sixteen species, but many others are known to exist, at least in
the islands. There are three species of Echiniscus, one of Milnesium, nine ofMacroliotus,
and three of Diphascon.
Seven of these species are known to be widely distributed over the world : they are,
E. arctomys (?), M. tardigradum, M. echinogenitus, M. oberhausen, D. chilenense,
D. alpinum, and D. scoticum.
Three species are not yet known outside the Antarctic, but the adjacent lands are
too little known to permit us to build anything on the restriction. The species are
E. meridionalis, M. meridionalis, and M. poZorts.
Two species are only known in the southern hemisphere. M. furciger is common
in the Antarctic region south of Cape Horn, and in the adjacent sub- Antarctic localities
of Tierra del Fuego and South Georgia. On the other side of the world it occurs in
New Zealand, where it is found in both the principal islands, and in the North Island
attains to a sub-tropical latitude. M. asper is only found in the Antarctic to the
south of Cape Horn and in the neighbouring South Georgia.
Three species appear to have a bi-polar distribution. E. wendti is in both polar
regions, and is also recorded for Germany and doubtfully for Scotland.* M. arcticus
is in both polar regions, and is also doubtfully recorded for New Zealand, Australia,
* It has now been found (September 1910) on the summit of Snowdon, in Wales.
ANTARCTIC TARDIGRABA 108
Africa, Scotland, and Canada. M. antarcticus is in the South Polar region, and though
not yet known in the Arctic, it comes pretty near to it in Sweden, where it has
recently beeii found by Carlzon (1). The other six bi-polar species are cosmopolitan,
or very widely distributed.
DISTRIBUTION : LOCAL AND GENERAL,, OF ALL KNOWN
ANTARCTIC TARDIGRADA.
Antarctic
Sub-Antarctic
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K. wendti .
—
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X
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E. meridionalis .
—
—
X
Milnesiwn tardigradum
—
—
X
—
X
X
X
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X
X
X
X
X
X
Macrobiotus furciyer
—
—
X
X
X
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X
—
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3[. echinoyenitits
—
—
X
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X
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X
X
X
^f. meridionalis .
X
M. polaris .
X
M. oberhciiiseri .
X
—
—
X
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X
X
X
X
X
X
—
X
X
M. arcticws
X
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— .
—
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J/. asper .
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X
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M. iiuirrayi
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M. aitlarcticiis .
—
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.
DipJutscon cluleiiense
—
—
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1). alpinum
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D. scoticum
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X
The local distribution within the Antarctic is very remarkable. In the three
portions of the Antarctic from which Tardigrada are known the species are all
different in each locality, with the single exception of D. alpinum, which occurs in
both Victoria Land and the Graham Land region. From the uniformity of the
conditions round the Antarctic coast it would be expected that the fauna would
be uniformly distributed. It is true that there is a great difference in latitude,
amounting to not far short of 1000 miles, between the regions studied in East and in
West Antarctica. The climatic conditions do not correspond to this great difference
of latitude, though the Graham Land region is somewhat warmer.
It might be suggested that the Tardigrada have peopled the different districts
independently by migration from the nearest lands to the north. A glance at the
map will show how improbable this is. Migration across those broad oceans must be
excessively difficult for Tardigrada, even considering the few small islands which
afford intermediate resting-places. Migration of such animals is involuntary, and
104 J. MURRAY
can scarcely be effected by any other agency than the winds. The prevalent storm
winds on the Antarctic coast are southerly. The Tardigrada of the different regions
do not correspond more closely with the nearest northern lands than with other
regions, except to some small extent to the south of Cape Horn, where access to the
Antarctic from the north is easiest. In Victoria Land all the species of Tardigrada
yet found are unknown in New Zealand, with the one exception of D. alpinum, and
M. furciger, an Antarctic species found in New Zealand, is absent from Victoria
Land.
Undoubtedly migration along the Antarctic coast is easier than migration to it
from the north. In that case the birds, such as the Skua gulls, could assist in the
process. If the Antarctic has been peopled from the north, the Cape Horn to
Graham Land route is the likeliest to have been followed, and the numerous islands
would make it easier.
There is little evidence that any species have been developed through long
isolation in the Antarctic. M. furciger and M. asper might have originated thus
and spread to the north, but it is quite as likely to have been the other way round.
There are, of course, the three species peculiar to the Antarctic, but they may any
day be found on the other side of the world, as happened with M. antarcticus.
Contrasting the Antarctic Tardigrada with the Rotifera of the same region, we
arrive at the following considerations. In both groups there are about the same
number of Antarctic species known. In the Tardigrada the few peculiar species are
not very peculiar, and their modifications do not appear to render them better fitted
for the conditions than other species. None of them is strikingly abundant in the
region. If there is a dominant species in any region it is M. arcticus. The peculiar
species of Bdelloid Rotifera are more peculiar, and their peculiarities seem more
likely to have arisen through long isolation, in adaptation to the conditions, and the
extraordinary abundance of some of them (e.g., Philodina gregaria and Adineta
grandis] bear this out.
The different life-conditions of these two groups must, however, be borne in mind.
The Bdelloids reproduce only parthenogenetically, so that there is no check by cross-
breeding to variation. The Tardigrada are sexual, so that if fresh migrants arrived,
even at long intervals, the development of new forms would be retarded.
One of the most curious features in the Antarctic Tardigrade fauna is the absence
of the genus Echiniscus from the Antarctic Continent. No Echiniscus is yet known
on the continent, or anywhere south of the Antarctic Circle. In the North Polar
region there are at least a dozen species found within the Arctic Circle, and almost as
many even in Spitsbergen and Franz- Josef Land, in latitude 80° or higher. Is it
that the Ecliinisci are less resistant to cold than the Macrobioti 1 In the Antarctic
Islands, the South Orkneys and South Shetlands, &c., there are many species of
JEchiniscus.
The relation of the Antarctic Tardigrade fauna to that of other regions of the
ANTARCTIC TARDIGRADA 105
earth is indicated in the Table by the following figures. Of the 16 species, 10 occur
also in Europe, 9 in the Arctic, 8 in Australasia, 8 also in the sub-Antarctic islands,
7 in North America, 5 in Asia, Africa, and Oceanic Islands, and 4 in South America
(excluding Tierra del Fuego, which is taken into the sub- Antarctic region).
These figures are of little value, on account of the paucity of our knowledge. As
they stand, they indicate the closest affinity with Europe, and after that with the
Arctic, the sub-Antarctic islands, and Australasia, -
The Antarctic species which extend into Europe and the Arctic are identical in
these two regions, with the addition of M. antarcticus to the European list.
Of the eight species in the sub- Antarctic region, and the eight in Australasia,
there are five which are common to the two regions.
BEIT. ANTARCT. EXPED. 1907-9. VOL. I.
TARDIGRADA OF NEW ZEALAND
Collecting in New Zealand. — Tardigrada were collected in a great number of
localities — fairly representative of all the varied climates of the country — in both of
the large islands : in the moist sub-tropical bush of the extreme North, beyond
Auckland — in the volcanic district of Rotarua — in the cold lake district of the South
Island — and among the Alpine ranges of Mount Cook.
New Zealand stretches from nearly 34° S. latitude to nearly 47° S., and Stewart
Island passes the 47th parallel by a few miles. The districts visited stretched
between 36° and 47° S.
On the arrival of the Nimrod in the end of 1907 there was one month available
for work in New Zealand. After a day or two spent in the neighbourhood of
Christchurch a party set out for the Hermitage at Mount Cook. Dr. Mackay was
bent on mountain-climbing : Dr. Michell assisted me in collecting.
From the railway terminus at Fairlie two days were occupied in getting to the
Hermitage. The first day's journey was made on foot, ending at a guest-house
on the shore of Lake Tekapo, whose green, milky water made a strange outlandish-
looking landscape. The chalky water was examined for pelagic animals, which were
present but not abundant.
Next day we went by motor-coach to the Hermitage. The hostel stands at an
elevation of more than 2000 feet above sea-level. The valley bottoms of the Tasman
and Hooker Rivers had the aspect of Scottish Moorland, and abounded in similar
mosses, peat-mosses and others, albeit the flowering plants were very different. No
Scotch thistle is so formidable a foe as a New Zealand " Spaniard."
The home-like character of this moorland was only disturbed by the wide
desolate beds of the glacial rivers, miles across at places, a vast stony waste,
intersected, now when the rivers were low, by innumerable branching channels.
The moorlands of Mount Cook were not nearly so prolific in forms of microscopic
life as similar places in Scotland would be, although this district yielded the greater
part of our New Zealand species. The Sphagnum was particularly unproductive.
This may be due to the fact that the mosses have to endure much greater heat, and
are dried up for longer periods.
While I collected in the valley and at moderate elevations, my companions
climbed some of the peaks in the neighbourhood, and they were always mindful to
107
108 J. MURRAY
bring mosses from the highest points at which they grew. Together they climbed
Mount Wakefield, over 6000 feet high, and got moss near the top. Dr. Michell
ascended to a similar height on the Sealy Range.
Dr. Mackay made the first ascent of the hitherto maiden peak of the Nun's Veil,
just under 9000 feet in height. From that trip we got the most interesting biological
results of our whole work in New Zealand. Dr. Mackay got some moss at about
6000 feet, and some lichen near the top. The moss was a little scrap of a
Dicranum, and I am told that it is a previously unknown species. In the axils of its
leaves there were several species of water-bears, two of which are new to science
and are the only peculiar species obtained in New Zealand.
Lake Wakatipu was next visited. The microfauna was much the same as in the
Mount Cook district. The water of this lake was not milky, but beautifully clear
and cold. In the plankton of the lake a species of the rotifer Pedalion was found.
A journey was made by coach through the beautiful Otira Gorge to the west
coast. As soon as we crossed the pass and got on to the western slope we found
ourselves in a different world. The whole country was densely wooded, the tree-ferns
giving a tropical and unfamiliar look to the bush. Everything was festooned with
luxuriant moss. Incidentally, the torrential rains which promote the luxuriance of
the vegetation poured upon us all the time that we stayed on the coast.
There was little opportunity for studying the moss fauna on the spot, though a
light microscope was carried with us. The fauna was poor, especially in Tarcligrada,
but .one or two rotifers were got which are peculiar to New Zealand.
The available time being short, we set out on a coasting steamer for Wellington.
At every port we visited, from Hokotika to Nelson, we had a few hours ashore, and
spent them in the bush, picking mosses from the trees. No attempt could now be
made to examine them, and they were sent off from time to time by mail for future
study.
Foggy weather delayed us, and when we reached the North Island our time was
almost spent. In the few days remaining it was desired to get as far north as
possible, so as to include in our study of the fauna as great a range in latitude as
might be. We went to Auckland, and on the advice of Mr. Cheeseman, of the
Auckland Museum, proceeded to the Waitakerei Range, a short distance to the
north-west of the town. Here we found the bush of similar character to that of the
South Island at Hokotika, but if anything moister and still more abounding in ferns.
It was a happy hunting-ground for the student of mosses and moss faunas,
but we were not permitted to linger hunting there. In our limited time we.
collected half a dozen kinds of water-bears, which had all been previously obtained
in the South Island.
When the Nimrod came back out of the south the first green-clothed land we
came to was Stewart Island. We had a whole day ashore there. The bush in that
uninhabited region was almost impenetrable, and after spending some hours and
TARDIGRADA: NEW ZEALAND 109
getting only a few hundred yards through it, without encountering any tnoss worth
collecting, I came out on the shore again. There was a little islet with precipitous
sides, cut off from the main island at high water. On the summit of the islet there
was only some scrubby bush and deep cushions of some large mosses. These were
not of sorts usually favourable to microscopic life (they were large Dicrana, and
Hylocomium and the like), and we did find them very unproductive, but among other
things we got five kinds of Tardigrada.
No collecting was done in the South Island during the second visit, in 1909, but
a week was spent in the volcanic region of Rotorua, in the North Island. The
country had the look of being habitually parched and dusty, and was not mossy,
though there were mosses enough in shady nooks by springs and waterfalls. Hot
springs and cold springs were examined, the lakes were netted, and the mosses were
washed, but no water- bear of any genus appeared to reward our exertions.
Two members of the Expedition, Mawson and Mackintosh, fetched me moss from
a considerable elevation on the slopes of the volcano Ngauruhoe, but this also was
unproductive.
Summary of previous knoivledge. — Scarcely anything is known of the Tardigrada
of New Zealand. Captain Hutton's Index, 1904 (10), makes no mention of the
group. The only paper I know on the subject is Professor Ric liters' " Moosfauna
Australiens " (37), in which three species are noted from the North Island, and from
Bare Island, a little islet near Napier, on the east coast. They are Echiniscus
gladiator, E. novaizeelandice and Macrobiotus hufelandii.
We found all these species again, but curiously enough we did not get the type of
E. gladiator, but only the spineless variety exarmatus, hitherto unknown outside of
Scotland.
In February 1907 Mr. D. J. Scourfield gave me some moss which he had received
from Gisborne, N.Z. In this I found a single example of Macrobiotus nodosus, a
species discovered shortly before in Africa. The species did not occur in our
collections from New Zealand, but it was obtained by Captain Davis, when the
Nimrod called at the Macquarie Islands on the homeward voyage.
LIST OF SPECIES FOUND
Hchinismts muiabilis, Murray. M. dispar, Murray.
E. novcezeelandice, Richters. M. arcticus, Murray ?
E. gladiator, Murray. M. nodosus, Murray.
E. velamimis, sp. n. M. sattleri, Bichters.
Milnesium tardigradum, Doyere M. papillifer, Murray.
Macrobiotus hufelandii, Schultze. M. ornatus, Richters.
M.furciger, Murray. M. annulalus, Murray.
M. echinogenitus, liichters. Dipbascon chilenense, Plate.
M. harmsworthi, Murray. D, alpinum, Murray.
M. montamis, sp. n. D. scoticum, Murray.
M intermedius, Plate.
Four Species not identified (3 Echiniscus, 1 Mctcrobiolus).
110 J. MURRAY
NOTES ON THE SPECIES
Genus — Echiniscus, Schultze
A : SPECIES HAVING SEGMENTS V. AND VI. DISTINCT
Echiniscus mutabilis, Murray
The commonest Echiniscus in New Zealand, occurring in many localities in both
of the large islands, but not in Stewart Island, the Aucklands, or the Macquaries.
I was formerly inclined, relying on second-hand descriptions of E. arctomys,
Ehr., to believe that E. mutabilis was closely related to that species. Reference to
Ehrenberg's original figure (4) and description (3) shows that they have nothing in
common. They even belong to different sections of the genus, which will probably
become distinct genera eventually. E. arctomys has segments V. and VI. (Richters)
completely fused, and E. mutabilis has these segments distinct. Any records which
I have made of E. arctomys in previous papers are erroneous — they all refer to
E. mutabilis (see p. 126, footnote).
Echiniscus novwzeelandice, Richters (37) (Plate XV. Fig. 5)
Specific characters. — Segments V. and VI. separate, V. a pair, obscurely divided,
bearing two stout dorsal processes somewhat near the median line : first and second
median plates divided by transverse line : all finely dotted : no fringe on the
fourth legs.
Description. — Professor Richters contents himself with a rather brief diagnosis, so
a fuller one is given here. The animal varies considerably, in size and other respects.
It is the closest relative of E. mutabilis, and it is doubtful if they could be
distinguished, except by the characteristic spines on V. The colour varies from
brick-red to yellow. The number of plates is difficult to state, on account of the
obscure separation of some of them, and the sub-division of others. I prefer to
regard those sub-divisions as not constituting distinct plates, and thus consider a
typical animal in which V. and VI. are distinct as having twelve plates, of which
three are median, and there are three pairs. There are species in which the plates
of the third pair (segment V.) are completely fused.
Apart from the ten processes on the head, which appear to be common to all
Echinisci* there are usually no processes on the body of E. nonezeelandice except
the two stout spines on V. These vary greatly in size, sometimes reaching almost
as far as the posterior border of plate VI., sometimes reduced to short cones. In
one example they measured 30 n in length.
These processes are often forked. There are often short cones at the postero-
lateral margins of the plates of the second pair (d Richters). There may be rudi-
* Except only E. ImberUs, which is said by Prof, Richters (38) to lack the six processes near the mouth
TARDIGRADA: NEW ZEALAND 111
ments of similar cones at the angles of the preceding segments (b and c Richters).
There is a slender spine on the first leg, and a blunt palp near the base of the fourth,
as in many other species. Seta a at the base of the head is generally nearly straight
— the auricle at its base is fairly large. The legs are long. No barbs were detected
on the inner claws of the fourth leg, but they would be readily overlooked if as small
as in E. mutabilis.
It is usually a small species, but attains to a length of 280 // or more, exclusive of
the fourth legs. The eggs are two to four in number. Two have been seen in a skin
which measured only 100 M> so that the species evidently, like E. mutabilis, grows
after attaining to maturity. The eggs measured 50 ^ in length.
The species was described by Richters (37) in the same year (1907) in which we
collected it. It was only known in New Zealand till 1909, when I obtained it in
Australia, and later a variety in Hawaii. This ^variety (see p. 151 and Plate XIX.
Fig. 35) completely links E. novcezeelandice with E. mutabilis, as the dorsal
processes on V. are reduced to mere angles on the posterior margins of the plates.
The figure (Plate XV. Fig. 5) shows the forked process and little lateral cones (d),
although these peculiarities were not observed in New Zealand. The figure is from
an Australian specimen.
Habitat. — North Island, Waitakerei Range, north-west of Auckland ; South
Island, Mount Cook district, elevation 2000 to 3000 feet ; shore of Lake Wakatipu
at Kingston.
There are only about a dozen species in that section of the genus which has
segments V. and VI. separate. It is only necessary to compare E. novcezeelandice
with those which have processes on the posterior border of V. (third pair). Of these
there are three : E. islandicus and E. borealis have numerous spines and seise,
E. imberbis has a long seta at d, and a dorsal spicule over c.
A species from the Canary Islands, figured but not named by Heinis (8) has the
spines of V. precisely like those of E. novcezeelandice, but it has setse c, d, and e.
•B: SPECIES HAVING SEGMENTS V. AND VI. COALESCED
Echiniscus gladiator, Murray (12)
We did not find the type of the species, but the variety exarmatus, Murray (18), was
got in moss brought by Drs. Mackay and Michell from an elevation of about 5000 feet
on Mount Wakefield, in the Mount Cook district of the South Island. This spineless
variety was previously known only in the Shetland Islands.
Professor Richters got the type from Bare Island, close to the coast of the North
Island.
There is one character, by which E. gladiator and its variety exarmatus may be
distinguished from all other species, which has not been referred to in the descriptions
J. MURRAY
or shown in the figures. The paired plates meet in the middle line only close to the
anterior border, and gape behind. A pair of plates, showing the gap, is figured
among Canadian Tardigrada below (Plate XX. Fig. 51).
Echiniscus velaminis, sp. n. (Plate XV. Fig. 6)
Specific characters. — Size moderate ; plates nine, two pairs, two median, surface
with fine irregular pits, very large on the last (lumbar) plate ; c, d, and e are setae,
d shorter, e very long ; dorsal setse over c and d ; fringe of few large blunt processes ;
claws all without barbs.
General description. — Length 270 ^ seta a 75 n, c 125 /x, d 80 M, e 250 /* ; setee d
has sometimes a short curved spine at its base. None of the plates has the surface
markings interrupted by lines or bands. The markings are fine on all the plates
except the last, where they are very coarse. They are very unequal in size, and look
like perforations. The teeth of the fringe are very few in number (5 or 6), and
are large and very obtuse. There is a blunt palp at the base of the fourth leg.
Habitat. — Among mosses from the Nun's Veil Mountain, Mount Cook district,
elevation about 6000 feet ; collected by Dr. Mackay.
Remarks. — E. velaminis belongs to that group of species which have segments V.
and VI. (Kichters) united, and which possess one or more long lateral setae in
addition to the one (a) at the base of the head, which is present in all known species.
Though there are no species in that group so near as to require careful discrimination,
it will be necessary to compare it with a considerable number of species, many of
which are very insufficiently described. Several species have the same number of
lateral setae, but they are differently arranged, and there are other distinctions.
E. testudo has the setae a, b, c, and e, the dorsal setae over c is lacking, and the
" fringe " has many smaller teeth. E. blumi has'setae a, b, c, and d, and the outer
claws are barbed. E. crassus has setae a, b, c, and d, e is a blunt process, the fringe
has small blunt teeth, and the surface is coarsely granular. E.filamentosus has setre
a, b, c, and e (as in testudo), and on each side, according to Plate, two dorsal spines
over the second and one over the third leg. E. muscicola has setae also, a, b, c, and e,
and there is a long dorsal seta over c, but none over d.
Several species have one lateral seta more than velaminis, besides other differences.
E. creplini has small spines near the bases of setae b, c, and d ; E. quadrispinosus and
E. scrofa have some of the plates sub-divided, and the fringe with many sharp teeth.
A number of species have one seta less than velaminis, and differ in other respects.
E. bellermanni has small spines at the postero-lateral angles of the segments ;
E. merokensis has the outer claws barbed ; E. longispinosus lacks seta e, and has the
fringe of slender spines ; E. meridionalis has d a spine, and a dorsal spine over it ;
E. granulatus has the dots real granules.
The forked or double seta of velaminis is reminiscent of E. aculeata (23), but
TARDIGRADA: NEW ZEALAND 113
according to Plate the forked process should be c, there are no lateral setae except a,
and there is no mention of fringe. I do not consider forked processes as of any specific
importance. They occur as " sports " in various species, and very often only the
process of one side is forked. The lateral processes, c and d, are those most commonly
forked.
Echiniscus, sp. ? (Plate 1TV. Fig. 7)
Size moderate, 220 ^ in length ; plates nine, V. and VI. joined, two pairs, two
median. Lateral processes, a, c, d, e — a a seta 120 M long, c a seta of 90 M, d and e
short curved spines of 20 n. Dorsal processes, over c a seta of 90 n.
Lack of information about surface texture, fringe and claws, makes identification
impossible.
Habitat. — Among the moss Thuidium at the Hermitage, Mount Cook.
No affinity with any known species can be suggested. There is no species which
has c a long seta, and d and e short spines.
Echiniscus, sp. ? (Plate XV. Fig. 8)
Very small, length 130 M ; plates nine, V. and VI. joined, two pairs, two median.
Lateral processes, a, c, d, e — a a curved seta, 50 p. long, with a small blunt "auricle"
at its base, c and d short curved spines of 10 and 15 M, e a curved seta of 70 M. The
fringe of the fourth legs has short blunt processes. The claws were not seen.
The surfase markings were small and regular, but could not be definitely stated
to be either pits or papillae. The posterior borders of the second paired plates and of
plate VI. between the slits, showed an undulation like that of E. perarmatus (20) so
that in all likehood the dots are papillae.
This little animal has a very distinctive form. The paired plates have an
anterior narrow portion forming a distinct roll, which shows on the outline, and the
posterior edge diverges widely laterally. No other species besides perarmatus
shows the dots on the edge of some of the plates.
It is very probably a distinct species, but the small size indicates that it may be
a larva, and as the claws were not seen the study cannot be completed.
Habitat. — Among New Zealand moss, note of the locality lost.
There is no close relationship to any known species. In this section of the genus
there is none which has a and e long setae, and c and d short spines.
Echiniscus, sp. ? (Plate XV. Fig. 9)
Small, length ISO/*; plates nine, V. and VI. united, two pairs, two median.
Lateral processes, a, c, d, andV- « a curved seta of 70 M, c a long slender spine of
50 M with bulbous base, d a short curved spine of 25 /&, e a seta of 65 /*• Dorsal
BRIT. ANTARCT. EXPED. 1907-9. VOL. I. Q
J. MURRAY
processes — over c a broad -based curved spine of 35 /j. — over d triangular process
12 /a. long and 6 M across the base.
There are no notes as to surface markings, fringe, or claws, and so the species
cannot be identified.
Habitat. — Among moss from the hills at Lyttelton, South Island.
The animal has a certain resemblance to E. oihonncB. The relative proportions
of the processes a, c, d, and e are similar, and the dorsal processes correspond. It
differs in lacking process b and the small spicules at the angles of the plates.
Milnesium tardigradum, Doyere (2)
Found in several localities in the three principal Islands (North, South and
Stewart Island). The short claws of several examples which were studied had three
points each, except those of the first legs, which had two points each. The cyst was
found in Mount Cook district. It was of oblong form, with rounded ends, and
measured 230 ^ in length. The gullet and pharynx were present, but no claws or
other recognisable organs.
One specimen from the Lyttelton Hills had the thick claws of the first legs
unbranched, those of the last legs three-pointed, and of the second and third legs
some with two and some with three points.
Ascends to an altitude of 5000 feet on Mount Wakefield, Mount Cook district.
Genus — Macrobiotus, Schultze (42)
A: SPECIES WHICH LAY ROUGH EGGS, FBEE
Macrobiotus hufelandii, Schultze (42)
The commonest species in New Zealand, as in most other countries. It occurred
in all three islands.
Wherever I have recorded M. hufelandii it is on the authority of Professor
Richters, who has established Schultze's name for the water-bear which is commonest
in Europe, and apparently almost everywhere else.
Schultze's own description is insufficient to allow of the identification of the
animal with any certainty.
The characteristic egg has been regarded as the best character for identifying
Richters' hufelandii, but it is now known that other species have similar eggs
(M. hufelandioides, see p. 138).
Macrobiotus furciger, Murray (15)
Habitat. — Near Auckland, North Island; shores of Lake Wakatipu, South
Island. Identified by the eggs.
TARDIGRADA: NEW ZEALAND 115
This is one of the most interesting species found in New Zealand. Its distribution
is peculiar. So far as our knowledge goes it is solely a southern species. It is found
in a number of islands in the Cape Horn region, from Tierra del Fuego to the
Antarctic Continent at Graham Land (Richters, 38). It was discovered by the
Scottish Expedition in the South Orkneys. New Zealand is the lowest latitude
recorded for it. Its known range is from about 36'30° S. (near Auckland, N.Z.)
almost to the Antarctic Circle.
It has been considered as a southern representative of M. hufelandii. In most
of the localities it is not associated with that species — it takes its place — only in
New Zealand and Tierra del Fuego have the two species been found together. A
peculiarity in its distribution is its absence from the Victoria Land region of the
Antarctic, though it is so abundant on the other side of the continent.
Macrobiotus echinogenitus, Richters (27)
In the South Island only. As we learn more about Tardigrada it appears that
there are several species which have stellate eggs, more or less resembling those of
M. echinogenitus, some of them probably not to be distinguished from them unless
they contain well grown embryos. The separation of some of these (for example,
M. areolatus and M. harmsivorthi) makes it easier to understand M. echinogenitus
itself, which formerly seemed to vary to such an extent that it was difficult to get a
clear conception of it. M. areolatus, and some related species not yet described,
have eggs the surface of which is reticulate. The shells are double-skinned and the
space between is divided by septa into hexagonal chambers. From these hexagons
the processes spring, at such distance apart that they are always separated by the
width of one cell.
M. harmsivorthi has the egg processes close together, and the claws united as
in M. hufelandii (Richters). The true M. echinogenitus has the egg-shell without
reticulation, the conical processes almost or quite close together, and the claws
joined at the bases only. Further study is necessary to clear up the whole group
satisfactorily (see p. 89, footnote).
Macrobiotus harmsworthi, Murray (19)
South Island and Stewart Island. The egg cannot be distinguished from some
forms of those of M. echinogenitus. In examples from the Mount Cook district the
identity was established by finding eggs in the body, which, when freed from the
enclosing membrane, showed the dose-set acuminate processes.
116 J. MURRAY
Macrobiotus montanus, sp. n. (Plate XV. Figs. lOa-lCkZ)
Specific characters. — Large, brown ; gullet wide ; pharynx with three short rods
and a " comma " ; claws of hufelandi type, united for half their length, those of each
pair equal and placed side by side, but only one with a strong supplementary point ;
egg spherical, covered with hemispherical processes, which almost meet at their bases.
Detailed description. — Length about 500 M and upwards : diameter of egg, over
the processes, 75 to 80 M. Old examples are deeply coloured with a dull brown
pigment similar to that of M. hufelandii. The claws are thick and strong, and are
nearly equal : in most species of the hufelandi group they are more or less unequal,
often markedly so. The gullet is wide, and the teeth strong and curved ; the end of
the gullet in the pharynx expands into a prominent flange, beyond which the
apophyses are fixed. The three thickenings in the pharynx are just about equal
in size ; they are twice as long as broad, and are rounded at the ends ; the " comma "
is rather an obscure one.
Habitat. — Among moss gathered by Dr. Mackay on the Nun's Veil Mountain,
in the Mount Cook district, South Island, at an elevation of about GOOO feet, on the
occasion when that peak was ascended for the first time, December 1907.
Remarks. — M. montanus is one of a large group of species, closely related to
M. hufelandii. All of them possess wide gullets and strong teeth. The pharynx
has either three distinct rods, or the two next the gullet are united, usually showing
traces of their component rods. The claws are united for a considerable distance,
usually half-way or more, and one of each pair has one or two supplementary points.
The eggs are spherical and are ornamented with processes of various form.
The majority of the species have eggs very different from those of M, montanus.
It is only necessary to compare it critically with one or two species which approach it
very closely. Those are M. meridionalis, Richters (40), and M. polaris, Murray (see
the preceding section on Antarctic Tardigrada in this paper, p. 98).
According to Professor Richters, M. meridionalis has the processes on the egg
nipple-shaped, which he explains in a letter as being a hemispherical base surmounted
by a narrower portion. This form of egg I know from the Scottish mountains, though
we did not find it in the Antarctic.
The egg of M. polaris differs markedly from both. The surface of the shell is
reticulate, like that of M. areolatus (see Canadian Tardigrada, p. 167 of this paper).
The double shell encloses a series of compartments, showing as polygons at the
surface. From certain polygons arise the processes, at such distance apart that they
are separated always by one polygon. The processes vary from round to acuminate.
The polygons are relatively much smaller than in M. areolatus. M. polaris has
moreover, much more slender and very unequal claws, and the gullet is relatively
narrower. M. montanus was found to be alive after the moss containing it had been
TARDIGRADA: NEW ZEALAND 117
dry for two and a half years. Only a small proportion of the individuals revived when
the moss was moistened. The relation of the egg to the animal was demonstrated by
the presence in some of them of young in which the pharynx and claws could be seen.
Eggs of Macrobiotus having close-set processes, nearly or quite hemispherical,
have been figured by Richters and myself. In his " Eier der Tardigraden " (32),
Plate V. Fig. 4, Richters figures one such as apparently a variety of M. echinogenitus.
In "Scottish Alpine Tardigrada" (14), Plate IH. -Fig. 10, I show an egg which in
all probability is that of M. nwntanus. A similar egg occurred in Nova Zembla,
and is figured in "Arctic Tardigrada" (19), Plate XLV. Fig. 4.
According to Doyere (2), confirmed recently by Richters in a letter, M. ober-
hiiuseri has an egg of similar form, with rounded processes.
Macrobiotus intermedius, Plate (23)
In many localities in the South Island ; Hills at Lyttelton, Otira Gorge, near Lake
Wakatipu ; Mount Cook district, at an altitude of 5000 feet on Mount Wakefield ;
near Auckland, North Island.
The eggs were found near Lyttelton. They had not the typical top-shaped
processes, expanded upwards. They were narrower at the top, like truncate cones,
of a height equal to the breadth of base, and with a very slight indication of
expansion at the top. The processes were separated by spaces rather greater in
diameter than their bases. The exposed surface of the shell between the processes
was finely papillose.
Professor Richters informs me in a letter that he knows a variety of M. inter-
medius which has processes very like those of M. hufelandii. This variety I have
also seen (see Australian Tardigrada, p. 139, in this paper). The variety from
Lyttelton is like it, but the processes are shorter, and not expanded into a disc
above.
Macrobiotus dispar, Murray (16)
Habitat. — Many localities in the South Island ; a pond on the Moraine of the
Miiller Glacier ; on the Vegetable Sheep (Haastia), Mount Wakefield, elevation
about 5000 feet; on the Nun's Veil peak, elevation 6000 feet (collected by
Dr. Mackay) ; among tree-moss from the moist bush of the west coast. The simplex
condition also occurred.
It is curious that, though usually dwelling in ponds, the species occurred only
once in a pond in New Zealand. In all other instances it was got among moss, or
moss-like vegetation (Haastia).
The examples from the Nun's Veil had the two dorsal processes of the type,
which are often lacking.
118 J. MURRAY
B : SPECIES WHICH LAY THICK-SHELLED EGOS, WITH EMBEDDED RODS
Macrobiotus an- liens, Murray ? (19)
Habitat. — Mount Cook district, South Island.
In the absence of the egg there is some doubt about the identification. The
claws are of the Diphascon type. Apart from M. oberhduseri, which is easily
distinguished by the colour and the pharynx, there are very few species having claws
of this type, and M. arcticm is the commonest of them.
C : SPECIES WHICH LAY SMOOTH EGGS IN THE CAST SKIN
Macrobiotus nodosus, Murray (20)
Habitat. — Near Gisborne, North Island ; Macquarie Islands.
The species was collected by the expedition in the Macquaries, but it had been
previously found in moss from Gisborne, given to me by Mr. D. J. Scourfield. Its
distribution is peculiar. It is known in Africa, New Zealand, the Macquaries,
and Fiji.
Macrobiotus sattleri, Bichters (26)
Habitat. — Hills beside Lake Wakatipu, South Island.
Macrobiotus papillifer, Murray (12)
Habitat. — On Myriophyllam in a torrent fed by melting snow, Black Birch
Creek, Mount Cook district, South Island.
Macrobiotus ornatus, Richters (24)
Habitat. — Among bog-mosses, near Birch Hill Creek, and Myriophyllum, Black
Birch Creek, Mount Cook district. In the latter place a cyst was found.
Macrobiotus annulatus, Murray (12)
Habitat. — Birch Hill Creek, and in the bush of the west coast, near Westport,
South Island. At Birch Hill Creek it was abundant, but it was never seen carrying
the eggs in the characteristic way. The examples from Westport had the papillae
very small.
Macrobiotus, sp. ? (Plate XV. Fig. 12)
Macquarie Islands. A large species with claws of the Diphascon type. It could
not be identified because it was in the condition known as simplex (i.e., it had no
TARDIGRADA: NEW ZEALAND 119
teeth and no rods in the pharynx). It is pretty certain that it is distinct from
all known species. The position of the pharyngeal bulb indicates that it is a
Macrobiotus, not a Diphascon. There are only a few species having claws of the
Diphaacon type, and none of them have such strong claws as this animal. They
more resemble those of some of the larger northern species of Diphascon, such as
D. spitzbergense (27). The length was 550 /*.
Diphascon chilenense, Plate ? (23) (Plate XV. Figs. 11«, lib)
This species is noted from the Mount Cook district and from Stewart Island. It
is recorded as doubtfully D. chilenense on account of the characters of the claws,
which appear to be intermediate between the true Diphascon and the hufelandi
types. The two pairs are not very unequal in size, and the claws of each appear to
be united for some distance above the base. As the animal was a very small one,
and the claws relatively very small, it could not be satisfactorily determined whether
the structure of the larger pair was essentially that of the Diphascon type, which is
shown in Fig. 12 on the same plate.
The pharynx (Fig. lla) has four round nuts in each row of thickenings, besides
the apophyses on the end of the gullet. This seems to be one more than in any
other Tardigrade, unless we regard the fourth nut as homologous with the " comma."
Plate figures four nuts, but does not show the apophysis.
The example here figured was obtained by Dr. Mackay from a height of about
6000 feet on the Nun's Veil, a peak nearly 9000 feet high.
Diphascon alpimtm, Murray (14)
South Island only (Otira Gorge). Though discovered in Scotland, the species is
best known as a southern and Antarctic species. It occurs on both sides of the
Antarctic, on the Continent at Graham Land, on the adjacent islands, and also in
Victoria Land.
Diphascon scoticum, Murray (11)
In Stewart Island only. The species is found in both Polar Regions and in
several places between. In the north it attains a high latitude in Spitsbergen and
Franz- Josef Land. In the south Richters has found it in Possession Island (recorded
as D. crozetense), and in the South Shetlands. It is also in Australia, Hawaii, and
Canada.
REMARKS ON THE TARDIGRADE FAUNA OF NEW ZEALAND
Its composition. — For a country of such extent, so isolated, and offering such a
variety in climate and conditions, the Tardigrade fauna appears poor in species, and
120
J. MURRAY
remarkably deficient in peculiar forms. It cannot, of course, be supposed that our
knowledge of the fauna is nearly adequate to permit of definite conclusions on such
points, but it might be expected that in an examination extending over two months,
during which a great variety of localities were visited, we would have detected signs
of a rich fauna or of peculiar forms, if the fauna were rich or peculiar. Further work
may yet bring such forms to light.
Altogether 25 species were observed, of which 21 were identified. These were
4 species of Echiniscus, 13 of Mctcrobiotits, 1 of Milnesium, and 3 of Diphascon.
The unidentified species were 3 of Echiniscus and 1 Macrobiotus. Two species, an
Echiniscus and a Macrobiotus, are described as new species.
About one-half of the species are common and widely distributed. The others
are in varying degrees local, and several have a very peculiar distribution.
The accompanying table of local and general distribution shows the relation of
the Tardigrade fauna to that of other parts of the world.
DISTRIBUTION : LOCAL AND GENERAL
Australasia
New Zealand
05
5
OQ
HI
02
03
t— t
m
Ed
d
3
j
Ed
,
0
0
o
d
H
*""*
E-i
K
JH
w
AH
«jj
o
s
w
o
EC
M
tf
o
Ed
Ed
H
K
EH
I
O
H
0
Si
Ed
<y
u
i
O
*H
1
s
K
3
s
<
»
i
«
^
EC
02
S
«!
i
fe
02
0
Echiniscus mutabilis
X
X
X
X
X
X
X
—
—
X
—
K. novcezedandice
X
X
—
. —
X
X
—
—
—
• — •
—
—
—
E. gladiator .
X
X
—
—
—
—
X
—
—
X
—
—
—
Milnesium tardigradnm
X
X
X
X
X
X
X
X
X
X
X
X
Macrobiotus hufelandii
X
X
X
—
X
X
X
X
X
X
X
X
—
M. furciger
X
X
—
—
—
—
—
—
—
—
X
—
X
M. echinogenilits
X
X
. —
—
X
X
X
X
X
X
X
X
X
M. harmsivorthi
—
X
X
— .
X
X
X
—
X
X
—
X
—
! SUlTtfOCt/TtlfiS ,
—
X
—
—
—
X
X
X
X
X
X
X
M. nodosus
X
X
X
,
Diphascon scoticum
—
—
X
—
X
X
X
—
—
X
X
X
X
Nineteen species were collected in the islands, 3 in Fiji and 16 in the Hawaiian
group (Sandwich Isles), but only 15 were identified. Adding M. samoanus,
* All these records, except the Australian one, are wrong ; the animals recorded as E. arctomys are
either E. iiiutabilis or E. suillus (see p. 126 and footnote).
158 J. MURRAY
discovered by liichters in Samoa, and E. arctomys, recorded by him for Oahu, we
have 17 species known from the Pacific Islands.
The only instance of a species known to be common to two islands is M, hufelandii
in Oahu and Hawaii. This fact is of no importance, as only Oahu is at all well
known.
There are 7 species of Echiniscus (besides 1 not identified) all from Oahu. No
Echiniscus is known in the other islands. There are 8 identified species of Macro-
biotus, 5 from Oahu, 2 from Fiji, and 1 from Samoa, the ubiquitous Milnesium, and
Diphascon scoticum.
There are eight of the species which are almost cosmopolitan in distribution, and
which are therefore of little importance in the study of island faunas. They indicate
that migration to these remote islands is not difficult.
The other nine species are of very restricted range. None of them is known in
more than three of the regions, though only M. samoanus is confined to the islands.
E. novcezeelandiai occurs in Oahu, Australia, and New Zealand ; E. intermedius
in Oahu, Australia, and Canada ; E. viridis in Oahu and Scotland ; E. spinulosus in
Oahu, Europe, and the Arctic ; E. perarmatus in Oahu and Africa ; M. occidentals
in Oahu, Australia, and Canada ; M. crassidens in Oahu, Australia, and Africa ;
M. nodosus in Fiji, New Zealand, and Africa.
The proportion of the island species found in other countries is as follows : Of the
17 species, there are 11 in Australia, 10 in Europe and Africa, 9 in the Arctic, 8
in N. America, 7 in New Zealand and Asia, 5 in S. America, and 4 in the Antarctic.
While the indicated affinity is closest with Australia, it is scarcely less with the
distant Europe and Africa. The presence of so many species in common with such
diverse and distant lands, and the scarcity of peculiar species, show the population of
these islands to be a heterogeneous lot, recruited by casual immigration from all sorts
of places.
Our information is too scanty to allow of even this conclusion being made with
any confidence, as it is quite possible that further work may bring to light a peculiar
insular fauna of water-bears. At present there is no indication of such, except the
solitary M. samoanus.
CANADIAN TARDIGEADA
Collecting in Canada. — The only opportunities which offered for collecting in
Canada were the pauses in a hurried journey across the continent. At Victoria,
British Columbia, where the steamer called for a few hours, plenty of moss was found
on the sea-shore, close by the wharf. This proved afterwards to be the most
productive got in Canada. At Vancouver a whole day was spent, but the time
available was not sufficient to go farther afield than the Stanley Public Park. This
park, in its half-wild state, and everywhere clothed in moss, was very suitable for
the collection of microscopic animals.
Thereafter, right across the continent, there was nothing to be looked for but
chance collecting at any stations where the train stopped for more than a few
minutes. Through the Selkirks and the Rockies, at all the stations where the train
stopped long enough to allow time for meals, I generally managed to get my pockets
filled with moss.
Accident assisted science. Twice on the journey the train was held up for some
time. At Sicamous, floods had destroyed a little bridge, and a heavy engine had got
overturned. While a new loop of railway was being built round the obstruction
there were two days available for collecting. It was a beautiful spot, on the shore
of a lake, and moss was abundant, but somehow it proved very unproductive. We
were afraid to make long excursions up the neighbouring hills, as we did not know
when the train might go on.
A few days later, after crossing the Prairies and just after passing the Lake of
the Woods, there was a train-wreck in front of us, on a high bank between two
lakes. This delayed us for several hours. The microscope was again got out, the
dry moss was washed in the lake, and the sediment examined. This time we were
luckier, and got many interesting things.
Half a day was spent in Ottawa, and much moss was collected in the public parks
and roadsides. This also proved good.
The only records of Canadian Tardigrada with which I am acquainted are in
Richters' " Moosfauna Australiens, &c.," 1908 (37). He records three species from
Vancouver : Echiniscus gladiator, Milnesium tardigradum, and Macrobiotus hufe-
landii. For the whole North American continent I only know one other record,
Packard's insufficiently described Macrobiotus americanus (21). Some half-dozen
species are recorded for South America.
159
160 J. MURRAY
LIST OF SPECIES COLLECTED
Hchiniscus gladiator, Murray. M. oberJtauseri, Doyere
E. sylvanus, sp. n. M, intermedius, Plate.
E. intermedius, Murray. M. articus, Murray?
E. canadensis, sp. n. M. sattleri, Richters.
E. oihonnce, Richters ? M. tuberculatus, Plate.
E. bisetosus, Heinis? M. canadensis, sp. n.
Milnesium tardigradum, Doy6re. M. virgatus, sp. n.
Macrobiotus hufelandii, Schultze. Dipfiascon chilenense, Plate.
M. echinoyenitus, Richters. D. alpinum, Murray.
M. areolatus, sp. n. D. scoticnm, Murray.
M. harmsworthi, Murray. D. canadense, sp. n.
M. occidentalis, sp. n.
Eight species not identified (8 Echiniscus, 5 Macrobiotus).
NOTES ON THE SPECIES
Genus— Echiniscus, Schultze (43)
The eight species collected all belong to that section of the genus in which
segments V. and VI. are completely fused into one plate. It is rare to find a
district possessing so many species without including one in which V. and VI. are
separate. E. nmtabilis is the commonest species in that section, but it is not yet
recorded from either North or South America.
Echiniscus gladiator, Murray (12) (Plate XX. Fig. 51)
Stanley Park, Vancouver, British Columbia. Professor Richters had previously
found it in moss from Vancouver (37).
The British Columbian form differs from the type in that the plates are covered
by wide but very low bosses. The figure, in which these bosses are indicated by
black lines, inevitably exaggerates their prominence, but it shows their size in
relation to the plates.
The paired plates of E. gladiator and its variety exarmatus differ from those of
all other known Echinisci in that they only touch in the middle line for a short
distance near the anterior border, and diverge behind, as shown in Fig. 51. This
character has not been indicated in previous figures.
Echiniscus sylvanus, sp. n. (Plate XX. Fig. 49)
Specific characters. — Size moderate ; colour yellow ; plates nine, two pairs, two
median ; V. and VI. joined ; seta a thick, short ; no other dorsal or lateral processes,
other than those on the head ; dots of moderate size, and seeming to be perforations ;
each plate of the pairs divided in two by lines ; lumbar plate faceted and trefoliate ;
fringe on fourth legs ; inner claws barbed.
Detailed description, — Length 275 /*, exclusive of fourth legs. The palps at the
TARDIGRADA: CANADA 161
mouth are prominent and the cirri short. Seta a measures about 50 M in length. It
is thicker than in any known species except E. cornutus, Richters, but it scarcely
tapers at all, and has a blunt rounded end. The " auricle " at its base is large and
of triangular form. The first median plate is separated from the plate in front of it
by a broad space. The second median is divided into two dotted portions by a
transverse plain band. Each plate of the pairs is divided into two unequal parts by
a line which runs parallel with its anterior border, the narrow part in front forming
a prominent roll which continues round the side and shows in the outline. The
granulation is of the sort which makes the plates seem cribrose. The dots seem to
perforate the plate ; they are unequal in size and are separated by irregular spaces.
The lumbar plate has four facets, the posterior one obscurely divided into two.
The processes of the fringe are few (about eight to ten) ; they are narrow and
acute, and are separated by small spaces. The claws are large, about 25 /u in length.
The inner cla,ws of the fourth legs have small decurved barbs near their bases.
Habitat. — Among dry moss from the woods on the shore of the Lake of the
Woods, Ontario.
E. sylvanus belongs to a group of species of which E. arctomys may be taken as
the type. These have nine or ten plates, segments V. and VI. have completely
coalesced, and there are no processes, dorsal or lateral, on the body, after seta a.
This section of the genus includes ten forms previously known, and three others, in
addition to E. sylvanus, are described in this paper. Two of these forms (macromastix
and exarmatus) were described merely as varieties.
Most of these species differ in conspicuous characters from E. sylvanus. The
thick seta a alone separates it from all of them. Other distinguishing points are
here given briefly. Four species have seta a extremely long (macromastix, wendti,
reticulatus, tessellatm], four species have no fringe (elegans, intermedius, arctomys,
exarmatus), two species have totally different surface markings (spiculifer, bigranu-
latus). There remain three species which bear a closer resemblance to E. sylvanus.
E, viridis and E. macronyx have, like E. sylvanus, very large claws. E. viridis has
the surface dots very large, and seta a very short and fine. E. macronyx has
extremely fine and close dots, no barbs on any claws, and only the fourth legs have
large claws.
E. kerguelensis is nearest to E. sylvanus. It has shorter claws and seta a is
slender. It is also considerably smaller (165 n, according to Richters, but Australian
examples larger, 225 n; sylvanus, 275 /").
Echiniscus intermedius, Murray (Plate XX. Figs. 52a, 526)
Characters of Canadian variety. — Small, hyaline or greyish. Mouth cirri with
large conical bases ; seta a long, no other processes on body. Three median plates,
first and second divided into two equal portions by transverse lines. Plates finely
BRIT. ANTARCT. EXPED. 1907-9. VOL. T. Y
162 J. MURRAY
dotbad with pjlluiid d)ts. Lumbar plate not trefoliate, faceted in three panels, the
lines separating the lateral from the median panels going right up to the third
median plate. No fringe on fourth legs, or barbs on claws. Red eyes. Larva with
two claws, otherwise like the adult.
General description. — Length 175 M, exclusive of the legs, larva 100 M, seta a (of
adult) 60 n. The anterior portion of the head is rounded, and more like a head of
Macrobiotus than Echiniscus, lacking the lateral knobs and the beak usual in the
genus. The base of the head is wider than the front of the next segment, so that
there is a slight neck, rather more pronounced in the larva. The first and second
median plates are each divided into two portions, as is common in that section of the
genus which has V. and VI. separate. The third median often appears to be divided
in exactly the same manner, but the second portion is closely joined to the
lumbar plate. Both parts of the divided median plates are dotted. The plates
of each pair appear to be divided into two by a broad band, which is, however,
dotted.
E. intermedius is a very distinct species. It has V. and VI. united, yet has the
median plates transversely divided, which is only usual in species having V. and VI.
separate. It appears to consistently lack the normal red colour of Echiniscus, though
that is not a safe character. The faceting of the lumbar plate is unusual in that the
angle of junction of the three facets continues to the anterior edge of the plate. The
absence of a trefoil on the lumbar plate is very unusual. I know of no other species
where this character is so distinctly marked. Some have been figured without the
cuts which make the trefoil, but unless the authors emphasise the absence it is likely
that they have been overlooked.
The eggs have only been seen in a form, probably of specific value, from Hawaii
(see p. 151).
Habitat. — Among moss from the sea-shore at Victoria, British Columbia,
June 1909, abundant.
This variety, with fine pellucid dots, is not known anywhere else, but in Queens-
land there occurs the type in which the plates have a broad reticulation instead of
the pellucid dots (see p. 129). Curiously enough, there is a form linking these
extremes in Hawaii. It is reticulate, like the Queensland form, but the reticulation
is smaller, and is formed by the edges of shallow pits (see p. 151).
Echiniscus canadensis, sp. n. (Plate XX. Fig. 47)
Specific characters. — Large, red ; plates nine, two pairs, two median, V. and VI.
fused, dots round, regular, close ; no lateral processes except seta « ; dorsal processes
— a long setse over c, sometimes a shorter seta over d ; fringe of sharp spines on the
fourth leg ; inner claws with decurved, outer with straight, spines near the base.
Detailed description. — Length 300 M and upwards, exclusive of the fourth legs ;
TARDIGRADA: CANADA 163
seta a about 75 n, dorsal seta over c about 100 to 150 M. Body very thick
dorso-ventrally. The dots on the plates are of moderate size. They are circles which
touch at their edges, and which often show a central dot. They look like very flat
granules. The paired plates are each divided by a furrow into two parts. It could
not be seen whether the dots ceased in this furrow. Most of the specimens, even
very large ones, had no processes on the body except the dorsal seta over c ; only
a few had the smaller one over d. The lumbar plate is obscurely trefoliate. The
spines of the fringe are triangular, and are usually separated a little at their bases.
The two-clawed larva has been seen, and from two to four shortly oval eggs in
the cast skin. The barbs of the inner claws are fairly large and high up on the
claw. The outer claws have not been seen with more than one straight spine near
the base.
Habitat. — Among moss growing on the sea-shore, but little above the high-water
mark, Victoria, British Columbia. Very abundant in some pieces of the moss.
It is not necessary to compare E. canadensis with very many species in order to
discriminate it. There are very few species known which have straight barbs on the
outer claws. Among them there are none which are destitute of lateral processes
(other than a). Indeed it is very rare for any species to have dorsal and no lateral
processes.
The Echinisci with straight barbs on the outer claws are four in number —
E. granulatus, E. blumi, E. oihonnce, and E. merokensis. The types of E. granulatus
and E. oihonnce are not described as having outer barbs, but the barbs have been
observed in Scottish examples. All of these species have from two to four lateral
processes (exclusive of seta a), while E. canadensis has none. The dots of E.
granulatus are distinct papillse. E. merokensis and E. blumi have, according to
Richters' figures, much coarser dots. E. oihonnce has four lateral spicules, over b, c,
d, and e, in addition to the larger spines and setae.
Echiniscus oihonnce, Richters ? (27) (Plate XX. Fig. 48)
This is a very doubtful identification, and the Canadian animal is only provisionally
united with oihonnce because we know so little as to the limits to the variation in the
length of the setse and spines. There are points of striking resemblance to oihonnce,
as well as important differences.
This form differs from oihonnce in lacking seta b, and in the great elongation of
process d (which is a spine in oihonnce). It resembles it in having all the correspond-
ing processes except seta b, in having the dorsal process over d a broad triangle, and
in the little spicules near the bases of the lateral seta3.
Description. — Of large size ; length 300 p- exclusive of head and foot. Plates nine,
two pairs, two median. Surface punctate with large dots, which appear to be
perforations, and are separated by spaces wider than themselves. The three lateral
164 J. MURRAY
setae c, d, and e are nearly equal to one another and about 200 p. in length. The
dorsal seta over c is about 120 /x long. The fringe on the fourth leg consists
of short triangular teeth. The claws measure 30 /t in length. The inner ones
have small decurved barbs, which are placed very high, little below the middle of
the claw.
Habitat. — Stanley Park, Vancouver, British Columbia.
In Scottish examples of oihonncc, which are typical in all other respects, the outer
claws of the fourth legs bear straight spines near their bases, although Professor
Richters makes the absence of such spines one of his specific characters. Although
the presence of these spines may be of some positive value, since they are possessed
by very few species, their absence cannot be relied upon as a specific character. In
E. granulatus (or the animal which I identify as that species) the young which have
j>assed the two-clawed stage have no outer barbs, which are acquired at later moults,
and increase in number to three in large examples. The same progressive develop-
ment of the outer barbs occurs in E. blumi, which may also have as many as three of
them on one claw.
The character is thus shown to be a mark of age, but as the great majority of the
species do not have them at any age, when present they make a good confirmatory
character.
The Canadian examples, though of large size, had no barbs on the outer claws, in
this respect conforming to Richters' type. It differs mainly in lacking seta b, and in
the equalisation of the lateral processes c, d, and e.
Echiniscus bisetosus, Heinis ? (7) (Plate XX. Fig. 50)
This is a somewhat doubtful identification, though the animal is certainly very
near Heinis's species. The differences are not very serious, and as our animal was
small, and therefore probably immature, the processes which it lacks might be
acquired later.
Description. — Colour red; length 175 M, exclusive of legs. Plates nine, two
pairs, two median, dots obscure. Lateral process one, the seta a, 50 M in length.
Dorsal processes — over c a seta of 60 M, over d a small curved spine of 8 /*. Fringe
of small triangular teeth on fourth leg. No barbs seen on any claws.
Habitat. — Among moss from the sea-shore at Victoria, British Columbia, one
example only.
According to Heinis, E. bisetosus has also a short spine c, and decurved barbs on
the inner claws. His figure shows a minute lateral spicule d, of which I find no
mention in the text.
It might be suggested that this small animal could be a young example of
E. canadensis (Fig. 47) which was found abundantly at the same place. That
species has, however, at all ages conspicuous barbs.
TARDIGRADA: CANADA 165
Echiniscus, sp. ? (Plate XX. Fig. 44)
Description. — Large, red; length 300 /u, exclusive of legs. Plates ten,V. and VI.
united, two pairs, three median. Lateral processes four — a a seta of 80 p., c a seta of
120 /u, d a seta of 70 n, e a seta of 90 M. Dorsal processes — over c a curved spine
of variable length, 10 to 25 /a and upwai'ds — over d a small spicule. The mouth palps
are large and stout, and the cirri short. The auricle at the base of a is small and
rounded. The fringe on the fourth leg is crenate. The inner claws of the fourth leg
have decurved barbs of moderate size. The dots on the plates are very fine, and
appear to be granules. An example of 210 M in length had claws of nearly 25 M.
Skins with two eggs have been found. The larva with two claws, found
associated with this animal, had only the setae a and c. All the dorsal processes
may be lacking.
Habitat. — Among moss from the sea- shore at Victoria, British Columbia, several
examples, larva, and eggs.
It rarely happens that an Echiniscus with V. and VI. united has three distinct
median plates. Sometimes the skin between IV. and V. is dotted, but there are no
lines marking the boundaries of a plate. In this animal, and another figured on the
same plate (Fig. 46), the third median is distinct.
The lateral setaj are the same in number as in E. testudo, E. muscicola, E.fila-
mentosus, &c., but they are differently arranged. E. velaminis has the same number
of setae, and their positions are the same, but it has the dots of a totally different
character, no barbs on the claws, and much larger processes on the fringe.
Although the finding of eggs shows that the animal is mature, it cannot be
positively identified with any known species, nor yet accepted as distinct. The
variability of the dorsal processes confirms what has been already stated as to the
unreliability of that character. It is a good example of those forms which may be
regarded as distinct species, but which lack sufficiently good characters, other than
the arrangement of the spines and setae.
Echiniscus, sp. ? (Plate XX. Fig. 46)
Desertion. — Small, red ; length 180 M, exclusive of legs. Plates nine, V. and VI.
united, two pairs, three median. Lateral processes four — a a seta of 80 M, 6 a seta of
40 M, c a seta of 100 p., d a seta of 100 /". Dorsal processes — over c a seta of 40 /u,
over d a spine of 15 p.. Mouth palps relatively large, and cirri long. Lumbar plate
trefoliate. Fringe of few large blunt processes. A blunt palp at the base of the
fourth leg. The inner claws of the last legs with small decurved barbs. Dots on
the plates small and uniform, apparently granules.
Habitat. — Stanley Park, Vancouver, British Columbia.
In the number and the surface texture of the plates this animal resembles that
166 J. MURRAY
figured on the same plate (Fig. 44). It has the same number of lateral setae, but
they are a, b, c, and d, instead of a, c, d, e.
It has a close resemblance to E. blumi, Richters (27.) The lateral setae are the
same in position and relative sizes. The differences are — the lack of barbs on the
outer claws, the finer granulation, and the blunt processes of the fringe. The
absence of barbs on the outer claws is of little importance as a negative character,
since it varies with age. There is no indication that our animal is mature. Richters
figures E. blumi with very coarse granulation, and with the fringe of sharp spines.
Many examples had no dorsal processes.
Echiniscus, sp. ? (Plate XX. Fig. 45)
Description. — Size moderate; length 212 /x. Plates nine, V. and VI. united, two
pairs, two median. Dots small, some appearing as perforations, irregular, of two
sorts, a larger dark, a smaller pellucid. Lateral processes four — a a curved seta of 70 /« ;
c, d, and e long, broad spines of 50, 50, and 80 M respectively. Dorsal processes —
over c, a flat spine of 40 /x — between this spine and the lateral one a spicule. The
lateral and dorsal spines are all rough. There is a fringe of obtuse processes on the
fourth leg. It was not ascertained if there were barbs on any claws.
Habitat.-^-Among moss from the sea-shore of Victoria, British Columbia, one
example only.
This appears to be a distinct species, but it is not sufficiently known, as only one
example was observed, and that might not be mature. There is no species known
with similar roughened spines. In E. duboisi they are spinulose, not rough, and the
animal is otherwise very different.
Genus — Milnesium, Doyere (2)
Milnesium tardigradum, Doyere (2)
This, the most generally distributed of all water-bears, was only observed in
Ontario, where it occurred in two localities, near the Lake of the Woods and in
Ottawa. Length, up to 800 M. Four eggs seen in the body.
Genus — Mctcrobiotus, Schultze (42)
A : SPECIES HAVING ROUGH EGGS
Macrobiotiis hufelandii, Schultze (42)
Abundant in all the Canadian localities visited ; eggs also plentiful. In the
Rocky Mountains it attained a length of 1200 /.-, being the largest water-bear I
have measured.
TARDIGRADA: CANADA 167
Macrobiotus echinogenitus, Richters ? (27)
This identification, being made from the egg alone, is somewhat uncertain. The
egg (Plate XXI. Fig. 58) is like that of M. areolatus (Fig. 53d), but is smaller,
and the spines are not separated by any space.
Habitat. — British Columbia and Ottawa.
For a discussion of the affinities of this species with M. areolatus and other
related species, see below, under M. areolatus.
Macrobiotus areolatus, sp. n. (Plate XXI. Figs. 53a-53e)
Synonym : M. echinogenitus, Richters, var. areolatus, Murray (19)
Specific characters. — Large, dark brown ; gullet wide ; teeth strong, bent near the
furca ; pharynx shortly oval, with apophyses, three narrow equal rods, and no comma ;
egg large, bearing very large conical processes, which are separated at their bases,
the surface between marked with irregular polygons which form a symmetrical
pattern.
General description. — The largest example measured was 700 ^ in length. The
egg measures about 100 /u. without the spines, 200 n over the spines, which, however,
vary considerably in length. Old individuals, as in most large species of the genus,
are strongly pigmented with a dingy brown colour, which is not disposed in such
regular bands as in M. oberhciuseri, but which still falls into obscure bands, prob-
ably caused by the disposition of the muscles and other structures in the skin.
The young are colourless and transparent. The gullet is somewhat constricted
towards the mouth and enlarged towards the pharynx, and bears the usual apophyses
on its end. The three linear rods are slightly curved and are nearly equal in length.
The stomach consists of few large cells. There is a pair of dark eyes. The claws
are in two similar pairs, which are united at the bases only, and diverge widely.
One claw of each pair is longer, and has supplementary points.
Four unripe eggs have been seen in the body together. The shell of the egg is
thick, and of two layers. The reticulation which appears on the surface is produced
by the edges of septa which cross the space between the two layers. The areolation
appears to be originally regular hexagons, of which alternate ones bear processes,
The intermediate hexagons are each divided by a transverse septum, producing two
irregular pentagons. The turgidity of the processes further distorts and obscures
the regularity of the original hexagons.
The areolation varies considerably. In one variety (Fig. 53f) the intermediate
hexagons are undivided and equal the processes in basal area. Some forms have all
the areolue rounded, and circular or elliptical.
Owing to imperfect understanding of M. echinogenitus this species was at first
168 J. MURRAY
united to it as a variety. Subsequent experience of the animal in many parts of the
world established confidence in the constancy of its peculiarities. The absence of a
" comma " in the pharynx cannot be considered an important specific character, but
when we find it constantly associated with an areolated egg, while M. echinogenitus
has a conspicuous comma, and non-areolated egg, the character adds weight to the
other specific distinctions.
Habitat. — Among moss, Kooky Mountains, British Columbia, and Ottawa,
June 1909. Widely distributed over the world — recorded from Spitsbergen, Scotland,
India, Tropical and South Africa, Australia, New Zealand, &c. Richters (39) has
recorded it from Ascension and Comoro.
Arctic examples have attained to a larger size, but the Canadian eggs are the
largest yet seen.
The rods in the pharynx are relatively shorter and broader in the young
(Fig. 53 is drawn from a young hyaline example). In the adult they are linear and
curved.
The characters of the egg sufficiently distinguish the species from all others
hitherto described. There are some species, not yet described, which have the
surface areolate in the same manner, but the processes of different form.
If the egg is not found, the identification is less certain. The presence of three
equal linear rods, without a comma, and of divergent claws, united at the base only,
differentiates it from the typical forms of hufelandii, harmsworthi, and other related
species. According to Richters, however, M. hufelandii may have three equal rods,
and the claws may be joined at the base only. The other species probably vary in
the same manner.
Since Richters' original diagnosis of M. echinogenitus was evidently too compre-
hensive, and included several forms which we are now, after extended experience,
enabled to separate, it will be well to take a review here of the group of related
species.
Richters, in his description, recognised two forms which he distinguished as a
and b, but he did not bestow separate names on them ; a has three " bacilla " and a
" comma " in the pharynx, l> has two equal bacilla, and a comma. Both have
V-shaped claws and the eggs are exactly similar in form, but that of a is much
larger.
M. harmsworthi was at first separated from echinogenitus on account of the form
of the claws alone, which were joined for about half of their length, as in M. hufe-
landi. Richters considers that the claws vary in the amount of union, just as those
of hufelandi do, but assents to the separation of M. harmsworthi, which he supposes
to be his echinogenitus a,* on the characters of the pharynx.
M. areolatus is also supposed by Richters to be included in echinogenitus a, but
* See Richters' "Tardigraden aus den Karpathen," Zool. Anzeiy., Bd. 3C, July 1910, p. 7. This
paper appeared after our Bibliographical List was completed, so could not be included in its proper place.
TARDIGRADA: CANADA 169
that has a " comma " in the pharynx, and the egg is figured with the processes close
together at their bases, so that there is no room for the " areolation."
There remains M. echinogenitus b, which, with the separation of a, becomes the
type of the species. It has the two bacilla equal, a not very common arrangement,
and the claws united at the base or for a short way above the base ; the surface of
the egg is not " areolate." A great many of the records of this species must be
regarded as doubtful. Many of these were made before the related species areolatiis
and harmsworthi were discriminated, so that unless details of the pharynx are given
it cannot be known to which form a record refers. It was till recently supposed that
the eggs of echinogenitus and harmsworthi could be distinguished. The original
specimens of the eggs of harmsworthi had processes much more shortly acuminate than
those of the typical echinogenitus, but some have recently been seen with points
almost as long. Records of echinogenitus have been commonly made (at any rate in
my papers) on the strength of finding the egg alone. Although the eggs with short
points may be usually harmsworthi and those with long points echinogenitus, there
will always be doubt about these records unless the eggs are so ripe as to show
pharynx and claws (see p. 89, footnote).
M. polaris has the surface of the egg "areolate " in the same manner as in one
form of areolatus (Fig. 53e), and the processes may be of the same form. It is,
nevertheless, a totally different egg, being much smaller, and with the polygons
relatively much smaller, and therefore more numerous.
Macrobiotus harmsivorthi, Murray (19)
Habitat. — Near the Lake of the Woods, and in the public parks, Ottawa.
The identification was made from eggs which contained young in which the
pharynx and claws could be seen. The egg measured 60 /a. without the processes,
and 80 n over the processes. These were even shorter than in the type, being
considerably broader than long. The claws of a pair were unequal, and united for
about half the length of the larger one.
A very similar egg, but of larger size, was got in Ottawa. It measured 80 /*,
without the spines, and 120 n over the spines. The pharynx and claws were
not seen.
Macrobiotus occidentalis, sp. n. (Plate XXL Figs. 54a-54e)
Specific characters. — Large, orange-red ; stomach and eggs darker red ; skin
hyaline, dotted ; gullet narrow ; teeth moderately slender, curved ; pharynx shortly
oval, with apophyses and two rods, the second shorter, and no " comma." Claws of
hufelandi type, united half-way. Egg spherical, covered with slender tapering
curved processes, which are separated at their bases.
General desertion. — Total length up to 800 p, pharynx of small example,
BRIT. ANTARCT. EXPED. 1907-9. VOL. I. 7,
170 J. MURRAY
42 fi long, claws 20 M- Eyes dark. The dots on the skin are irregularly placed,
but at very nearly equal spaces. They are not pigment spots, but appear to be
minute elliptical pieces of the same nature as the skin, set into it like buttons in a
cushion, and looking dark from the different refraction of the light. The reddish
colour is confined within the body fluid, and is resident chiefly in the fat-cells,
though the fluid is also more or less coloured. There are two layers of skin,
between which there is enclosed a colourless fluid, in which float numerous thin
hyaline oblong plates. This may be a pathological condition, but it is noteworthy
that it occurred in all the individuals examined, old and young, and in none of the
other species present.
The claws are like those of hufelandii, but more slender. Those of each pair are
unequal in length, and are united for half the length of the larger one, which bears
the usual two supplementary points.
The basal ridge, which in M. coronifer bears the " corona " of little spines, is
in this species irregularly dentate.
The processes of the egg are separated by interspaces greater than their own
diameter at the base. They have narrow conical bases, and taper to slender points
which are curved over.
The gullet is about 4 n in width, but is rather narrower below and expanded
towards the pharynx. The rod next the gullet is about three times as long as
broad, the second about twice as long as broad.
Habitat. — Among moss from the sea-shore, little above high-water level, at
Victoria, British Columbia ; very abundant, eggs also abundant.
M. occidentalis appears to have its closest affinities with that group of northern
species of which M. coronifer may be regarded as the type. Several of these species
have a spinose ridge in front of the claws, most distinct on the fourth legs. In
M. coronifer and M. granulatus the ridge bears spines — in M. crenulatus it is
wrinkled — in M. harms^vorth^ it is crenate or plain. In large examples of M.
occidentalis the ridge is dentate (Fig. 54d).
Several of the northern species have distinctive colouring, resident in the fat-cells.
M. coronifer and M. islandicus are bright yellow. The colour of M. occidentalis
is more inclined to red. As in M. coronifer, the colour is present even in the egg.
In the egg it is darker, and more distinctly red. It cannot then be resident in the
fat-cells. The body fluid itself is in old animals of a yellowish colour.
The processes of the egg have some resemblance to those of M. coronifer and
M. islandicus. The egg of coronifer is elliptical, and the spines are straight ;
that of islandicus is round, but it has two sorts of processes, spines like those of
coronifer and processes like those of granulatus. M. occidentalis has longer and
more slender spines than either, and they are variously bent and turned over at
the ends.
The pharynx is quite like those of coronifer and islandicus. On the whole the
TARDIGRADA: CANADA 171
species is nearest to islandicus, from which it is distinguished by the ridge without
spines, by the different egg-spines, and by the dotted skin.
In Australia and Hawaii there is a form, apparently belonging to this species,
agreeing with it in most characters, but not yet fully studied. The colour is a paler
yellow. The skin is dotted as in the type, but the specimens lacked the clear fluid
circulating between two layers of skin, and containing numerous hyaline plates. The
eggs were not seen in these countries.
Macrobiotus intermedius, Plate (23)
Habitat. — Near the Lake of the Woods, Ontario.
The egg had the typical top- shaped processes. It measured 50 ^ without the
processes, 58 n over them. The young squeezed out of the egg was 120 n in length,
and the round pharynx was 15 n in diameter. The processes of the egg were
separated by spaces greater than their own diameter, and the surface between them
was covered with regular pellucid dots.
Macrobiotus oberhauseri, Doyere (2)
Habitat. — Vancouver, British Columbia.
American examples were not papillose, as is so often the case in Africa.
B : SPECIES HAVING THICK-SHELLED EGGS WITH EMBEDDED RODS
Macrobiotus arcticus, Murray ? (19)
Habitat. — Vancouver, Rocky Mountains, Lake of the Woods.
No eggs were found in Canada, and without them there is some doubt about the
identification. The animal found in three of the localities visited agrees with
M. arcticus in having a narrow gullet, two short " bacilla " in the pharynx, and claws
of the Diphascon type.
The only other species known which has similar eggs is H. hastatus, Murray (18),
v.'hich was not observed in Canada.
C : SPECIES HAVING SMOOTH EGGS
Macrobiotus canadensis, sp. n. (Plate XXI. Figs. 6la-6ld)
Specific characters. — Small, hyaline ; gullet slender ; teeth abruptly enlarged
about the middle ; pharynx nearly round, with three short nuts, increasing in size
from first to third, comma very obscure or none ; claws widely divergent, but
approaching the Diphascon type ; one claw of each pair is longer and thinner than
the other, and that of one pair is very long and slender; eggs narrowly oblong,
smooth, laid in the skin at the moult.
General description. — Length 225 M. A pair of small dark ejes. The teeth are
172 J. MURRAY
straight for somewhat more than half their length from the points. They are then
enlarged and curved outwards to the furca. The pharynx is a little longer than
broad. The end of the gullet bears the apophyses. The nut next the gullet is
scarcely longer than broad ; the second is a little longer, and the third a little longer
still. The first is nearest the middle line, and the second and third diverge successively
farther from it.
The stomach is oblong and very slightly coloured. Its separate cells are not
conspicuous.
One egg measured 70 M by 36 M, but others were rather shorter. The newly
hatched young was 100 /u. in length.
There are few species with which it is necessary to compare M. canadensis very
carefully. The Macrobioti which lay smooth eggs are not so numerous as those which
lay rough eggs. There are two principal groups of them, one comprising species
which have warts or spines or distinct papillae on the body, the other species which
are smooth or very finely papillose. In the former group there are several which
have three short nuts in the pharynx ; in the second group, to which M. canadensis
belongs, there are only two which have the pharynx of this type.
M. tetradactyloides, Richters (35), and M. schaudinni, Richters,* are very similar
to M. canadensis. Both are, however, much larger and more robust animals.
Richters himself considers the size important in distinguishing M. tetradactyloides
from M. tetradactylus. M. schaudinni is nearly twice as long as M. canadensis, and
M. tetradactyloides is larger still. It is pretty certain that M. canadensis is really a
small animal, although some water-bears vary greatly in size. It was very common
in some parts of Canada, and yet it was uniformly small. The largest measured was
only 225 n in length, and many mature- looking animals, with eggs in the body, were
less than 200 M.
M. canadensis may be distinguished from M. tetradactyloides by the unequal
pairs of claws and by the greater inequality of the claws of each pair. It is more
difficult to separate from M. schaudinni, which has the pairs unequal. Richters
figures his species with much robuster claws.
In M. canadensis the claws are intermediate between the Diphascon type and the
V type. The long claw of each pair is very slender, and that of the larger pair is
almost bristle-like, as in M. oberhauseri. The bristle-like claw is set on to the lesser
claw of the same pair in the same manner as in Diphascon, but nearer to the base.
The two claws diverge very widely.
While usually smooth, it sometimes happens that the body is finely papillose on
the posterior half, or more than half, though never all over.
The nuts of the pharynx are only slightly rounded at the ends, not enough to
destroy their quadrate character.
Habitat. — Victoria, British Columbia, and the Rocky Mountains, abundant.
* " Tardigraden-Studien," Ber. Senckbg. Nnturf. Ges., 1909, p. 32.
TARDIGRADA: CANADA 173
Macrobiotus tuberculatus, Plate (23)
Habitat. — Victoria, British Columbia.
The examples were rather larger than the average, but not so large as M. nodosus.
They were colourless, and the knobs did not show on the outline in dorsal view.
Macrobiotus sattleri, Richiers (26)
Habitat. — Rocky Mountains.
M. sattleri is either variable in the characters of the pharynx, or there are several
forms which have the same peculiar skin-markings.
One form has three short rods in the pharynx. The Australian form (see p. 141)
has two unequal rods. The Canadian form has two equal rods, each about four times
as long as broad. No comma was seen. The pairs of claws are unequal, and the
claws of the larger pair are united for a short distance above the base.
Macrobiotus, sp. ?
Four eggs in a skin of 200 M in length. Eggs smooth, of about GO n by 45 M. Claws
Vs, equal pairs of equal claws, joined at the base only.
Without knowledge of the pharynx identification is not possible. It is certainly
different from any of the species recognised. The claws prove that it is not a
Diphascon.
D : EGGS UNKNOWN
Macrobiotus virgatus, sp. n. (Plate XXI. Figs. 55a-55c)
Specific characters. — Large and robust ; pigmented with a warm brown colour
arranged in longitudinal bands ; gullet very wide, and teeth strong ; pharynx
shortly oval, with apophysis, and two rods and a nut in each row, the nut between
the two rods ; dark eyes ; claws very thick, of the hufelandi type, very unequal,
united for half the length of the longer one, which bears two thick supplementary
points.
Detailed description. — Length, up to 750 /j.. The colour is a warm brown, not
unlike that of M. oberhciuseri, but less inclined to purple. It is arranged in three
principal longitudinal bands, one median and two lateral. These latter may be
subdivided into narrower bands, and there are some thin transverse bands.
The gullet is as much as 9 or 10 M in diameter. The teeth penetrate its wall at
about the middle of its length. The end of the gullet in the pharynx has a slight
projecting rim. The rods are of unusual proportions — the first and third are about
four times as long as broad, the second varies from little longer than broad to twice
as long as broad.
174 J. MURRAY
The claws are quite like those of hufelandii, but they are more unequal and even
thicker. The egg is unknown.
Habitat. — Victoria, British Columbia ; Ottawa, Australia (?), Franz- Josef Land (?).
The records for Australia and Franz-Josef Land are very uncertain, as they
depend on the characters of the pharynx only. There were claws on the only
specimen from Franz-Josef Land, but as they were only seen in profile, little can
be said about them. They do not appear to be so thick or so far united as in
M. virgatus.
If the Franz-Josef Land specimen is M. virgatus, then the eggs are smooth and
are laid in the skin. Judging from the characters of the teeth, gullet, and claws, it
might be expected that the animal would prove to belong to the hufelandi group,
with rough eggs. There are very few species which combine the characters of
smooth eggs and hufelandi claws. M. rubens is the best instance known to me.
The peculiar proportions of the " bacilla" in the pharynx, one short between two
long, distinguish M. virgatus from all related species. Only M. augusti, which is not
closely related, has a rather shorter middle rod.
It is rarely permissible to describe a Macrobiotus of which the egg is unknown.
In this case the association of so many distinct characters — the colour, the pharynx,
the claws — seemed to justify a breach of the rule. When the eggs are found there
should be no difficulty in demonstrating their connection with this species.
Macrobiotus sp. ? (Plate XXI. Fig. 60)
Description. — Size moderate, length 300 M. Colour grey ; no eyes. Gullet of
moderate width. Teeth strongly curved, with large furca. Pharynx shortly oval,
with conspicuous apophysis, and two "bacilla" in each row. The first, next the
gullet, is three times as long as broad. It is divided by a constriction into two
equal parts. The second is not quite twice as long as broad. There is no comma.
The claws resemble exactly those of M. canadensis (Fig. 61c). The pairs are
unequal and diverge widely. Each pair has one claw longer than the other, and the
long claw of the larger pair is very slender, almost bristle-like. The long claw is
attached to the shorter claw near its base, showing a slight approach to the
Diphascon type.
The association of several distinct characters marks this as a distinct species, but
as only one example was seen, and the egg is unknown, it is left in the meantime
unnamed.
Habitat. — Rocky Mountains
TARDIGRADA: CANADA 175
E : UNIDENTIFIED EGGS.
Macrobiotics, sp. ? (Plate XXL Fig. 57)
A large egg, measuring 105 n over the processes, which are low cones, rounded
at the ends, and papillose all over. The processes stand close together, without any
interspaces.
This closely resembles an egg figured by Eichters in the " Moosrasen des
Gaussbergs " (35), Plate XX. Fig. 7, as probably related to M. echinogenitus. It is
probably a distinct species.
Macrobiotus, sp. ? (Plate XXI. Fig. 56)
A large egg, measuring 100 M over the processes, 80 M without the processes.
These are narrow conical pegs, rounded at the ends. They are separated by spaces
about equal to the diameter of the pegs at the base.
A similar egg is figured in " Tardigrada of the South Orkneys " (15), Plate IV.
Fig. 14, as perhaps a form of M. echinogenitus. Another somewhat like it is figured
in "Arctic Tardigrada" (19) Plate XLV. Fig. 2, as perhaps M. islandicus, Richters.
The processes are not rounded, but many are a little expanded at the tip, showing an
approach to hufelandii.
The Canadian egg is probably a distinct species.
Macrobiotus, sp. ? (Plate XXL Fig. 59)
A small egg, 80 M over the spines, 68 /« without them. The processes are small
cones, slightly acuminate, acute. They are separated by spaces rather less than the
diameter of the processes at the base. The surface exposed between the processes is
marked by regular pellucid dots.
Several species have eggs like this, and they cannot be distinguished with any
certainty unless they are found containing young.
M. dispar has such an egg, but rather larger ; that of M. pullari is somewhat
smaller. The egg of M. ascensionis is considerably smaller, and has the processes
more closely set.
Genus — Diphascon, Plate (23)
Diphascon chilenense, Plate (23)
Habitat. — Rocky Mountains.
Diphascon alpinum, Murray (14)
Habitat. — Rocky Mountains : near the Lake of the Woods, Ontario.
176 J. MURRAY
Diphascon scoticum, Murray (11)
Habitat. — Stanley Park, Vancouver : Rocky Mountains.
Diphascon canadense, sp. n. (Plate XXI. Figs. 62o-62c)
Specific characters. — Small, without eyes, hyaline. Gullet long, slender.
Pharynx round, with apophysis, two rods and a " comma " in each row, the first
about twice as long as broad, the second about as long as broad. Claws typical for
the genus.
Detailed description. — Length up to 250 M. Form narrow and elongate.
Pharynx 25 M in diameter. Gullet 50 to 60 p. in length, very slender, little over 1 /*
in diameter. The largest pair of claws is about 12 to 15 M in length. The long
claw is very slender, the short one somewhat thick. Both claws of the smaller pair
are slender — they are unequal and joined at the base only. The eggs were not
seen.
Habitat. — Moss from the sea-shore at Victoria, British Columbia, fairly abundant.
Among species having the gullet slender and the pharynx short it is only
necessary to compare D. canadensis with D. oculatum (13), to which it is closely
related. It was at first supposed to be a blind form of oculatum, till other slight
differences were noticed.
D. oculatum is larger, and has a pair of dark eyes. The pharynx is not so
nearly round, and the two thickenings in each row are equal, and scarcely longer
than broad. The claws of the shorter pair are shorter and thicker.
The eye-spots are not considered to be trustworthy as specific characters, though
I know of no variation in the genus Diphascon. When, however, we consider the
slight but constant differences in the pharynx and claws, the species appears
sufficiently distinguished from D. oculatum.
REMARKS ON THE CANADIAN TARDIGRADE FAUNA
Its composition. — Of the 31 species studied 23 were identified — the other 8
require further study. There are 9 species of Echiniscus, 17 of Macrobiotus, 4 of
Diphascon, and 1 Milnesium. Seven species are considered to be new to science
(2 Echiniscus, 4 Macrobiotus, and 1 Diphascon}.
The list is such a one as might be expected as the result of a very limited amount
of work on any continental area. The different genera arc represented in about the
average proportions, and there are no very peculiar forms. The most curious fact in
the composition of the list is the total absence from it of species of Echiniscus of
that section of the genus which has plates V. and VI. quite separate. As E. muta-
bilis is one of the most cosmopolitan water-bears the absence from our collections
must be considered as accidental.
TARDIGRADA: CANADA 177
In the accompanying table the distribution of the Canadian species, both in
Canada and over the world, is shown in thirteen columns.
DISTRIBUTION : LOCAL AND GENERAL
Canada.
02
pa
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P
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K
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S
CO
J
0
ri
J
M
3
^
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E
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£
03
M
*"'"
0
9
OJ
o
O
O
*
0
O 0
£
B —
a
-O
*3*
3
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H
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1
•
o
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^N
•
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02
^
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*
o
Echiniscus gladiafor
X
—
. —
—
—
X
—
—
—
X
—
—
—
E. sylranus ...
—
—
X
—
—
—
—
—
—
—
—
—
E. intermedius
X
—
—
—
—
—
—
X
—
X
—
—
7-T J •
&. cannctensi-s
E. oihonnce ...
X
—
—
—
X
—
—
X
—
—
X
—
E. bisetosus ...
X
—
—
—
X
—
—
—
—
—
—
—
Milnesium tardigradiim
—
—
X
X
X
X
X
X
X
X
X
X
X
Macrdbiotus hufelandii .
X
X
X
X
X
X
X
X
X
X
X
X
—
if. echinogenitus .
X
—
—
X
X
X
X
X
X
X
X
X
X
M. areolatus.
X
X
—
X
—
X
X
X
X
—
X
X
—
M. harmsiuorthi
—
—
X
X
—
X
—
X
X
X
X
X
—
M. occidentalis
X
—
—
—
—
—
—
—
X
—
X
—
—
M. intermedius
—
—
X
—
X
X
X
X
X
X
X
X
—
M, oberhciuseri
X
—
—
. —
X
X
X
X
—
—
X
X
X
M. arcticus ...
X
X
X
—
—
X
—
X
X
X
—
X
X
M. tuberculatus
X
—
—
—
—
X
—
X
—
—
—
X
—
M. sattleri ...
X
—
—
—
X
X
—
X
X
X
—
—
M. virgatus ...
X
X
X
X
Diphascon chilenense
—
X
—
—
X
X
X
—
X
X
—
X
X
D. alpinum ...
—
X
X
—
X
X
—
—
—
X
—
X
X
D. scoticum ...
X
X
—
—
X
X
—
—
X
X
X
X
X
D. canctdense.
The table brings out some interesting facts in distribution. About nine of
the species are cosmopolitan, or very widely distributed. These are Milnesium, M.
hufelandii, M. echinogenitus, M. areolatus, M. intermedius, M. oberhauseri, M.
arcticus, D. chilenense, and D. alpinum. These occur in six or more of the nine
great regions into which the surface ol the earth has been divided for the purpose of
this comparison.
The others are more restricted in their range. The seven new species described
in this paper are not all confined to Canada ; three of them are already known else-
where. M, areolatus (which was described some years ago as a variety of M. echino-
genitus} is even among the cosmopolitan species. M. occidentalis, though discovered
and fully studied in Canada, appears to have a " Pacific" distribution, as it has since
been found in Honolulu and Australia. Another species having the identical range
is Echiniscus intermedius, first discovered in Australia, and subsequently in Hono-
lulu and Canada. In this case the species appears under a slightly different form in
all three localities.
BEIT. ANTARCT. EXPED. 1907-9. VOL. I. 2 A
178 J. MURRAY
The relation of the Canadian Tardigrade Fauna to that of other regions is shown
in the following figures. Canada has 16 species which occur also in Europe, 14 in
Australia, 14 in the Arctic, 8 in the Pacific Islands, New Zealand, and Asia, 9 in
Africa, 7 in the Antarctic, and 6 in S. America. From these figures it would seem
as if the relation with Europe were closest, and with S. America most distant, but it
is scarcely necessary to point out that these figures probably indicate the condition
of our knowledge of the different regions rather than the real affinities. Of course,
since the part of Canada explored, and the part of Europe of which the Tardigrada
are best known, are both in the north temperate zone, we might expect, and there
may really be, a close affinity between the Tardigrada of the two regions. The
occurrence of seven species in Canada which are as yet unknown in Europe suggests
that the affinity is not very close. No confirmation • of this can be drawn from the
large number of European species which are unknown in Canada, as Canada is an
almost unexplored country.
In Canada the productiveness of British Columbia in relation to the other districts
visited is very striking. All the collecting was hurriedly done, in Victoria no less
than elsewhere, yet the moss from the sea-shore there yielded sixteen out of the
twenty-three species noted for Canada.
SUMMARY OF RESULTS
THE work of the Expedition on Tardigrada has resulted in the enumeration of
fifty species, distributed in five genera. This may seem a small number, considering
how many countries were visited, and the great range of climatic conditions which
they present, from tropical heat to polar cold.
These fifty species are, however, about half of the known Tardigrada, and
considerably more than half of the land and fresh-water species. And in addition to
the fifty species identified, a considerable number more (something like twenty-three
species) were observed and described, which, while not sufficiently known to be
identified or pronounced to be new, are recognised as distinct from any of the species
in the list.
It must be remembered, too, that it is only within the last few years that
naturalists have begun to discriminate the species of Tardigrada carefully, and to
realise that they are fairly numerous. For half a century after the discovery of the
first water- bear by Go'ze in 1773 naturalists supposed that there was only one, or at
most two, species. From Schultze's time (1834), when Tardigrada began to be more
carefully looked at, for another half- century, scarcely a dozen species were dis-
tinguished. Even in 1888 Plate's monograph (23) admitted only twenty-five species,
in six genera. Eight of the species and two of the genera were Plate's own
discoveries. One of his genera, and at least four of his species, would now be
disallowed.
Only with the arrival of Richters in the beginning of the present century were
the Tardigrada looked at with sufficient care, and satisfactory diagnoses inaugurated.
About the same period it began to be appreciated what great facilities were afforded
by mosses for the collection and study of microfaunas, and in consequence the
Tardigrada and other microscopic animals of distant countries became known.
Among the Tardigrada collected by the Expedition there are seventeen species
recognised as new to science, and one new generic type was found in Australia.
In the accompanying table is given a list of all the species collected, and their
distribution is shown in all the countries visited, and also the distribution over the
world.
Distribution. — An analysis of the list brings out some interesting facts. There
were observed in all 73 species, of which 33 were identified as known species, 17 are
179
180 J. MURRAY
DISTRIBUTION OF ALL SPECIES COLLECTED BY THE EXPEDITION.
ANTARCTIC.
NEW ZEALAND.
AUSTRALIA.
PACIFIC ISLANDS.
CANADA.
S. AMERICA.
EUROPE.
•J
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«
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o
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:3
r£
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cT
i
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0
VICTORIA ET VANCOUVEK
MTS SELKIRK, MKS ROCHEUSES
ONTARIO
Heleopera petricola ....
„ sordida ....
„ sphagni ....
,, sylvatica
Hyalospheuia cockayni
,, subflava
Lesquereusia .....
Lieberkiihnia wageneri
Nebela bigibbosa ....
„ caudata .....
+
4-
—
—
+
—
—
+
+
—
—
+
—
+
—
—
—
+
+
+
+
4-
4-
+
+
+
+
—
+
—
—
i
—
+
—
—
—
4-
—
_l_
+
-I-
+
—
—
+
—
—
„ collaris .....
,, dentistoma ....
„ flabelluluin ....
,, griseola .....
„ lageniformis ....
„ longicollis ....
„ martial! .....
,, militaris ....
„ minor .
,, tincta .....
„ tubulosa ....
„ vas
Pannnlina cyathu.s ....
Phryganella hemisphaerica .
,, nidulus ....
Placocysta jurassica ....
,, spinosa ....
Plagiopyxi.s callida ....
+
—
—
+
4-
4-
+
i
+
—
+
4
4-
—
—
-4-
4-
—
4-
4-
4-
4-
4-
4-
4-
4-
4-
4-
+
+
+
+
+
+
—
+
_L
+
4-
4-
+
+
+
+
4-
4-
—
—
4-
• —
+
4-
—
—
+
—
+
—
—
— _
+
4-
—
—
4-
4-
4-
—
4-
+
+
—
+
—
4-
4-
—
4
—
+
+
_i_
4-
4-
+
+
+
+
+
+
4-
4-
—
—
—
+
4-
—
—
+
4-
+
+
+
+
—
—
+
4-
4-
—
Pontigulasia bryophila
„ compressa
Quadrula irregularis ....
„ symmetrica ....
Sphcnoderia dentata ....
„ fissirostris
Trinema complanatum
„ enchelys ....
„ lineare .....
Acanthocystis .....
4-
—
—
—
+
4-
4.
+
j_
4-
4-
+
—
+
+
+
4-
4-
E
4-
4-
+
+
+
+
+
+
+
+
+
4-
4-
4-
—
4-
4-
4-
222 E. PENARD
CONSIDERATIONS GENERALES SUR LES ESPECES RENCONTREES
Amoeba muralis. PENARD
Arch, ftir Protistenkunde, B'1 17, p. 267. 1909
Cette amibe, de faible taille, et peu apparente en elle-meme, a dfi probablement
se trouver dans plusieurs des re'coltes ; mais ce n'est qu'a 1'etat vivant et actif qu'on
pent esperer pouvoir la reconnaitre. Dans la mousse du Pare National, & Sydney,
il s'en est rencontre un individu, bien vivant, qui par ses principaux caracteres,
plasma tres-dense, opalescent sur les bords, prolongements pseudopodiques tres-courts
rayonnant sur les cote's, noyaux tres-petits et nombreux, repondait bien a ce que
nous savons de cette espece. Les noyaux, pourtant, montraient une structure un
peu speciale ; au lieu du nucleole central et unique d^crit dans le type, on en
trouvait plusieurs, repartis sans ordre dans un sue nucleaire tres-compact.
Amoeba radiosa. EHRENBERG
Abh. K. Akad., Berlin, p. 39. 1830
Podostoma filigerum. CLAP. ET LACHM. Mem. Inst. Nat. Genevois, vol. 6, p. 441. 1859
Cette amibe n'est apparue qu'une seule fois, dans les mousses recoltees aux
environs de Sydney. Elle y etait represented par cette forme tres-petite, a pseudo-
podes droits et tres-fins, d'une longueur extraordinaire, pour laquelle Claparede et
Lachmann ont cree le nom de Podostoma Jiligerum, mais que 1'on s'accorde en
general aujourd'hui a regarder comme une simple variete" de A. radiosa.
II n'en a e"te trouve qu'un seul individu, mais typique, vivant, actif et largement
ddploye. Peut-etre serait-on en droit de supposer que cette amibe se trouvait la
en qualite" d'etrangere, apportde avec 1'eau dans laquelle la recolte avait ete lavee.
Je crois la chose a peine vraisemblable ; mais le fait est possible pourtant.
C'est ici du reste, pour le dire en passant, la seule fois que je croie devoir
indiquer comme douteuse la provenance des organismes qui font 1'objet de ce
mdmoire.
Amoeba sphaeronucleolus. GREEFF
Biolog. Centralbl', Bd 11, p. 639. 1891
Les Amoebiens nus ne sont guere determinables autrement que sur le vivant ;
mais il faut faire une exception pour les amibes terrestres, ou " pelliculeuses," qui,
dejk mortes ou a I'e'tat de vie latente, plissees, informes, montrent cependant encore
d'une maniere tres-nette les caracteres distinctifs du groupe special auquel elles se
RHIZOPODES D'EAU DOUCE 223
rattachent. Mais quant aux caracteres de V espece, ils restent beaucoup moins
distincts, et dans les diverses recoltes ou se sont trouvdes des amibes a pellicule,
j'ai dii me contenter, pour la diagnose " sphaeronucleolus," de constater les signes
distinctifs suivants : taille faible, noyau unique et arrondi.
Dans les mousses des Montagues Rocheuses, les individus se sont parfois
montres vivants, mais jamais actifs ; ils etaient enkystes, on plutot, ils revetaient
1'etat special qui dans ce groupe represente 1'errkystement ; plasma retracte, arrondi,
sous la protection de la membrane propre ou pellicule caracteristique, qui, en fait,
a la valeur d'une enveloppe veritable.
Amoeba terricola. GEEEFF
Arch, fur Mikrosk. Anat., B'1 2, p. 300. 1866
Beaucoup plus nettement reconnaissable que la precedente, 1'Amoeba terricola
s'est rencontree dans un grand nombre de stations, souvent morte, ou bien aussi sous
cette forme retractee qui dans cette espece represente 1'enkystement ; plus rarement
elle etait vivante (Montagues Rocheuses, Victoria), mais alors, malades en apparence,
les animaux n'ont jamais montre les signes d'un reveil certain.
Dans un individu,qui semblait vouloir sortir de son e"tat de repos et montrait
deja quelques petites vesicules contractiles, le noyau renfermait, dissdmines parmi
les nucleoles caracteristiques, un certain nombre de grains d'excretion brillants. C'est
la un fait que j'avais constate quelquefois deja, a Geneve, sur des individus qui
avaient e"te soumis a un jeune prolong^.1
Amphitrema stenostoma. NUSSLIN
Zeitsch. fur Wiss. Zool., Bd 40, p. 717. 1884
Cette espece, caracte"ristique des tourbieres a sphagnum, est assez rare en general ;
mais quand on la rencontre, c'est presque toujours en grandes quantites. Tel etait bien
le cas a Katoomba, oil ce rhizopode ne s'est montre que dans une petite touffe de
sphagnum jointe aux mousse ordinaires. Les individus etaient nombreux, encore
verts et pourvus des zoochlorelles caracteristiques de cette espece, morts cependant, ou
peut-etre seulement enkystes. Aucun d'eux, en tout cas, n'est revenu a la vie active.2
1 Je crois avoir aussi, a deux reprises, rencontre Amoeba alba, qui se distingue de la precedente par sa
forte taille et par la presence de noyaux tres petits et en nombre considerable. Mais il ne m'a pas ete possible
d'arriver a des conclusions certaines.
2 Pendant les quelques semaines qu'a dur6 mon examen de ces recoltes, il a fait tres-chaud, et les
rhizopodes, ou bien revenaient a la vie dans les quarante-huit heures, ou bien n'y revenaient pas du tout.
De plus, ceux-la meme qui reprenaient vie mouraient tres-rapidement. A cette regie, il y a eu cependant des
exceptions.
E. PENARD
Amphizonella violacea. GEEEFF
Arch, fur Mikr. Anat., Ba 2, p. 323. 1866
Cette espece s'est montree dans trois stations, au Mont Cook, dans les Alpes
australiennes, a Katoomba. Les individus, partout rares, etaient morts, reconnaissables
pourtant a des traits certains ; mais pales, jaunatres, et presque entierement de"pourvus
de la teinte violette qui caracttSrise cet organisme.
Arcella arenaria. GREEFF
Arch, fiir Mikr. Anat., B'1 2, p. 330. 1866
»
C'est la un des rhizopodes les plus caracteristiques des mousses, dans lesquelles
il est bien rare de ne pas le trouver. II manque pourtant. dans quelques-unes des
recoltes ; peut-etre y existe-t-il en fait, et a-t-il passe" inapercu ; mais eh tout cas,
il y serait fort rare, car, lors de la confection de mon tableau general, etonn^ de ne
pas avoir a 1'indiquer, j'ai tenu a contr61er le fait en examinant ce qui me restait
de mousses, mais sans rien trouver de nouveau.
Presque partout ce rhizopode se trouvait a 1'etat de coquilles vides. Dans les
Alpes australiennes, beaucoup d'individus vivaient, mais aucun n'a deploye de pseudo-
podes. A I'lle Macquarie, cette espece se trouvait represented par une variete plus
claire et plus grande que le type habituel.
Arcella vulgaris. EIIRENBERG
Abh. Kais. Akad. Wiss., Berlin, p. 40. 1830
Cette espece n'habite pas les mousses, ou elle est normalement remplact'e par
1'Arcella arenaria. Aussi bien n'est-ce pas dans les mousses qu'elle a etc recontree,
mais dans un paquet de Conferves prises dans un marecage, et qui se trouvaient
jointes aux mousses de la meme station (He Macquarie).
L 'Arcella vulgaris est connue par un grand nombre de varietes ; a 1'ile Mac-
quarie, c'etait une petite forme, de 75 p. de diametre, dont 1'enveloppe, tres-claire,
couverte de dessins alveolaires bien nets et reguliers, dtait relativement peu rennee,
se rapprocbant par la de V Arcella discoides, qui, elle aussi, fait le desespoir des
naturalistes par ses nombreuses varietes.
Les individus, qui se trouvaient la en grand nombre, etaient toujours a, 1'etat
de coquilles vides.
RHIZOPODES D'EAU DOUCE 225
Assulina muscorum. GREKFF
Sitzungsber., Marburg, p. 117. 1888
Cette petite espece, caracteristique avant tout des mousses, oil elle ne manque
presque jamais, s'est en effet rencontree un peu partout dans les recoltes, presque
toujours a 1'etat de coquilles vides et souvent de"colorees.
Assulina seminulum. (EHRENBERG), LEIDY
Difflugia seminulum. EHRBG. Monatsberichte K. Akad. Wiss., Berlin, p. 379. 1848
Assulina seminulum. LEIDY. Freshw. Rhiz. N.A., p. 225. 1879
Ce rhizopode est surtout caracteristique des Sphagnum, mais on le trouve
aussi, frequemmeut, dans les mousses belles et bien fournies des for6ts. Sa taille
normale est, le plus souvent, de 80 n environ, mais il arrive a de"passer quelquefois de
beaucoup cette mesure. A Ngauruhoe, par exemple, c'etait une grande forme, d'un
brun-rouge vif, et qui mesurait jusqu'h, 115 /u en longueur; tl Vancouver, dans les
mousses, la forme et la taille etaient normales, mais dans les Sphagnum, on y trouvait
cette belle et grande varidte discoide, de 110 ^ 115 /", et a dessins particulierement
bien marques, que dans le temps j'avais cru devoir considerer comme une espece
speciale (^4. scandinavica).
Aiverinzewia cyclostoma. (PENARD), SCIIOUTEDEN
Hdeopera cyclostoma. PENARD. Faune Rhizopodique Leman, p. 390. 1902
Awerinzewia cyclostoma. SCHOUTEDEN. Ann. biol. lacustre, t. i. p. 357. 1906
Quelques individus seulement, a 1'dtat de coquilles vides, revetues encore de leur
teinte caracteristique d'un violet noiratre, se sont montres dans la rdcolte de Ottawa.
Bullinula indica. PENARD (emend)
Bulinella indica. PENARD. Journ. R. Micr. Soc., p. 274. 1907
La premiere mention de ce Rhizopode, Fun des plus interessants parmi ceux des
mousses, et le prototype de tout un groupe special voisin du genre Diplochlamys, ne
date que de trois annees a peine, et son histoire est assez curieuse en elle-meme.
Trouv(5 en 1906 par J. Murray, dans des mousses qui provenaient de 1'Himalaya,
puis de'crit par moi-meme en 1907, sur la demande de Mr. Murray, ce rhizopode sem-
blait devoir rester tres-rare ; mais a peine 1'espece venait-elle d'etre publiee, que
Mr. Murray la retrouvait dans les Sphagnum de 1'Ecosse, puis dans des mousses venant
226 E. PENARD
de 1'Ouganda. En 1909, je la re*coltais enfin moi-meme, en grande abondance, dans
une tourbiere du Jura suisse (La Pile), et alors dans un etat de vie et de fraicheur
qui me permettait d'en faire une etude biologique qui n'avait pas 6t6 possible
jusque la.1
Au retour de 1'Antarctique, Mr. Murray a re"colte cette espece dans toutes les
localites visite"es ; elle ne manque, en fait, qu'aux lies Auckland, lesquelles d'ailleurs
n'etaient representees dans les recoltes que par quelques fragments d'ecorce revetus d'une
mousse rase.
Partout la Bullinula revetait son apparence caracteristique, mais cependant elle se
montrait quelque peu differente suivant la nature des particules (paillettes siliceuses
plus ou moins nombreuses, petits grains de quartz, fibres de mousses, diatomees) qui
recouvraient 1'enveloppe, comme aussi sous le rapport de la taille, qui dans cette espece
varie dans une mesure assez forte.
Mais a cet dgard, il est un detail sur lequel il me faut insister un instant, et sur
lequel Mr. Murray, en 1909, et a propos des mousses de 1'Ouganda, attirait deja mon
attention : A cote de la forme normale, grande et relativement allongee, ellipsoidale, de
190 a 200 M en ge'ne'ral, et qui arrive jusqu'a 250 M dans des cas exceptionnels, on en
trouve parfois une autre, presque ou meme quelquefois tout-a-fait ronde, et beaucoup
plus petite, de 130 a 170 p. de diametre (Fig. 1).
C'est bien la e"galement ce que montraient les recoltes qui font 1'objet du memoire
actuel ; la petite forme ronde s'est retrouvee dans une grande partie des collections ;
a Katoomba, a Kotorua, a Oahu, a 1'ile Macquarie, elle etait seule, et la grande
forme manquait; a 1'ile Stewart, dans les Alpes australiennes, a Sydney, les deux
formes se trouvaient melangees ; au Queensland, & Ngauruhoe, a Victoria, dans les
Montagues Rocheuses, seule la grande forme etait representee.
II faut voir, semble-t-il, dans cette petite forme ronde une variete speciale ; mais
entre cette variete et le type, on trouve, dans les stations ou elles vivent ensemble,
les transitions les plus diverses, de sorte qu'on peut se demander s'il n'y aurait pas
la une espece unique, mais assez variable, et qui tendrait a se fixer sous deux
formes extremes.
La premiere description de ce Rhizopode date de Juin 1907; mais d'apres ce
que Mr. Murray m'apprend aujourd'hui, il parait que ce nom de Bullinella a ete deja
applique a un genre de mollusques.
II faut done nous hater, pendant que cet organisme reste encore peu connu,
d'apporter la rectification necessaire ; et peut-etre la denomination generique de
BULLINULA au lieu de Bulinella suffira-t-elle pour contenter les zoologistes.
1 Cette eiude fera 1'objet d'un chapitre dans un des prochains numoros de la Revue Suisse de Zoologie
(Notes sur quelques SarcodinOs, 3° partie, vol. 19, 1911).
RHIZOPODES D'EAU DOUCE 227
Centropyxis acaleata. STEIN
Sitzungsber. Bohm. Akad. Wiss., B'1 10, p. 43. 1857
Cette espece est toujours rare dans les mousses, et il u'en a ete trouve que
quelques exemplaires dans quelques-unes des collections. A Vancouver (Sphagnum]
elle etait representee par cette petite forme trcs-plate, claire, transparente et mem-
braneuse, dont CASH a fait une variete speciale, var. s-pinosa (British Freshw. Rhizop.,
Vol. I. p. 135, 1905). A Oahu, c'etait egalement une petite forme, tres-rapprochee
de la m6me varie'te'.
Centropyxis delicatula. PENARD
Faune Rhizopodique Leman, p. 308. 1902
Quelques exemplaires seulement, a 1'etat de coquilles vides, an Queensland et
a Fiji.
Centropyxis horrida, spec, nova
Fig. 2, a et b
En meme temps qu'il m'envoyait ses re"coltes, Mr. Murray attirait mon attention
sur une Centropyxis de 1'ile Stewart, qui lui paraissait nouvelle, et qui en efFet me
semble certainement devoir etre decrite comme une espece a part.
L'enveloppe elle-meme, dans son ensemble, n'a rien qui doive la faire distinguer
de la Cent, aculeata, connue d'ailleurs par de nombreuses varietes. Aplatie sur son
axe dorso-ventral, et plus fortement a sa partie anterieure ou se trouve 1'ouverture
buccale excentrique et infere, elle est jaunatre, formde d'une chitine relativement
tres-claire dans laquelle sont empatees des paillettes siliceuses, minces, tres peu
distinctes, puis souvent aussi des particules de nature organique, et des filaments
brunatres arraches aux mousses.
Mais ce qui distingue cette forme de toutes les autres du meme genre, c'est
1'existence, non pas de cornes veritables, mais d'une sorte d'aile ou de carene, plate,
dechiquetee, creusee de lacunes que parfois viennent combler des fragments siliceux,
et qui borde toute la peripherie de 1'enveloppe, a 1'exception toutefois de la partie
anterieure, qui reste lisse aux environs de la bouche.
De distance en distance, alors, cette carene lamellaire est munie de prolongements,
tantot arrondis a leur sommet, sinueux, lobes, capricieux dans leur forme, tantot (et
BRIT. ANTAIICT. EXPED. 1907-9. VOL. I. D
228 E. PENARD
le plus souvent) etires en longues Opines acerees, droites ou recourbees ; plus rarement
enfin, les Opines sont remplacees par de simples expansions anguleuses (Fig. 2b).
Toute cette structure donne 1'impression d'une exsudation, qui se serait figee
apres avoir coule ; il semble que la coquille a etc collee a quelque chose, aux feuilles
des mousses, d'ou la pluie ne pourrait alors plus la detacher ; et peut-etre cette
impression premiere correspond-elle a la realite ; ce qui contribuerait a le faire croire,
c'est que les longues epines laterales, au lieu d'etre normalernent dirigees par leur
pointe vers la partie posterieure de la coquille, comme c'est le cas dans les autres
Centropyxis et dans les Khizopodes en general, montrent des directions qnelconques,
et mSme semblent se recourber de preference vers 1'avant ; c'est la en tout cas un
fait anormal, pen en rapport avec les besoins de I'animal a 1'etat actif.
La Centropyxis horrida dtait abondante a 1'ile Stewart. La seule station ou
elle se soit retrouve"e a e"te Katoomba, dans les Montagues bleues, ou trois individus
seulement se sont montres.
Centropyxis laevigata. PENARD
Faune Rhizopodique Leiuan, p. 306. 1902
Cette espece est rare ; il n'en a ete vu qu'un nombre tres-restreint d'individus,
a Rotorua, a Ngauruhoe, et a Katoomba.
Corycia aculeata. GREEFF
Pseudochlamys aculeata. GREEFF. Sitzungsber., Marburg, p. 332. 1888
Deux exemplaires seulement, nettement reconnaissables, a dents relativement
courtes, ont ete trouves a 1'ile de Suva, Fiji.
Corycia flava. GREEFF
Amphizonella flava. GREEFF. Arch. f. Mikr. Anat., Bd 2, p. 329. 1866
Cette espece, en general commune, surtout sur les vieux murs ou dans les mousses
rases qui garnissent les troncs des arbres, s'est rencontree dans un grand nombre de
stations, toujours typique dans ses caracteres, mais assez variable de taille suivant les
localites. Dans cette espece, 1'enveloppe mesure de 80 a 100 M en general; mais a
Katoomba, elle arrivait a 145 M, et jusqu'a 150 M dans les Montagnes Rocheuses.
Partout les animaux ^taient vivants, mais aucun d'eux n'a montre de pseudopodes ;
dans le genre Corycia, d'ailleurs, on ne les voit presque jamais.
RHIZOPODES D'EAU DOUCE 229
Corycia penardi. PENARD, AWERINZEW
Cor. Coronata, var. simplex. PENARD. Faune Rhizop. Leman, p. 179. 1902
Cor. Penardi. AWERINZEW. Trudui Petersb. Ob., torn. 36, p. 143. 1906
Cette espece, pen commune en general, ne s'est montree que dans pen de stations,
et toujours represented par quelques individus settlement, 3 a Vancouver, 1 a Suva,
1 au Mont Cook. A Ottawa, elle etait pourtant moins rare.
Corythion dubium. TARAKEK
Abhand. Kon. Bohm. Gesellschaft (6), B'1 2. 1882
Ce petit Rhizopode n'est pas rare en general dans les mousses et les Sphagnum ;
mais on ne le reconnait que dimcilement, par suite de sa ressemblance presque parfaite
avec une vari^te bryophile de Trinema enchelys. Dans le Corythion, la coquille est
formee d'ecailles toutes petites, de 3 a 4 /* seulement de longueur, elliptiques ou
vaguement rectangulaires avec angles arrondis, et disposees sans beaucoup d'ordre les
unes a cote des autres ; dans le Trinema, les ecailles sont beaucoup plus grandes,
rondes, et imbriquees dans un ordre g^ometriquement regulier ; mais comme, dans
ces deux formes, les elements siliceux ne sont que tres-difficilement visibles un a un,
la distinction n'est pas toujours aise"e.
A 1'lle Macquarie, comme aussi au Mont Cook, cette espece revetait une forme
speciale (Fig. 3«), tres-large, arrondie en un cercle presque parfait, et la taille, de 62
a 63 M de longueur, etait de beaucoup superieure a celle du type habituel, laquelle ne
mesure guere que 35 a 40 M.
A Ngaurnhoe, a 1'ile Stewart, a Oahu, au Queensland, on trouvait une variete plus
curieuse encore (Fig. 3b) ; 1'enveloppe, de 40 a 50 n de longueur, tres-large, a forte
ouverture buccale, se montrait revetue sur tout son pourtour (sauf a la bouche) d'une
rangee reguliere d'aiguillons aceres. Ces aiguilles, du reste, n'^taient pas siliceuses
au meme titre que les ecailles de la coquille ; elles etaient composees de cette meme
matiere chitino'ide, incolore, qui cimente les ecailles entre elles, et se dissolvaient
immediatement dans 1'acide sulfurique concentre bouillant.
C'est la, me semble-t-il, une varie'te fixee, et que Ton pourrait appeler var.
spicata.
Cyphoderia ampulla. EHRENBERG
Monatsberichte K. Akad., Berlin, p. 199. 1840
Cette espece n'est pas muscicole, mais on la rencontre quelquefois dans les
Sphagnum. Au Mont Cook, il s'en est trouve* deux exemplaires, typiques, et a dessins
230 E. PENARD
ardolaires relativement tres-grands, 3 M environ ; puis a Vancouver, une seule coquille
vide ; dans ces deux recoltes, il faut le remarquer, il y avait du Sphagnum. A Oahu,
par centre, ou s'est egalement montre'e une coquille vide, le sphagnum manquait ; mais
on y voyait de belles mousses bien fournies, 1'une d'elles tres-claire, longue et soyeuse,
et dans ces conditions-Ik les mousses renferment quelquefois des organismes sphagno-
philes en general.
Difflugia arcula. LEIDY
Fresh w. Rhiz. N. America, p. 116. 1879
Cette difflugie, — qui n'est pas une difflugie en realite, mais un Rhizopode
tres-voisin des genres Bullinula et Diplochlamys,1 — est 1'une des especes les plus
caracteristiques des Sphagnum et des mousses, et s'est rencontree dans la plupart des
recoltes de Mr. Murray. Tout en restant toujours bien typique dans son apparence
generale, elle varie passablement, d'une localite a une autre et souvent aussi dans une
meme localite, sous le rapport de la taille, de la hauteur comparee a la largeur, des
elements qui recouvrent 1'enveloppe. Quelquefois aussi, dans une meme station, on
trouve deux formes 1'une a cote de 1'autre, une grande et une petite, que ne relie
aucune transition. Tel a ete le cas a Oahu, a 1'ile Macquarie, comme aussi an
Queensland.
Dans plusieurs des recoltes, les animaux se sont montrds vivauts. Ce rhizopode,
en effet, est done" d'une resistance tout-a-fait remarquable ; mais jamais aucun individu
n'a ddploye de pseudopode. Le contraire, du reste, eut ete" tres-dtonnant, car dans
cette espece, les pseudopodes, quelquefois mentionne's par suite d'une confusion avec
un autre organisme (Centropyxis laevigata), n'ont jamais ete vus en re"alite, et parais-
sent decide"ment manquer.
Difflugia bacilli/era. PENARD
Me"m. Soc. Phys. Hist. Nat., Geneve, vol. 31, p. 146. 1890
Seulement trois coquilles vides, a Vancouver. Cette espece est tres-rare, et peu
caracteristique ; peut-6tre eut-il mieux valu, des 1'origine, 1'envisager simplement
comme une varie"te" de Diff. pyriformis.
1 Votr un des prochains numeros de la Revue Suisse de Zoologie, tome 19, 1911.
RHIZOPODES D'EAU DOUCE 231
Difflugia constricta. EHRENBERG
Abh. Kon. Akad. Wiss., Berlin, p. 410. 1841
C'est la le rhizopode le plus constant dans les mousses, et qui, en fait, s'est
montre dans toutes les stations (il manque dans 1'Antarctique proprement dit, ce
qui n'est pas sans etonner). II est toujours et parfout represente par de nombreuses
varietes, petites ou grandes, longues ou larges, plus ou moins comprime'es, etc.,
et tout cela s'est trouve" dans les recoltes de Mr. Murray.
Difflugia fallax. PENARD
M(kn. Soc. Phys. Hist. Nat., Geneve, vol. 31, p. 144. 1890
Cette petite espece n'a ete rencontree que dans les Alpes australiennes, puis a
Sydney, sous la forme de rares coquilles vides. Elle est toujours difficile a deter-
miner, et vu le mauvais etat des exemplaires examines, peut-etre faudrait-il ne
1'indiquer ici que suivie d'un point d'interrogation.
Difflugia lucida. PENARD
M6m. Soc. Phys. Hist. Nat., Geneve, vol. 31, p. 145. 1890
Ce rhizopode est de faible taille, peu apparent, et passe facilement inapercu ;
mais grace a sa forme special e et a 1'aplatissement considerable de la coquille, on
le reconnait facilement. Dans la plupart des recoltes, il s'est montre rare, repre"-
sente par quelques coquilles vides seulement ; a Katoomba, dans la mousse comme
aussi dans le sphagnum, les exemplaires en etaient plus nombreux, et souvent
vivants, mais alors enkystds.
Difflugia manicata. PENARD
Faune Rhizopodique Le"man, p. 226. 1902
Quelques individus seulement, dans le Sphagnum du Mont Cook ; ovoides, de
90 M de longueur, peu typiques de forme ; mais grace a leur couronne de grosses
particules quartzeuses en arriere du peristome, on pouvait les identifier sans difficulte.
E. PENARD
Difflugia pyriformis. PERTY, var. bryophila. PENARD
PERTY. Zur Kenntniss Kleinstcr Lebensformen, p. 187. 1851
PENARD. Faune Rhizop. Leman, p. 221. 1902
La Diff. pyriformis, le plus corumun peut-etre de tous les rhizopodes d'eau douce
en general, manque, a 1'etat typique, dans les mousses (sauf, bien entendu, dans
les mousses submergees, qui peuvent heberger toute la faune habituelle des etangs) ;
elle y est remplacde, cependant, par une variete plus petite, var. bryophila, pas tres
commune en general, et qui ne s'est rencontree que dans quelques-unes des recoltes.
Diplochlamys fragilis. PENARD
Arch, fur Protistenkunde, B'1 17, p. 272. 1909
Parmi les differentes especes qui composent le genre Diplochlamys, il en est
deux, D. fragilis et D. Leidyi, qui sont fort rapprochees 1'une de 1'autre, et ne
peuvent guere etre distinguees autrement qu'a 1'etat vivant ; dans les recoltes de
Mr. Murray, les rares individus rencontres etaient ou morts, ou enkystes, et si la
determination du genre est certaine, peut-etre faudrait-il mettre un point d'interro-
gation a cote du nom de 1'espece.
(?) Diplochlamys gruberi. PENARD
Arch, fiir Protistenk., Bd 17, p. 282. 1909
Les restrictions qui viennent d'etre faites a propos de la Diplochlamys fragilis
sont peut-etre plus ne"cessaires encore dans le cas actuel. Un representant de ce
genre, qui semblait ne pouvoir 6tre autre chose que D. gruberi, s'est montre* dans
plusieurs des recoltes, mais c'est seulement dans les mousses d'Oahu que j'ai cru
pouvoir m'assurer d'une determination a peu pres certaine.
Diplochlamys timida. PENARD
Arch, fiir Protistenk., Bd 17, p. 275. 1909
Cette espece, de taille plus faible que les prece"dentes, et qui s'en distingue
en outre par des caracteres assez prdcis (noyau, enveloppe interne, revetement
externe), a pu, au contraire de ces deux dernieres, dtre de'termine'e avec certitude.
A Ottawa, au Queensland, on la trouvait vivante, mais retractee dans son
enveloppe.
RHIZOPODES D'EAU DOUCE 233
Diplochlamys vestita. PENAED
Amoeba vestita. PENARD. M<kn. Soc. Phys. Hist. Nat., Genkve, vol. 31. 1890
Diploch. vestita. PENARD. Arch, fur Protistenk., Bd 17, p. 279. 1909
Beaucoup plus grande que toutes les especes precedentes, dont elle differe
egalement par la nature de son enveloppe et par la presence de noyaux tres
nombreux et tres-petits, la Dipl. vestita s'est montree a Vancouver et a Katoomba.
Dans cette d era i ere localite, cependant, le seul exemplaire rencontre etait en trop
mauvais etat pour qu'il fut possible de 1'identifier avec certitude.
Euglypha alveolata. DUJARDIN
Hist. Nat. Zooph. infusoires, p. 252. 1841
1-^ Euglypha alveolata ne se rencontre guere dans les mousses, mais par centre elle
est assez frequente dans les Sphagnum. C'est bien dans le sphagnum aussi qu'elle
a ele trouvee, au Mont Cook, ; Vancouver, dans les Alpes australiennes ; a File
Macquarie, par centre, le sphagnum manquait, mais on y voyait une toufFe d'une
mousse claire, longue et tres foumie, laquelle, comme nous 1'avons deja vu, a fourni
plusieurs organismes sphagnicoles en general.
Dans les trois premieres de ces stations, 1'espece etait representee par cette varidtd,
normalement prolongee en arriere de quatre longues Opines, que Leidy, par exemple,
figure dans la PI. 35 (Fig. 15 a 17) de son grand ouvrage, et qui & 1'heure actuelle est
gdneralement indiqude comme Euglypha brachiata (voir page suivante, remarques &
propos de cette espece). Mais cette forme elle-meme est assez variable quant au
nombre des epines qui la caracterisent. Dans les Alpes australiennes, comme aussi
au Mont Cook, la plus grande partie des coquilles, reconnaissables pourtant (grace a
leur forme, aux denticulations buccales, etc.) comme devant se rattacher a cette variete
spdciale, etaient lisses, sans dpines. A Vancouver, on trouvait une seule aiguille, sur
le cote, ou bien deux, ou trois, ou bien encore 1'une des aiguilles e"tait terminale, pro-
longeant d'une longue e"pine le fond de la coque.
Quant a YEuylypha alveolata de 1'ile Macquarie, c'etait une varietd speciale, de 75
a 105 M de longueur, a coquille l^gerement comprimee (a section transversale elliptique),
ornee de dessins areolaires largement marques, et munie a la bouche d'ecailles pourvues
en avant d'une ligne de denticulations tres-petites et serrees.
Euglypha brachiata. LEIDY
E. brachiata. LEIDY. Freshw. Rhiz. N. Amer., p. 220. 1879
non PENARD. Faune Rhizopodique Leman, p. 504. 1902
Peu de Rhizopodes sont aussi variables — en apparence au moins — que ceux qui se
rapportent au genre Euglypha ; et pour cette raison meme, il est peu de genres dans
234 E. PENARD
lesquels on ait reuni sous une meme denomination spdcifique une aussi grande quantite
de formes differentes, et dont plusieurs, sans aucun doute, representent en realitd des
especes bien autonomes.
Mais il est aussi, dans ce genre Eugtypha, des especes bien nettement distinctes,
faciles h, caracteriser, et avant toutes les autres on peut citer sous ce rapport VEuglypha
brachiata, sous sa forme typique, celle que Leidy figure a la PI. 37 de son grand ouvrage
(Fig. 5 a 10). Et pourtant, elle aussi reste encore meconnue ; sous le nom specifique
de brachiata, on s'accorde en general aujourd'hui a reunir toutes les formes dont les
epines, au lieu de constituer chacune un element distinct en soi, ne sont qne le pro-
longement effile d'une des dcailles ordinaires et regulierement imbriquees qui con-
stituent 1'ensemble de la coquille.
Cette generalisation, comprehensible dans 1'etat actuel de nos connaissances, est
regrettable ; et quant a Euglypha brachiata, il y a Ik une grave erreur ; re"parer. Parmi
les naturalistes qui ont cite cette espece, bien pen, en realite, me semblent avoir eu sous
les yeux la forme type de LEIDY ; aucun meme peut-etre, a la seule exception de G. S.
WEST/ qui, lui, 1'a e"videmment trouvee (Capel Curig, N. Wales, et small lakes East of
Recess, W. Ireland), et qui dit a cette occasion : " This is another rare species of the
genus, which I have only obtained twice, and in both instances from submerged
sphagnum." Pour mon compte, ce n'est que tout recemment, dans les sphagnum de
Vancouver, que j'ai pu voir cette espece telle qu'il faut la comprendre aujourd'hui ; il
s'en est tout-a-coup montre un exemplaire, une coquille vide en parfait e"tat, puis, apres
bien des recherches, j'ai fini par en trouver un second.
Comme la description de LEIDY, bien que parfaitement exacte, est un peu ecourt(5e,
il ne sera pas inutile de donner ici quelques renseignements suppldmentaires sur cette
Euglypha, peut-etre la plus elegante dans ses formes de toutes celles que Ton connait
actuellement (Fig. 4, a et &).
La coquille, allongee, legerement pointue en arriere, s'elargit de Ik graduellement
pour acquerir son plus grand diametre un peu en arriere du milieu de la longueur, puis
s'inflechit vers 1' avant, en ddcrivant une Idgere courbe rentrante, et vient se terminer en
une bouche relativement dtroite, formee de 7 ecailles (7 en tout cas dans 1'individu
particulierement examine).
Comme toujours dans le genre Euglypha, ces ecailles de la bouche sont denticulees
en avant, et ici, la disposition de ces dents est assez curieuse : une large dent mediane,
presque triangulaire, puis de chaque cote une dent plus petite, allongee en forme de
d^ k coudre, puis une plus petite encore, et enfin 1'ebauche d'une troisieme.
En arriere de cette premiere rangee buccale, la seconde ligne d'dcailles porte
encore des denticulations, puis viennent les ecailles lisses, ovales, tres-belles, qui se
continuent sur toute la coque, et, en rapport avec la forme meme de cette derniere, plus
grandes sur les parties renHees, plus petites sur les parties etroites, meme rondes au
fond de la coquille, de maniere a ne jamais violenter la belle imbrication reguliere.
1 Linnsean Society's Journal, Zool., vol. 28, p. 328, 1901.
RIIIZOPODES D'EAU DOUCE 235
Quant aux longs "bras" caracteristiques de cette espece, et que LEIDY indiqne
comme etant au nombre de 2 ou de 4, plus rarement de 6, 1'un des individus que j'ai
pu examiner en portait 2, 1'autre 4. Ce ne sont la pas autre chose que des ecailles,
tres-fortement modifies ; par leur base, elles s'inserent entre les ecailles ordinaires, et
font partie du revetement general ; puis elles s'elancent au dehors, et par une large
courbe elles vont s'epanouir en am ere, cornme deux ailes ou deux longs bois de renne.
Vus de cdte, ces appendices se presentent ccnmrfle de simples aiguilles recourbees
en alene ; mais si Ton s'arrange a les voir d'en haut, par exemple, on constate que ce
sont des lames, larges, minces, et, comme j'ai pu m'en assurer sur un " bras " casse,
creusdes en gouttiere l (Fig. 46).
Ces longs appendices fonctionnant comme des ancres, doivent etre d'une reelle
utilite pour retenir 1' animal entre les feuilles des Sphagnum; peut-etre m^me est-ce pour
cette raison que dans les localites oil on le trouve on a peine a en obtenir quelques rares
individus ; mais dans d'autres circonstances, ces ramures doivent ge'ner considerable-
ment 1'animal, et Ton peut se demander si cette espece si peu repandue n'est pas en
train de disparaitre, comme out disparu d'autres animaux plus eleves en organisation,
par suite d'une trop grande complication de leurs organes a 1'origine protecteurs.
Euglypha ciliata. EHRENBERG
Monatsber. K. Akad. Wiss., Berlin, p. a 79. 1848
Cette espece est toujours commune dans les mousses, et de fait, elle a ete vue
un peu partout. A Ngauruhoe, it 1'ile Macquarie, dans les Alpes australiennes, on
en trouvait une variete tres-hirsute, a epines fortes et nombreuses, disposees en
desordre les unes dans les autres.
Euglyplia compressa. CARTER
Ann. Mag. Nat. Hist., vol. 13, p. 32. 1864
Commune en general dans les mousses et surtout dans les Sphagnum, cette
espece n'a manque nulle part, sauf a Fiji, oil les circonstances, nous 1'avons vu,
etaient un peu particulieres. Dans plusieurs des recoltes (He Stewart, M* Cook,
Sydney), on la rencontrait sous deux formes, une grande et une petite, sans qu'il
apparut de transitions. A File Stewart, presque tons les exemplaires se rattachant
a la grande forme etaient depourvus d'aiguilles ; au Mont Cook, 3a grande forme,
tres-belle, rappelait d'assez pros la Placocysta spinosa, avec laquelle du reste cette
espece a ete souvent confondue. A Sydney (National Park), Ton trouvait une
varidte superbe, de 145 /u de longueur.
1 LEIDY ecrit Ji ce sujet : " Rarely, I have seen them straight, with thickened ends and oar-like, as seen
in Fig. 7."
BRIT. ANTARCT. EXPED. 1907-9. VOL. T E
236 E. PENAHD
Euglypha cristata. LEIDY
Proc. Acad. Nat. Sci., Philad., p. 226. 1874
non AWERINZEW. Rhizopodienstudien, Ann. de Biol. Lacustre, torn. 1, p. 323. 1906
Cette jolie petite espece, toujours facile a reconuaitre, ne s'est rencontree que
dans une senle des recoltes, dans les Sphagnum de Katoomba, Mes Bleues, et cela
sous la forme d'une seule coquille vide, parfaitement typique d'ailleurs.
Euglypha fill/era. PENARD
M&n. Soc. Phys. Hist. Nat., Geneve, vol. 31, p. 179. 1890
Ce petit rhizopode, characterise surtout par ses aiguilles fines, longues et tres-
droites, pen nombreuses, disposees de distance en distance sur 1'arete de la coquille,
s'est montre dans la seule station de Creel, Alpes australiennes.
Euglypha lacvis. PERTY
Mittheil. Naturf. Ges., Bern., p. 163. 1849
On rencontre pour ainsi dire toujours, dans les mousses, toute une serie
d' 'Euglypha tres-petites, lisses, a dessins alveolaires a peine reconnaissables, et que,
faute de mieux, on s'accorde generalement a considerer comme representant V Euglypha
laevis de Perty.
Tel a etc" le cas dans les recoltes de Mr. Murray. Dans plusieurs stations, on
voyait egalement, k cote de la forme habituelle, cette variete extremement petite
(36 M) que j 'avals decrite en 1890 * comme var. minor.
/
Euglypha strigosa. LEIDY
Proc. Acad. Nat. Sci., Philad., p. 172. 1878
Difficile a distinguer de V Euglypha ciliata dont elle differe par un revetement
d'aiguilles tres-courtes et tres-serrees, comme aussi par sa forme un pen speciale
et la structure particuliere du peristome,2 V Euglypha strigosa s'est montree dans
quelques-unes seulement des recoltes. Dans les Alpes australiennes, la coquille etait
pourvue d'aiguilles relativement tres-fortes et tres-serrees. A Vancouver, dans la
mousse lavee depuis trois jours, il s'en est rencontre un individu vivant, qui courait
gaiement au milieu des debris, avec pseudopodes deployes.
1 Mem. Soc. Phys. Hist. Nat., Geneve, vol. 31, p. 182.
2 Penard, Faune Rhizopodique Lenian, p. 503.
RHIZOPODES D'EAU DOUCE 237
Heleopera petricola. LEIDY
Freshw. Rhiz. North Am, p. 165. 1879
Cette espece s'est rencontree dans un nombre relativement restreint de stations,
sous la forme de coquilles vides ou d'individus enkystes, et souvent (Australian Alps,
Mt. Cook, Vancouver) avec cette teinte rosee que- prend volontiers la coquille. A
Suva (Fiji) comme aussi a Sydney (National Park), c'e"tait une petite variete incolore,
qui peut-etre aurait droit a une denomination specifique particuliere.
Heleopera sordida. PENARD
Revue Suisse de Zoologie, torn. 18, fasc. 3. 1910
Cette Heleopera, a coquille petite, large et renflee, d'un jaune sale, et dont la
description toute recente a ete donnee d'apres des echantillons re"coltes en 1909 dans
les tourbieres du Jura suisse, ou les individus ont e"te etudies vivants et en etat
d'activite, s'est rencontree dans les Sphagnum de Vancouver, puis au Pare National
de Sydney, mais toujours a 1'etat de coquilles vides.
Heleopera sphagni. LEIDY
Difflugia (Nebela) sphagni. LEIDY. Proc. Acad. Nat. Sci., Philad., p. 157. 1874
Heleopera picta. LEIDY. Freshw. Rhiz. N. Amer., p. 162. 1879
Heleopera sphagni. CASH et HOPKINSON. Brit. Freshw. Rhiz., p. 143. 1909
Cette grande et belle espece s'est montree dans cinq des recoltes, toutes, il faut
le remarquer, de 1'hemisphere sud. Elle etait partout bien reconnaissable, mais
partout aussi un pen differente du type figure par LEIDY, et tel que, pour mon
compte, je 1'ai e'galement trouve en Europe.
La coquille, jaunatre pale, relativement tres-elargie, portait ici des dessins
areolaires extremement petits, tels qu'on n'est guere habitue" a les voir. Elle variait,
suivant les exemplaires, entre 92 et 110 M. A File Macquarie, c'etait une forme
tres-belle, de 130 a 140 M en longueur ; souvent les individus se trouvaient a 1'etat
enkyste, et Ton pouvait reconnaitre au plasma une teinte verdatre qui semblait
montrer que la aussi la symbiose est un phdnomene ordinaire.
Heleopera sylvatica. PENARD
M<5m. Soc. Phys. Hist. Nat., Gen., vol. 31, p. 168. 1890
Caracteristique des mousses proprement dites, ce rhizopode manque le plus
souvent dans le Sphagnum. Distinct des autres especes du genre par sa taille tres-
238 E. PENARD
faible (60 M en general), par sa transparence, 1'imbrication de ses ecailles, comme par
d'autres caracteres moins e"vidents, il s'est montre bien normal dans les differentes
stations ou il a 6te recueilli. A Oahu, cependant, la taille etait relativement
tres-faible.
Get organisme resiste bien a la dessication, et s'enkyste volontiers ; c'est sous
cette forme vivante, mais enkystee, qu'il a etc souvent trouve".
Hyalosphenia cockayni, spec, nova
Fig. 5, a et &
En m'envoyant quelques petits fragments de mousses recueillis par le Dr.
Cockayne, botaniste du Gouv. de Nile. Zelande, sur le tronc d'un arbre aux iles
Auckland, Mr. Murray attirait mon attention sur un Ilhizopode qu'il croyait nouveau.
Ce devait 6tre la, pensait-il, une Nebela ; mais a mon avis, cet orgauisme doit plutot
rentrer dans le genre Hyalosphenia, dont il serait meme un representant assez
typique.
Entre ces deux genres, du reste, il n'existe pas de differences bien sensibles ; on
s'accorde a considerer comme Hyalosphenia les formes dont la coquille est ou parait
etre lisse, reservant le nom de Nebela pour celles dont 1'enveloppe est revetue d'ecailles
ou de plaques. Mais dans le fait, cette distinction est assez subtile, et dans la plupart
des Hyalosphenia, on distingue souvent, chez certains individus ou dans certaines
localites, des reticulations dues a la presence d'elements siliceux fort petits, noyes dans
1'epaisseur de 1'enveloppe.1
Dans la H. cockayni 1'enveloppe, d'un jaune chamois tres-clair, transparente,
comprimee, a peu pros pyriforme d'apparence, et une fois et demi aussi longue que
large, se montre tout d'abord lisse et unie ; mais en 1'examinant avec attention, on
finit par y reconnaitre, dans tous les individus mais sur les uns plus distinctement
que sur les autres, la presence d'elements siliceux, generalement tres-petits (quelques M),
ronds ou plus souvent ovales, et se touchant les uns les autres sur toute la surface de
1'enveloppe. Dans plusieurs exemplaires, j'ai trouve quelques-uns de ces elements
siliceux qui n'etaient pas autre chose que les ecailles buccales d'une Euglypha, ct
meme, grace a la structure et a la disposition des denticulations caracteristiques, on
reconnaissait E. compressa.2
La structure de notre H. cockayni est done a peu pres identique a celle de
H. papilio ; mais la forme est toute differente : les bords lateraux de la coquille, au
1 Daus //. tincta (Nebela tincta, AWERINZEW), on peut trouver des stations oil tons les individus
montreront une enveloppe parfaitement lisse a premiere appareuce, tandis que dans d'autres locality's, plus
nombreuses, cette euveloppe se verra tout entiere recouverte d'ecailles bien nettement definies.
2 C'est pour cette raison, meme, que j'ai porte sur la liste des especes recoltees cette Euylypha qui en
rdalite n'y a pas etc" trouvee en tant qu'individu ; mais il fallait bien qu'elle existat, puisque notre Hyalo-
sphenia en avait amasse les tScailles.
RHIZOPODES D'EAU DOUCE 239
lieu d'aller tout droit et d'une seule venue jusqu'a 1'extremite, se creusent brusquement,
a gauche et a droite, et vers le tiers anterieur de 1'enveloppe, d'une encoche, d'un arc
rentrant, et ces deux encoches droite et gauche donnent & toute cette enveloppe une
apparence assez elegante, qui rappellerait celle d'un violon.
En avant, 1'ouverture buccale, plus etroite que dans H. papilio, est un pen renflee
en levre, arquee sur la face large, creuse"e sur la face etroite.
Sur les cotes de cette partie anterieure retreeie, qu'on pourrait appeler le tube
buccal, on voit souvent une ou meme deux perforations ou pores, mais qui n'existent
pas necessairement. En arriere, on ne distingue aucun pore, rien d'analogue a ce qui
existe chez H. papilio.
Tons les exemplaires rencontres 1'ont ete soit a 1'etat de coquilles vides, soit
enkystes, le plasma figurant une masse ovo'ide. D'apres la couleur et 1'apparence de
ce plasma, il semblerait qu'a 1'etat de vie active on trouve, ici aussi, normalement des
phenomenes de symbiose.
C'est, comme nous venous de le voir, aux lies Auckland que cette espece a ete
decouverte tout d'abord ; plus tard, elle s'est retrouvee dans les Sphagnum du Mont
Cook, representee par une seule coquille vide, tres-grande, de 115 M; enfin, dans les
environs de Sydney, elle s'est revue encore, pas tres-rare ; les exemplaires mesuraient
100 M environ.
Aux iles Auckland, la longueur variait entre 89 et 100 M, pour un largeur de
45 a 55 n. L'enveloppe est assez fortement comprimee, et son epaisseur arrive a peine
a la moitie de sa largeur.
Hyalosphenia sub/lava. CASH
Brit. Fresh w. Rhiz. and Helioz., vol. 2, p. 87. 1909
Fig. G, a, b, et c
Dans les mousses provenant d'Eumundi, Queensland, et plus specialement dans la
touffe de sphagnum qui s'y trouvait jointe, s'est rencontrde, avec une abondance
relative, cette petite Hyalosphenia, ddcrite tout demierement, comme provenant soit
des tourbieres d'Irlam Moss pres de Manchester, soit des monts Pentland (Mr. Evans)
en Ecosse.
Bien que le Rhizopode trouve au Queensland puisse etre assimile sans trop de
peine a celui des lies Britanniques, il n'en est pas moins vrai que Ton y constatait des
differences de detail assez sensibles. L'enveloppe, plus petite (59 M au maximum, et
generalement 53 M, en regard de 65 a 70 /« qu'indique CASH), relativement plus large,
un pen moins comprimee, etait termine"e par une ouverture buccale assez petite, ovale
ou presque ronde, pourvue d'une sorte de bordure ou levre tres courte et a peine
indiquee, qu'entourait a son tour un petit cercle de grains faiblement marques ; cette
levre ne portait pas trace d'encoche sur les cotes, et aucun exemplaire n'a montre le
pore terminal posterieur dont parle CASH (mais qu'il ne figure pas).
240 E. PENARD
L'enveloppe, jaunatre, semblait parfaitement lisse a premiere apparence, mais un
examen attentif y decelait la presence de granulations serrees, ou elements siliceux
tres-petits noye"s dans la masse chitineuse.
Presque tous les exemplaires n'ont ete trouves qu'a 1'dtat d'enveloppes vides ;
quelquefois, un reste de plasma, inerte et en mauvais etat, laissait supposer qu'il y
avait peut-etre eu la des zoochlorelles symbiotiques.
Lesquereusia . . . spec.
Les elements siliceux qui forment la coquille dans les Lesquereusia sont tellement
caracteristiques, que je n'ai aucune hesitation a indiquer la presence de ce genre dans
le sphagnum de Vancouver, bien qu'il n'ait e"te trouve qu'un tout petit fragment de
coquille. Meme, d'apres la forme trapue et courte des quelques ecailles siliceuses
vermiculaires qui constituaient ce fragment, je croirais volontiers que c'etait la
Lesquereusia epistomium, PENARD. Mais ce serait la en tout cas une identification un
peu aventureuse.
Licberkiihnia wageneri. CLAP. ET LACIIMANN
Mum. Inst. Nat., Geneve, vol. 6, p. 465. 1859
Apres avoir observe, a Vancouver comme dans les Alpes australiennes, des
organismes inertes et peu reconnaissables que je croyais pourtant devoir rapporter a
cette espece, j'ai fini par constater sur quelques individus la presence de ce long
" pedoncule " interne, caracteristique du genre, et qui ne peut montrer autre chose
qu'une Lieberkithnia. La taille, la presence de petits globules en grand nombre qui
sans doute reprdsentaient les noyaux, indiquaient la Lieb. ivageneri, et non pas
L. paludosa qui ne possede qu'un seul noyau tres-gros, ou bien quelquefois en
renferme plusieurs, mais encore volumineux et en nombre tres-restreint.
Nebela bigibbosa. PENARD
Mem. Soc. Phys. Hist. Nat., Geneve, vol. 31, p. 161. 1890
Cette curieuse Nebela, si typique avec ses deux brides laterales invaginees, et qui
fut trouve'e pour la premiere fois aux environs de Wiesbaden en 1890, puis en 1902
dans les mousses du Spitzberg, en 1903 dans les forests du Valais, enfin en 1909 a la
montagne des Voirons pros de Geneve, s'est encore revue dans les collections Murray,
a Vancouver et a Victoria ; rare d'ailleurs et representee par des coquilles vides, mais
bien uettement caracterisees.
KHIZOPODES D'EAU DOUCE 241
Nebela caudata. LEIDY
Proc. Acad. Nat. Sci., Philad., p. 58. 1876
Cette espece rare, decouverte par Leidy en 1876 dans les tourbieres du New
Jersey, et qui, a ma connaissance n'a etc trouvee en Europe qu'une seule fois, dans le
Loch Ness en Ecosse,1 semble etre beaucoup plus commune dans I'hemisphere sud.2
Les recoltes de Mr. Murray 1'ont montree dans une demi-douzaine de localites, soit
a 1'etat de coquilles vides, soit enkystee, le plasma figurant une masse spherique
protegee par un epiphragme.
Ce qu'il y a de plus caracteristique dans cette espece, ce sont ses cornes, ou
prolongements creux, au nombre de 3, 4, 5, qui terminent la coquille en arriere. Mais
ces cornes sont bien souvent rudimentaires, a moitie formees, ou bien manquent en
partie, quelquefois meme tout-a-fait.
Au Queensland, la forme etait normale, et presque toujours les individus portaient
les prolongements caracteristiques, le plus souvent au nombre de 4 ou 5. A Ngauruhoe,
les individus montraient le plus gdneralement une ou deux cornes seulement, et plus ou
moins rudimentaires ; a 1'ile Macquarie, on trouvait toutes les transitions possibles
entre un simple rudiment d'une seule corne et des prolongements normaux. A
Victoria, B.C., a Vancouver, presque tons les individus dtaient lisses, depourvus de
prolongements quelconques, et persoune, a premiere vue, n'aurait hesite- a considerer
ces coquilles comme autre chose que ce qu'elles etaient vraiment ; il y avait la, en
apparence, une simple variete de Nebela dentistoma.
Nebela certesi. CERTES, spec, nova
Neb. collaris. CERTES. Var. Cap Horn, a et b. Mission scientif.
Cap Horn, torn. 6, Zool., p. 14
Fig. 7, a et b
En meme temps qu'il decrivait sa curieuse Nebela martiali, CERTES designait
sous les noms de Neb. collaris, var. o Cap Horn et Neb. collaris, var. b Cap Horn,
deux rhizopodes d'une forme et d'une apparence speciale, et caracterises par la
presence d'un " sillon " qui "part de 1'ouverture et va en s'attenuant jusqu'a la
moitie de la longueur du col, orne de perles irregulierernent disposees de chaque
cote du sillon au nombre de 3 a 7 et meme plus." La variete b se distinguait en
outre par la presence de " quatre ouvertures triangulaires symetriquement disposees,"
et situdes a la base du col.
1 J MURRAY, in Proc. Koy. Soc., Edinburgh, 1904-5, vol. 25, p. 6H.
2 Elle s'est retrouvue dans les mousses rapporttes par le Dr. Bruce de Gough Island, Ocean atlantiquc
Sud, an cours du voyage de la Scotia.
242 E. PENARD
II n'est pas difficile de reconnaitre la unc Nebela qui s'est rencontrde d'abord a
Katoomba (Montagues Bleues), puis aux environs de Sydney, partout tres-rare et
reprdsentde par des euveloppes vides et souvent en mauvais dtat, mais suffisantes
pour une determination exacte.
II est dtrange, a vrai dire, que CERTES n'ait cru devoir considerer ce lihizopode
que comrae une varietd de Nebela collaris, clout il s'dloigne en rdalite de tres-loin ;
mais, si j'en puis juger d'apres quelques-unes des figures doimees par 1'observateur
francais, il est probable que certaiues coquilles malformdes et en mauvais dtat
lui auront paru constituer des formes de transition qui Fauront induit en erreur.
En meme temps, ces deux varidtes a et b de CERTES ne concernent bien
certainement qu'un meme organisme, car les quatre " ouvertures triangulaires "
existent toujours en principe, mais souvent si rudimentaires et si inddcises a la vue
qu'on a peine a les trouver.
Pour moi, c'est la une espece tout-a-fait caractdristique, et qui, me semble-t-il,
doit porter le nom du consciencieux savant francais.
La Nebela ccrtesi est une espece d'assez grande taille, ou plutot, faudrait-il dire,
d'une taille assez allongee, mesurant 145 a 150 M en general; mais la coquille, fine,
mince, etroite et delicate, en fait un organisme assez chdtif en apparence, et qui
passe facilement inapercu.
Elle a la forme d'une bouteille, etroite, au col allonge, et, assez fortement
comprimde, elle prdsente une face large et une face etroite. Cette enveloppe est
jaunatre. tres-claire, formee d'une matiere chitinoi'de dans laquellc sont empatds
des elements siliceux ovales ou arrondis, tres-petits, ou melds de plus gros. La
bouche est bordde d'un epaississement, ou levre saillante et arquee en avant sur
la face large, creusee d'une faible gouttiere sur la face etroite. Mais ce qu'il 'y a
de plus curieux dans cette espece, c'est la presence d'un dtroit sillon, qui, partant
du peristome et au milieu de chacune des levres, descend tout droit le long du col,
pour disparaitre a la moitid de la longueur de ce col.
C'est la un veritable canal, creusd dans 1'epaisseur de la paroi, et bordd par
centre, sur 1'un de ses cotes, d'un epaississement qui simule un batonnet; aussi,
sur des coquilles brisdes, trouve-t-on quelquefois que la ddchirure s'est faite juste
le long du sillon, c.a.d. sur la ligne de moindre rdsistance.
A droite et a gauche du canal, on remarque egalement toujours deux ou
plusieurs lignes de petites perles brillantes.1
Plus bas, en arriere du col, on voit, a gauche et a droite, et sur chacune des
faces, une petite ouverture, irrdguliere de contour, mais si faible qu'elle ne se
distingue guere que sur des coquilles et bon dtat. Ce ne sont pas la, du reste, de
simples perforations, mais 1'indice d'une sorte dc tube, plus ou moins rudimentaire,
qui traverse la coquille de part en part.
1 Dans le bauuic du Canada, cos perles no so distinguent presque plus, et le sillou buccal lui-mt'me
devient peu visible.
RI-IIZOPODES D'EAU DOUCE 243
Aucun des individns rencontres ne 1'a ete autrement qu'a- 1'etat de coquille vide ;
ou bien aussi Ton trouvait qnelques restes de plasma mort, a 1'abri d'un diaphragme
epais et feuillete, tel qu'on le connait dans les Nebelides en general.
Nebela collaris. EHRENBERG
Diffliu/ia collaris. EHRBG. Monatsb. Akad._Wiss., Berlin, p. 218. 1848
Nebela collaris. LEIDY. Proc. Acad. Sci., Philad., p. 162. 1879
Cette espece est fort variable, et souvent difficile a determiner. En general,
on ne la trouve guere repre'sente'e dans les mousses proprement dites que par de
petites formes, dont certaines (Neb. minor, parvula] ont dte" elevees au rang
d'especes ; et dans les rdcoltes de Mr. Murray, la forme type s'est montree tres-rare ;
on ne la trouvait guere qu'au Mont Cook. Dans les Montagnes Rocheuses, comme
au Queensland, c'etaient de petites varie'tes ; a Ngauruhoe, a Vancouver, la taille
dtait forte, et la forme rappelait plutot la Neb. bohemica de Taranek, dont les
caracteres ne sont peut-etre ni assez prdcis ni assez constants pour autoriser une
separation d'avec le type. A Waiata-Rua, une varie'te tres -petite montrait ce
meme type bohemica, tandis qu'a Oahu Ton avait affaire a une forme tres-large,
discoide, qu'ou eut pu sans trop de difficulte rapporter a Nebela tincta.
Nebela dentistoma. PENARD
Mem. Soc. Phys. Hist. Nat., Geneve, vol. 31, p. 162. 1890
Nebela crenulata. CASH. Tran. Manch. Micr. Soc., p. 50. 1891 (1892)
A'-sez peu repandue en general, cette Nebela ne s'est guere trouvee qu'au Mont
Cook et a Vancouver, dans les Sphagnum ; puis a 1'lle Macquarie, ou, nous 1'avons
deja vu, dans une grande mousse claire et serree, plusieurs especes se sont rencontre'es
qui sont sphagnicoles en general.1
Nebela flabellulum. LEIDY
Proc. Acad. Nat. Sci., Philad., p. 157. 1874
Cette espece sera toujours difficile a determiner exactement, par le fait que
plusieurs Nebelides (Nebela collaris, galeata, tincta, &c.) ont une tendance a
1 L'histoire de cette espece est assez curieuse : decrite en 1890 sous le nom de N. dentistoma, elle devint
C7i 1893 Neli. crenulata; j'avais era en effet devoir operer cette rcforme, pour obeir aux lois de la nomenclature
qui, parait-il, repoussent tout adjectif form6 d'un nom grec et d'un nom latin a la fois. Mais en adoptant ce
tenne de crenulata, j'ignorais totalement qu'il existait dcja, en fait, une Nebela crenulata, creee par CASH en
1892, et qui, bien plus, par une curieuse coincidence, se trouvait etre justement cette meme espece que je
venais de debaptiser. Aujourd'hui, il faut parait-il en revenir au premier nom specifique de Neb. dentistoma
(ainsi uommce d6ja dans le volume de Cash & Hopkinson, 1909); un auteur n'a plus le droit de changer un
nom d'espece, mgme forme coiitre toutes les regies.
BRIT. ANTARCT. EXPED. 1907-9. VOL. I. F
244 E. PENARD
revetir la forme tres-large qui constitue le caractere spdcial de N. flabdhdum, et
dans certaines stations ne se trouvent meme plus guere que sous cette forme-la.
L'espece typique, telle qu'elle est indiquee et figuree p.e. par CASH et HOPKINSON
dans le 2e vol. des British Freshwater Khizopods (p. 119, PI. 28, Fig. 9-11), ne s'est
rencontree que dans les Sphagnum de Vancouver.
Nebela griseola, spec, nova
Fig. 8, a ct 1}
Dans le sphagnum de Katoomba s'est montre, en exemplaires assez nombreux,
un petit Rhizopode que Ton croirait a premiere vue devoir rapporter au genre Difflugia,
mais qui, grace a la nature particuliere de son enveloppe, doit etre attribue au genre
Nebela.
La coquille, grisatre, tres-peu transparente, de 70 a 75 M en longueur, et qui
rappellerait celle de certaines varietes de Difflugia pyriformis, est faite d'une matiere
chitinoi'de, une pellicule sur laquelle sont colic's, series les uns centre les autres, des
elements siliceux, arrondis, ovales, droits (diatome'es modifie'es), ou bien quelque
peu recourbe"s, vermiformes ; le tout, enfin, formant un revetement qui tient a la
fois du genre Nebela et du genre Lesquereusia.
Elle est pyriforme, a peu pres deux fois aussi longue que large, et peut etre
indiquee comme comprimee, car elle presente une face large et une face etroite ;
mais cette compression est si faible que, dans certains individus examines en section
transversale, on se demande s'il y a une difference quelconque entre les deux axes
de la figure elliptique que Ton a sous les yeux ; plus souvent, cependant, la difference
est nettement sensible, le grand axe etant au petit comme 6 est a 5, ou m6me quel-
quefois comme 5 est a 4.
A la bouche, 1'enveloppe s'eVase legerement, figurant un bourrelet plus ou moins
prononce, et qui dans certains cas m'a paru resulter d'un reploiement sur elle-meme
de la membrane garnie encore de ses elements de recouvrement.
L'ouverture buccale est petite, elliptique dans son contour, ou meme, tres-
souvent, parfaitement circulaire.
La Nebela griseola ne s'est montr^e, dans les recoltes de Mr. Murray, qu'tl 1'etat
d'enveloppes vides ; mais pourtant, je puis ajouter ici quelques renseignements con-
cernant le plasma.
Si cette espece, en effet, est aujourd'hui ddcrite pour la premiere fois, elle n'est
pas absolument nouvelle, en ce sens qu'elle a 4te" recoltee, par moi-meme, le 30
Juillet 1909, dans les mousses de Glendarary Ho., au promontoire d'Achill, cote
occidentale de 1'Irlande ; mais les resultats de mes recherches ne devant etre publies
que dans un avenir peut-etre encore assez ^loigne, la station des Montagues Bleues
vient, de fait, premiere en date.
RHIZOPODES D'EAU DOUCE 245
L'enveloppe renferme alors un plasma clair, limpide, dans le sein duquel on
trouve des grains d'amidon, des elements de reserve pour la confection d'une nouvelle
coquille, puis une, on deux, ve"sicules contractiles, et enfin le noyau ; ce dernier,
spherique, relativement petit (12 M), consiste en une masse d'un bleu tendre, ren-
fermant un amas central, irregulier, de nucleoles pales et tres-petits.
Tres-timides, les animaux n'ont jamais deploye de pseudopodes ; ils se collent
souvent au substratum, par leur pdristome un pen glutineux.
En Irlande, la coquille etait identique & celle de Katoomba, mais plus grande,
en general, de 80 a 85 n au lieu de 70 a 75 /u qu'elle avait en Australie.
Nebela lageniformis. PENARD
Mem. Soc. Phys. Hist. Nat., Geneve, vol. 31, p. 159. 1890
La Nebela lageniformis est 1'i.m des representants les plus caractdristiques des
mousses, aussi les rdcoltes de Mr. Murray 1'ont-elles montree a peu pres partout.
Cependant cette espece, qui jusqu'ici etait connue pour posseder des caracteres de
forme tres-nets et peu sujets a de serieuses variations, s'est montree, dans tout
1'hemisphere sud, extremement variable, et cette variabilite affectait uniquement la
partie anterieure de la coquille, le col, qui dans le type figure une sorte de tube
aplati, pose directement sur la partie renflee et arrondie de 1'enveloppe.
Au Mont Cook, on trouvait toutes les transitions entre la forme normale et
une autre a col tres-elargi a sa base ; a Waiata-Rua, il en dtait encore de inline, et
les extremes aiTivaient si loin qu'a certains exemplaires on ne trouvait plus de col
du tout, et qu'on aurait cru a une espece speciale, ou a une variete de Neb. collaris.
A 1'ile Stewart, le col etait toujoura tres-large a sa base, un simple retrecissement
graduel de la partie anterieure de la coquille ; a 1'ile Macquarie, avec la forme typique
mais variable, on en trouvait une autre, a col etroit, droit, nettement indique ; a
Ngauruhoe, enfin, c'etait une grande forme, de 140 a 160 /u, a col tubulaire, etroit,
prolongeant tout droit la coquille large et arrondie.
On constatait egalement d'assez grandes variations sous le rapport de la taille ;
tandis que dans la regie, la coquille mesure de 120 a 125 /w, on trouvait 140 k 160 M
h, Ngauruhoe ; et par contre, a Katoomba, une petite variete, melee a la forme type,
n'arrivait qu'a 85 /u.
Nebela longicollis. PENARD
Mem. Soc. Phys. Hist. Nat., Geneve, vol. 31, p. 158. 1890
En 1890, j'avais decrit sous le nom de Neb. longicollis un Rhizopode a coquille
tres-allongee, presque tubulaire, tres peu comprimee ; mais plus tard, j'avais cru
devoir rattacher cette _espece a la Neb. americana de TARANEK. Or, a ce qu'il me
semble aujourd'hui, la Neb. americana de TARANEK n'est pas autre chose que la Neb.
E. PENARD
lagenifonnis, PKNARD; seulement, la chose n'est pas certaine, car TARANEK indique la
coquille comrue non comprimee, ronde en section transversale ; mais . . . je serais
porte a croire que cette determination de " drehrund " repose stir une erreur d' ob-
servation.
Reste maintenant la Neb. barbata de LEIDY, a laquelle on rapporte quelquefois
la Neb. longicollis; mais, on le sait enfin aujourd'hui, la Neb. barbata existe, espece
bien autonome, avec sa forme speciale, son recouvrement de cils.
II me faut alors reprendre aujourd'hui 1'espece abandonnee, pour distinguer sous
le nom de Neb. longicollis une forme incolore, claire et transparente, en forme de
bouteille tres-allonge"e, presque tubuleuse, faiblement comprimee, & e"cailles fortes et
nettes, qui dessinent des areoles tres-marquds. A la bouche, les ecailles font parfois
saillie, ou bien elles y sont noyees dans un bourrelet jaunatre, souvent un peu creuse
sur la face e"troite.
Mais, il faut 1'avouer, cette espece est extremement polymorphe ; tantot droite,
tant6t recourbe"e en virgule, 1'enveloppe est quelquefois si courte, que s'il n'existait
pas toutes les transitions possible (par ex. au Mont Cook), on croirait devoir la rapporter
a la Nebela vitraea.
Au Mont Cook, dans le sphagnum, cette espece n'etait pas tres-rare ; a Waiata-
Rua, a Katoomba, il ue s'en est rencontre que quelques exemplaires.
Nebela martiali. CERTES
Mission scient. du Cap Horn, torn. 6, Protoz., p. 14. 1889
Fig. 9, a et I
CERTES a donne, en 1889, une bonne description de cette curieuse espece, qu'il
avait recue du Cap Horn, et qui n'a pas 6t6 revue depuis.
Comme dans la Neb. certesi dont il a 6t6 question plus haut, nous trouvons
sur le col un certain nombre d'ornements en forme de perles, tres-petites, disposees
ici comme un collier, en plusieurs rangees et juste en arriere de la bouche.
CERTES indique, comme traits caracteristiques, " six ouvertures ou pores disposes
symetriquement deux a deux ; la premiere paire a moitie" du col, les deux autres
perpendiculaires a la premiere, a la base du col."
Mais les choses sont un peu differentes en realite : la premiere paire dont parle
CERTES, se rapporte en effet a deux perforations, tres-faibles et souvent meme invisibles,
sur les deux cotes du col ; mais les deux autres paires representent chacune un tube,
qui traverse la coquille de part en part, et dont les deux ouvertures ont ete prises par
CERTES chacune pour une simple perforation. Cos tubes, dont les orifices se trouvent
eux-memes au fond d'un leger etranglement de 1'enveloppe, s'ouvrent au dehors en une
large lumiere, et figurent a la vue deux grands yeux enfonces ; la coquille, vue la
tete en bas, semble etre un animal fantastique, une sorte de chouette, qui rcgarde
fixement.
RHIZOPODES D'EAU DOUCE 247
Cette enveloppe, jaunatre, est couverte de petites plaques serrees, pen apparentes,
generalement elliptiques, quelquefois entremelees de plus grandes, qui semblent avoir
e'te' volees aux Trinema.
A 1'ile Macquarie, les exemplaires, tres-rares, mesuraient 125 M ; a Ngauruhoe, une
seule coquille vide a ete trouvee ; dans les Alpes australiennes, cette nebe"lide etait
commune, tres-belle, mais toujours a 1'dtat de coquilles vides, lesquelles, beaucoup plus
grandes qu'a Macquarie, mesuraient de 165 a 180-?*—
CERTES indique pour cette espece une longueur de 550 M, une largetir de 290 M ; il
doit y avoir la un simple lapsus calami.
Nebela militaris. PENARD
Me'm. Soc. Phys. Hist. Nat., Geneve, torn. 31, p. 164. 1890'
Cette jolie petite espece, tres-delicate, et toujours assez rare, s'est rencontre'e a
Katoomba, a Ngauruhoe, a Oahu. Dans cette derniere station, la taille dtait tres-faible,
51 M en longueur.
Nebela minor. PENARD
Arch. Sci. Phys. Nat, Geneve (3), vol. 29, p. 181. 1893
Cette petite Nebela, a coquille tres-claire et tres-transparente, couverte en general
de plaques rondes bien regulieres, et qui pent etre consideree comme un diminutif de la
Neb. collar is, mais semble pourtant avoir droit au titre d'espece, etait abondante dans
une mousse claire et bien fournie provenant de Oahu. Elle s'est retrouvde, beaucoup
plus rare, a Ottawa.
Nebela tincta. LEIDY
ITyalosphenia tincta. LEIDY. Fresh w. Rhiz. N.A., p. 138. 1879
Nebela tinda. AWERINZEW. Trudui, S. Petersb., vol. 36, p. 354. 1906
Cette espece, qui sous sa forme typique, avec enveloppe fortement comprimee,
r6gulierement arrondie, transparente et lisse, represente nettement une Hyalosphenia,
est assez variable pourtant, et dans beaucoup de stations se montre couverte de dessins
qui 1'ont fait reunir (a tort, peut-etre) au genre Nebela. Elle s'est rencontrde dans un
assez grand nombre des recoltes.
Nebela tubulosa. PENARD
Mem. Soc. Phys. Hist. Nat., Geneve, vol. 31, p. 159. 1890
Cette grande espece, assez rare, n'est apparue que dans la seule station de Van-
couver, bien typique, avec ses ecailles tres-petites, sa teinte speciale d'un brun chocolat
clair, et sa taille de 205 a 225 M en general.
248 E. PENARD
Nebela vas. CERTES
Mission scientifique du Cap Horn, torn. 6, Zoolog., p. 15. 1889. Fig. 10
Cette belle espece, inconnue jusqu'ici en Europe comme aussi aux Etats-Unis.
mais qui semble etre tres-repandue sur tout le territoire du Pacifique et probablernent
dans 1'hemisphere sud tout entier, a etc decrite, avec une bonne figure, par CERTES en
1889, d'apres des exemplaires provenant du Cap Horn.
Comme forme, elle rappelle a s'y meprendre 1'une des varietes de la Difflugia
(Pontigulasid) vas de LEIDY, et plus particulierement celle qui, elevee au rang d'espece,
est connue comme Pontigulasia spectabilis, PENARD. Mais ce n'est la qu'une conformite
d'apparence, et, comme CERTES 1'a deja remarque, " ce n'est pas la premiere fois que
Ton rencontre des especes differentes construisant chacune leur coquille sur le meme
type." L'enveloppe, ici bien plus fortement comprimeX revet la structure typique des
Nebela. Le plasma, le noyau, le mode d'enkystement, le diaphragme qui ferme le col
lors de 1'enkystement, tout cela est d'une Nebela.
La Nebela vas s'est trouvee dans la plupart des recoltes de Mr. Murray, toujours
bien caracteristique, mais variable dans ses dimensions, suivant la localite. Dans
plusieurs stations (Mont Cook, ile Macquarie), outre la forme normale qui generalement
mesure de 160 a 165 M de longueur, on en trouvait une autre. plus claire, plus delicate,
et beaucoup plus petite; a WaTata-Rua, cette petite forme, de 95 /u seulement, existait
seule ; il semble y avoir Ik une variete" speciale, qu'aucune transition ne relie au type.
La coquille est jaunatre, et formee dans la regie de plaquettes arrondies bien
nettes. Mais il y a sous ce rapport un fait interessant a noter : dans cette espece,
1'animal semble montrer une predilection toute particuliere pour les ecailles des
Euglypha ; il les capture, les incorpore a la masse de ses "plaques de reserve," et les
emploie en aussi grand nombre que possible. Mais, ce qu'il y a de curieux, c'est que
toutes les ecailles qui dans YEuglypha avaient etc" buccales, c'est-a-dire que Ton trouve
munies des ornements et denticulations caracteristiques, se voient sur le col de la
Nebela, et jamais sur le corps meme de la coquille. En outre, presque toujours, les
denticulations de ces ecailles parsemees un peu partout sur le col, sont dirigees en
avant ; il y a done eu un choix, tant dans le placement des ecailles que dans leur
orientation.
Mais ce n'est pas tout : s'il y a, dans cette capture des ecailles d'Euglypha, en
quelque sorte une habitude acquise par la Nebela, cette habitude peut faire defaut dans
certains cas ; par exemple, tandis qu'a Ngauruhoe, a Vancouver, la plupart des coquilles
portaient de ces Ecailles speciales, et que dans les autres stations beaucoup d'individus
n'en montraient pas, elles manquaient toujours a Tile Macquarie, au Mont Cook, a
Waiata-Rua, bien que les Euglypha (E. ciliata, E. compressa) y fussent aussi
nornbreuses qu'ailleurs.
RHIZOPODES D'EAU DOUCE 249
La plupart des representants de cette espece ont ete trouvds a 1'etat de coquilles
vides, ou bien aussi le plasma, encore vivant, se voyait enkyste ; mais aucun individu
n'est revenu a la vie active. Dans un exemplaire provenant de Ngauruhoe, cependant,
et garde pendant 5 jours dans un verre de montre, j'ai pu examiner le plasma, qui se
trouvait rempli de petits grains d'amidon, puis renfermait encore de grosses spherules
jaunatres, d'aspect cireux (re"sidus de nourriture ?), et un beau noyau, de 28 n de
diametre, du type habituel aux Nebela, c'est-a-dire- renfermant un certain nombre de
nucleoles dissemines sans ordre dans une masse (sue nucldaire) grisatre, pateuse, pleine
de granulations tres-petites.
Parmulina cyathus. PENARD
Faune Rhizopodique Le"man, p. 207. 1902
Ce petit organisme, tres-peu apparent de sa nature, et qui lorsqu'il n'est pas en
tres-bon etat de forme reste facilement inapercu, ne s'est rencontre d'une maniere
evidente qu'a Sydney (National Park) et a Ottawa. Dans ces deux stations, 1'animal
etait vivant, mais enkyste", c'est-a-dire simplement retracte en boule sous la protection
de son envelqppe.
Pkryganella hemisphaerica. PENARD
Pseudodiffluffia hemisphaerica. PENARD. Mern. Soc. Phys. Hist. Nat., Geneve,
torn. 31, p. 169. 1890
Phryganella hemisphaerica. Faune Rhizop. Leman, p. 421. 1902
Le plus commun peut-etre de tous les Rhizopodes des mousses, et qui en effet
n'a manque nulle part, sauf aux lies Auckland, d'ou il n'a ^te rapporte qu'un simple
fragment d'ecorce garni d'une petite mousse rase.
On la rencontre presque toujours et partout a 1'etat de coquilles vides ; et presque
toujours aussi, on y reconnait deux formes, 1'une petite, tres-variable d'ailleurs, et qui
semble se rapporter a ces divers petits rhizopodes que sous le nom de Difflugia
globulosa, LEIDY, represente a la PI. XVI. de son grand ouvrages (Fig. 11 h, 20) ; puis
une plus grande (40 a 50 M), que dans cette meme planche de Leidy on trouve repre"-
sentde par les Fig. 23 et 24. CASH identifie cette esp6ce, peut-^tre avec raison, a
Difflugia acropodia, HERTWIG ET LESSER.
A I'lle Macquarie, une tres-grande variete arrivait a 100 M.
Phryganella nidulus. PENARD
Faune Rhizopodiqne Leman, p. 419. 1902
Cette grande espece est apparue dans la rdcolte du Mont Cook ; mais encore faut-il
faire des reserves sur 1' exactitude de la determination. En eifet, la seule coquille ren-
250 E. PENARD
contree, vide, prdsentait des caracteres particuliers ; de 190 M de diametre, hemisphdrique,
pourvue d'une ouverture plutot petite et dechiquetee, elle etait munie sur tout son
pourtour, a la face ventrale, d'une sorte de rebord ou d'aile circulaire, qui peut-etre
n'avait d'autre signification que celle d'un mucilage durci, destine" a souder la coquille
sur le substratum.
Placocysta jurassica. PENARD
Revue Suisse Zool., vol. 13, p. 611. 1905
Ce Rhizopode, qui jusqu'ici n'avait, & ma connaissance, ete indique que dans une
tourbiere du Jura suisse (la Pile), a refait cette annee son apparition, clans le sphagnum
de Vancouver, ou il n'etait pas rare ; parfaitement typique d'ailleurs, et avec la taillc
normale de 70 a 75 /« en moyenne, mais pourvu d'aiguilles plus longues et plus serrees
que celles des exemplaires du Jura.
Placocysta spinosa. LEIDY
Proc. Acad. Nat. Sci., Philad., p. 226. 1874
Dans ce mSme sphagnum de Vancouver s'est egalement montree, beaucoup moins
abondante que la precedente, la Placocysta spinosa, bien typique, mais revetue d'une
profusion tout-a-fait anormale d'aiguilles, qui poussaient serrees, souvent en faisceaux,
tout le long de 1'arete laterale. Ces aiguilles, dgalement, etaient moins larges, moins
aplaties que dans le type ordinaire, et leur extremite se voyait tronquee il angle droit.
Les exemplaires rencontres montraient entre eux de grandes differences sous le
rapport de la longueur, et variaient de 104 a 140 /«. II est rare que dans 1'espece type
on ait trouve jusqu'ici moins de 130 M.
Plagiopyxis callida. PENARD
Revue Suisse de Zoologie, tome 18, fasc. 3. 1910
Cette espece, ddcrite tout rdcemment d'apres des exemplaires trouvds aux environs
de Geneve, rappelle a premiere vue une des nombreuses varietes de Diffiugia constricta;
mais c'est Ik en realite vin type tout different, un organisme qui par la structure toute
particuliere de son plasma, lequel s'dpaissit k sa surface de maniere k constituer une
veritable enveloppe interne, se rapproche de tres-pres des genres Diplochlamys et
Bullinula.
Elle s'est renccntree dans plusieurs re'coltes, mais rare partout sauf a 1'ile Stewart,
oil elle atteignait jusqu'a 135 M, une taille relativement tres forte. Celle du type mesure
rarement plus de 120 M.
RHIZOPODES D'EAU DOUCE 251
A Sydney, les individus etaient vivants, et j'ai pu m'assurer que le plasma
etait le meme qu'a Geneve, avec ses myriades de petits grains pales, ses particules
nutritives jaunatres ou Ton reconnait des parcelles arrachees aux mousses, et son noyau
caracteristique.
Plagiopyxis labiata, spec, nova
Fig. 1 1
Dans les mousses des environs de Sydney, de Creel, de Victoria (B.C.), s'est montre,
toujours tres-rare, un petit rhizopode qui a premiere vue pouvait etre piis poui une des
nombreuses formes de Difflugia constricta, mais qui en realite doit rentrer dans le
genre Plagiopyxis.
Plus petite que dans 1'espece precedente, la coquille, de 80 a 88 M de diametre,
brunatre, chitino'ide avec pierres de recouvrement, hdmisphe'rique avec face dorsale
sure'levee, est pourvue d'un orifice allonge en fente, ventral, excentrique, tres-rapproche
du bord anterieur. Le peristome, alors, simplement arrondi sur 1'un de ses bords (bord
anterieur), est prolonge sur 1'autre bord d'une levre plus ou moins proeminente, une
sorte de languette vaguement triangulaire, qui plonge en avant vers 1'inte'rieur de la
coquille, et tient lieu ici de ce plancher caracteristique qui dans la Plagiopyxis callida
s'enfonce jusque bien bas dans 1'interieur.
Le plasma, qu'il ne m'a etc possible d'examiner que sur un seul individu, est
conforme a celui de 1'espece prece'dente, completement bourre de grains pales extra-
ordinairement petits, puis renfermant des parcelles de nourriture jaunatres, et un noyau
spherique, pourvu a son interieur d'un gros nucleole granule. II n'a pas e"te trouve de
vesicule contractile, laquelle ne s'observe d'ailleurs que tres-rarement dans ce genre.
Quant aux pseudopodes, il en est probablement ici de meme que dans la Flag, callida,
oil Ton pent examiner bien des centaines d'animaux pleins de vie sans les voir se
deVelopper.
Pontigulasia bryophila. PENARD
Faune Rhizopodique Leman, p. 324. 1902
Ce rhizopode, qui rappelle Difflugia pyriformis dont il se distingue avant tout par
1'etranglement caracteristique du col, ne s'est montre qu'au Mont Cook et a Vancouver,
sous la forme de quelques coquilles vides.
Pontigulasia compressa. CARTER
Difflugia compressa. CARTEK. Ann. Mag. Nat. Hist. (3), vol. 13, p. 22. 1864
Pontig. compressa. CASH et HOPKINSON. Brit. Freshw. Rhiz., vol. 2, p. 62. 1909
Cette espece s'est rencontree a Waiata-Rua, puis au Mont Cook, toujours en
individus isoles, qui revetaient la forme typique, et atteignaient jusqu'a 190 M en
longueur.
BRIT. ANTARCT. EXPED. 1907-9. VOL. I. G
252 E PENARD
Cette Pontigtila&a a ete signalee dans ces dernieres annees par differents
auteurs comme 6tant identiqne a Pontiff, bigibbosa, PENARD, laquelle habite les
lacs profonds. A differentes reprises, en 1909 et 1910, je 1'ai moi-meme trouvee
dans les Sphagnum de la Suisse, et en effet, je n'y trouve pas de difference impor-
tante avec Pont, bigibbosa. II est done regrettable que cette derniere ait, par
suite de 1'ignorance de ce qu'etait en realite la Difflugia coniprestsa de CARTER,
ete eleve'e au. rang d'espece distincte. Mais il n'en reste pas moins vrai qu'entre
la Pont, compressa, CARTER, et la grande forme lacustre, la difference est tres-
grande ; dans le Leman, les individus atteignent facilement 250 M en longueur,
avec une largeur si considerable qu'elle egale tres-souvent la longueur, et dans des
cas exceptionnels va meme jusqu'a la depasser.
Quadrula irregularis. ARCHER
Quart. Journ. Micr. Sci., vol. 17, p. 103. 1877
Cette espece est assez rare dans les mousses ; aussi ne s'est-elle trouvee repre-
sentee dans les collections de Mr. Murray que dans une seule station, Suva dans
1'archipel des Fiji (nous 1'avons pourtant dejk rencontrtSe, on se le rappelle, dans
les petits lacs du Cape Itoyds). Mais a Suva, les conditions etaient un pen
speciales ; les mousses avaient ete" recoltees au bord d'un marecage, puis dans les
fosses pres de la route ; aussi cette espece n'etait-elle peut-etre pas la bryophile.1
Quadrula symmetrica. F. E. SCHULZE
Arch. f. Mikr., Anat., Bd 11, p. 329. 1875
C'est dans les Sphagnum, il Vancouver et dans les Alpes australiennes, que
s'est rencontree cette espece, laquelle n'habite pas les mousses proprement dites.
II ne s'en est montre qu'un nombre extremement restreint d'individus ; mais
j'ai pu faire, cependant, a deux reprises et sur des coquilles provenant de Vancouver,
certaines experiences qui ne seront pas hors de place ici :
On sait que LAGERHEIM (Geol. Foren. Forhand, torn. 24) a fait en 1902 la
curieuse constatation que les plaques caracteristiques de la Quadrula irregularis
etaient, non pas siliceuses, mais formees d'une combination de calcium. En 1893,
puis plus tard encore, je confirmais de mon cote les observations de LAGERHEIM,
en montrant qu'en tout cas ces plaques etaient solubles dans 1'acide sulfurique
concentre a chaud. Awerinzew, en 1907 (Zool. Anzeiger, B'1 31, No. 8 confirme
a son tour la chose, et se demande en meme temps si, malgre la grande analogic
qui existe dans la structure de la coquille, il ne faudrait pas voir dans Quad.
1 Voir pages suivantes pour les observations sur la nature des plaques dans cette espece
RHIZOPODES D'EAU DOUCE
irregularis et Quad, symmetrica deux rhizopodes tres-eloignes 1'un de 1'autre, et
appartenant en realite a des genres differents.
Cette hypothese est tres-naturelle en elle-meme, et pour mon compte, c'est
ainsi que, depuis 1893, je m'expliquais la chose ; mais encore fallait-il connaitre
d'une maniere certaine la composition des plaques dans la Quad, symmetrica.
Uepuis longtemps j'avais pu m'assurer, sur une coquille appartenant a une grande
variete speciale provenant des Montagnes Kocrreuses, que les plaques etaient
siliceuses ; mais c'etait la, apres tout, une variete ; il fallait faire des experiences
sur le type.
Or, en experimentant sur la Quad, symmetrica de Vancouver, qui represente
bien le type normal, j'ai pu constater que les plaques sont purement siliceuses ;
1'acide sulfurique bouillant les laisse parfaitement indemnes, et on les retrouve
bien nettes egalement apres Faction du chalumeau. Je serais done assez dispose
a adopter 1'opinion d'AwERixzEW, qui suppose une simple convergence dans deux
orgauismes generiquement differents.1
Peut-etre me sera-t-il permis, a ce propos, d'ajouter quelques mots a propos
d'experiences de m6me nature faites sur la Quadrula irregularis :
Dans cette espece, 1'acide sulfurique concentre froid dissout a peine, tres-
difRcilement, les plaques; a chaud, il les dissout de suite; 1'acide chlorhydrique
dissout les plaques, mais laisse intacte la pellicule fondamentale, sur laquelle on
voit encore, nettement imprimees, les marques des plaques disparues ; 1'acide acetique,
par centre, n'a aucune action ; il laisse les plaques absolument indemnes. Pour
dissoudre les plaques dans cette espece, il faut done en tout cas un acide fort, et
ce n'est pas la, sans doute, un simple carbonate de chaux.
Sphertoderia Ji&sirostris. PENAKD
Mem. Soc. Phys. Hist. Nat., Geneve, vol. 31, p. 187. 1890
Une seule coquille vide, au Mont Cook.
Trinema complanatum. PENARD
M&n. Soc. Phys. Hist. Nat., Geneve, vol. 31, p. 187. 1890
Cette petite espece, tres-frequente en general dans les mousses, s'est rencontree
dans quelques-unes des recoltes.
1 AWERINZEW se fonde d'ailleurs, et a juste titre, non pas seulement sur la composition des plaques, mais
encore sur la forme de 1'ouverture buccale, ainsi que sur la structure differente du plasma.
254 E. PENARD
Trinema enchelys. DUJARDIN
Triiieme, Duj. Ann. Sci. Nat., p. 205. 1836
Tres-frequente dans les mousses, la Trinema enchelys s'est montree dans un
grand nombre de stations, tres-variable comme toujours, et ofFrant surtout deux
formes bien accusees, 1'une d'elles belle, grande, a fortes ecailles (la forme typique
de LKIDY, Fig. 1, 2, 3, 4, de sa PI. 39); 1'autre, particuliere aux mousses, toute
differente en realite de la premiere (PENARD, Faune Rhizop. Leman, p. 527, Fig. 5),
et qui rappelle de fort pres, a premiere vue, le Corythion dubium de TARANEK.
Trinema lineare. PENARD
Mem. Soc. Phys. Hist. Nat., Geneve, vol. 31, p. 187
Cette espece, toute petite, chetive et qui passe facilement inapergue, a ate" notee
dans bon nombre de stations, soit enkystee, soit a I'd tat de coquilles vides.
CONCLUSIONS
LES Rhizopodes d'eau douce rapportes par le biologiste du Nimrod forment
aujourd'hui la liste assez respectable de 79 especes ; assez respectable en effet,
car, a une ou deux exceptions pres, les collections etaient representees par des
mousses, et seule la faune des mousses s'y est trouvde, presque au complet. Les
Sphagnum, egalement, se sont montrds reprdsentes dans quelques-unes des recoltes,
et la comme partout, ont permis d'etudier quelques especes particulieres, qui manquent
aux mousses proprement dites.
II est bien evident que les marecages, les etangs, les flaques d'eau douce, auraient
fourni un supplement considerable d'informations ; on eut trouve* des Arcelles, des
Difflugies, et bien d'autres choses encore ; mais dans les circonstances ou operait
Mr. Murray, il ne faliait guere songer a ce genre de recoltes ; et d'ailleurs, on peut
se demander si 1'absence, dans ces collections, de Rhizopodes purement aquatiques
est vraiment a, regretter.
Tels qu'ils sont, en efFet, les resultats acquis montrent une homoge'neite peut-
etre encore plus significative que s'ils eussent ete fournis par differents milieux.
Cette faune speciale et pourtant si riche encore, apporte avec elle certains
renseignements, quelque confirmation ;i ce que nous savions deja ; comme la faune
aquatique, elle est cosmopolite ; mais comme elle aussi, elle ne Test pas d'une
maniere absolue ; certaines especes sont localisees, et leur rencontre peut donner lieu
a des conclusions d'un interet special.
La Nebela mis, par exemple, decrite il y a longtemps deja comme provenant
du Cap Horn, et que le Prof. Richters a retrouvee dans toute une se"rie de locality's :
Australie, N"<! Zelande, Hawaii, Java, lies Chatham, Staaten Isld, Falklands,
Kerguelen, St. Paul, etc. etc., s'est montree commune dans 1'hemisphere austral et
sur le territoire du Pacifique tout entier ; on la trouve encore a Vancouver, mais elle
ue semble pas penetrer dans 1'interieur du continent americain. Les Nebela certesi,
Nebela martiali, ces formes si curieuses, que CEKTES avait egalement observees
au Cap Horn, ct qui n'avaient pas e"te vues depuis, ont refait leur apparition, et
se sont montrees, dans le Pacifique, avec leurs caracteres nettement marquds ;
Hyalosphenia cockayni semble dans le Pacifique sud tenir la place de I'Hyalosph.
papilio en Europe et aux Etats-Unis ; et la Centropyxis horrida possede une aire
de dispersion peut-etre plus localisee encore.
Toutes ces donnees, sans doute, n'apportent pas avec elles une bien grande
certitude ; les Rhizopodes d'eau douce sont trop insuffisamment connus pour que
255
256 E. PENARD
Ton soit en droit de tirer de leur presence des conclusions evidentes ; mais ils ont
pourtant leur mot a dire, eux aussi, dans les questions de geographic zoologique.
On les a trop negliges jusqu'ici, et, il faut bien le dire, il est pen de naturalistes
qui semblent se reiidre compte de 1'importance generale qu'ils pourraient bien avoir
en effet. Certains de ces organismes, il est vrai, se montrent sous des aspects si
divers, ou sous la forme de varietes si nombreuses, qu'ils semblent n'etre la que
pour deconcerter 1'observateur ; mais beaucoup d'autres, les plus nombreux meme,
sont parfaitement nets et fixes dans leurs caracteres ; tout au moins ne varient-ils
que dans une faible mesure, souvent beaucoup moins, meme, que tant d'etres plus
Sieves en organisation, Entomostraces, Insectes, Le'pidopteres, qui pourtant sont
consideres comme ayant une certaine importance dans les questions de faunistique
generale.
Dans le compte-rendu donne tout dernierement dans " Nature " l du volume de
CASH et HOPKINSUN sur les Rhizopodes d'eau douce, on trouve exprimee dans les
termes suivants 1'opinion qui, me semble-t-il, est a peu pres celle de la majorite des
naturalistes: "To the working microscopist who is anxious to find names for the
varieties he discovers in the fresh-waters that he visits, the book will doubtless be
of some value, for it gives him . . ." &c. Mais tout cela n'est pas tout-a-fait
juste ; pour les Rhizopodes, il est vrai, et plus pour eux peut-etre que pour tout autre
groupe animal, les etudes systematiques sont necessaires, et le seront longtemps
encore ; mais non pas pour collectionner des noms ; c'est pour arriver a des faits
qu'il faut cataloguer ; quand la systematique sera mieux fixe^e, on reconnaitra a ces
organismes une importance plus grande qu'on ne 1'a soupconnee jusqu'ici.
Et alors, si cette importance etait reconnue, si dans chaque expedition aux pays
uouveaux le natural iste decidait de songer a ces organismes un peu plus qu'on ne
1'a fait jusqu'a ce jour, si les mousses et les sphagnum etaient consideres comme
devant avoir leur juste part dans la repartition entre botanistes et zoologistes, quoi
de plus aise que de les re"colter? quelques petits paquets de vegetaux sees, et vous
aurez toute une faune, Rotiferes, Tardigrades, Nematodes, Rhizopodes et d'autres
choses encore, que souvent vous pourrez rapporter vivants, et qui se trouveront avoir
leur importance dans les re"sultats biologiques obtenus.
Sous ce rapport aussi, me semble-t-il, les collections recueillies par Mr. Murray
renferment un enseignement. Pour moi, j'ai pris a cette etude un interet tout par-
ticulier, et en rendant compte du resultat de mes recherches, c'est avec autant de
plaisir que de reconnaissance que je remercierai Sir Ernest Shackleton et Mr. James
Murray d' avoir bien voulu me confier ces collections.
1 No. 2118, vol. 83, June 2, 1910.
LISTE BIBLIOGRAPIIIQUE
CASH, J., AND HOPKINSOX, J., "The British Freshwater Rhizopoda and Heliozoa." Printed for the Ray
Society. Vol. ii. 1909.
CERTES, A., " Mission scientifique du Cap Horn," torn, vi., Zoologie, 1889.
LEIDY, J., '-Freshwater Rhizopods of North America," Report of the United States Geol. Survey, vol. xii.,
1879.
LENHESFELTJ, R. V., " Australian Freshwater Rhizopods," Proceed. Linnaean Soc. of New South Wales,
vol. x., 1885, p. 724.
MURRAY, JAMES, " On Collecting at Cape Royds," British Antarctic Exped., 1907-9. Biology, vol. i.
part i.
,, "On Microscopic Life at Cape Royds," British Antarctic Exped., 1907-9. Biology,
vol. i. part ii.
PENARD, E., " On Some Rhizopods from the Sikkim Himalaya," Journ. Roy. Microsc. Soc., 1907,
pp. 274-78.
,, " Faune Rhizopodique du Bassin du Leman," Geneve, 1902.
RICHTERS, F., " Fauna der Moosrasen des Gaussbergs und einiger siidlichen Inseln," Deutsche Siidpolar
Exped., 1901-3, Bd. 9, Zool.
„ " Moosbewohner," Schwedische Siidpolar Exped., 1901-3, Bd. 6, 1908.
,, " Moosfauna Australiens, etc.," Zool. Jahrbuch. : AU.fiir Syst., Bd. 26, 1908, p. 196.
„ " Moosfauna-Studien," Her. der Senclcbg. Naff. Ges. Frankfurt-a-M., 1908, p. 22.
SCHEWIAKOFF, W., " Ueber die geographische Verbreitung der Siissw. Protozoen," Mem. Acad. Sci.,
Petersbourg, vii. Ser., torn. xli. No. 8.
WHITELEGGE, T., " Invertebrate Fauna of Port Jackson and Neighbourhood," Journ. and Proceed. Roy.
Soc. New South Wales, vol. 23, 1899, p. 296.
„ "List of the Freshwater Rhizopods of New South Wales," Proceed, of the Linnivan
Soc. of New South Wales, Ser. 2, vol. i., 1886, p. 497.
259
PLATE XXII
H
260
EXPLICATION DE LA PLANCHE XXII
FIGURE 1. — Bullinula indica, petite vane" to ronde.
FIGURE 2. — Oentropyxia horrida. (a) Forme type ; (b) forme de"pourvue d'epines.
FIGURE 3. — Corythion dubium. (a) Varidte' orbiculaire ; (b) varie"te spicatum.
FIGURE 4. — Euglypha brachiata. (a) Aspect general de la coquille ; (6) la meme vue
d'en haut, pour montrer 1'aplatissement des " bras."
FIGURE 5. — Hyalosphenia cockayni. (a) Vue par le cote* large : sur le col on remarque
deux ecailles buccales d'une Euglypha ; (b) vue par le c6te" e"troit.
BRIT. ANTARCT. EXPED. 1907-9
VOL. I. PLATE XXII
PENARD: RHIZOPODES
3
E. Peiiard del.
261
PLATE XXIII
262
EXPLICATION DE LA PLANCHE XXIII
FIGURE 6. — Hyalosphenia subftava. (a) Vue par le c6td large ; (b) cote etroit ; (c) la
coquille vue d'en haut par 1' orifice buccal.
FIGURE 7. — Nebela certesi. (a) Cdte1 large ; (b) cdte etroit.
FIGURE 8. — Nebela griseola. (a) Coquille vide; (b) une autre, vue d'en haut.
FIGURE 9. — Nebela martiali. (a) C6te" large; (b) cote" etroit.
FIGURE 10. — Nebela vas. Sur le col, ou voit trois ^cailles buccales d'Euglypha.
FIGURE 11. — Plagiopyxis labiata. La coquille vue par la face ventrale.
BRIT. ANTARCT. EXPED. 1907-9
VOL. I. PLATE XXIII
PENARD: RHIZOPODES
a
6
V
11
a
10
E. Penard del.
BRITISH ANTARCTIC EXPEDITION 1907-9
UNDER THE COMMAND OF SIR E. H. SHACKLETON, C.V.O.
REPORTS ON THE SCIENTIFIC INVESTIGATIONS
VOL. I
BIOLOGY
EDITOR - - JAMES MURRAY
PART VII
FRESHWATER ALG^E
BY W. WEST, F.L.S., AND G. S. WEST, M.A., D.Sc., F.L.S.
(WITH THREE PLATES)
LONDON
PUBLISHED FOR THE EXPEDITION BY WILLIAM HEINEMANN
21 BEDFORD STKKKT. W.C.
1911
ISSUED SEPARATELY, DECEMBER 1911
PRICE TIIIIKE SHILLINGS NET
REPORTS ALREADY ISSUED
VOL. I. PART I., ON COLLECTING AT CAPE ROYDS
BY JAMES MURRAY PRICE Is. 6d. NET
VOL. I. PART II., ON MICROSCOPIC LIFE AT CAPE ROYDS
Bv JAMES MURRAY PRICE 5s. Od. NET
VOL. I. PART III., ANTARCTIC ROTIFERA
BY JAMES MURRAY PR,CE 5s. Od. NET
VOL. I. PART IV., MUSCI
BY JULES CARDOT PRICE Is. Od. NET
VOL. I. PART V., TARDIGRADA
BY JAMES MURRAY PRICE 10s. Od. NET
VOL. I. PART VI., RHIZOPODES D'EAU DOUCE
BY EUGENE PENARD PR,CE 3s. Od. NET
PART VII
FRESHWATER ALG^E
BY W. WEST, F.L.S., AND G. S. WEST, M.A., D.Sc., F.L.S.
With Plates XXIV to XXVI
INTRODUCTORY REMARKS
UP to the present our knowledge of Antarctic Freshwater Algae is very scanty. The
first records were by Hooker and Harvey* of certain forms collected during the
voyage of H.M. ships Erebus and Terror from 1839 to 1843. Wildemanf has given
a brief account of the Algae collected by Racovitza on the Antarctic expedition of
the s.y. Belgica, and Van HeurckJ has worked out the diatoms of this expedition
very fully. Wille§ has also recorded two Algse from Cape Adare, South Victoria Land
(rather more than 71° S. lat.), collected on the Borchgrevink Antarctic Expedition.
Other papers dealing more or less with Antarctic freshwater algae are those of Reinsch
on Algae collected in Kerguelen Land|| and in South Georgia.** These islands, how-
ever, do not extend beyond 55° S. lat., and they possess an Alga-flora which cannot
be regarded as of an Antarctic type.
The algal material collected by the different members of the expedition, but mostly
by Mr. James Murray, is probably fairly representative of the Alga-flora of the
Antarctic continent in the vicinity of Ross Island. The Algae were found on the
ground, on stones, and in the lakes, and owing to the severity of the weather conditions
they are completely frozen for at least nine months of the year. In some cases it
appears that the lake does not become free from ice for several years, and the Algae
have thus to retain their vitality in a frozen state for an extended period. Many of
* J. D. Hooker and W. H. Harvey, " Algoe in J. D. Hooker's Botany of the Antarctic Voyage," Flora Antarc-
tica, Part I., London, 1847.
f E. de Wildeman, " Note preliminaire sur lea Algues rapportees par M. E. Racovitza, naturaliste de 1'expedition
antarctique beige," Bull. Acad. Soy. Belgique, 1900.
{ Van Heurck, " Diatomees in Result. Voyage du s.y. Belyica en 1897-99," Anvers, 1909, pp. 1-128, Pis. I. -XIII.
§ N. Wille, " Antarktische Algen in Mittheilungen tiber einigc von C. E. Borchgrevink auf dem antarctischen
Festlande gesanimelte Pflanzen," Nyt Mag. f. Naturvidenskab, Bd. 40, Heft III. 1902.
|| P. F. Reinsch, " Species ac. gener. Nov. Algarum aq. dulc. Expedit. Vener. transit, hieme 1874-75 in insul.
Kerguelensi a cl. Eaton collectis," Journ. Linn. Soc. lot., xv., 1876.
** P. F. Reinsch, " Spec, et gen. nov. Algarum ex insul. Georgia Austr.," Ber. Deutsch. botan. Oes., vi., 1888.
BEIT. ANTAKCT. EXPED. 1907-9. VOL I. PART 7, ISSUED DECEMBER 1911 I
0
264 W. AND G- S. WEST
them were actually collected by being thawed out of the ice, in which they could be
seen embedded. Others were obtained from under the ice of the lakes, and from the
lake- bottom, by digging through a great thickness of ice, in one case as much as 15 feet.
Yet others were exposed on the surface of the ice as the latter was removed by ablation
during the blizzards.
The smaller ponds completely thawed in the brief summer period, but they did
not contain so many Algae as the more or less frozen lakes.
It is evident from the foregoing remarks that the Antarctic Freshwater Algae have
a very severe struggle for existence, and evidences of this can be seen in many ways.
Both slowness of growth and interrupted growth are noticeable in the Prasiolas, and
the frustules of the more delicate diatoms, such as Tropidoneis Icevissima, exhibit
considerable deformity.
The Algae observed in the collections were as follows :
Genera Species
Chlorophyceae ..... 6 . . 16
Bacillarieae ..... 16 .. 30
Myxophyceae 11 . . 39
33 84
Thus the Chlorophyceae were but feebly represented, and it should be remarked
that no Conjugatae were observed. The Myxophyceae (or Blue-green Algae) were the
most frequent, although the diatoms were very numerous in some of the lakes. Many
representatives of both these groups were mixed up in the sediment of the ponds and
lakes, and on the surface of the latter, often embedded in the ice, were very extensive
sheets of Myxophycese. These sheets were often of a brilliant blue-green colour, but
were frequently bleached, owing most probably to the intensity" of the light, and
were usually of a warm brown colour.
There was a conspicuous absence of red or yellow snow in the region visited by
the expedition. A small amount of yellow snow was observed by Mr. R. E. Priestley,
of the Western Geological Party, but the collection was lost with much of the rest of
the impedimenta on that memorable escape of the members of this party from a
drifting ice-floe.
Special attention is directed to the great salinity of the water of Green Lake, from
which locality quite a large percentage of the species recorded in this paper were
obtained. These species, which included a number of Algae generally regarded as
freshwater types, were certainly living in water in which there was a much greater
degree of concentration of salts than in ordinary sea- water. A number of typical
maiine diatoms occurred in this lake, but other species which were here plentiful
also occurred in other lakes and ponds in which the water was not strongly saline. We
have not, however, as yet any information as to the degree of salinity of Green Lake
in summer, when the ice is all melted.
FRESHWATER ALGM 265
Compared with the Alga-flora of Arctic land-areas in a similar latitude, that of
the Antarctic continent is distinctly poor. The Green Algse are relatively fewer, and
there is a lack of diversity among the species of diatoms. Both of these facts may
be directly due to the undoubted salinity of most of the lakes near the coast. Such
a condition would be prejudicial to the growth of most Green Algse, and the majority
of freshwater species of diatoms would be excluded for the same reason. The most
truly freshwater of the diatoms observed were_jthpse from the ponds on the lower
slopes of Mt. Erebus. The comparative poorness of the Antarctic Alga-flora may
also -be in part due to the greater severity of the climate combined with the remoteness
of the Antarctic continent from other continental areas.
BRIEF ACCOUNT OF THE COLLECTIONS
Most of the collections were made by Mr. James Murray in the near vicinity of
the winter quarters of the Expedition at Cape Royds on Ross Island. One was made
by the leader of the Expedition at Hut Point, Ross Island, and two others on the
mainland of South Victoria Land. The following data will include brief statements
as to the habitats and the principal Algse found in each collection.
I. Ross ISLAND
The camp at Cape Royds was at 77° 32' S. lat. and 166° 12' E. long. All the
collections were made within a few miles of this, except the one at Hut Point, lat.
77° 50' S. The numerous ponds and lakes near the camp yielded quite a number of
interesting Algse, and special attention should be drawn to the numerous Blue- green
Algse, some of which occurred in extensive sheet- like expansions.
1 Pony Lake, Cape Royds
April 22, 1908, and January 4, 1909; coll. J. Murray. Close to winter quarters.
Never clear of ice except in small parts between November 28, 1908, and the end of
January 1909. Temperature of water never much above freezing-point, the highest
record being 35° F. on December 4, 1908.
In January the material was distinctly green, and consisted of quantities of
Chlamydomonas subcaudata and C. intermedia forma antarctica among the numerous
trichomes of Phormidium autumnale.
In April the material (obtained from under the ice) was darker in colour and
contained fewer Algse.
In both months filaments of the curiously contorted Phormidium antarcticum
were common, also Pleurococcus frigidus.
2 Ponds, Cape Royds
A number of small ponds in which the temperature about the end of December rose
to 60° F. Coll. J. Murray.
266 W. AND G. S. WEST
(a) December 1908. Great quantity of Nostoc antarctica with many diatoms.
(b) December 1908. Large quantity of Nostoc antarctica with several Green
Algse.
(c) January 26, 1909. Half a mile from camp. Mostly Ulothrix subtilis var.
mriabilis, with several blue- green filamentous forms, including Oscillatoria terebri-
f or mis forma tennis, and 0. amphibia var. robusta.
3 Green Lake, Cape Royds
Half a mile from camp. Coll. J. Murray. Three collections were made:
(a) November 29, 1908. Some open water on the lake.
(b) January 3, 1909. Temperature of open water 36° F.
(c) February 2, 1909. Temperature of open water 35° F. On this day the lake
became frozen over again.
The water of this lake is very saline, and there is always a little water under the
ice. Owing to its remarkable salinity, the fluid obtained from under the ice at the
time when it was thickest did not freeze until the temperature was reduced to — 7° F.
On June 26 the temperature under the ice was 21° F., and on August 6, 1908, it was
8° F.
The lake contained twenty-six species of Algse, among which the Myxophyceae
predominated. These Blue- green Algae were free-floating in the briny liquid, no
extensive sheets being developed. The Green Algse occurred mostly in February.
Eleven species of diatoms were observed, almost all of which were marine forms.
Two of them, however, Navicula muticopsis and N. (Pinnularia) globiceps, occurred
plentifully in other situations which were not so saline in character.
We append a complete list of the species of Algse which lived in association under
the conditions of great salinity prevailing in this lake :
Ulothrix tenerrima Kiitz. forma antarctica.
U. cequalis Kiitz. forma.
Pleurococcus angulosus Lagerh. forma.
PI. antarcticus sp. n.
PL dissectus (Kiitz.) Nag.
Trochiscia aspera (Reinsch) Hansg.
Coscinodiscus lentiginosus Jan.
Hemiaulus ambiguus Jan. var.
Triceratium arcticum Btw.
Fragilaria obliquecpstata V. Heurck.
„ „ forma maxima V. Heurck.
Navicula muticopsis V. Heurck.
2V. (Pinnularia) globiceps Greg.
Trachyneis aspera (Ehrenb.) Cleve.
Tropidoneis Iwvissima sp. n.
FRESHWATER ALG^ 267
Cocconeis litigiosa V. Heurck.
Lyngbya limnetica Lemm.
L. murrayi sp. n.
Phormidium fragile (Menegh.) Gomont.
Oscillatoria deflexa sp. n.
0. chlorina Kiitz.
0 limosa Ag.
Chroococcus cohcerens (Breb.) Nag.
Chr. minutus (Kiitz.) Nag.
Chr. minor (Kiitz.) Nag. forma minima West.
Microcystis chroococcoidea sp. n.
Asterocystis antarctica sp. n.
4 Penguin Rookery, Cape Royds
February 3, 1909. On the sloping ground below the rookery, largely permeated
by the drainage from this nesting- place, were masses of the Hormidium- stage of
Prasiola crispa. Amongst the filaments were vast numbers of Navicula muticopsis.
A small pond in the rookery, which was never observed with water in it, contained
quantities of Prasiola crispa forma aspera.
5 Small, almost dried-up lake, Cape Royds
Not far from the Penguin Rookery. [No further data.] Material consisted of a
great quantity of Ulothrix cequalis, amongst which were Phormidium retzii and large
numbers of Navicula muticopsis,
6 Moraines near camp, Cape Royds
March 12, 1908 ; coll. J. Murray. Material collected in underground pockets,
believed to have formed part of an old lake-bottom. No Algae of any kind were
observed in this collection.
7 Recent geological deposit containing subfossil diatoms
Coll. J. Murray. Material obtained from the summit of a small hill about 200
feet high, near Cape Royds. Believed by the geologists to be the bottom of an old
tarn.
Contained four species of interesting diatoms, three of which were found in the
ponds and lakes, the remaining one being new. The species were: Navicula muti-
copsis V. Heurck, N. globiceps Greg., N. peraustralis sp. n., and Stauroneis anceps
Ehrenb. var. amphicephala Kiitz.
8 Blue Lake, Cape Royds
December 10, 1908 ; coll. J. Murray. This was the largest lake of the district,
situated half a mile east of the camp. It was never even partially melted. The
268 W. AND G. S. WEST
material was obtained from under fifteen feet of ice in one place and from one to
three feet in another. The ice of this lake yielded water of such purity that it
could be used in place of distilled water in chemical experiments.
The material consisted of thin, tough sheets of Myxophycese, of a deep blue-green
colour and 0.5-1 mm. in thickness. The greater part of the material consisted of
Phormidium glaciale, with smaller quantities of Phormidium inundatum and Lyngbya
martensiana. A species of Calothrix was also present in small quantity. Among the
blue- green sheets were Pleurococcus antarcticus forma robusta and Chlamydomonas
nivalis (in the resting state).
No diatoms were observed in the lake.
On the ground not far away from the margins of the lake was an abundance of
Prasiola crispa in the Schizogonium-stage and with quantities of the earlier Pleuro-
coccws-like stages. Amongst it were large numbers of Navicula muticopsis.
9 Coast Lake, Cape Royds
One mile from the camp. Some open water on November 28, 1908, and all
clear of ice early in January 1909 ; completely frozen over again early in February.
Temperature of water at the edge of lake : December 4, 1908, 47° F. ; January 2, 1909,
40° F. ; January 18, 1909, 45° F. This was the best lake for animal life.
(a) September2, 1908. Expanded sheets of a deep blue-green colour, often bleached
and about 1 mm. in thickness, consisting of two species of Phormidium, more especially
Ph. glaciale.
(b) January 1909; coll. J. Murray. Algae were all Myxophyceae, but all free-
floating. No sheet-like expansions observed. No diatoms. Mostly species of Oscil-
latoria, such as 0. deflexa, 0. cortiana, 0. formosa, 0. tennis, 0. subproboscidea, and
0. limosa, with small quantities of Lyngbya kutzingii, Microcystis stagnalis, and
Chroococcus minutus var. obliteratus.
10 Clear Lake, Cape Royds
April 1908 ; coll. J. Murray. This lake was one mile north of the camp, and
was never seen clear of ice. The temperature of the water was just above freezing-
point, the highest reading being 35° F. under nine feet of ice.
(a) Material obtained from the bottom of the lake at a depth of 17 feet. It was
black and smelt offensively, but contained many species of ciatoms. Also small
quantities of Pleurococcus antarcticus, the cells of which contained one or two small
globules of a fatty oil.
(b) Material melted from the ice and consisting of layers of Algae. Some were
sheets of Phormidium glaciale with a slight admixture of Oscillatoria cortiana ; others
were compactly interwoven layers of Ulothrix cequalis.
11 Deep Lake, Cape Royds
October 1908 ; coll. J. Murray. Two miles south of camp, lying in a deep
FRESHWATER ALGM 269
gully, and of unknown depth. The lake was never seen clear of ice, but round the
stones at the margin a little water was found in which Algae occurred. The tem-
perature of the water was just over freezing-point. The material consisted of thin
papery sheets of Phormidium angustissimum, densely studded with colonies of
Pleurococcus dissectus.
12 Ponds on the slopes of Mt. Erebus
There were a number of these ponds from whieh^two collections were obtained.
(«) March 28, 1908 ; coll. D. Mawson, from ponds rather near the camp. The
material consisted of extensive cartilaginous patches of Lyngbya erebi, some 3 to 5 mm.
in thickness. Enclosed in this tough stratum were also threads of Lyngbya kiUzingii
and Oscillatoria limosa, together with colonies of Pleurococcus dissectus and a few
specimens of Tabellaria flocculosa, Cocconema pusitta, Navicula radiosa, and Melosira
varians.
(b) December 31, 1908 ; coll. J. Murray, from ponds about two miles from the
camp. The great mass of material consisted of grey papery patches of considerable
extent and of 0.5 mm. in thickness, the nature of which was difficult to determine.
It is probable that most of it consisted of Leptothrix or other allied filamentous bacteria.
Amongst these colourless threads were various sparsely scattered Myxophycese,
including Phormidium autumnale and Oscillatoria producta. Colonies of Pleurococcus
dissectus occurred on both sides of the grey expanded sheets.
One tube contained a quantity of Nostoc antarctica.*
13 High moraines on Mt. Erebus
Altitude from 300 to 400 feet, and situated about three miles from the camp.
Coll. J. Murray.
(«) December 1908. Ulothrix implexa, with large numbers of Diatoms, especially
Navicula muticopsis.
(b) January 1909. Quantities of Prasiola antarctica attached to the stones.
Amongst it were Gloeocapsa shutUcworthiana and Aphanocapsa montana, the latter
covering quite large areas of the stones.
14 Pond at Cape Barne
December 12, 1908 ; coll. J. Murray. Two miles south of the camp. Tem-
perature of water 54° F. The pond was covered from side to side (from 20 to 30
yards) with an unbroken sheet of Myxophycese about 12 mm. in thickness. The mass
was kept floating by oxygen bubbles generated by the activity of the Algae.
This stratum, which was of a totally different character from the blue- green sheets
in Blue Lake, Coast Lake, or Deep Lake, consisted of Oscillatoria sancta and Phormidium
autumnale.
* Attached to some half-decayed thalli of this Nostoc were a few small colonies of the fungus Sporodinia asper-
gillus Schroet. The specimens, which were carefully compared with British ones, were found to be typical in all
respects. This is probably the farthest south record for any member of the Mucorinse.
270 W. AND G. S. WEST
15 Lake at Hut Point
Coll. Sir E. Shackleton. Not seen clear of ice. The material was in yellow-
brown or greenish-brown sheets, and consisted mostly of a Leptothrix, in which
matrix were Phormidium inundatum, Lyngbya shackletoni, and Calothrix epiphytica.
II. SOUTH VICTORIA LAND
Only two collections were brought from this part of the mainland of the Antarctic
continent.
16 Lake on west side of McMurdo Sound
January 1909 ; coll. R. E. Priestley. This lake was close to the " Stranded
Moraines " and twenty-five miles from the camp at Cape Royds. Lat. 77° 45' S. It
is said to be a large lake, but there were no data as to the temperature of the water
or how long clear of ice. Water swarming with red Rotifers. Yellowish snow on
shores, believed to be caused by Rotifers, but specimen lost.
The main mass of the material consisted of Oscillatoria priestleyi, amongst which
were scattered trichomes of Phormidium autumnale and Oscillatoria deflexa. Also
two species of Chroococcus. Numerous diatoms occurred, of a very similar nature to
those found in the lakes of Ross Island.
17 Five and a half miles south of Cape Irizar
On the western side of the Ross Sea. October 23, 1908 ; coll. Prof. T. W. E.
David. Lat. 75° 40' S. Situated 120 miles from the camp at Cape Royds.
The material consisted of Prasiola crispa, mostly in the Hormidium- and Schizo-
gonium- stages.
FRESHWATER ALG^E 271
SYSTEMATIC ACCOUNT OF SPECIES OBSERVED
CHLOROPHYCE^E
ULOTRICHACE.P
Genus — ULOTHRIX Kiitz.
1 Ulothrix subtilis Kiitz.
Phyc. Germ. 1845, p. 197 ; Tab. Phycol. II, 1852, t. 85, f. 1 ; Rabenh. Flor. Europ.
Alg. III., 1868, p. 365.
Var. variabilis (Kiitz.) Kirchner, Alg. Schles. 1878, p. 77. U. variabilis Kiitz.
Spec. Alg. 1849, p. 346
Crass, fil. 6-7 M ; cellulis diametro lj- 2-plo longioribus.
Hob. In pond, Cape Royds.
The specimens were a trifle thicker than the average for this plant, but they are
referable to no other species of the genus. Some of the more elongated cells had
formed " thin- walled aplanospores," which were germinating while still within the
original mother- cell. It is possible that this may have been the germination of
incompletely formed zoogonidia, a phenomenon which occurs not infrequently in
U. zonata, but the material did not admit of a determination of this point.
2 Ulothrix tenerrima Kiitz.
Phyc. gen. 1843, p. 253, t. 9, f. 1 ; Phyc. Germ. 1845, p. 197 ; Spec. Alg. 1849,
p. 346 ; Rabenh. Flor. Europ. Alg. Ill, 1868, p. 366. U. tennis Kiitz. Spec. Alg.
1849, p. 346.
Forma antarctica. (PI. XXIV, Figs. 5-7)
Forma filis brevissimis, flexuosis vel subcontortis, libere natantibus inter algas
varias Myxophycearum ; cellulis plerumque brevibus, diametro J-l-plo longioribus.
Crass, fil. 7-8 M.
Hob. Green Lake.
The extreme shortness of the filaments and their bent character are the distin-
guishing features of this form. It should be emphasised that this form was growing
in very saline water, with a much greater concentration of salts than in sea- water.
3 Ulothrix implexa Kiitz.
Spec. Alg. 1849, p. 349 ; Tab. Phycol. II, 1852, t. 94, f. 2 ; Hazen in Mem. Torr.
Bot. Club, xi, No. 2, 1902, p. 153, t. 21, f. 1, 2.
Crass, fil. 11.5-12.5 p.; cellulis plerumque paullo inflatis, diametro f-l-plo
longioribus.
Hab. High moraines, Mt. Erebus.
BRIT. ANTARCT. EXPED. 1907-9. VOL. I. K
272 W. AND G. S. WEST
This is normally a submarine species, and although in its present habitat there
would doubtless be a substratum containing a considerable percentage of saline
constituents, yet this locality was the least saline of any from which species of
Ulothrix were obtained. Some of the diatoms amongst the mass of Ulothrix were
typically submarine.
4 Ulothrix cequalis Kiitz.
Phyc. Germ. 1845, p. 197 ; Spec. Alg. 1849, p. 347 ; Tab. PJiycoL II, 1852, t. 89, f. 1.
Hormiscia cequalis Rabenh. Flor. Europ. Alg. Ill, 1868, p. 363.
U. filis exacte cylindricis, subrectis vel leviter flexuosis, cellulis diametro 0.7-1 .3-plo
longioribus; chromatophora parietali subparva, sublobata, plerumque unilateral! ter
disposita, pyrenoidibus nullis.
Diam. cell. 14-20 M; long. cell. 10-25 M- (PL XXIV, Figs. 1, 2)
Hob. Dried-up Lake, Cape Royds.
As the characters of Ulothrix cequalis are still somewhat obscure, we have given
a concise description and figures of the Antarctic specimens. We attach no importance
to the absence of pyrenoids from the chloroplasts, as this condition is frequent
throughout the genus when the chloroplasts are small and only occupy about half
the cell.
Both Ulothrix cequalis and U. tenerrima have been found in the snow- flora of
Spitzbergen. Cf. Lagerheim nNuova Notarisia, ser. vi, 1895, p. 6 (sep.).
Forma filis paullo tenuioribus, chromatophora majori cum pyrenoide singulo vel
rare pyrenoidibus birds.
Diam. cell. 13-16 M; long. cell. 11.5-18 M- (PL XXIV, Figs. 3, 4)
Hob. Green Lake.
This form was not uncommon among various Myxophyceae.
PRASIOLACE./E
Genus — PBASIOLA Ag.
5 Prasiola crispa (Lightf.) Menegh.
Kiitz. Phyc. gener. 1843, p. 295 ; Gay, Recherches sur la developpement et la
classif. Alg. Vertes, Paris, 1891, p. 86.
Hob. On the ground and in a small pond, Cape Royds ; also covering a consider-
able area of the ground below the Penguin Rookery. Near Blue Lake, on the ground.
Also on the ground 5j miles south of Cape Irizar.
Nearly all the specimens of the genus Prasiola examined from the Antarctic region
were without doubt forms of P. crispa.
In the vicinity of Cape Royds the Hormidium- and Schizogonium- stages were
iquent. The cells measured 9-12 /u. in diameter, and the filaments were mostly
rather short. In the Penguin Rookery, where the necessary conditions for the growth
of this Alga were fully realised, it occurred in great abundance. The filaments showed
FRESHWATER ALGJE 273
every sign of being subjected to very rigorous conditions. The small amount of
growth from one of the old cells of a preceding season could be well seen in most of
the specimens. (Vide PL XXIV, Figs. 12-14)
Among the filaments in the H or midium- stage, which formed a felty covering over
the ground near Blue Lake on December 11, 1908, were all the early stages of develop-
ment which so much resemble both small and large colonies of Pleurococcus. They
have, however, several distinguishing features,- one of which is in the form and
disposition of the chloroplast.
The expanded thalli were abundant from several places near Cape Royds and
from 5j miles south of Cape Irizar, South Victoria Land. All stages in the.develop-
ment were frequent, and were very similar to those figured by Wille (in Nyt Mag /.
Naturvidenskab, Bd. 40, Heft III, 1902, t. 3) from another Antarctic region. The
spacing of the cells was exactly as in P. crispa. (PL XXIV, Figs. 8, 9)
It would appear that the great mass of the Antarctic Prasiola differs in no way
from the northern P. crispa. The fully expanded and plicate thalli attained a breadth
of 13 mm., the cells varying from 6 to 9 M in diameter.
Var. aspera var. n. (PL XXIV, Figs. 15-18)
Filis in statu Hormidio modo visis ; membrana exteriori cellularum incrassata et
lamellosa, marginibus valde et irregulariter asperis ; cellulis ssepe brevissimis.
Crass, fil. 12-18 /x; diam. cell. 9-11 /*; long. cell. 1.4-7.5 p.
Hob. In small pond, Cape Royds.
This peculiarly rough variety certainly does not agree with the published descrip-
tions and figures of Prasiola crenulata. (Kiitz.) Gay ( = Hormidium crenulatum Kiitz.
Phyc. Germ. 1845, p. 193 ; Ulothrix crenulata Kiitz. Tab. Phycol. II, t. 97, f. 2).
6 Prasiola antarctica Kiitz.
Spec. Alg. 1849, p. 473 ; Tab. Phycolog, V, t. 40, f. 4 ; Rabenh. Flor. Europ. Alg.
Ill, 1868, p. 311.
Long, thall. 2-4 mm.; lat. thall. 1.5-3 M; diam. cell. 4-7 /*. (PL XXIV, Figs.
10, 11)
Hob. High moraines on Mt. Erebus, attached to stones.
Although most of the Prasiolas examined from this Antarctic region were without
doubt forms of P. crispa the one on the " high moraines " of Mt. Erebus was different
in the growth of its thalli and in the spacing of the cells. The spaces between the
cell- groups were wider and the cells themselves were not so closely aggregated. This
Prasiola agrees in all essential particulars with that described and figured by Kiitzing
as P. antarctica ; and, moreover, the habitat on the moist stones of Mt. Erebus is
precisely like that given by Kiitzing (" in rupibus madidis ") for this species, whereas
P. crispa was only observed on the ground or submerged.
Under the circumstances we are compelled to regard the Mt. Erebus specimens as
Prasiola antarctica, and to disagree with Wille' s assertion, based on a single Antarctic
274 W. AND G. S. WEST
sample from Cape Adare, that there is no difference between P. antarctica and P. crispa.
Wille's figures show that the specimens he examined were ordinary forms of P. crispa,
but we fail to see how that in any way affects the specific distinctness of P. antarctica.
VOLVOCACE.E
Genus — CHLAMYDOMONAS Ehrenb.
7 Chlamydomonas nivalis (Bauer) Wille
"Alg. Notizen XI," Nyt Magazin f. Naturvidenskab, Bd. 41, Heft 1, 1903,
pp. 147-149, t. 3, f. 44, 45 ; t. 4, f. 25. Sphcerella nivalis (Bauer) Sommerfeld, 1824.
Diam. cell. 19-23.5 ^
Hob. Blue Lake and Deep Lake.
Only the resting stage of this Alga was observed. The brillant red cells, full of
fatty- oil reserve, occurred in small groups among Pleurococcus antarcticus forma robusta
in the stratum of Phormidium glaciate.
8 Chlamydomonas subcaudata Wille
I.e. p. 118, t. 3, f. 12-18.
Long. cell. 31-34 /* ; kt. 16.6-18 /u. (PI. XXIV, Figs. 25-29)
Hob. Pony Lake.
This rather large species of the genus was fairly common amongst Chlamydomonas
intermedia forma antarctica. It possesses very distinctive characters, and the
Antarctic form scarcely differs in any way from that originally described by Wille
from Aalesund. The protoplast fully occupied the middle portion of the cell, but
both in front and behind it did not extend close up to the cell- wall. The posterior
extension of the cell- wall was generally acute, but no specimens were seen in which
it was subcaudate as in Wille's Fig. 12. Nearly all the cells were very slightly
asymmetrical about the longitudinal axis.
The chloroplast is massive, with a very large pyrenoid in its hinder half. The
pyrenoid is in the median line and immediately in front of it is the nucleus. The
nucleus is thus slightly nearer the centre of the cell than in Wille's specimens. (Of.
Wille's Figs. 12-15.) In all the specimens seen the chloroplast appeared to be finely
pitted, giving rise to flattened tooth-like marginal projections such as those figured
by Wille in his end view of the cell (Wille, Fig. 16), but in the Antarctic specimens it
was not possible to ascribe this appearance to longitudinal grooves. We would prefer,
however, not to press this point, as it is often difficult to clearly make out the finer
details of a chloroplast in preserved material.
Many examples were in the process of formation of daughter- cells, four of which
were produced in the mother- cell. The division was evidently by two oblique planes
crossing each other almost at right angles. ( Vide PI. XXIV, Figs. 28, 29)
FRESHWATER ALG/E 275
9 Chlamydomonas intermedia Chodat
In Bull, de VHerb. Boiss. torn, ii, p. 590, t, 22, 23 ; Wille, I.e. p. 142, t, 4, f. 15.
Forma antarctica. (PI. XXIV, Figs. 19-24)
Forma minor, stigmata juxta pyrenoidem.
Long. cell. 12-15 /*; lat. 8-10 ^.
Hob. Pony Lake.
Very well-preserved specimens of this species-were abundant among Phormidium
autumnale. The cells were ellipsoid, or ovoid- ellipsoid, and somewhat smaller than
in the form described by Chodat, but otherwise they were very similar. The proto-
plast completely occupied the confines of the delicate cell- wall, and the chloroplast
was large and cup- shaped, with a prominent pyrenoid situated in the median line
toward the hinder end of the cell. The pigment spot (stigma) was of variable shape,
generally elongated and somewhat curved, and was disposed towards one side of the
cell, either near the pyrenoid or half way between the pyrenoid and the insertion of
the cilia. Two daughter- cells evidently arose by longitudinal division of the mother-
cell. (Figs. 23, 24.)
One cell was observed with two pyrenoids (Fig. 24). This individual was without
doubt active when fixed in formalin, but it is possible that the division of the pyrenoid
had occurred as an antecedent to cell- division.
PLEUROCOCCACEjE
Genus — PLEUROCOCCUS Menegh.
10 Pleurococcus pachydermus Lagerh.
In Wittr. et Nordst. Alg. Exsic. No. 447; in Botaniska Notiser, 1882, p. 55.
Forma cellulis plerumque globosis sed hinc inde anguloso-globosis e mutua pres-
sione; crass, membr. cell. 2 p.; diam. cell, matur. 17.5-19.5 fj, cell. juv. (gonid.)
8-8.5 M. (PL XXIV, Figs. 45, 46)
Hob. Green Lake.
All these cells contained a single parietal chloroplast of considerable extent,
furnished with a single pyrenoid. Iodine solution stained this pyrenoid deeply, and
also brought out the cell- nucleus. The latter stained well after treatment with
osmic acid, and is seen to be situated internal to the chloroplast and rather to one side
of the cell.
Forma stipitata. (PI. XXIV, Figs. 47, 48)
Cellulis solitariis, libere natantibus, " stipitatis."
Diam. cell. 18-20 /a; long. stip. 7.5-26 M; lat. stip. 1.8-3.1 M.
Hub. In pond, Cape Royds.
The cells of this curious form possessed a cylindrical and more or less elongated
appendage at one side. This appendage was generally curved and quite solid, and
27(5 W. AND G. S. WEST
had more the appearance of a short stalk than anything else, although the cells were
quite free and unattached.
11 Pleurococcus frigidus sp. n. (PI. XXIV, Figs. 40-44)
Cellulis globosis, 20-24 ^ latis, singulis vel plerumque aggregatis, intra stratum
Myxophycearum nidulantibus ; membrana cellularum 1.4 /< crassa; chromatophora
singula magna, indistincte limitata, cum granulis amylaceis numerosis vel pyrenoidibus
conspicuis 2-3. Propagatio gonidiis globosis 2 vel 3 intra cellulas maternas ortis;
diam. gonid. 10-14 i*.
Hob. Pony Lake.
This species was more or less abundant in a matrix consisting mostly of Phormidium
autumnale and other Blue-green Algae. It appears to belong to that section of
Pleurococcus for which Klebs founded the special genus Chlorosphcera, but so far as
could be ascertained from the preserved material multiplication takes place by the
division of the contents of the mother- cell into two or three portions, which then
become rounded non-motile gonidia. As these grow in size they gradually throw off
the old wall of the mother- cell. Each cell possesses a single nucleus, placed in a rather
excentric position.
In the specimens collected in April 1908 the massive chloroplast contained numerous
minute granules of starch. In those obtained from the same locality in January
1909 the chloroplast possessed either 2 or 3 distinct pyrenoids and no scattered starch
granules.
The cells are considerably larger than those of Pleurococcus angulosus Menegh.,
and the chloroplast is not reticulated.
12 Pleurococcus antarcticus sp. n. (PI. XXIV, Figs. 49-51)
Cellulis globosis, 20-37 (plerumque 32) p. latis, singulis vel in familiis parvis aggre-
gatis inter algas Myxophycearum libere natantibus, membrana cellularum usque ad
2.2 /a. crassa, homogenea vel nonnunquam indistincte lamellosa ; chromatophora magna
parietali, indistincte limitata, ssepe cum globulis oleariis conspicuis. Propagatio ignota.
Hob. Green Lake and Clear Lake.
This is a large species, and, like the preceding, probably belongs to the section
" Chlorosphcera (Klebs) Hansg." Nothing was seen which in any way gave a clue as
to its methods of propagation. The chloroplast is massive, but its limitations could
not be de6nitely made out from the available material. It was generally partly
contracted from the wall at one side of the cell, and this space frequently contained
one, or more rarely two, globules of a fatty oil, such as occurs in Pleurococcus rufescens.
Forma robusta. (PI. XXIV, Figs. 52-54)
Cellulis majoribus, diam. 35-78 n, intra stratum Myxophycearum nidulantibus,
saepe dense aggregatis ; membr. cell. 2.5-6 p. crassa et lamellosa ; chromatophora
seepe distincte parietali, globulis oleariis nullis.
FRESHWATER ALG.E 277
Hob. Blue Lake.
These are the largest Pleurococcus cells we have yet seen.
13 Pleurococcus dissectus (Kiitz.) Nag.
Gatt. einzell. Alg, 1849, p. 65, t. 4, f. 3; Protococcus dissectus Kiitz. Spec. Alg.
1849, p. 199 ; Tab. Phyc. p. 4, t. 3.
Diam. cell. 4.5-9 p. (usque ad 16 n in cellulis solitarus). (PI. XXIV, Figs. 30-39)
Hab. Ponds on Mt. Erebus, attached to tire -tough sheets of Lyngbya Erebi.
Green Lake, among Ulothrix ccqualis forma. Blue Lake and Deep Lake, attached to
the sheets of Myxophycese.
The specimens agreed well with forms of this species, and especially with the
figures given by Suringar (Observ. Phyc. in Flor. Batav., Leovardice, 1857, p. 56, t. 4,
f. D) of "Protococcus dissectus /3 cuneata." They were, however, submerged (though
frozen), whereas the usual habitat of PI. dissectus is a subaerial one., The division of
the cells, partly radial, partly tangential, and partly oblique, is very characteristic.
This species often forms a one- layered stratum, in which the cells remain angular
by compression. This condition is almost a definite state of the plant. (Vide
PI. XXIV, Fig. 39).
Genus — TKOCHISCIA Kiitz.
14 Trochiscia aspera (Reinsch) Hansg.
In Hedwigia, 1888, p. 128 ; G. S. West, Treatise Brit. Freshw. Alg. 1904, p. 303,
c. figs. 82 A-F. Acanthococcus aspera Reinsch, 1886.
Diam. cell. 15.5-22 M.
Hab. Green Lake.
PALMELLACE.E.
Genus — GLCEOCYSTIS Nag.
15 Gloeocystis sp.
Diam. cell. 5-12 ^, c. integ. gelat. 14-30 /j..
Hob. In pond, Cape Royds.
From the available material it is not possible to exactly identify this Alga. Wille
has recently accepted certain views of Gerneck, and stated that the various Algse
included in Nageli's genus Gloeocystis are merely developmental stages of other green
Algse (c/. Wille in Engler and Prantl, Naturlich. Pflanzenfam. Chloropliycece, 1909,
p. 31). This view we do not entirely accept, as there is no positive proof that all
these Algse are states of species belonging to other genera, and Gerneck' s experience
is much too limited to warrant such a wide statement. This is the more emphasised
when one finds that some of Gerneck' s recently proposed genera can scarcely be
regarded as specifically distinct, let alone as valid generic types.
The greatest suspicion always attaches itself to Algse found only in culture
vessels. Such forms may in many cases be mere states of other Algse which remain
278 W. AND G. S. WEST
for a longer or shorter period in a profoundly modified condition owing to the unnatural
and generally abnormal conditions of the culture.
It is very probable that several species of Glaeocystis, including G. gigas (Kiitz.)
Lagerh., to which the Antarctic plant nearly approaches, are independent organisms
with a life- history of their own.
BACILLARIE^E
COSCINODISCACE.E
Genus — MBLOSIRA Ag.
16 Melosira distans (Ehrenb.) Kiitz.
A. Schmidt, Atlas Dial. t. 182, f. 4 ; 0. Miiller in Engler's Botan. Jahrbuch.
xxxiv, 1904. p. 271.
Diam. cell. 9,4-10 /y.
Hob. In pond, Cape Eoyds.
17 Melosira varians Ag.
W. Sm. Brit. Diat. II, p. 57, t. 51, f. 332; V. Heurck, Synops. Diat. Belg. t. 85,
f. 10, 11, 14, 15.
Diam. valv. 17-19 /*.
Hob. Ponds on Mt. Erebus.
Genus — COSCINODISCUS Ehrenb.
18 Coscinodiscus lentiginosus Jan.
A. Schmidt, Atlas Diat. t. 58, f. 11.
Diam. valv. 120 /u.
Hob. Green Lake, not uncommon.
This interesting species has been found to be fairly frequent in soundings made
by the s.y. Belgica from 62° 11' to 70° 53' S. lat.
Genus — TRICERATIUM Ehrenb.
19 Triceratium arcticum B right w.
Diam. valv. 230-258 fi.
Hal). Green Lake.
The specimens were typical, agreeing exactly with the figures in A. Schmidt's Atlas
Diat. t. 79, f. 12 and 12«, and also with examples mounted and named by J. Tempere.
FRESHWATER ALGM 279
Genus— HEMIATJLUS Ehrenb.
20 Hemiaulus ambiguus Janisch
Var. V. Heurck, Diatomees, Result. Voyage du s.y. "Belgica," Anvers, 1909, t. 8,
f. 110.
Diam. valv. 60 /u.
Hob. Green Lake.
Dr. Van Heurck was apparently somewhat doubtful concerning the exact
determination of his specimen (I.e. p. 36). Those observed from the above locality
were in exact agreement with his figure.
TABELLARIACE.E
Genus — TABELLARIA Ehrenb.
21 Tabellaria flocculosa (Roth.) Kiitz.
W. Sm. Brit. Dial. II, p. 45, t. 43, f. 316. V. Heurck, Synops. Dial. Belg. t. 52,
f. 10-12.
Long. valv. 26-27 n ; lat. max. ad med. 9.5-10 fj.
Hob. Ponds on slopes of Mt. Erebus.
In his Synopsis of Naviculoid Diatoms, Part I., 1894, p. 12, Cleve states that " of
Tabellaria flocculosa, one of the most frequent diatoms of Europe, not a trace has
been found either in S. America from Ecuador to Argentina, or in Australia or New
Zealand." Recently, however (G, S. West, " Alg. Yan Yean Reservoir," Journ. Linn.
Soc. bot. xxxix, 1909, p. 30), this cosmopolitan species has been shown to be abundant
in at least one district of Victoria, and the examples from Mt. Erebus were in every
way typical. Moreover, the specimens were in no way fragmentary, but had been
fixed in the living state, although the colonies were not observed to consist of more
than two cells, possibly owing to the rigorous conditions of existence.
FRAGILARIACE^
Genus — FRAGILAEIA Lyngb.
22 Fragilaria tenuicollis Heib. var. antarctica var. n. (PI. XXVI, Fig. 128)
Var. polis valvso valde productis et leviter subcapitatis ; striis non radiatis, 16 in
10 ft.
Long. valv. 37 M; lat. 3 /u.
Hob. Clear Lake.
In outward form the valves were very similar to F. tenuicollis var intermedia
(Grun.) V. Heurck [ = F. intermedia Grun. ; V. Heurck, Synops. Diat. Belg. t. 45, f. 9"!,
but they did not possess the median unilateral central area which is characteristic of
that variety.
BRIT. ANTAKCT. EXVED. 1907-9. VOL. I. L
280 W. AND G. S. WEST
23 Fragilaria virescens Ralfs
V. Heurck, I.e. t. 44, f. 1.
Long. valv. 40 n; lat. 8.5 M.
Hob. Ponds on Mt. Erebus.
24 Fragilaria obliquecostata V. Heurck
Diatomees, Result. Voyage du s.y. " Belgica" Anvers, 1909, p. 25, t. 3, f. 38.
Long. valv. 47 /u ; lat. 8 n ; costis 6 in 10 ju.
Hob. Green Lake, frequent.
Forma maxima V. Heurck, I.e. f. 40.
Long. valv. 75 M; lat. med. 8.5 /x> costis 5j in 10 M.
Hob. Green Lake, rare.
The valves were not so large as those described and figured by Van Heurck, but
they possessed the same slightly tumid median portion.
ACHNANTHACE.E
Genus — ACHNANTHES Bory.
25 Achnanthes brevipes Ag. var. intermedia (Kiitz.) Cleve
Synops. Navic. Diat. II., 1895, p. 193 ; A. subsessilis Kiitz., 1833.
Long. valv. 31-33 /* ; lat. 8.8-9.3 ». (PL XXVI, Figs. 126, 127)
Hob. Moraines near camp, Cape Royds. Clear Lake.
In outward form the valves were similar in form to those of A. brevipes var. inter-
media, although not quite so elliptic, and occasionally there was a faint indication
of a median constriction. The markings agreed well with the general disposition of
the punctate costse of this most variable species. We figure the upper and lower
valves for future comparison with other Antarctic forms.
Striations on lower valve 13 in 10 n, strongly radiate; on upper valve 12 in 10 M,
transverse in the middle of the valve, but rather irregularly bent towards the poles.
COCCONEIDACE^E
Genus — COCCONEIS Ehrenb.
26 Cocconeis litigiosa V. Heurck
Diatomees, Result. Voyage du s.y. "Belgica," Anvers, 1909, p. 18, t. 2, f. 28.
Long. valv. 79 /m ; lat. 54 n ; striis 9 in 10 p.
Hob. Green Lake, rare.
Only the inferior valves were seen, and although a little smaller than Van Heurck' s
original specimen, they possessed precisely the same striation. Moreover, they
FRESHWATER ALGM 281
possessed a similar transversely expanded, asymmetrical central area, and the lunate
terminal areas. The latter feature is also possessed by C. pellucida Hantzsch (cf.
A. Schmidt's Atlas Dial. t. 195, f. 1-6.)
NAVICULACE.E
Genus — TROPIDONEIS Cle_ve.
27 Tropidoneis Icevissima sp. n. (PL XXVI, Figs. 115-120)
T. parva, delicatissima et laevissima; valvis oblongo-linearis, diametro 8-11-plo
longioribus, in parte mediana marginibus parallelibus, apices versus leviter et gradatim
angustioribus, polis obtuse rotunda tis, raphe recta sed juxta polos levissime curvata
nodulo centrali in staurum transversum producto, stauro angustissimo plerumqu
valvae margines versus paullulo dilatato, alis (vel carinis) carentibus, striis non visis
Cellula in aspectu cingulato anguste oblongo-rectangulari, medio constricta, lateribu?
convexis et angulis rotundatis.
Long. valv. 49-98 M ; lat. med. 8-9.5 /*.
Hob. Clear Lake, Green Lake, and lake on west side of McMurdo Sound.
This diatom was one of the most abundant species in the Antarctic collections
occurring in great quantity in several of the lakes, both freshwater and strongly saline
It belongs to the section Orthotropis of the genus Tropidoneis, having a raphe
which is practically straight. The valves are very convex (consult Fig. 120), with a
slight thickening along the central axis, but they are entirely destitute of wings (or
keels). The great convexity of the valves, the slight indication of the bending of the
raphe near the poles, and the form of the girdle- view are all characters which place
the diatom in the genus Tropidoneis. The markings, if any are present, must be
exceedingly fine. Carefully prepared valves were examined most minutely with a
Leitz oil-immersion lens giving a magnification of 2000 diameters, but no trace of
striations could be discovered.
The central stauros is very narrow and reaches to the edges of the valve, where it
is slightly dilated. The general character of this stauros is very similar to that of
the pelagic Navicula (Stauroneis) Biblos Cleve (Le Diatomiste, Mar. 1892, p. 77, t. 12,
f. 9, 10), but the other characters of the diatom are very different. A stauros of a
similar kind is found in several species of Stauroneis, and there is a slight resemblance
between Tropidoneis Iwvissima and Stauroneis spicula Dickie (vide Van Heurck,
Synops. Diat. Belg. t. 4, f. 9), but the valves are not so attenuated and the striations
(if present) are not nearly so strong.
282 W. AND G. S. WEST
Genus — STAURONEIS Ehrenb.
28 Stauroneis anceps Ehrenb.
W. Sm. Brit. Diat. I, t. 19, f. 190 ; V. Heurck, Synops. Diat. Belg. t. 4, f. 4, 5.
Var. amphicephala Kiitz.
Long. 37-43 M ; lat. 8-10 n ; striis 22 in 10 ^
Hob. Recent geological deposit containing sub fossil diatoms. Also lake on west
side of McMurdo Sound.
The specimens seen were rather small, but otherwise typical.
Genus — TEACHYNEIS Cleve.
29 Trachyneis aspera (Ehrenb.) Cleve.
Synops. Navic. Diat. I, 1894, p. 191.
Long. 254 ^; lat. 48 ^.
Hob. Green Lake.
This handsome diatom was seen very sparingly, and we have not placed it definitely
under one of the numerous described varieties synopsised by Cleve. In outward form
the valves would agree with var. genuina Cleve, but in general appearance of markings
it agrees best with var. oblonga (Bail.) Cleve [vide figure of Stauroptera oblonga Bail,
in A. Schmidt's Atlas Diat. t. 48, f. 16], which is known from Australia, South America,
and the Island of Kerguelen. The alveolate striae were closer together (7 in 10 /«)
near the poles of the valves than they were in the median part (6 in 10 M).
Genus — NAVICULA Bory.
30 Navicula perlepida Grun.
" Diatom. Franz Josefs-land," in Denk. Akad. Wiss. Wien, xlviii, 1884, p. 104,
t. 1, f. 44.
Long. 16-18 n; lat. med. valv. 2.6-2.7 n.
Hob. Clear Lake.
This minute diatom agrees with Grunow's description of Navicula perlepida, and
also with his Fig. 44a. His dimensions are long. 20-34 /x ; lat. 2-3.3 M, so that the
Antarctic specimens, although of the same breadth, are scarcely so long as the Arctic
ones.
In general form and size the valves of this diatom are very similar to species of
Achnanthes of the section Microneis.
31 Navicula glaberrima sp. n. (PI. XXVI, Fig. 125)
N. minutissima et laevissima; valvis rhomboideis vel rhomboideo-ellipticis cum
lateribus psene rectis, polis levissime rotundatis, raphe rectissima, nodulis terminalibus
conspicuis ; striis non visis.
FRESHWATER ALG.E 283
Long. valv. 19 M; lat. 5.1 M.
Hob. Lake on west side of McMurdo Sound.
This minute Navicula was rather scarce amongst quantities of other diatoms,
especially Hantzschia elongata, Tropidoneis Icevissima, and Navicula muticopsis.
Careful examination of the valves with a one- twelfth- inch oil- immersion lens revealed
no trace of striations. Other examples of apparently smooth species of Navicula are
N. perlepida Grun., 2V. lineola Grun., and 2V. wwfo-P.antoesek.
2V. glabemma should be compared with 2V. palpebralis var. minor (Greg.) V. Heurck
(Synops. Diat. Belg. t. 11, f. 11 ; = 2V. minor Greg.), from which it differs in being
smaller and more angular, and in having no striations.
32 Navicula rhynchocephala Kiitz. forma
Long. 51 M; lat. 10 n; striis 12 in 10 /j..
Hob. In pond, Cape Royds.
The form noticed was apparently one of the intermediate states between the more
typical form (with subcapitate poles) and the var. amphiceros (Kiitz.) Cleve, Synops.
Navic. Diat. II, 1895, p. 15. The poles were slightly swollen and more obtuse than
the figure of var. amphiceros given by 0. Miiller in Engler's Bot. Jahrbuch. xlv, 1910,
t. 2, f. 20. The form was very similar in outline to " 2V. rhynchocephala var." in Cleve
et Grun. Beitr. Arctisch. Diat. 1880, t. 2, f. 33.
33 Navicula muticopsis V. Heurck
Diatomees, Result. Voyage du s.y. " Belgica," Anvers, 1909, p. 12, t. 2, f. 181.
Long. 14.6-23 /*; lat. 8-9.1 /*; striis 13-14 in 10 M. (PL XXVI, Figs. 121-124)
Hob. In ponds, and on the ground in the Penguin Rookery, Cape Royds. Also
in recent geological deposit containing diatoms. Clear Lake, Green Lake, Pony
Lake, and near Blue Lake. Lake on west side of McMurdo Sound.
This was the most abundant of all the Antarctic diatoms collected by the
Expedition. It occurred in nearly all the ponds and lakes, often in quantity, but
was most abundant in the Penguin Rookery at Cape Royds.
Considerable range of variation was exhibited by the valves, mostly in the character
of the poles, which were sometimes decidedly subcapitate, whereas at other times
they were merely " produced." The actual polar margin was frequently sub truncate,
but more often much more rounded. The central area was of variable form, from
a transverse stauros to a rounded area of considerable size. The striations in all
the specimens examined were 13 or 14 in 10 /*, and consisted of 3-5 puncta each,
except in the median part of the valve and at the poles. A solitary unilateral punctum
was present in the central area of some specimens, but no trace of it existed in others.
In all cases the median portion of the valve possessed subparallel sides.
The two following forms may be distinguished:
Forma evoluta. Cellulse paullo elongate, infra polum unumquemque distincte
sed leviter constrictse ; striis 13-14 in 10 ^.
284 W. AND G. S. WEST
Forma reducta. Cellulee breviores, polls minus productis et distincte truncatis ;
striis 13 in 10 /u.
Both of these forms were common over the whole area in which collections were
made.
34 Navicula muticopsiforme sp. n. (PI. XXVI, Fig. 131)
N. minutissima, valvis subellipticis, diametro circiter duplo longioribus, parte
mediana parva cum marginibus parallelis, polis longe cuneiformibus et obtuse
rotundatis, platea centrali magna et transverse dilatata, platea axiali lata, striis 17
utrobique (16 in 10 /z), punctatis, in parte mediana (ad versus plateam centralem)
striis tribus brevibus e puncto singulo elliptico formatis, ceteris radiatis e punctis
duobus formatis.
Long. 11.3 M; lat. 5.3 M.
Hob. Clear Lake.
This minute species has a slight resemblance to the more reduced forms of
N. muticopsis V. Heurck, but differs in the cuneate poles of the valves and in its striae.
There is a relatively large central area, caused by the shortening of three striae on
each side. These striae are reduced to a single elliptical punctum. The remaining
striae, which are somewhat divergent, are disposed in seven pairs in each half of the
valve. They leave a rather wide axial area, and each striation consists of two some-
what elongated puncta.
The species should be compared with N. mutica Kiitz. and N. seminulum Grun.,
from both of which it differs in outward form and in the striation of the valves.
35 Navicula peraustralis sp. n. (PI. XXVI, Fig. 132)
N. parva, valvis anguste lineari-lanceolatis, diametro circiter 4j-plo longioribus,
lateribus triundulatis inflatione mediana majori, polis inflato- capita tis et levissima
angularibus, platea centrali parva et elliptica, platea axiali angusta, striis validis
18 in 10 /u, leviter radiatis, a plateis lateralibus duobus interruptis.
Long. 47.7 /u; lat. 11.1 p..
Hob. Recent geological deposit containing subfossil diatoms.
This species is similar to those of the N. mesolepta group, but does not agree with
any of them. The two lateral areas interrupting the striations are also very
characteristic. In general outline it is not unlike N. semicruciata Ehrenb., as
figured in A. Schmidt's Atlas Diat. t. 44, f. 45.
36 Navicula (Pinnularia) globiceps Greg.
Cleve, Synops. Name. Diat. II, 1895, p. 77.
Long. 23-25 M; lat. 9-9.5 /m; striis 18 in 10 M. (PI. XXVI, Fig. 135)
Hob. Green Lake and Clear Lake. Recent geological deposit containing diatoms.
In one half of the central area was a single isolated punctum. The striations in
the median part of the valve were interrupted by two longitudinal lines, and in the
region of the capitate poles by one longitudinal line.
FRESHWATER ALG/E 285
37 Navicula (Pinnularia) murrayi sp. n. (PI. XXVI, Fig. 129)
N. parva, valvis linear!- ellipticis, diametro circiter 4-plo longioribus, polis obtusis
laevissime dilatatis, platea centrali transverse expansa, marginem utrinque psene
attingente, cum puncta solitaria asymmetrica, platea axiali angusta, striis 14 in 10 M,
punctulatis, in to to radiatis, in parte mediana 4 (circ.) utrobique multe brevioribus.
Long. 45 /a.; lat. 11.5 n-
Hob. In pond, and moraines near the camp,~Cape Royds.
This species stands nearest to N. reinhardtii Grun. (cf. V. Heurck, Synops. Dial.
Belg. p. 86, t. 7, f. 5, 6 ; = N. vernalis Donkin), but differs in the form of the central
area, with its solitary unilateral puncta, in the shorter striae bordering the central
area, and in the much closer stria tions (which are 14 in 10 M as compared with 9 in
10 fi). The striations are also radiate throughout, and never transverse at the poles,
which are very slightly dilated.
Var. elegans var. n. (PI. XXVI, Fig. 130)
Var. minor, polis valvae angustioribus, productis et subcapitatis ; striis 15 in 10 yu.
Long. 31 n; lat. 8.1 /x.
Hob. Pond, Cape Royds.
This variety should be compared with N. salinarum Grun. (in Cleve et Grun.
" Beitr. Arctisch. Diat.," Kongl. Sv. Vet.-Akad. Handl. Bd. 17, No. 2, 1880, p. 33,
t. 2, f. 34), from which it differs in many respects, although the spacing of the striae
is the same.
38 Navicula (Pinnularia} cymatopleura sp. n. (PI. XXVI, Figs. 133, 134)
N. minutissima, valvis sublinearis, diametro 4J- 5§-plo longioribus, lateribus
triundulatis, undulo mediana levissime majori, polis subcapitatis, platea centrali
magna et longitudinaliter elliptica, platea axiali lata, striis brevibus et laevibus,
20-21 in 10 m, leviter radiatis, in parte mediana (adversus plateam centralem)
brevioribus.
Long. 17-27 /«; lat. 4-5 a.
Hub. Clear Lake. Lake on west side of McMurdo Sound.
This minute species occurred abundantly in the above-mentioned lakes. It is
one of the N. mesolepta group, but the poles are much too wide for any form of
2V. mesolepta, and the striations are too fine. The iindulate sides are not quite parallel,
as there is a gradual and almost imperceptible attenuation. N. rupeslris Hantzsch is
of a similar shape, and there is a similar reduction of the striations in the middle of
the valve (vide A. Schmidt's Atlas Diat. t. 45, f. 45-48), but the striations oi N. cymato-
pleura are finer and shorter than in any of these forms.
It should be carefully compared with Pinnularia gracillima Greg., P. appendiculata
Ag., and P. undulata Greg. The latter is figured in Cleve's " Diatoms of Finland,"
Acta Soc. pro fauna et flora Fennica, VIII, No. 2, 1891, t. 2, f. 8, and is at once seen to
be quite a different diatom.
286 W. AND G. S. WEST
39 Navicula (Pinnularia) shackletoni sp. n. (PI. XXVI, Figs. 136-138)
N. minuta, valvis anguste elliptico-lanceolatis cum polls valde productis levissime
subcapitatis et obtusis, platea central! minuta, platea axiali angustissima, striis
10-12 in 10 M, leviter radiatis, validis et non punctulatis, in medio striis duobus
utrobique adversus plateam centralem delicatissimis et valde indistinctis. Cellula
in aspectu cingulato anguste lineari-oblonga.
Long. 25-29 p.; lat. 4-5 M.
Hob. Clear Lake, abundant.
This minute species should be compared with N. lanceolata Kiitz. var. tenella
(A. Schm.) Cleve [ = N. tenella A. Schmidt, Atlas Diat. 1876, t. 47, f. 45], from which
it is distinguished by its distinctly produced poles, its more distant and less radiate
striae, and by the two faint median striae on each side of the central area.
N. cryptocephala Kiitz. is also a similar species, but the lateral margins of the
valves are more ventricose and the poles more distinctly demarcated from the elliptic
body of the valve than in N. shackletoni. Moreover, the striae are as a rule much
finer.
In general outline N. shackletoni is not unlike N. El Kab forma lanceolata
0. Mil Her (in Hedwigia, xxxviii, 1899, p. 311, t. 12, f. 19), but the striation is much
coarser and differently disposed.
Var. pellucida var. n. (PI. XXVI, Figs. 139-141)
Var. platea centrali majori, leviter transverse dilatata, striis minus robustis, in
medio duobus utrobique brevioribus. Cellula in aspectu cingulato conspicue latiori.
Long. 25-29 p.; lat. 4.5-4.8 M.
Hob. In pond, Cape Eoyds, common.
This appears to be a well-marked variety, but the distinctions are hardly such as
to warrant specific separation. The much greater width in the girdle view was a
constant and noteworthy feature.
40 Navicula radiosa Kiitz.
V. Heurck, Synops. Diat. Belg. p. 83, t. 7, f. 20. Pinnularia radiosa W. Sm.
Brit. Diat. p. 56, t. 18, f. 171. P. acuta W. Sm.
Long. 74 ft; lat. 13 p.; striis 11-12 in 10 p..
Hob. Ponds on Mt. Erebus.
The specimens were not altogether typical as the apices were as acute as those of
the var. tenella (Breb.) Cleve, Navic. Diat. II, 1895, p. 17, but the striations were
rather coarser than in that variety.
FRESHWATER ALG.4S 287
COCCONEMACE.E
Genus — COCCONEMA Ehrenb.
41 Cocconema pusilla (Grun.) nob.
Cymbella pusilla Grun., 1875 ; Cleve, Synops. Name. Dial. I, 1894, p. 162.
Long. valv. 31 M; lat. 5.2 ^; striis 17 in 10 n,
Hob. In ponds, Mt. Erebus.
NITZSCHIACE.&
Genus — NITZSCHIA H assail
42 Nitzschia subtilis (Kiitz.) Grun. var.
Long. 39.2 /u; lat. 3.1 M; punctis 16 vel 17 in 10 ^
Hub. Clear Lake.
This Nitzschia seems undoubtedly to belong to N. subtilis, and resembles very much
the var. glacialis Grun. (" Diat. Franz Josefs- land," Denk. Akad. Wiss. Wien. xlviii, 1884,
p. 107, t. 1, f. 64). It is, however, very slightly more attenuated and the carinal
puncta are further apart. It should also be compared with N. subtilis var. paleacea
Grun. (cf. V. Heurck, Synops. Diat. Belg. t. 68, f. 9, the right-hand figure).
Genus — HANTZSCHIA Grun.
43 Hantzschia amphioxys (Ehrenb.) Grun.
Long. 56 fj. ; lat. 8.6 //.
Hah. Lake on west side of McMurdo Sound.
Var. capitellata Grun. in Cleve et Grun. " Arctisch. Diat.," K. Sv. Vet. -Akad.
Handl. Bd. 17, No. 2, 1880, p. 103.
Long. 72 M; lat, 13 n.
Hal>. In pond, Cape Royds.
44 Hantzschia elongata (Hantzsch) Grunow
H. Amphioxys var. elongata Grun. in V. Heurck, Synops. Diat. Belg. t. 26, f. 21-24.
Long. 168-190 /* ; lat. 9.5-11 M; carin. punct. 7-8 in 10 /u.
Hob. Clear Lake. Lake on west side of McMurdo Sound, very common.
SURIRELLACE.E
Genus — SURIRELLA Turpin.
45 Surirella angusta Kiitz.
Long. 29 M ; lat. 9.3 /x ; costaa 6 in 10 M.
Hub. In pond, Cape Royds.
BEIT. ANTAECT. EXPED. 1907-9. VOL. I. il
288 W. AND G. S. WEST
This diatom was seen very sparingly, but the specimens were quite typical, and
agreed exactly both in the poles and the costae with those in A. Schmidt's Atlas Dial.
t. 23, f. 40.
MYXOPHYCE7R
NOSTOCACB2E
Genus — NOSTOC Vaucher
46 Nostoc antarctica sp. n. (PI. XXV, Figs. 58-67)
Thallo initio globoso, parvo, pallide eerugineo, demum irregulariter expanso,
fuscescente, tenuissimo et membranaceo, crispato, usque ad 4.5 cms. lato, brunneo ;
trichomatibus in thallo membranaceo densissime contorto-intricatis, confertissimis ;
cellulis parvis, globosis vel nonnunquam ellipsoideis, confertissimis tanquam paren-
chymaticis; heterocystis subglobosis, diametro trichomatis duplo latioribus, rare
solitariis sed plerumque 2-5 seriatis ; sporis ellipsoideo-globosis, episporio glabro.
Diam. cell, veget. 2.5-3 M; diam. heterocyst. 6-7 M; diam. spor. 6.5-7 M.
Hob. In ponds, Cape Royds. Also in ponds, Mt. Erebus.
This is the most extraordinary Nostoc we have yet examined. Except for its
brown colour, the adult thallus presents all the appearance of a Monostroma. It is
exceedingly thin, with a crisped or pleated margin, and the trichomes are so densely
intricate as to give rise to an apparent parenchyma. In fact, we know of no species
of the genus with anything like this density of interlaced trichomes. The hetero-
cysts are rarely single, but more often 2-5 seriate, a character in common with
N. commune Vauch.
N. antarcticum is perhaps nearest to N. minutum Desmaz. (vide Born, et Flah.
" Revis. Nostoch. Heter." Ann. Sci. Nat. 7° serie, Bot. vii, 1888, p. 209), from which
it differs in the much greater dimensions of the adult thallus, the much denser
disposition of the trichomes, the spherical cells, and the greater diameter and seriate
character of the heterocysts.
NOTE. — A species of Nostoc was observed from Blue Lake, forming rounded
colonies up to 800 M diameter. Diam. cells 4 M. The material was only fragmentary
and insufficient for accurate identification.
OSCILLATORIACE^i
Genus — PLECTONEMA Thuret.
47 Plectonema notatum Schmidle
In Allg. lot. Zeitschr. 1901 Nos. 3 and 4, p. 4, Figs. 8, 9 on p. 2.
Crass, fil. 3-3.8 M, trich. 1.6-2 M.
Hob. In ponds, Mt. Erebus.
The specimens observed from the Antarctic were more in agreement with
FRESHWATER ALG.E 289
Plectonema notatum than with any of the other small species of the genus. The
false branches were rare and solitary, and the cells were sometimes twice as long as
broad.
Genus— LYNGBYA C. Ag.
48 Lyngbya shacUetoni sp. n. (PL XXV, Figs. 68, 69)
L. filis sparsis inter stratum algarum Myxophycearum, subrectis ; vaginis firmis
achrois distincte lamellosis; trichomatibus Isete aerugineo-coeruleis, inter cellulas non
constrictis, massa intercellular! inflate- torulosa et refringenti crebro interruptis,
apicibus rectis ; cellulis diametro trichomatis quadruple brevioribus, cytioplasmate
minute granulate; cellula apicali elongato-conica et pallidissime seruginea.
Crass, fil. 12-12.5 /*, trich. 8.5-9.5 M ; long. cell. 2-2.4 M.
Hob. Lake at Hut Point.
This species should be carefully compared with L. nigra Ag., which is the only
species with which it might be confused. The sheaths of L. shackletoni are much
thicker than those of L. nigra and are evidently lamellose ; the trichomes are
also frequently interrupted by lens- shaped masses of brightly coloured, highly
refractive material, and the apical cell, although markedly conical, has no calyptra.
Moreover, the end of the trichome is never capitate.
49 Lyngbya martensiana Menegh.
1837 ; Gomont " Monogr. des Oscillar.," Ann. Sci. Nat. 7C serie, Bot. xvi.,
1892, p. 165, t. 3, f. 17.
Crass, fil. 8-9 M, trich. 6-7 M.
Hob. Blue Lake.
50 Lyngbya cerugineo-coerulea (Kiitz.) Gomont,
l.c. p. 166, t. 4, f. 1-3.
Crass, fil. 7 M, trich. 5.7.
Hob. Ponds on Mt. Erebus.
51 Lyngbya murmyi sp. n. (PI. XXV, Figs. 70, 71)
L. filis inter algas varias Myxophycearum libere natantibus, stratum non formatis,
flexuosis ; vaginis tenuibus arctissimis et achrois ; trichomatibus laete serugineo-
coeruleis, inter cellulas non constrictis, apicibus rectis, non attenuatis, obtusis vel
rotundato-truncatis; cellulis diametro trichomatis Ij-lf-plo longioribus, cytio-
plasmate homogeneo cum granulis magnis 1 vel 2 prope dissepimenta utrobique
aggregatis.
Crass, fil. 3.1-3.3 /*; long. cell. 5-6 /*.
Hal). Green Lake.
This species probably stands nearest to L. versicolor (Wartm.) Gomont, but is
distinguished by its free-floating habit, its much thinner sheaths, which are not
agglutinated, and by the coarsely granulate dissepiments.
290 W. AND G. S. WEST
52 Lyncjbya Kiitzingii Schmidle
in Allg. hot. Zeitschr. 1896, iii, p. 58 ; Lemm. Alg. Brandenburg, 1907, p. 136.
Leibleinia martensiana Kiitz.
Crass, fil. 2-3 M.
Hob. In ponds on Mt. Erebus. Coast Lake. Blue Lake.
Var. distincta (Nordst.) Lemm. in Engler's Botan. Jahrbuch. xxxv, 1905, p. 620.
L. martensiana var. distincta Nordst. Alg. aq. dulc. et Char. Sandvic. 1878, p. 4.
L. distincta (Nordst.) Schmidle, 1896. L. subtilis West in Journ. Roy. Micr. Soc.
1892, p. 29, t. 10, f. 58.
Crass, fil. 1.5 n.
Hob. In pond, Cape Royds.
In his " Sylloge Myxophycearum " Dr. Achille Forti places L. subtilis as a distinct
species, owing perhaps to an error in his description. His " articulis diam. ad 2-plo
longioribus" should be " articulis diam. ad 2-plo brevioribus"
53 Lyngbya limnetica Lemm.
in Botan. Centralbl. Bd. 76, 1898, p. 154; Alg. Brandenburg, 1907, p. 135 et
p. 102, f. 8.
Crass, fil. IM; long. cell. 1-1.4 /«.
Hob. In pond, Cape Royds. Green Lake.
54 Lyngbya erebi sp. n. (PI. XXV, Figs. 72a-72«!)
L. strato valde expanse, cartilagineo, 3-5 mm. crasso, obscure serugineo vel vix
colorato ; filis flexuosis, densissime tortuoso-intricatis ; vaginis tenuibus arctis, validis
et achrois, crebro vacuis; trichomatibus angustissimis pallide serugineis, inter
cellulas non constrictis, apicibus rectis, obtusis et non attenuatis ; cellulis diametro
trichomatis paullo brevioribus, contentu homogeneo.
Crass, fil. et trich 0.9 M ; long. cell. 0.6-0.8 n.
Hab. In pond on Mt. Erebus.
This narrow Lyngbya formed a thick and very tough cartilaginous stratum,
almost destitute of colour, owing to a great extent to the enormous number of
empty sheaths of which it was composed. The sheaths themselves, although
so thin, are both strong and persistent, and form a densely contorted mass
difficult to tease out. The trichomes are very pale aBi-ugiiious green, and show no
signs of granulation.
In size L. erebi compares with the smallest of the plankton- Lyngbyas, but is
entirely different from them in habit.
It should be compared with Phormidium glaciale, from which it is distinguished by
the nature of its stratum and its strong persistent sheaths, both of which characters
give an entirely different aspect to the plant.
FRESHWATER ALG^E 291
Genus — PHORMIDIUM Kiitz.
55 Phormidium autumnale (Ag.) Gomont
I.e. 1892, p. 187, t. 5, f. 23, 24; Johs. Schmidt, " Danmarks blaagroenne Alger,"
Bot. TidssJcrift, Bd. 22, Heft 3, 1899, p. 348. Ph. uncinatum (Ag.) Gomont, l.c. p. 204,
t. 5, f. 21, 23.
Crass, trick. 4.6-7 M. (PI. XXV, Figs. 77-85)
Hob. Pony Lake. Coast Lake. Ponds on Mt.-Erebus.
This Alga appears to be frequent in the Antarctic regions, and was recorded from
the Falkland Islands by Hooker and Harvey as long ago as 1847. It occurred in great
abundance in Pony Lake, and the ends of the trichomes were furnished with precisely
similar small clusters of Bacteria as one so constantly observes in European specimens.
It not only occurred in pure masses, but was frequent in the strata of other species
of Phormidium.
As pointed out by Johs. Schmidt, there is no question of the identity of Ph.
autumnale and Ph. uncinatum. There are no characters by which it is possible to
discriminate between them.
56 Phormidium retzii (Ag.) Gomont
l.c. p. 195, t. 5, f. 6-9.
Crass, trich. 6-6.5 /«.
Hob. Dried-up lake, Cape Royds, among Vlothrix cequalis Kiitz.
57 Phormidium inundatum Kiitz.
Spec. Alg. 1849, p. 251 ; Tab. Phycol. I, p. 32, t. 45, f. iii; Gomont, l.c. p. 192,
t. 4, f. 31, 32.
Crass, trich. 3.5-4.1 /«.
Hob. Among Calothrix sp. in lake at Hut Point. In pond at Cape Barne,
abundant among Oscillatoria sancta.
A slightly thicker form (crass, trich. 4.6-5 M), but well within the limits of size for
this species, also occurred in Blue Lake.
58 Phormidium fragile (Menegh.) Gomont
l.c. p. 183, t. 4, f. 13-15.
Crass, trich. 1.2-1.6 M.
Hob. Pony Lake, Green Lake, and Coast Lake.
Forma tenuis. (PL XXV, Fig. 76)
Crass, trich. 0.9-1 n.
Hab. Coast Lake.
59 Phormidium glaciale sp. n. (PI. XXV, Figs. 73«-73d)
Ph. strato valde expanso, laete aeruginoso vel aeruginoso-cceruleo, usque ad 2 mm.
crasso ; fills flexuosis densissime tortuoso-intricatis, vaginis in muco hyalino diffluen-
292 W. AND G. S. WEST
tibus; trichomatibus angustissimis, laete eeruginosis, inter cellulas plus minusve
constrictis, apicibus rectis non attenuatis nee capitatis; cellulis tarn longis quam
latis vel paullo longioribus, contentu homogeneo.
Crass, trich. 0.8-0.9 M; long. cell. 0.8-1.1 M.
Hob. Blue Lake, Coast Lake, and Clear Lake ; in great abundance and forming
extensive sheets.
This species was one of the most abundant of the Blue- green Algae which occurred
in certain of the frozen lakes. Most of the specimens were obtained by melting the
blue- green sheets from the ice, where they must remain frozen sometimes for several
years. It is allied to Ph. angustissimum, but forms a thicker stratum, has slightly
thicker and more contorted trichomes, and has shorter cells.
60 Phormidium angustissimum W. & G. S. West
" Welw. Afric. Freshw. Algae," Journ. Bot. Aug. 1897, p. 72.
Crass, trich. 0.6-0.7 M ; long. cell. 2-4 /j..
Hob. Deep Lake, forming thin papery sheets of considerable extent and about
0.1-0.2 mm. in thickness.
This species was only observed from this one lake, forming extensive sheets
similar to those formed by Ph. glaciale, but much thinner. The trichomes are densely
intricate and flexuose, but are not so contorted as those of Ph. glaciale, and the cells
are much longer than in that species.
61 Phormidium antarcticum sp. n. (PI. XXV, Figs. 74, 75a-75
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